https://cpw.cvlcollections.org/files/original/a415a8ba8c5646a253c966b5310d6b22.pdf 20ad5caa20015ec9a9222d9bad93a509 PDF Text Text Colorado Division Wildlife Research April 1995 of Wildlife Report JOB PROGRESS State of: Project: Colorado Period 1 of Habitat Colorado 01 January Bird Research 24 : Job Evaluation in Eastern Covered: Authors: upland W-167-R Work Plan: Job Title: REPORT through Thomas E. Remington Development 31 December and Warren for Ring-necked Pheasants 1994 D. Snyder Personnel: C. E. Braun, J. J. Brim, T. J. Davis, M. A. Etl, K. M. Giesen, E.T. Gorman, W. R. Hanson, L. K. Haynes, R. W. Hoffman, T. J. Legg, J. L. Mekelburg, M. A. Porter, T. E. Remington, B. J. Rosenbach, G. G. Ruhser, J. C. Ruhser, W. D. Snyder, M. L. Trujillo, J. D. Weiland, B. T. Weinmeister, J. A. Yost, D. J. Younkin. ABSTRACT Expenditures under the Pheasant Habitat Improvement Program (PHIP) increased from about $220,000 in 1993 to $278,000 in 1994. We estimated Pheasants Forever Chapters contributed an additional $71,664 in labor, fuel, equipment rental, etc. to facilitate PHIP habitat developments. Most habitat developments were sorghum plantings (333 of 613), although, as in past years, the majority of PHIP expenditures went towards establishment of plum thickets, most of which contained juniper windbreaks. PHIP has now resulted in the establishment of 347 such thickets in it's first three years. Severe drought through most of the growing season stunted sorghum growth and resulted in some plum seedling mortality. As a result, cover value of sorghum plantings for ring-necked pheasants (Phasianus colchicus) was poor, with an average height of only 3 dm, and a height-density index of only 0.49 dm among 88 plots measured. Average counts of crowing males did not differ (£ = 0.85) between treatment and control blocks, not surprising given the poor quality of survival cover planted in 1993. Counts increased from about 14 calls per station in 1993 to 17 in 1994. Hunter pressure increased substantially from 1993, while harvest rates declined from 0.19 birds per hour to 0.05 birds per hour in 1994. One hundred and fifty-six hens were captured and radiomarked, bringing the sample of birds available for survival estimates to 206 including survivors from the 1993 trapping effort. A minimum of 36% of radio-marked hens survived from fall 1993 to fall 1994, suggesting survival was fairly good. No differences in survival between treatment and control blocks were apparent. ��3 EVALUATION OF HABITAT DEVELOPMENT FOR RING-NECKED PHEASANTS IN EASTERN COLORADO Thomas E. Remington and Warren D. Snyder INTRODUCTION Pheasants are pursued by more small game hunters than any other small game species in Colorado (83-88% of small game license buyers). In a recent survey, 74% of pheasant hunters rated their hunting trips in Colorado as poor (45%) or fair (29%), while only 10% rated their trips as very good or excellent. Lack of birds and places to hunt were identified as the most significant reasons why some hunters did not hunt pheasants in Colorado. Small game license sales have declined by about 70,000 (40%) in the last 10 years. It is apparent that if the Division of Wildlife is going to turn this decline around pheasants will be a key species. Presumably, recruitment and retention of hunters will increase if the quality of pheasant hunting is improved, i.e., increases in pheasant numbers and places to hunt. Previous research has indicated that over-winter survival of pheasants is the most critical factor limiting pheasant populations. The Pheasant Habitat Improvement Program (PHIP) was created to establish overwinter survival cover within historically good pheasant range in eastern Colorado. The program was conceptually designed to overcome significant obstacles to developing habitat, mainly a lack of manpower and a burdensome contractual system (costs of administering contracts exceeded costs of developments). Under PHIP, the Division of Wildlife contracts with individual Pheasants Forever chapters in eastern Colorado to contact landowners and develop habitat on private lands following specific guidelines. Each chapter develops contracts with individual landowners and pays them when the habitat work is completed and verified. Division of Wildlife personnel inspect a subsample of habitat developments and verify completion and compliance with guidelines. P. N. OBJECTIVES To determine if habitat developments offered through the Pheasant Improvement Program increase pheasant survival, breeding density, harvest within selected northeast Colorado study areas. SEGMENT Habitat and pheasant OBJECTIVES 1. Work with Pheasant Forever Chapters, management personnel, and landowners to develop habitat within treatment sites and elsewhere in the primary pheasant range in northeast Colorado. 2. Monitor hen pheasants previously radiomarked with mortality-sensing transmitters within treatment and control blocks to compare survival, nesting success, and use of habitats through 1994. Trap and radiomark additional hens as necessary to increase sample size to 100 hens each in the treatment and control blocks in fall 1994. 3. Conduct pheasant during April-May 4. Monitor control hunting sites. 5. Conduct evaluations 6. Prepare an annual crowing 1994. pressure counts within and pheasant of the quality progress report. all treatment harvest of annual within and control treatment plantings blocks and as survival cover. �6 Table 2. Pheasant habitat planted and/or contracted by Pheasants Forever chapters during 1994 in northeastern and east-central Colorado through the Pheasant Habitat Improvement Program. Number Plantings Habitat/Contract Sorghum Plantings Northeast Coloradoa Phillips County Yuma County Washington County Subtotal 46 110 169 63 34.1 428.0 _1 ___b_l 73 464.2 1,692 20,030 73 $21,795 7 6 27.0 15.5 165.0 207.5 $1,005 590 6,640 $8,235 40.0 15.0 $400 150 90 $640 9 Tall Wheat Stubble Retention Northeast Colorado Yuma Washington county Subtotal Shrub Thickets and Windbreaks Northeast Colorado Phillips County Yuma county Washington county Kit Carson Subtotal Total $7,075 14,640 30,322 1 100 $53,137 333 Disturbance Tillage (Annual Forbs) Northeast Colorado Phillips County Yuma County Subtotal Plantings/acres .zs 37 4 1 _3 8 a Colorado 64.0 14.2 27.4 14.9 27.3 ~ ~ 162 613 Expenditures Northeast __2_& 25 41 36 46 Custom Tillage (Contracted Site Preparation) Phillips County Custom 34 Phillips County Fertilizer 36 Yuma county 26 Washington County ~ 149 Subtotal Total Payment 181.5 369.5 765.8 27.5 1,344.3 __.§. Switcharass Plantings Northeast Colorado Phillips County Yuma County Subtotal Acres 90.6 $23,263.48 52,981.59 36,992.01 58,029.62 12,319.32 $183,586.02 2,170.6 361.3 140.6 71.0 166.5 739.4 5,420.00 844.00 1,252.50 3,020.00 $10,536.50 $277,929.52 Chapter Logan County wet snowfall and high winds. This fall, favorable hunting forecasts in Denver newspapers may have induced additional hunters to come out as well. Harvest rates were low, averaging 0.05 birds per hour across all contacts. Thus, on average hunters would have had to hunt about 19 hours to kill a rooster. This compares to an overall harvest rate of about 0.19 birds per hour (requiring about 5.2 hours to harvest a rooster) in 1993. Harvest rates were, on average, higher within treatment areas (0.07 birds/hour) than control areas (0.04 birds/hour), but high variability precluded statistical significance. There seemed to be a preponderance of inexperienced and/or less committed hunters in 1994 which may have lowered success rates. �7 Table 3. Sorghum, switchgrass, within and immediately adjacent Colorado, 1994. Block Cover type Holyoke SE and disturbance tillage plots established to the 9 treatment blocks, northeastern Number plots Number acres Sorghum 17 62.0 Mailander sorghum 24 64.0 Kurtzer Sorghum Switchgrass Subtotal 15 3 18 52.5 6.25 58.75 Fleming sorghum Switchgrass Subtotal 21 3 24 86.0 8.5 94.5 Pauli Sorghum Switchgrass Dist. tillage Subtotal 19 3 2 24 59.5 18.75 3.5 81. 75 Clarkville Sorghum 30 123.0 Y-W Sorghum Dist. tillage Subtotal 22 3 25 69.0 19.0 88.0 Kuntz sorghum Dist. tillage Subtotal 11+8 1 12 Otis Curve sorghum 12 56.5 171 9 6 610.0 33.5 24.5 186 667.0 Total Sorghum Total Switchgrass Total Disturbance Grand tillage totals 38.5 2.0 40.5 a Eight or more additional sorghum plots were established by T. Kuntz without receiving PHIP payments. The Kuntz block was subsequently dropped from evaluation due to severe hail and drought conditions. Sorghum and disturbance tillage plots were very popular with hunters where these provided adequate cover to hold birds. Most hunting parties were not large enough to hunt quarter sections of wheat or CRP effectively. The value of PHIP sorghum and disturbance tillage plots as accessible places to hunt where birds might concentrate can not be discounted. Pheasant Trapping and Survival Pheasant trapping began on 3 October and continued until 7 December. Trapping was completed a month sooner than in 1993 because two crews were used continuously, we had a better idea where to find birds, and the weather did not delay us as much. We placed radios on 156 new hens, and replaced old transmitters on an additional 6 hens which were re-captured. One of these recaptured hens had lost its' transmitter from 1993. Eighteen roosters were captured and banded, but no transmitters were placed on males. Fifty transmitters were still functioning (as of 1 Oct) on surviving hens trapped and marked in 1993, bringing the total of hens available for assessment of survival in 1994-95 to 206. �8 Table 4. Vegetation characteristics of sorghum plots planted within treatment blocks in 1994 and sampled in January-March 1995, northeastern Colorado. Treatment block Holyoke SE Mailander Kurtzer Pauli Fleming Clarkville Y-W Kuntz Otis Curve Mean N 10 10 10 10 11 10 10 7 10 VOR (dm) 0.26 0.09 0.19 0.57 0.68 0.11 0.89 0.68 0.84 sx 0.13 0.04 0.05 0.17 0.25 0.03 0.25 0.19 0.12 Height (dm) sx 4.00 0.34 3.60 0.38 3.13 0.46 3.43 0.26 2.72 0.46 2.43 0.25 2.70 0.51 2.41 0.29 2.68 0.38 88 0.48 0.11 3.01 Sorghum 28.38 15.94 14.03 21.37 21. 79 15.68 29.88 25.63 32.31 0.19 Cano:QY cover (%) Forbs sx 4.67 15.99 18.93 2.89 2.56 17.02 1.84 10.79 2.91 15.29 2.32 7.02 7.09 13.57 3.93 10.72 4.39 19.53 2.23 22.78 Table 5. Pheasant crowing census data among treatment northeastern Colorado, spring 1994. 14.32 and control sx ___ 2.04 2.69 4.49 2.13 3.13 1.45 2.38 3.79 2.55 1.39 blocks, Count Block 1 2 3 Average of 2-3 countsa Highest count High count/ stationb Treatments Holyoke SE Mailander Kurtzer Clarkville Pauli Y-W Co. Line Fleming Kuntz otis Curve 17.3 9.8 7.1 10.6 15.3 10.8 14.9 23.0 15.0 12.3 12.4 16.7 28.0 17.6 17.1 15.5 19.9 13.9 15.4 19.6 18.5 16.3 Average 18.9 27.0 17.9 16.8 20.6 16.7 28.7 18.3 17.1 15.5 23.0 15.0 15.4 19.6 16.7 28.0 17.6 17.1 15.2 20.1 12.9 13.9 16.0 16.7 20.6 14.2 17.1 ± 2.6 18.6 ± 4.3 20.0 ± 4.3 Controls Paoli NE Haxtun NE Paoli South St. Pete Kelly Yuma Co. Lonestar Platner Wash-West Average 16.4 13.7 14.5 32.8 8.4 24.5 14.4 7.3 19.6 13.6 15.0 15.1 28.8 14.3 17.1 12.6 29.3 20.2 13.0 13.0 17.1 7.8 24.5 18.3 13.9 14.2 30.8 11.4 20.8 13.5 7.6 17.1 29.3 20.2 15.0 15.1 32.8 14.3 24.5 14.4 7.8 ± 7.1 19.3 30.0 21.5 18.2 18.6 36.4 15.3 25.5 17.6 9.2 ± 8.1 a The lowest count was excluded when wide variance among counts occurred. b Obtained using the highest count per station among counts before 21.4 ± 8.2 averaging. �9 Survival of hens radio-marked between October and December 1993 was generally good, although survival rates have not been calculated. A minimum of fifty of 138 (36%; 154 marked in 1993 minus 16 which lost transmitters) hens survived to 1 October 1994. This number underestimates survival because the fate of birds that we lost contact with is unknown. It is likely that a small percentage of radios failed. We put considerable effort into locating missing birds both on the ground and through frequent flights over the study area, and have reasonable confidence that relatively few radios were working and not located. It is reasonable to assume that radios we lost contact with, particularly those with a long history of contact, were destroyed by predators chewing on the canister, biting off the antenna, or carrying them into a den. LITERATURE CITED Remington, T. E., and W. D. Snyder. 1994. Evaluation of habitat development for ring-necked pheasants in eastern Colorado. Colorado Div. Wildl., Prog. Rep., Fed. Aid Proj. W-167-R. Apr. 1-19. prepared Thomas E. Remington LjS, SjS Researcher IV by 1t!MMtJ ~ Warren D. Snyder LjS, SjS Researcher IV �Table 6. Pheasant 13 November 1994. Hunters Block hunters Hours/ Hunter contacted, Flush/ Hunter Inside Bag/ Hunter hunter effort, Birds/ Hour and hunter Hunters success Hours/ Hunter Flush/ Bag/ Hunter Hunter Outside Block in Treatment and Control Birds/ Hour Block blocks, 12- Birds/ Hour Combined Treatments Holyoke SE Mailander Kurtzer Clarkville Pauli Y-W County Fleming Kuntz Otis Curve 6 1.33 1.17 0.0 0.0 26 56 16 0.96 2.36 1. 78 0.12 0.68 1. 25 0.0 0.11 1. 50 0.82 3.57 1.28 3.34 0.64 0.55 2.27 0.12 0.70 0.72 14 11 15 41 20 Controls Paoli NE 37 Haxtun NE 32 Paoli South 6 St. Pete 53 Kelly 17 Yuma Co. Lonestar Platner Wash-West 23 43 32 26 2.08 5.00 2.42 0.46 4.12 4.77 2.79 5.21 2.33 3.83 3.28 49 7 3.96 1.71 0.92 1.39 0.33 0.16 0.37 0.44 0.0 0.045 0.246 6 25 0.087 0.049 0.064 0.083 0.0 0.093 0.0 0.074 0.173 0.29 0.14 0.18 0.47 0.02 0.05 0.095 0.22 0.131 0.019 0.015 4 5 12 0 12 4.25 2.20 1. 58 0.0 5.58 5.75 0.40 0.58 0.0 1.83 0.50 0.20 0.42 0.0 0.33 0.118 0.091 0.263 0.0 0.060 0.105 0.150 0.166 0.019 0.037 0.27 1.34 1. 50 0.75 0.88 0.12 0.19 0.0 0.11 0.06 0.108 0.090 0.0 0.047 0.129 38 14 0 29 7 5.50 2.71 0.0 1.62 3.21 0.79 1.79 0.0 0.62 1.43 0.03 0.07 0.0 0.17 0.14 0.005 0.026 0.0 0.106 0.044 0.023 0.067 0.0 0.063 0.066 2.13 0.74 0.41 0.65 0.35 0.12 0.09 0.08 0.084 0.024 0.034 0.015 14 11 8 13 2.77 3.09 2.50 3.65 0.57 0.55 0.38 0.08 0.14 0.0 0.13 0.0 0.052 0.0 0.050 0.0 0.075 0.021 0.037 0.011 o �11 Appendix A. SHRUB THICKETS PHIP Specifications AND SUPPLEMENTAL - 1994 WINDBREAKS Shrub (plum) thickets are the priority item. Small windbreaks, if planted, must be associated with a thicket and will not be funded if planted alone. Plantings will be eligible for funding only in farmed areas and must be within 0.1 mile of cultivated cropland. Plantings must remain'undisturbed for at least 10 years. Maximum planted another Funded: No more than 1 thicket (with or without wind barrier) can be per 80 acres. Each thicket/windbreak must be at least 1/4 mile from thicket/windbreak. Size: Shrub thickets must be at least 1/10th acre (4,300 ft2) and no larger than 2/10ths acre (8,800 ft2) in size, and must include at least 8 rows (excluding windbreak rows). Twelve hundred (1,200) feet is the maximum linear feet of fabric funded per thicket. Supplemental windbreaks, if planted, must be placed on the north and west side of the thicket and must include no more than 900 linear feet total if straight and 1,200 linear feet total if L-shaped. They must include at least 3 rows, one of which must be juniper or cedar. spacing between the thicket and the windbreak should approximate 100 feet (range 60 ft. minimum; 120 ft. maximum). Payment Rate: Payment will be at $0.55 per linear foot of fabric (6 ft. wide) to the maximums listed above if completed by PF Chapters. Actual costs not to exceed $.60/ft. if contracted to the State Forest Service or a local SCD. The maximum payment rate for approved private contractors will be $0.64 per linear foot of fabric. Supplemental payment rates/linear foot will be: $0.03 for application of polymer, $0.05 if 8 ft. wide fabric is used, and $0.01 for band application of an approved herbicide along the exterior edge of the fabric to reduce weed competition. Use of fertilizer will not be funded. Labor for landowners planting their own plots will not be funded. Planting Dates: Between March 20 and May 15. Pre-Plant Treatment: Sites must be tilled, preferably the fall prior planting. Tillage must be to bare soil with little residue remaining be deep enough to kill existing vegetation. Approved Species: (potted) . American Plum (bare root), Rocky Mt. Juniper, to and must E. Red Cedar Between-row Soacinq: A maximum of 10 feet will be permitted (6 to 8 feet spacing is recommended) for shrub thickets. A maximum of 15 feet will be permitted for wind barriers. In-row Spacing: A maximum of 8 feet will be permitted for shrub thickets (6 feet is recommended for plums) and 8-12 feet will be permitted for evergreens within wind barriers. Mulching: Woven polypropylene fabric is required for all plantings. fabric width is 6 ft. (3 ft. on each side of the row). Drip systems be cost shared. PERENNIAL GRASS AND GRASS-LEGUME Minimum will not PLANTINGS switchgrass provides tall cover that stands well over winter and has high value for pheasants. Small unfarmed tracts, currently in short, sodded grasses, are recommended for revegetation to switchgrass. Other shorter, cool-season grass-legume mixtures may be used in roadsides where snowdrift is a problem. This practice is funded only in farmland (not rangeland) settings. �12 PHIP SPECIFICATIONS - 1994 Payment Rate: $50.00 per acre as a one-time payment for sites up to 10 acres. For each additional acre (in sites larger than 10 acres [40 acres maximum]) the rate is $35.00. An additional $15.00 per acre will be paid for breaking out sod in heavily sodded sites and supplemental discing prior to planting switchgrass (this does not apply to roadsides). Preplant Soil Preparation: Adequate tillage to complet~ly destroy existing perennial vegetation and to establish a moist, weed-free, firm seed bed is required. Interseeding is not approved. A preemergent herbicide (e.g. Ally at 1/10 oz/acre) is recommended when planting switchgrass. Planting a tallsorghum mix (for which payment is available) is recommended the first year. switchgrass can be seeded into the residual sorghum without tillage during the subsequent spring but application of Ally herbicide is recommended. Plantinq Procedures: Planting procedures outlined in the Division's Game Information Leaflet #113 should be considered when planting switchgrass. In general, about 20 pure live seeds/ft2 (2 - 3 lbs/acre) should be planted using a drill with double-disk furrow openers, I-inch depth bands, and packer wheels. If a herbicide is not used, up to 1 lb/acre of an adapted dryland alfalfa and up to 1/2 lb of sweet clover should be added. Approved Species: In plots, switchgrass should comprise at least 75% of the live seed (alfalfa and sweet clover are approved additions). Within roadsides, switchgrass is the priority species where snowdrit't is not a problem. Other approved warm-season grasses include bluestems and Indian grass. Where these can not be used the tallest wheatgrasses (tall, intermediate, or standard crested) the roadside site will allow, should be used in combination with alfalfa (1 to 2 lbs/acre). Planting Dates: Warm-season grasses including 15: Cool-Season Grass-legume Mixtures: March switchgrass: 15 - July 15 March 15 to May Plot Duration: Grass and grass-legume plantings must remain ungrazed and undisturbed for at least 7 years. Roadsides should remain unmowed unless essential to reduce snowdrift. If essential, mowing should be delayed until after 1 August and restricted to the road shoulder. Prescribed burning, thinning tillage, or other renovation treatments to rejuvenate grass stands may be applied after 7 years. Grass stands that are relatively thin provide taller, better cover for pheasants. Legumes provide nitrogen and increase growth and quality when added to mixtures. DISTURBANCE TILLAGE AND TALL WILD ANNUALS Wild sunflowers, kochia, pigweed, and other tall annuals which attain 4 - 6 ft. height stand better through winter than other herbaceous vegetation, and provide excellent cover for broods, protection from blizzards and predators, and supplemental food. This is the most effective and least expensive approach for increasing pheasants and other upland game birds. Fallow land that is left idle usually converts to annual grasses or dog-hair stands of weeds by the 2nd year following tillage. Thus, at least one tillage each spring is usually needed to promote growth of tall annuals and a second thinning tillage is sometimes needed. Maximum Funded: 14 acres/0.25 section, ac. should be at least 0.25 mi. apart. 28 acres/section. Plots larger than 3 Funding Rate: $30.00/year for patches 0.1 to 0.5 acres in size, patches larger than 0.5 acres are considered 1 acre. $40.00/acre/year for sites up to 5 acres (7 ac. in pivot corners). $30.00/acre/year for additional acres up to 10 (6th-10th acres). Seeding wild sunflower or other approved wild annuals at 2 to 4 lbs. per acre will be funded at direct seed costs (see seed sources below. �PHIP SPECIFICATIONS 13 - 1994 Plot Dimensions: Short, relatively wide patches, inundated by drifting snow, are preferred. which Placement: Adjacent to woody cover when possible. contain weeds and above average moisture are ideal. perennials should be avoided. will not be easily Draw bottoms that sites containing already noxious specifications: Initial tillage with a disk plow or mold-board plow is needed in sites containing perennial grass to destroy all perennial cover, preferably, immediately after the ground has thawed in early March. Large clod size is preferred to retain thin stands of annual forbs. Initial tillage in subsequent years should be conducted prior to May 1 A secon~ thinning tillage may be used prior to the 1st of June. Spring tillage is needed each year to retain tall annuals. Annual grasses usually dominate if tillage is not used each spring. Wild sunflowers and annual ragweeds can be drilled or broadcast and harrowed at low rates to help establish tall annuals, if they are not already present. Known sources in Colorado include the Arkansas Valley Seed Company - Denver & Longmont, and Sharpe Bros. Seed Company - Greeley. Retention: Tall annuals must remain undisturbed through March of the following year. Sites should be prepared for the next year's growth during early to mid April if weedy cover exists. ANNUAL SURVIVAL PLANTINGS - SORGHUMS & DRYLAND CORN APPLICATION: On CRP, Annual Set Aside, and other cropland or tilled wasteland. When applied within CRP fields, SCS specifications for CP-12 must be used (see supplement). Dryland corn is primarily applicable in centerpivot corners next to irrigated corn. MAXIMUM FUNDED: 1 plot/SO-acre field, 2 plots/160 acres, ac.)/section. Plots must be at least 1/4 mile apart. 4 plots (28 PAYMENT RATE: $40.00/acre/year for 1-5 acres (7 acres within center pivot corners) and $25.00/acre/year for additional acreages in tracts larger than 5 acres (12-acre maximum). $30.00/acre/year if planted after June 15th. $6.00/acre for application of 30 lbs of nitrogen/acre. $15.00 per acre will be paid for breaking out sod in CRP or heavily sodded sites and supplemental discing prior to planting. Landowners can not be paid for labor/tillage on their own land other than for breaking out sod. PLACEMENT: Plots should be placed within placed crosswise to prevailing winds. or near cropland and SPECIFICATIONS: Preplant Soil Preparation: Initial treatment: Adequate tillage to destroy existing perennial vegetation in early spring prior to annual growth. Subsequent years: as needed prior to April is recommended. Preferably minimum tillage shredding of old materials 25. Annual application of nitrogen at 30-40 lbs./ac. Plot Dimensions: Minimum total plot width shall be 150 feet. Wider strips are preferred to reduce impacts of drifting snow. (See restrictions on dimensions in CRP). Row Spacinq: Sorghums - 15 to 30 inches; Dryland corn - 30 to 36. �14 PHIP SPECIFICATIONS - 1994 Seed Specifications: Sorghum Patches - At least 60% (75% preferred) of an adapted tall forage sorghum that will stand well with minimal lodging and will mature before frost. Up to 40% can be adapted varieties of grain sorghum. These can be mixed or planted in separate rows (i.e., 2 rows of grain sorghum to 6 rows of forage sorghum. These sorghums should equal a minimum of 75% of the total weight. Maximum amounts for other grains include: Dryland corn (25%), sunflowers (10%) and proso millet (10%). Addition of 1 to 2 Ibs./ac. of wild sunflower seed is recommended (Source: Arkansas Valley Seed Company - Denver). Dryland Corn Plots - Early maturing dryland varieties adapted to NE Colorado. Seed from these varieties that is one year removed frqm purchased hybrid can be used to reduce seed cost. Planting Dates & Rates: Sorghums - Between April 25 and June 15; Mid to late May is recommended. Plantings conducted after June 15 will be assessed a SlO/acre payment reduction. Sorghums should be planted at 4-8 Ibs./acre (30-inch rows) and at higher rates if drilled. Dryland corn - Between April 25 and May 15. Plantings after June 1 will not be accepted for payment. Seeding for dryland varieties should be from 10,000 to 13,000 seeds/acre. At least one cultivation is needed for corn and fertilizer should also be used. Plot Duration: 1 year. Sorghum plantings must remain undisturbed through March of the following year. Dryland corn must be left standing through March unless harvested. Harvesting may be conducted after March 15 of the following year. SUPPLEMENT FOR SORGHUM PLANTINGS WITHIN CRP SCS Notification: The CRP contract must be amended at the local SCS office prior to implementing CP-12 and breaking out food plots within CRP. This requires filling out a one-page form at your SCS office. The ASCS must be advised of the change for their records. Dryland corn is not approved for plots within CRP. Once a winter cover-food plot is broken out within CRP it must remain as such until the end of the CRP contract. Payments will be made annually based on seeded acres. If the farmer wishes to discontinue this practice he must reestablish grass (required by the ASCS). Reimbursement will be at S40.00/acre to cover reseeding grass. Maximum Funded: 1 plot/80-acre least 1/4 mile apart. field, 2 plots/160 acres. Plots must be at Maximum Size: The maximum size is 3 acres per site. CRP fields must at least 40 acres to be eligible for a CP-12 food plot. contain Plot Dimensions: Plantings may be up to 200 feet wide (100 ft in sandy soils). Typical 3-acre plots measure 198 x 660 feet. Where a 100 ft maximum is required a 30 ft wide buffer of untilled grass is left between two 99 x 660 ft parallel strips to obtain a 3 acre plot. Smaller plots should have reduced length to retain at least the 150 ft. minimum width. For example, a plot 99 ft wide x 440 ft. long equals 1 acre and two adjacent plots will exceed the minimum width requirement. Placement: Preferably within 50-100 yds of edge and near cropland, but location can vary depending on soil, wind, and moisture, and location of other winter covers if they occur. Sorghum plantings are not permitted in soils containing free lime (shows effervescence), or soils that are deep sands or choppy sands. �PHIP SPECIFICATIONS RETENTION OF STANDING WHEAT! 15 - 1994 TALL STUBBLE Application: Where winter wheat on Set Aside (ACR) acres is left standing under the ASCS wildlife food plot option. This option must be approved at your ASCS office. The objective is to provide taller, more secure cover for night roosting, feeding, loafing, and escape by pheasants through summer, fall, and winter. A primary concern with respect to pheasants is that unharvested wheat often does not stand well over winter. If the heads can be clipped (with ASCS approval) the resulting cover will be much more valuable to wintering wildlife. Payment Rate & Maximum Funded: (3) Funding to retain uncut wheat under the ASCS Wildlife Option on Set Aside tracts will be at $10.00 per acre up to 10 acres and $5.00 per acre for each additional acre up to 20 acres maximum. If the ASCS will permit clipping of heads to retain tall (>20 inch) stubble within Set Aside tracts payment rates will increase to $20.00 per acre for up to 10 acres and $10.00 for each additional acre to 20 acres. Maximum funded is 20 acres per quarter section and 40 acres per section. Specifications & Retention: Wheat or clipped stubble must remain standing through winter (ungrazed) until March 31 of the following year. Tall annual weeds, if present,_must be left standing and can not be treated with herbicides. Treatments will not be funded unless the entire stubble field is to be left undisturbed through the subsequent fall and winter. Placement: Standing cropland preferably wheat patches should within the southeast SUPPLEMENTAL PAYMENTS be near corn, sorghum, or other part of the stubble fields. FOR CUSTOM SITE PREPARATION Puroose: To prepare planting sites when the landowner does proper equipment or does not have time to prepare the site. not have the Treatment: Breaking out small tracts within CRP or sodded waste areas with a mold-board plow or heavy discing to completely destroy existing vegetation for reseeding to switchgrass or planting sorghum patches. Tillage must be to a depth of at least 6 inches. Payment Payment involve Rate: rate will be $15.00/acre two to three treatments. Equipment transportation will be SlS.00/hour. for adequate to and between site preparation small tracts. which Supplemental may payment ��17 Colorado Division Wildlife Research April 1995 of Wildlife Report JOB PROGRESS State of: Colorado Project: Job Title: Author: Upland W-167-R Work Plan: Period REPORT 1 : Job 25 Farming for Ring-necked Evaluation Covered: Thomas 01 January Bird Research through Pheasants 31 December - Program Development and 1994 E. Remington Personnel: C. E. Braun, S. M. DeMasso, T. E. Remington, and W. D. Snyder, Colorado Division of Wildlife; M. J. Manfredo, and J. J. Vaske, Colorado State University. ABSTRACT Implementation of an experimental fee hunting program for pheasants is on hold pending completion of the evaluation of the Pheasant Habitat Improvement Program (PHIP) under Work Plan 1, Job 24. Habitat developments proposed for the fee hunting program are being evaluated for effectiveness in increasing pheasant survival and harvest in the PHIP study; time constraints prevent doing both projects simultaneously. ��19 FARMING FOR RING-NECKED PHEASANTS - PROGRAM Thomas E. Remington DEVELOPMENT AND EVALUATION INTRODUCTION Small game license sales have declined markedly in Colorado over the past 10 years, from a high of about 200,000 in 1982 to a low of about 127,000 in 1993. Declines in participation in small game hunting are even more striking when considered as a percentage of Colorado's population. A recurring theme in surveys conducted to identify barriers to participation in hunting has been access to places to hunt, particularly places that have reasonably good hunting (Peterson and Manfredo 1993). Ring-necked pheasants (Phasianus colchicus) are the most commonly hunted small game species in Colorado (Braun et ale 1994). Declines in small game hunter numbers may be attributable, in part, to difficulties in acquiring access to places to hunt (pheasants) (Rounds 1975) and/or hunter dissatisfaction because of declines in pheasant populations (Farris and Cole 1981). Thus, pheasants were identified as a pivotal species in any strategy to reverse declines in small game hunter participation, and access to quality pheasant hunting was identified as a key management goal (Braun et ale 1994). Pheasant populations have declined in eastern Colorado because of a lack of survival cover and secure nesting cover (Snyder 1984, 1985, 1991) caused by intensive farming. Landowners are unlikely to alter agricultural practices to benefit pheasants or other wildlife, or in some cases allow hunting access, without significant financial incentives (Matulich and Bagwell 1979, Bishop 1981, Rasker 1989).The Pheasant Cooperative Program has been developed, so far only as a conceptual model, as a means to develop habitat, increase local pheasant populations, and provide hunting access to interested hunters for a fee (Remington 1993). The objective of this study is to ascertain small game hunter experience, satisfaction, and future interest in pheasant hunting in Colorado, determine hunter willingness to pay to hunt wild pheasants, and to predict the impact that fee rate and quality of hunting would have on rates of participation in a fee hunting program for pheasants. P. N. OBJECTIVES Develop a program to link hunters willing to pay for pheasant hunting opportunity with landowners willing to: 1) provide access for a fee, 2) develop habitat for pheasants within the program area, and 3) alter farming practices to make them more compatible with production and survival of pheasants. SEGMENT 1. 2. 3. 4. 5. OBJECTIVES Investigate feasibility of CDOW participation in forming a trial Community Cooperative fee hunting program in Burlington, Yuma, and/or other interested communities. Consider developing a detailed study plan for implementation and evaluation of this program depending upon feasibility analysis. Evaluate landowner and hunter interest with the program. Recommend changes to improve the program development. Prepare annual report. �20 LITERATURE CITED Bishop, R. C. 1981. Economic considerations affecting landowner behavior. Pages 73-87 in R. T. Dumke, G. V. Burger, and J. R. March, eds. Wildlife management on private lands. Wisconsin Chapter, The Wildl. Soc., Madison. Braun, C. E., K. M. Giesen, R. W. Hoffman, T. E. Remington, and W. D. Snyder. 1994. Upland bird management analysis guide, 1994-1998. Colorado Div. Wi1dl., Denver. 48 pp. Farris, A. L., and S. H. Cole. 1981. strategies habitat restoration on private agricultural Wildl. and Nat. Resour. Conf. 46:130-135. and goals for wildlife lands. Trans. North Am. Matulich, G. C., and G. Bagwell. 1979. On-farm pheasants enhancement potentials in irrigated agriculture. West. J. Agric. Econ. 4:99-109. Peterson, M. R., and M. J. Manfredo. 1993. are recreationists' perceived problems Colorado Div. Wildl., Human Dimensions Public access in Colorado: and preferred solutions? Persp. 15. 6pp. Rasker, R. 1989. Agriculture and wildlife: habitat management on farms in western state Univ., Corvalis. 241pp. an economic analysis of waterfowl Oregon. Ph.D. Diss., Oregon Remington, T. E. 1993. Farming for ring-necked pheasants - program development and evaluation. Colorado Div. Wildl., Prog. Rep., proj. W-167-R. Apr.: 25-36. Rounds, R. C. 1975. Public access to private Div. wildl. Spec. Rep. 2. 179pp. lands for hunting. Snyder, W. D. 1984. Ring-necked pheasant nesting ecology on the high plains. J. Wildl. Manage. 48:878-888. 1985. Survival J. Wildl. Manage. of radio-marked 49:1044-1050. 1991. Wheat stubble northeastern Colorado. Prepared by hen ring-necked as nesting cover for ring-necked Wildl. Soc. Bull. 19:469-474. Fed. Aid Colorado and wheat pheasants what farming in Colorado. pheasants in �21 Colorado Division Wildlife Research April 1995 of Wildlife Report JOB FINAL REPORT state of:. Colorado Project: W-167-R 3 Work Plan: Job Title: Period Personnel: Job: Kenneth 01 January Bird Research 18 Evaluation of Livestock Grazing Sage Grouse Nest Success. Covered: Author: Upland 1993 through and Residual 31 December Herbaceous Cover on 1994. M. Giesen C. E. Braun, K. M. Giesen, Division of Wildlife. D. B. LaBelle, P. D. Rosales, Colorado ABSTRACT Six strutting grounds in North Park, Colorado (Boettcher Junction, Coalmont, Delaney Butte, Lost Creek, Raven, and Spring Creek #1) were selected for documentation of hen movements to nests. A total of 71 hens was trapped and radio-marked on 4 strutting grounds (Boettcher Junction = 14, Coalmont = 22, Delaney Butte = 20, Spring Creek #1 = 15); no hens were trapped at Lost Creek or Raven strutting grounds because access was delayed by persistent snow. Forty-two hens were followed to nests in 1993; four hens were killed prior to nesting and 13 hens could not be located during the nesting season. No nests were located for another 12 hens. Movements from lek-of-capture to nest sites in 1993 ranged from 0.57 to 28.7 km with a mean movement of 3.52 ± 4.79 km (median = 1.88 km). Movements of 20 hens from lek-of-capture to nest sites in 1994 ranged from 0.66 to 7.79 km with a mean movement of 2.59 ± 2.09 km (median = 1.85 km). combined data from both years indicated nearly one-third (32.2%) of the marked hens moved farther than 3.0 km from the lek to nest. Nine hens in 1993 and two hens in 1994 were documented renesting and moved an average of 0.50 ±. 0.31 km between initial nests and renests. Sixteen hens followed to nests in both years moved an average of 0.79 ± 0.92 km between consecutive year nests. Nest success in 1993 was 22.0% and hen success was 26.2%. In 1994 nest success was 27.3% and hen success was. 30.0%. Most nest loss was due to depredation, primarily by ground squirrels (Spermophilus richardsonii). Shrub height, primarily sagebrush (Artemisia spp.) at nests averaged 54.6 ± 14.0 cm and 57.7 ± 14.5 cm in 1993 and 1994, respectively. Mean sagebrush canopy cover and density at nests were 35.3 ± 14.9% and 14,700 ± 6,000 plants/ha in 1993 and 42.8 ± 15.0% and 15,300 ± 6,200 plants/ha in 1994. No differences in habitat selected for nesting were observed by hens marked at different strutting grounds. ��23 EVALUATION OF LIVESTOCK GRAZING AND RESIDUAL ON SAGE GROUSE NEST SUCCESS Kenneth COVER M. Giesen INTRODUCTION Populations of sage grouse (Centrocercus urophasianus) in western Colorado have declined since the early 1980's (C. E. Braun, unpubl. data). This decline is reflected in the North Park, Colorado, sage grouse population by low numbers of male sage grouse on strutting grounds in spring and reduced harvests in fall. During this population decline nesting success of hens and production and survival of chicks has been below the long-term average. Considerable evidence suggests nesting success of sage grouse is related to amount of residual herbaceous cover in nesting areas which affects depredation of nests (Wallestad and pyrah 1974, Connelly et ale 1991, Ritchie et ale 1994, Gregg et ale 1994). The amount of residual herbaceous cover available to sage grouse in North Park is thought to be related to winter-spring precipitation and patterns of forage use by livestock (C. E. Braun, pers. commun.). The Bureau of Land Management reports that under optimum conditions, the best range sites in North Park produce only 800-1,200 poun~s of forage per acre, according to Natural Resource Conservation Service range site descriptions. When late-winter and spring precipitation levels are below average, as was the situation during 1990-1993, herbaceous forage production is less. The current pattern of livestock grazing in North Park may exacerbate the problem of lack of residual herbaceous cover needed for secure nesting cover for sage grouse in spring. Experiments to evaluate sage grouse nest success in relation to changes in residual herbaceous cover will provide information useful for evaluating and enhancing sage grouse nesting habitats in Colorado and elsewhere. It could also result in recommendations to experiment with new grazing systems and range enhancements. P. N. OBJECTIVES The primary objective of this study was to experimentally evaluate the relationships between residual herbaceous nesting cover and nest failure for sage grouse in North Park, Colorado. After nesting habitats associated with each strutting ground were identified, livestock grazing was to be experimentally excluded from parts of these nesting habitats (grazing exclosures) and nest success between grazing exclosures and control areas compared. Distribution of sage grouse nesting habitats in relation to strutting grounds was also identified and nest success was quantified. Specific objectives were to: 1. Trap and radiomark 20 hens at each of 6 selected strutting grounds and document their movements to nests. Identify nesting habitat adjacent to each study lek. 2. Ascertain nest failure. nest success 3. Measure vegetative selection for specific of radio-marked hens and ascertain structure at nest sites to ascertain nesting habitats. causes of possible 4. Experimentally exclude grazing from a portion of the nesting habitat associated with each strutting ground studied and compare subsequent nest success between nests in grazing exclosures and control areas. �24 STUDY AREA North Park in Jackson County, Colorado, is a large intermountain basin bounded on the east by the Never Summer and Medicine Bow ranges, on the west by the Park Range, on the south by the Rabbit Ears Range, and on the north by Independence Mountain. The topography of North Park is generally flat to rolling with elevations ranging from 2,400 to 2,800 m •. The vegetative community of the area is dominated by sagebrush on upland sites and grasses and sedges along native and irrigated meadows of the Canadian" Illinois, Michigan, and North Platte rivers. Approximately 90% of the sagebrush type is occupied by big sagebrush (Artemisia tridentata) (Smith 1966). Topography, soils, vegetation, and climate of the North Park area have been summaz Laed by Beck (1975), Petersen (1980), Emmons (1980), Schoenberg (1982), and Remington (1983). Six strutting grounds in North Park (Jackson County) were selected as sites for trapping and radio-marking hens based on criteria of access, consent of landowners or managers, lek size (number of males counted in 1992), and geographic distribution within North Park. These strutting grounds were Boettcher Junction and Delaney Butte leks in the northwest, Coalmont and Lost Creek in the southwest, Spring Creek #1 in the southeast, and Raven in the northeast. METHODS Spotlighting (Giesen et ale 1982) and walk-in funnel traps (Toepfer et ale 1988) were used to capture female sage grouse on or adjacent to selected strutting grounds in April and May 1993. Captured birds were placed in a burlap bag and processed within 30 minutes. Each captured hen was weighed on an electronic balance and banded with an aluminum band and red plastic bandettes. A Telemetry Systems solar or Holohil battery-powered transmitter was attached to hens using a poncho (Amstrup 1981) or necklace attachment. Transmitter packages weighed 12-18 gros, < 1.5% of the hens weight. Birds were released near the point of capture. Portable hand-held receivers and a 3-element Yagi antenna were used to monitor radio signals and locate hens once or twice weekly. Aerial tracking was used 4 times in May-June in an attempt to locate hens whose signals could not be detected from the ground. Hens were usually approached to within 20 m but not intentionally flushed when radio-tracked on the ground. Locations were plotted on a 1:24,000 scale U. S. Geological Survey topographic map in 1993 and distances and direction from capture site were measured. In 1994 a handheld Global Position System (GPS) receiver was used to ascertain capture locations and hen and nest locations. The 1993 map data were transformed to Universal Transverse Mercator (UTM) coordinates and movements and direction from capture site recalculated. Movements to nests by hens from different strutting grounds were examined using Kruskal-Wallis one-way analysis of variance by ranks and pair-wise multiple comparisons (test alpha = 0.20) to test for differences in median movements (Dunn 1964). Movements between first nests and renest locations were calculated for hens known to renest and distances between initial nests for 1993 and 1994 were compared for hens whose nests were located both years. Causes of nest loss were ascertained from evidence at the nest site (disturbance at the nest bowl, missing or broken eggs, etc.). Vegetation at nests was measured within a week of nest depredation, abandonment, or hatch. In 1994 vegetation at a randomly selected site within 400 m of each nest was measured for comparison of vegetative structure. At each nest or dependent random site heights of the nearest shrub, forb, and grass were measured to the nearest centimeter. A 10-m north-south transect was centered on the nest bowl or dependent site and the line-intercept of canopy cover (Canfield 1941) was measured for sagebrush (Artemisia spp.) and �25 other shrub species. Heights of all shrubs intercepted by the transect were also measured. Sagebrush density at the nest bowl or dependent random site and at 25 m in each cardinal compass direction was measured using a O.OOl-ha circular plot. Visual obstruction of vegetation was measured using a 3-sided cover board (Jones 1963) and measured from 0, 120, and 2400 aspects. RESULTS Trapping and Marking Seventy-one sage grouse hens were captured and radiomarked on or within 1.0 km of four strutting grounds in 1993. Most (n = 50; 70.4%) were captured by spotlighting and nearly all (n = 56; 78.9%) were adult. The number of radiomarked hens was 14, 22, 20, and 15 at Boettcher Junction, Coalmont, Delaney Butte, and Spring Creek #1 strutting grounds, respectively. Fewer hens were trapped on Boettcher Junction and Spring Creek strutting grounds because of high snowfall in April and persistent snowdrifts which delayed access for trapping. No hens were trapped on Lost Creek or Raven strutting grounds as access was delayed until after the peak of hen attendance. Signals were lost from 13 hens prior to nesting in 1993, 8 hens were shot by hunters during the 1993 hunting season, 10 hens were known to have been killed by predators prior to the 1994 nesting season" and another hen lost her radio in 1993. Of the possible 39 surviving hens in 1994, radio signals were detected from 23. Signals from 3 of these hens were lost during the nesting season or were inaccessible. Movements of Radio-marked Hens to Nest Sites Distances Moved to Nests. - Forty-two of 71 radio-marked hens were followed to nests in 1993 (Table 1). Four hens were killed by predators prior to nesting and 13 hens could not be radio-located within two weeks of being captured. Because of the intensive ground and aerial searches, it is likely that the transmitters on these hens failed or that these birds were killed by predators and the transmitters destroyed. Another 12 hens were radio-located at least twice weekly but no nests were located. It is likely that most of these hens attempted to nest but that nests were abandoned or depredated prior to incubation and no nest was discovered. Nine hens whose initial nests were depredated were documented to have renested; these data are presented separately. All 9 renesting hens nested within 1.0 km (range 0.05 - 0.90 km) of their initial nests which suggests that locations of initial nests and renests are not independent. Distances between capture site and nest location were similar between adult and yearling hens in 1993 (the samples of yearling hens from each strutting ground ranged from 0 - 6) so data were combined. Distances moved from capture site to nest sites were unique for each strutting ground. Movements from lekof-capture to nest site ranged from 0.57 to 28.69 km with a mean movement of 3.51 ± 4.79 km (median = 1.83 km). Overall, 15 of 42 hens (35.7%) moved farther than 3.0 km to nest (Fig. 1). Hens from Delaney Butte had the shortest movements between strutting ground and nest (1.69 ± 1.02 km) followed by hens from Boettcher Junction (2.35 ± 2.49 km), Coalmont (4.03 ± 3.66 km), and Spring Creek #1 (7.14 ± 9.67 km). Median movements were 1.57, 2.70, 1.47, and 4.29 km for Boettcher Junction, Coalmont, Delaney Butte, and Spring Creek #1 strutting grounds, respectively. The hens radiomarked at Spring Creek #1 strutting ground moved farther to nest sites than hens from Delaney Butte and Boettcher strutting grounds. No other significant differences were documented. �26 Table 1. 1993. Fates of radio-marked sage grouse Nesting strutting Ground n Boettcher Jct. Coalmont Delaney Butte spring Creek #1 Totals a Includes hens Hatched 14 22 20 15 2 3 5 1 71 11 2 hens killed hens in North Park, Colorado, fates of radio-marked Depredated Abandoned Lost hens Signal No Nest 5 1~ 9a 6 1 3 1 0 4 2 2 5 2 4 3 3 30 5 13 12 by predators prior to nesting. Twenty-three hens surviving into 1994 were monitored for nesting activities. Nesting fates of hens surviving into 1994 were similar to those observed in 1993 (Table 2). Movements between 1993 capture site and 1994 nest site were documented for 20 of 23 radio-marked hens. Radio signals from three hens were intermittent and these hens could not be located during the nesting season. Average distances moved from capture lek were 2.88 ± 2.80, 3.00 ± 2.19, 1.09 ± 0.20, and 3.44 ± 2.22 km for hens from Boettcher Junction coalmont, Delaney Butte, and Spring Creek #1 strutting grounds, respectively. The movement patterns among hens from the four strutting grounds in 1994 was similar to that documented in i993 with hens from Spring Creek #1 moving farther than hens from Delaney Butte strutting ground. Table 2. 1994. Fates of radio-marked sage grouse Nesting Strutting Ground Boettcher Jct. Coalmont Delaney Butte Spring Creek #1 Totals n hens 4 8 7 4 23 Hatched 0 3 2 1 6 Park, fates of radio-marked Depredated 4 2 3 3 12 hens in North Abandoned 0 1 1 0 2 Lost Colorado, hens Signal 0 2 1 0 3 No Nest 0 0 0 0 0 Distances between initial nests in 1993 and 1994 were documented for 16 hens. Movements were recorded for 3 hens from Boettcher Junction, 6 from Coalmont, 4 from Delaney Butte, and 3 from spring Creek #1 strutting grounds. The mean distance between consecutive year nest sites was 0.79 ± 0.92 km and ranged from 0.06 to 3.35 km (Table 3). Hens nesting successfully in 1993 (n = 5) tended to move less (0.37 ± 0.41 km) between consecutive year nest sites than unsuccessful hens (n = 11i 0.98 ± 1.04 km) although differences were not significant (Mann-Whitney u-test, P > 0.10). Furthermore, hens from Boettcher Junction strutting ground tended to move farther between consecutive year nests than hens from Delaney Butte strutting ground, although samples from each strutting ground were small. �27 Table 3. Distances between initial nest sites in consecutive years by sage grouse trapped at 4 strutting grounds in Jackson County, Colorado, 1993-94. Distance strutting Ground Boettcher Junction Coalmont Delaney Butte Spring Creek # 1 Totals n hens 3 6 4 3 16 ~ 1.41 0.84 0.36 0.66 0.79 between SD 1.68 0.93 0.48 0.33 0.92 nests (km) Range 0.40 0.09 0.06 0.29 0.06 - 3.35 2.61 - 1.07 - 0.94 - 3.35 Nine hens in 1993 whose initial nests were unsuccessful attempted renesting. The mean distance between initial and second nest was 0.49 ± 0.32 km (range = 0.02 - 0.90 km). Two hens were documented renesting in 1994 and they moved 0.25 and 0.76 km to their second nest. The origin of banding site had no effect on distances moved between initial nests and second nests. combined data from both years indicate a mean movement of 0.50 ±.0.3l km between initial and second nests. Hens that were unsuccessful in 1993 (n = 11) tended to move farther from their 1993 nest site to their 1994 nest site (0.98 ± 1.04 km) than hens that were successful in 1993 (n = 5, 0.37 ± 0.41 km) although the difference was not significant (Mann-Whitney u-test, P > 0.10). Direction of Movement. - Direction of movement between capture site and nest varied among the four strutting grounds (Figs 2-5). The strongest preference for direction of movement from capture site to nest was south and west, each with 21 movements recorded (combined years). The least common direction of movement was east with only 9 of 62 nests located east from the capture site. Nest Fates Only 11 of 42 hens whose nests were located in 1993 were successful in hatching one or more eggs from their nests for an estimated hen success of 26.2% (11/42 hens). Renesting was documented for 8 hens that lost their initial clutches; all renests were unsuccessful due to depredation. Overall nest success was 22.0 (II/50 nesting attempts). Most nest loss (34 of 40 nests, 85%) was due to depredation, primarily from Richardson's ground squirrels (Spermophilus richardsonii), although 6 nests were abandoned, possibly due to investigator disturbance. If we assume that the 12 hens which were not found on nests were also unsuccessful in nesting (they were observed regularly during May and June but were not seen on nests or with broods), then estimated hen success was to 20.4% (II/54 hens). Six of 20 hens whose nests were located in 1994 were successful in hatching one or more eggs from their clutch for an estimated hen success of 30.0% (6/20 hens). Two hens that renested after losing their initial clutches were also unsuccessful with their second nests. Overall estimated nest success in 1994 was estimated at 27.3% (6/22 nests). Richardsons's ground squirrels accounted for all depredation of initial and renest attempts in 1994 and two nests were abandoned prior to completion of incubation. Sixteen hens were documented nesting in both 1993 and 1994. Six of these hens were unsuccessful both years, 5 hens were successful in 1993 but unsuccessful in 1994, and 5 hens were unsuccessful in 1993 and successful in 1994. All these hens were at least two years of age in 1993 when initially captured. Combined data from both years indicated an overall hen success of 27.4% (16/62 hens). Overall hen success was highest among hens from Delaney Butte (7/20; 35%), followed by hens from Coalmont (6/20; 30%), Spring Creek #1 (2/10; 20%) and Boettcher Junction (2/12; 16.7%). �28 Vegetative Characteristics at Nest sites 1993 Nest Sites. - Vegetative characteristics were documented for 50 nest sites in 1993. These data included 10 nests from hens banded at Boettcher Junction, 16 nests each from Coalmont and Delaney Butte strutting grounds, and 8 nests from hens captured at Spring Creek. Vegetative heights at nest bowls averaged 55.7 ± 11.7 cm for shrubs, 6.4 ± 6.6 cm for fo~bs, and 20.8 ± 15.2 em for grasses. These vegetative heights above nests were similar among hens from all four strutting grounds (Table 4). Average VOR was 65.0 ± 15.8%. Although not quantified, nests were typically in sites (e.g. roadsides, mima mounds, irrigation ditches, other disturbed sites) having taller vegetative cover than adjacent areas. Table Park, 4. Vegetative height and VOR at 50 sage grouse Colorado, 1993. Values are presented as means Height Lek a SD Boettcher Jct. Coalmont Delaney Butte Spring Creek #1 Average 10 16 16 8 11.3 6.8 13.8 11.9 14.0 48.8 56.9 60.4 52.6 54.6 3.4 8.9 6.4 4.9 6.4 in North (cm) Forb x SD Shrub ~ nest sites 1 SD. ± 2.6 10.2 3.8 3.4 6.6 Grass ~ 13.9 19.9 27.2 18.6 20.8 VOR SD SD ~ 4.3 7.6 18.9 11.1 15.2 65.2 63.2 65.5 66.4 65.0 5.9 7.0 14.8 15.7 15.8 Live sagebrush canopy cover along 10-m nest transects averaged 35.3 ± 14.9%; dead sagebrush averaged 8.1 ± 7.4%. Average sagebrush height was 37.5 ± 16.0 cm and density averaged 14,700 ± 6,300 plantsjha (Table 5). There were no differences in sagebrush height, canopy cover, or density among nest sites from hens banded at different strutting grounds. of sagebrush at 10-m transects Table 5. Characteristics grouse nests in North Park, Colorado, 1993. Lek n % CanoQY ~ Boettcher Jct. Coalmont Delaney Butte Spring Creek #1 Average 10 16 16 8 34.3 35.4 35.7 35.4 35.3 Cover SD 14.6 17.0 13.0 17.3 14.9 Height ~ 29.2 38.2 44.0 38.0 37.5 (cm} SD 11.1 11.6 17.9 16.3 16.0 centered on sage Density (p.!_antsjha) SD ~ 18,900 15,200 11,800 14,600 14,700 7,700 6,300 5,300 3,100 6,300 1994 Nest sites. - Vegetative characteristics were documented for 21 nest' sites, including 2 renests, in 1994. Nests included 4 from Boettcher Junction strutting ground, 6 from Coalmont, 7 from Delaney Butte, and 4 from Spring Creek #1 strutting ground. Vegetative heights above nest bowls averaged 57.7 ± 14.5 cm for shrubs, 12.1 ± 12.1 cm for forbs, and 19.1 ± 10.1 cm for �29 grasses, and was similar among hens from different strutting grounds 6). Average VOR was 75.6 ± 12.0%. As in 1993, nests were typically having previous vegetative or soil disturbance. Table 6. Vegetative height and VOR at 21 sage grouse Park, Colorado, 1994. Values are presented as means Forb Shrub Lek !! Boettcher Junction Coalmont Delaney Butte Spring Creek #1 Average 4 6 7 4 SO K 45.2 59.3 58.1 66.8 57.7 n~st sites 1 SD. ± 4.8 13.5 15.2 17.1 14.5 K (Table in sites in North Grass SD 13.0 0.0 5.0 2.8 14.4 17.2 1.4 13.0 12.1 12.1 K VOR SO SD K 14.0 7.5 15.4 10.2 23.1 11.5 20.2 9.1 19.1 10.1 15.6 4.0 11.2 10.5 12.0 68.7 75.5 84.3 67.3 75.6 Live sagebrush canopy cover along 10-m nest transects averaged 42.8 ± 15.0 cm; dead sagebrush averaged 4.4 ± 5.4 cm. Average sagebrush height was 37.5 ± 14.9 cm and sagebrush density averaged 15,300 ± 6,200 plantsfha (Table 7). There were no apparent differences in sagebrush characteristics among nest sites from hens banded at the 4 strutting grounds. Compared to vegetative characteristics at dependent sites randomly located within 400 m of nests, sagebrush canopy cover and height, and VOR was greater at nest sites whereas sagebrush density did not differ (Table 8). Table 7. Characteristics of sagebrush at 10-m transects grouse nests in Jackson County, Colorado, 1994. n Lek % Canopy K Boettcher Junction Coalmont Delaney Butte Spring Creek #1 Average 4 6 7 4 Cover SD Height K centered on sage Density (cm) SD K SD 40.9 43.2 41.2 46.7 10.5 10.5 20.2 19.1 26.1 37.8 43.1 38.4 6.8 14.8 13.2 21.6 18,050 16,270 11,260 18,180 3,580 7,800 5,400 4,800 42.8 15.0 37.5 14.9 18,180 6,200 Comparison between 1994 nest transects and dependent random transects. - In 1994 I measured vegetation at a random dependent transect within 400 m of each nest. Because samples sizes of nests from each strutting ground were small, I pooled the data from all 4 strutting grounds (Table 8). Sagebrush canopy cover and VOR were both greater (P < 0.05) at nest sites than at dependent random sites. There were no differences (P > 0.10) in either sagebrush height along transects or sagebrush density between nest transects and dependent random transects. The greatest difference between nest transects and dependent random transects was in VOR suggesting nesting hens selected nest sites with greater than average herbaceous cover. �30 Table 8. Comparison of vegetative parameters measured at 21 sage grouse sites and dependent random sites in Jackson County, Colorado, 1994. Nest Vegetative Sagebrush Sagebrush Sagebrush VOR % Parameter canopy cover % height cm density, plants/ha Random SO X 42.8 37.5 15,300 75.6 nest SO X 15.0 14.9 6,200 12.0 28.2 26.8 15,800 27.9 14.3 14.9 6,700 27.3 Comparison of successful and unsuccessful nests. - Vegetative parameters at nest sites were compared between successful and unsuccessful nests for both 1993 and 1994. Oata were pooled for hens from all 4 strutting grounds within each year because of small samples of successful hens from individual areas. In 1993 there were no differences in the vegetative parameters measured among 11 successful and 39 unsuccessful nests (Table 9). Table 9. Comparison of vegetative parameters at successful sage grouse nests in Jackson county, Colorado, 1993. Successful X Sagebrush Canopy Cover (%) Sagebrush height (cm) Sagebrush density (plants/ha) VOR Shrub height above nest (cm) 3L4 33.0 13,200 68.6 52.5 (n SD 12.7 10.7 4,469 15.3 10.2 11) and unsuccessful Unsuccessful X 36.4 39.8 15,172 63.7 56.6 (n 39) SO 15.5 16.0 6,672 15.5 12.1 Vegetative parameters at successful nest sites were also compared to unsuccessful nest sites in 1994 (Table 10). Again no differences were detected. Examination of data indicated no differences in vegetation measurements between years although VOR tended to be higher in 1994, possibly due to the effects of above average precipitation in the spring of 1993 and its effects on residual vegetation the following year. �31 Table 10. Comparison of vegetative parameters sage grouse nests in Jackson County, Colorado, Successful ~ Sagebrush Sagebrush Sagebrush Canopy Cover (%) height (cm) density (plants/ha) VOR Shrub height above nest (cm) 43.8 32.7 16,133 74.7 54.5 at successful 1994. (n = 6) SD 12.1 11.4 7,914 11.8 15.6 and unsuccessful Unsuccessful ~ 42.3 39.4 14,967 75.9 58.9 (n = 15) SD 16.4 16.0 5,703 12.4 14.4 DISCUSSION Movements to Nests Previous studies of nesting sage grouse have documented a close relationship between strutting grounds and nest sites, with most nests within 3.2 km of leks (Patterson 1952, Schlatterer 1960, Gill 1965, Martin 1970, Braun et al. 1977). Prior to use of radiotelemetry, however, the spatial relationship between lek-of-mating and nest site was poorly known. Studies of female sage grouse using radiotelemetry indicate the range of movements of hens from lekof-capture to nest site recorded in this study was similar to that reported elsewhere (Montana,·Wallestad and pyrah 1974; Idaho, Wakkinen et al. 1992; Jackson County, Colorado, Petersen 1980). While the average movement by hens to nest sites is typically < 3.0 km, some hens move much farther to nest sites. The reasons for these longer movements are unknown because there is a lack of knowledge concerning seasonal home ranges of hens in relation to strutting grounds and nests. If strutting grounds are a focal point within female home ranges (Bradbury 1981), then nests should be clustered around strutting grounds where hens breed. However, a recent study (Wakkinen et al. 1992) indicates the distribution of sage grouse nests may be random with respect to strutting grounds. Consequently, spring home ranges of hens and nests should be randomly distributed within suitable habitats adjacent to leks. Therefore, the distribution of suitable nesting habitat is generally uniform around strutting grounds, then hens should select nest sites uniformly or randomly around leks. if Petersen (1980) and Schoenberg (1982) reported that adult hens in North Park nested farther from strutting grounds than did yearlings. Since adult hens attend strutting grounds earlier than yearlings (Petersen 1980), they might be expected to have first selection of nesting habitats and choose the better quality habitats. Longer movements by adult hens were not documented in this study, possibly due to small samples of yearlings marked. The longest movement between a capture site and nest site was of a yearling hen. Differences in hen movements among strutting grounds may be related to density of hens. If the number of hens attracted to a strutting ground is positively related to the number of displaying males, then the number of hens associated with Delaney Butte strutting ground was the least, followed by Boettcher Junction, Coalmont, and Spring Creek #1 (C. E. Braun, unpubl. data). If hens have exclusive nesting territories within their home range, and their home ranges include the strutting ground where they breed, then one would expect the shortest movements by hens to nests to occur at the smallest leks, and the farthest movements to occur at the largest leks. This relationship was �32 apparent observed ungrazed in this study. However, the shortest movements to nests were at Delaney Butte strutting ground, where hens were able to use nearby rangeland for nesting. Both the average movement between initial nest and renest location and distances between nests in consecutive years suggest that hens select nesting areas within their home ranges. This philopatry to nesting areas adjacent to leks indicates these areas should be protected from disturbance or habitat modification which may have deleterious effects on sage grouse nesting success. Nest Depredation The high rate of nest depredation or failure observed in this study is not unusual, and may have been affected by the study. Evidence from examination of hunter-harvested hens in North Park suggests nest success was 63% overall in 1993 (adults = 69%, yearlings = 40%) and 58% in 1994 (adults = 62%, yearlings = 44%) (C. E. Braun, unpubl. data). This difference suggests radiomarked hens may have been less successful than un-marked hens, and the effects of radiotracking and nest monitoring may have decreased their nest success. Excluding the 6 hens which abandoned their nests, the estimated hen success (30%) is half that as reflected in the hunter harvest. The estimates of nest success from wings collected in the hunter harvest may be biased by renesting hens. Since hens which were unsuccessful with initial and renesting attempts would not begin molting their primary feathers until the loss of their second nest, their primary molt may have been delayed and have been similar to or later than in hens successfully hatching their initial clutch. Renesting was documented in both years. The number of young as determined from the hunter harvest in 1993 (1.7 chicks/successful hen) was the lowest recorded in 20 years in North Park (C. E. Braun, unpubl. data) which may indicate that actual nest success as determined from wings collected in the hunter harvest may have been overestimated. However, in 1994 the number of young per successful hen was 3.1 which suggests hunter harvest data accurately reflected nest success. The major cause of nest depredation was the Richardson's ground squirrel. Other studies of nesting sage grouse in North Park have also documented high rates of nest depredation by this rodent (Gill 1965, Petersen 1980). Densities of ground squirrels in North Park are unknown, but ground squirrels and their burrows were commonly observed throughout the area. Fagerstone (1982) documented a positive relationship between ground squirrel densities and low herbaceous cover, which may occur with excessive livestock grazing. Thus, it is likely that ground squirrel densities have increased with high grazing levels in North Park and the recent drought which resulted in less forage for cattle. Increased ground squirrel densities and reduced vegetative cover for sage grouse nests could have increased the levels of ground squirrel depredation observed in 1993 and 1994. Habitat at Nest Sites Nesting habitat of sage grouse may be dependent on the quality of both herbaceous and shrub cover (Autenrieth 1981, Wallestad and pyrah 1974). Ritchie et ale (1994) suggested that herbaceous cover may be more important in determining sage grouse nest success than sagebrush height or cover. Results from this study, particularly the comparisons between nest transects and dependent random transects agree with this finding. North Park appears to have an abundance of sagebrush habitats of differing height classes, canopy cover, and density. However, the herbaceous understory in many areas is sparse. This is likely the result of several drought years exacerbated by livestock grazing. While predation may be the proximate cause of sage grouse nest loss, habitat at the nest site may be the ultimate factor determining nesting success. Because of the relationship between density of nesting cover and nest success in sage grouse (Connelly et ale 1991, DeLong et ale 1995) management efforts in North Park should be directed at improving herbaceous cover within sagebrush habitats. �33 While some studies have documented a positive relationship between nest success and vegetative cover (Wallestad and Pyrah 1974, Connelly et al. 1991) other studies (Autenrieth 1981, Ritchie et al. 1994) showed little or no differences. Intuitively one would expect that nest success would be higher where nesting cover was greater because on the evolutionary scale hens nesting is better cover would leave more progeny than hens nesting in poorer cover. The lack of difference in this study may have been the result of overall low nesting success by radiomarked hens because of observer'induced depredation. Additionally, nests in good vegetative cover may be relatively safe from avian predation but may not be secure from ground predators like ground squirrels which may use both visual and olfactory cues to detect nests. The high density of ground squirrels during this study may have resulted in above average nest depredation. LITERATURE Amstrup, S. C. 1981. 44:214-217. A radio-collar CITED for game birds. Autenrieth, R. E. 1981. Sage grouse management and Game. Wildl. Bull. 9. 238pp. Beck, J. Wildl. in Idaho. Manage. Idaho Dep. Fish T. D. I. 1975. Attributes of a wintering population of sage grouse, North Park, Colorado. M. S. Thesis, Colorado State Univ., Fort Collins. 49pp. Bradbury, J. W. 1981. The evolution of leks. Pages 138-169 in R. D. Alexander and D. W. Tinkle, eds. Natural selection and social behavior: recent research and new theory. Chiron Press, New York, N.Y. Braun, C. E., T. Britt, and R. O. Wallestad. 1977. Guidelines of sage grouse habitats. Wildl. Soc. Bull. 5:99-106. Canfield, R. 1941. Application of the line interception of range vegetation. J. For. 39:386-394. for maintenance method in sampling Connelly, J. W., W. L. Wakkinen, A. D. Apa, and K. P. Reese. 1991. Sage grouse use of nest sites in southeastern Idaho. J. Wildl. Manage. 55:521-524. DeLong, A. K., J. A. Crawford, and D. C. DeLong, Jr. 1995. Relationships between vegetational structure and predation of artificial sage grouse nests. J. Wildl. Manage. 59:88-92. Dunn, O. J. 1964. 252. Multiple comparisons using rank sums. Technometrics Emmons, S. R. 1980. Lek attendance of male sage grouse, North M. S. Thesis, Colorado State Univ., Fort Collins. 69pp. Park, 6:242- Colorado. Fagerstone, K. A. 1982. Ethology and taxonomy of Richardson's ground squirrel (Spermophilus richardsonii). Ph.D. Diss., Univ. Colorado, Boulder. 298pp. Giesen, K. M., T. J. Schoenberg, and C. E. Braun. sage grouse in Colorado. Wildl. Soc. Bull. Gill, R. B. 1965. Distribution in North Park, Colorado. Collins. 185pp. and abundance M. S. Thesis, 1982. Methods 10:224-231. for trapping of a population of sage grouse Colorado State Univ., Fort �34 Gregg, Jones, M. A., J. A. Crawford, M. S. Drut, and A. K. DeLong. Vegetational cover and predation of sage grouse nests Wildl. Manage. 58:162-166. R. E. 1968. A board to measure Wildl. Manage. 32:28-31. cover used by prairie Martin, N. S. 1970. Sagebrush control related to habitat occurrence. J. Wildl. Manage. 34:313-320. Patterson, R. L. 1952. Colo. 341pp. The sage grouse in Wyoming. Petersen, B. E. 1980. Breeding and nesting North Park, Colorado. M. S. Thesis, 86pp. Remington, T. E. 1983. Food selection, grouse during winter, North Park, State Univ., Fort Collins. 89pp. 1994. in Oregon. grouse. J. J. and sage grouse Sage Books, Inc., Denver, ecology of female sage grouse in Colorado State Univ., Fort Collins. nutrition, Colorado. and energy reserves of sage M. S. Thesis, Colorado Ritchie, M. E., M. L. Wolfe, and R. Danvir. 1994. Predation of artificial sage grouse nests in treated-and untreated sagebrush. Great Basin Nat. 54:122-129. Schlatterer, grouse E. F. 1960. Productivity and movements of a population of sage in southeastern Idaho. M. S. Thesis, Univ. Idaho, Moscow. 87pp. Schoenberg, T. J. 1982. Sage grouse Park, Colorado. M. S. Thesis, Smith, movements and habitat Colorado State Univ., selection in North Fort Collins. 86pp. E. L. 1966. Soil vegetation relationships of some Artemisia types North Park, Colorado. Ph.D. Thesis, Colorado State Univ., Fort Collins. 203pp. in Toepfer, J. E., J. A. Newell, and J. Monarch. 1988. A method for trapping prairie grouse hens on display grounds. Pages 21-23 in A. J. Bjugstad, Tech. Coord. Prairie chickens on the Sheyenne National grasslands. U. S. Dep. Agric., For. Servo Gen. Tech. Rep. RM-159. Wallestad, R. 0., and D. pyrah. hens in central Montana. 1974. Movement and nesting of sage grouse J. wildl. Manage. 38:630-633. Wakkinen, W. L., K. P. Reese, and J. W. Connelly. 1992. locations in relation to nests. J. Wildl. Manage. Prepared by Kenneth M. Giesen Sage grouse 56:381-383. nest �35 20 1993 15 10 en 5 z w :r: u.. o a: w to :2 o 20 ::> z 1994 BOETTCHER JCT. 15 ~ COALMONT ~ DELANEY BUTTE SPRING CREEK 10 5 o < 1.0 1.0-2.0 2.1-3.0 3.1-4.0 4.1-5.0 > 5.0 DISTANCE (km) Fig. 1. Movements of sage grouse hens from 4 strutting grounds to nests in Jackson County, Colorado, 1993 (top) an 1994 (bottom). �36 BOETICHER JCT. Fig. 2. Locations of 1993 (triangles) and 1994 (squares) nest sites of hens at Boettcher Junction strutting ground in Jackson County, Colorado in 1993. Letters indicate nests of the same hen in consecutive years. COALMONT 4km Fig. 3. Locations of 1993 (triangles) and 1994 (squares) nest sites of hens captured at Coalmont strutting ground in Jackson County, Colorado in 1993. Letters indicate nests of the same hen in consecutive years. �37 SPRING CREEK 4km Fig. 4. Locations of 1993 (triangles) and 1994 (squares) nest sites of hens captured at Spring Creek #1 strutting ground in Jackson County, Colorado in 1993. Letters indicate nests of the same hen in consecutive years. Fig. 5. Locations of 1993 (triangles) and 1994 (squares) nest sites of hens at Delaney Butte strutting ground in Jackson County, Colorado in 1993. Letters indicate nests of the same hen in consecutive years. �38 �Colorado Division Wildlife Research April 1995 of Wildlife Report JOB FINAL REPORT (Research) State of: Colorado Project: W-167-R Work Plan: Job Title: Period Author: 12 Richard 01 January Bird Research 17 Effects of Mycoplasma Infection Merriam's Wild Turkeys Covered: Personnel: : Job Upland 1992 through on Reproductive Performance of 30 June 1995 W. Hoffman Thomas A. Artiss, Clait E. Braun, Amanda S. Clements, Renzo Del Piccolo, Van K. Graham, John P. Gray, Anthony W. Hoag, Richard W. Hoffman, Robert T. Magill, and Walter J. Miller, Colorado Division Wildlife; William R. Davidson and Page M. Luttrell, Southeastern Cooperative Disease Study. ABSTRACT Interactions with domestic poultry have been implicated in the occurrence of Mycoplasma gallisepticum (MG) and M. synoviae (MS) in wild turkeys (Meleagris gallopavo). These organisms may suppress wild turkey populations through subtle changes in reproductive performance. Merriam's wild turkeys (M. g. merriami) commonly winter in large flocks around ranches where they take advantage of artificial food sources. Flocks existing under these conditions tend to be relatively tame and easy to trap and, therefore, are often targeted as a source of birds for relocation efforts. However, because these flocks frequently come into contact with domestic fowl, they may present a greater risk of disease dissemination if moved elsewhere. The objectives of this study were to evaluate the epizootiology of Mycoplasma infection in a population of Merriam's wild turkeys with a history of association with domestic fowl and to compare nesting effort, clutch size, nesting success, hatching success, and egg fertility between MGjMS seropositive and seronegative female wild turkeys. Trapping, testing, and radiomarking were conducted near Collbran in west-central Colorado. One hundred and eleven female wild turkeys and 31 domestic chickens were captured and surveyed for evidence of Mycoplasma infection by serologic and cultural methods. No clinical signs of Mycoplasma infection were apParent in any of the birds examined. However, 50 (45%) wild turkeys had positive rapid plate agglutination (RPA) reactions for MG (n = 5), MS (n = 27), or both MG and MS (n = 18); 40% of 50 adults and 49% of 61 subadults were classified as positive reactors. Weights did not differ between positive and negative reactors. Hemagglutination inhibition (HI) tests were uniformly negative for MG and MS and did not support the RPA results. In contrast, most chickens were strongly positive for MS according to the RPA (81%) and HI (58%) test results. Mycoplasma gallopavonis was the most common isolate identified from the wild turkeys. Mycoplasma gallinaceum was isolated from both the chickens and wild turkeys indicating possible contact transmission between the two groups. No pathogenic mycoplasmas were isolated from either group. Radio contact was maintained with 100 hens (47 adult, 53 subadult) into the nesting season of which 91 attempted to nest. Nesting effort was not a function of whether a bird tested positive or negative. Seventy-nine percent of the adults and 48% of the subadults were successful nesters in 1992 compared to 36 and 33%, respectively, in 1993. Seronegative adult hens were more successful than �40 seropositive adult. hens in 1993 but not in 1992. Positive and negative subadult hens nested with equal success. Clutch size of 1st nest attempts did not differ between years for adults or subadults nor did clutch size differ between age classes when years were combined. Clutch size was larger for 1st than 2nd nest attempts. Sample sizes were not adequate to examine annual differences in clutch size of 2nd nest attempts; however, when years were combined, no differences were apparent between age classes for 2nd nest attempts. Comparisons between positive and negative reactors revealed no differences in clutch size for 1st or 2nd nest attempts. Of 21 hens that renested, 13 (62%) were positive reactors; 13 of 29 positive and 8 of 26 negative hens available to renest actually produced a second clutch. Eightyseven percent of the 507 eggs examined from successful nests hatched, 6% were infertile, 5% contained fully developed but unhatched embryos, and 2% contained partially developed embryos. Hatching success and egg 'fertility did not differ between age classes. Likewise, hatching success and egg fertility did not differ between positive and negative reactors. Collectively, the serological test results were only suggestive of Mycoplasma infection. The availability of artificial foods may have masked the presence of disease and enhanced reproductive performance. It is also possible the organisms involved were less pathogenic, variant strains of MG or MS. RECOMMENDATIONS According to guidelines advanced by the Wildlife Disease Association suggest only seronegative turkeys should be considered for relocation efforts. This assumes there is a difference between seropositive and seronegative turkeys trapped from the same population. Data from this study do not support this assumption. Whereas it is important to continue to collect data to protect wildlife agencies against criticism of inadequate disease considerations, it is equally important that transplant programs not be hampered by overly restrictive disease monitoring guidelines. The turkey population monitored in this study took advantage of human-related food sources, thereby bringing them into contact with domestic fowl. This is a common situation in which wild turkeys exist in the western United States. To recommend that birds existing under these conditions not be used as transplant stock could curtail range expansion programs in some states, including Colorado. The following guidelines should be followed when trapping and transplanting turkeys known to be living in association with domestic fowl: 1. Pretest a sample of 5-10 birds prior to initiating a major trap and transplant program. Examine the birds for clinical signs of Mycoplasma infection such as swollen sinuses, nasal discharge, lameness, and lethargic behavior. Collect blood samples and test for M. gallisepticum and M. synoviae using the rapid plate agglutination and hemagglutination inhibition,:tests~ Swab the ,trachea and c~ltuJ;'efor-M. gallisepticum, M. synoviae, and M. gallopavonis~ If po s.sLb Le capture,' examine, and test some of the domestic fowl that are in contact with the'turkeys. . , 2. Positive RPA reactions, without confirmed isolation, clinical symptoms of infection, or supporting HI results, provide insufficient evidence to negate a transplant. However, when actually conducting the transplant it would be best to field test each bird and only move those that are RPA negative. This precaution is not necessary if pretesting shows no positive RPA reactions. 3. Regardless of the test results, turkeys living in association with domestic fowl should not be used to supplement existing populations should they be released into areas with a viable poultry industry. 4. nor Any birds with clinical symptoms of disease should be sacrificed and attempts made to isolate the Mycoplasma organisms responsible for the infection. �41 EFFECTS OF MYCOPLASMA INFECTION ON REPRODUCTIVE OF MERRIAM'S WILD TURKEYS Richard PERFORMANCE W. Hoffman The results of this study have been prepared as two manuscripts submitted for publication. The two manuscripts are: that have been Hoffman, R. W., M. P. Luttrell, and W. R. Davidson. 1995. Serological and cultural investigations of Mycoplasma infection in Merriam's wild turkeys living in association with domestic fowl. J. Wildl. Dis.: submitted. Hoffman, R. W., M. P. Luttrell, and W. R. Davidson. 1995. Reproductive performance of Merriam's wild turkeys with suspected Mycoplasma infection. Proc. Natl. Wild Turkey Symp. 7:000-000. Prepared by: Richard W. Hoff n L/S Sci Res/Scientist IV �42 �43 Colorado Division Wildlife Research April 1995 of Wildlife Report JOB PROGRESS state of: Colorado Project: W-167-R Work Plan: Job Title: Period 13 Personnel: Upland Job: 01 January Kenneth M. Giesen Clait E. Braun, Bird Research 10 Movements, Reproductive Success, Plains Sharp-tailed Grouse Covered: Author: REPORT and Habitat through 31 December Kenneth M. Giesen, Use by Introduced 1994 Colorado Division of Wildlife ABSTRACT Plans to transplant plains sharp-tailed grouse (Tympanuchus phasianellus jamesi) into eastern Colorado were temporarily postponed in 1994 because a suitable release site was not available. Additional habitat evaluations were conducted at the proposed release site near Raton Mesa in Las Animas County and preparations were made to transplant sharp-tailed grouse to this site beginning in 1995. ��MOVEMENTS, REPRODUCTIVE SUCCESS, AND HABITAT INTRODUCED PLAINS SHARP-TAILED GROUSE Kenneth USE BY M. Giesen INTRODUCTION Plains sharp-tailed grouse historically occurred in suitable foothills and riparian habitats along the Front Range of Colorado. Sharp-tailed grouse populations declined with human settlement and were extirpated from most of their range in eastern Colorado by the late 1800's. In recent years breeding populations were documented only in Douglas County, although winter migrants or transients have been reported from Yuma, Logan, and Weld counties (Hoag and Braun 1990). Plans to increase distribution and populations of plains sharp-tailed grouse in Colorado will rely primarily on transplants (Braun et al. 1992). While numerous transplants of prairie grouse have occurred, few have been successful (Toepfer et al. 1990, Rodgers 1992, Hoffman et al. 1992). Thus, it is desirable to document responses of sharp-tailed grouse to experimental transplants and evaluate parameters potentially affecting success including movements, habitat use, mortality, and reproduction. P. N. OBJECTIVES The objectives of this project are to assist with trapping and transplanting of plains sharp-tailed grouse into selected sites along the Front Range of Colorado and evaluate transplant success. Population characteristics of the transplanted population including movements and home range size, timing and causes of mortality, habitat use, and nest success will be compared to those described in the literature for native and transplanted prairie grouse. SEGMENT on prairie OBJECTIVES 1. Review literature habitat use. grouse introductions, movements, and 2. Coordinate landowners for plains 3. Transplant up to 50 plains sharp-tailed grouse from southeastern into suitable habitats along the Front Range of Colorado. 4. Radiomark up to 25 sharp-tailed grouse in the transplanted population and monitor movements, habitat use, reproduction, and mortality. 5. Conduct site. a pre-release 6. Prepare annual efforts with Wyoming Game and Fish personnel and affected in southeastern Wyoming to locate potential trapping sites sharp-tailed grouse. evaluation progress of the habitat at the selected Wyoming release report. METHODS Contact was made with personnel of the Wyoming Game and Fish Department to obtain the necessary permits for trapping plains sharp-tailed grouse in southeastern Wyoming for transplant into Colorado. Active dancing grounds in Wyoming were located as potential trapping sites (Toepfer et al. 1988, Schroeder and Braun 1991). Personnel of the SE Region of the Colorado Division of Wildlife prepared a vegetative cover map of the release site at Raton Mesa. �46 RESULTS Transplant of sharp-tailed AND DISCUSSION grouse Permission to release sharp-tailed grouse was not obtained from the 2 primary transplant sites, Rocky Flats and Fort Carson (Braun et al. 1992). Additional release sites in Larimer county near Livermore and in L~s Animas County near Raton Mesa were evaluated. Vegetation at both sites was cover mapped and landowner contacts were made to secure permission to transplant birds and conduct evaluations. No transplant occurred in the spring of 1994 as prerelease evaluations were not completed. A planned experimental fall release of up to 10 sharp-tailed grouse from southeastern Wyoming into the Livermore area in Larimer County, Colorado was cancelled because permission for capturing sharp-tailed grouse by nightlighting of CRP fields in Wyoming could not be obtained. Surveys of known dancing grounds in Wyoming indicated that attendance and display of sharp-tailed grouse was sporadic in October and no hens were observed. Plans have been initiated to trap up to SO plains sharptailed grouse from Wyoming for release into Colorado in 1995. LITERATURE CITED Braun, C. E., R. B. Davies, J. R. Dennis, K. A. Green, 1992. 'Plains sharp-tailed grouse recovery plan. Denver. 33 pp. and J. L. Sheppard. Colorado Div. Wildl., Hoag, T. W., and C. E. Braun. 1990. Status and distribution tailed grouse in Colorado. Prairie Nat. 22:97-102. of plains sharp- Hoffman, R. W., W. D. Snyder, G. C. Miller, and C. E. Braun. 1992. Reintroduction of greater prairie-chickens in northeastern Colorado. Prairie Nat. 24:197-204. Rodgers, R. D. 1992. A technique for establishing sharp-tailed unoccupied range. wildl. Soc. Bull. 20:101-106. grouse in Schroeder, M. A., and C. E. Braun. 1991. Walk-in traps for capturing prairie-chickens on leks. J. Field Ornithol. 62:378-385. Toepfer, J. E., R. L. Eng, and R. K. Anderson. 1990. Transplanting prairie grouse: what have we learned? Trans. N. Am. Wildl. and Nat. Resour. Conf. 55:569-579. __________, J. A. Newel, and J. Monarch. 1988. A method for trapping prairie grouse hens on display grounds. Pp. 21-23 in Prairie chickens on the Sheyenne National Grasslands. (A. D. Bjugstad, Tech. Coord.). u.S. Dept. Agric. For. Servo Gen. Tech. Rep. RM-159. Prepared by Kenneth M. Giesen Wildlife Researcher C �47 Colorado Division Wildlife Research April 1995 of Wildlife Report JOB PROGRESS State of: Colorado Project: W-167-R Work 17 Plan: Population Job Title: Period Covered: Author: Personnel: REPORT Clait Upland Job 7 Dynamics 01 January Bird Research of White-tailed through E. Braun and Kenneth 31 December Ptarmigan 1994 M. Giesen Kathy Martin, University of British Columbia; Clait Kenneth M. Giesen, Colorado Division of Wildlife E. Braun and ABSTRACT Long-term studies of populations of white-tailed ptarmigan (Lagopus leucurus) were continued at hunted (Mt. Evans) and unhunted (ROcky Mountain National Park) areas in Colorado through 1994. Breeding densities of ptarmigan decreased slightly at Rocky Mountain National Park (RMNP) and increased at Mt. Evans. Breeding population levels continue to be low at RMNP but have increased from the low in 1993 at Mt. Evans. This low was the result of overharvest in 1992. Nest success at RMNP in 1994 was good (62%) as was brood size (3.2 chicks/hen) but was poor at Mt. Evans (20%, 1.0 chicks/hen). Nest success was poor for the 4th consecutive year at Mt. Evans. However, apparent nest success in adjacent areas appeared good as recruitment of yearlings in spring 1994 was good. Harvest regulations at Mt. Evans remained restrictive in 1994 and no harvest was known to occur. With the 1/2 mile closure (permanent) along the Mt. Evans highway to all hunting, harvest of ptarmigan in the study area is expected to remain close to zero. ��49 POPULATION DYNAMICS Clait E. Braun OF WHITE-TAILED and Kenneth PTARMIGAN M. Giesen Long-term studies of trends in population size and investigation of reasons for fluctuations in size of tetraonid populations are lqcking. Studies on the population dynamics of unhunted and hunted populations of white-tailed ptarmigan were initiated in Colorado in 1966 and have continued essentially uninterrupted at 2 sites. Studies of the unhunted population (Rocky Mountain National Park) identified possible short-term cycles of 7-8 years with an amplitude of 25-30% between high and low breeding densities. Conversely, studies of the manipulated population (hunted) at Mt. Evans have not indicated any cyclic pattern and it would appear that controlled hunting may mask any long-term trend that may occur. This study is designed to examine the question whether white-tailed ptarmigan are truly cyclic and whether hunting affects the apparent oscillations. P. N. OBJECTIVES The goals of this investigation are to be able to predict the length and amplitude of cycles in white-tailed ptarmigan in Colorado, to examine the impact of hunting on cycles, and to clarify underlying c?uses of the apparent cycles. SEGMENT OBJECTIVES 1. Conduct breeding (May-Jun) and brood (Aug-Sep) censuses ptarmigan using tape-recorded calls of males (breeding) (broods). 2. Censuses will be conducted on previously established, defined study areas at Mt. Evans (hunted) and at Rocky Mountain National Park (unhunted). 3. Capture (noose poles) and band (aluminum and plastic color-coded bands) all unmarked white-tailed ptarmigan encountered on study areas at Mt. Evans and at Rocky Mountain National Park. 4. Individually identify all ptarmigan observed on study areas and Rocky Mountain National Park through use of binoculars. 5. Make hunting season and bag limit recommendations for Mt. Evans and collect hunting data through use of volunteer wing barrels and hunter field checks. 6. Compile data, analyze results, and prepare progress of white-tailed and chicks at Mt. Evans reports. STUDY AREA AND METHODS Areas investigated were Mt. Goliath-Mt. Evans in Clear Creek County and at Tombstone Ridge-Sundance Mountain to Fall River Pass in Rocky Mountain National Park in Larimer County. The physiography, geology, location, and vegetation of these study areas have been previously described (Braun 1969, 1971; Braun and Rogers 1971; Giesen 1977). Ptarmigan were located through use of tape-recorded calls (Braun et ale 1973), captured through use of telescoping noose poles (Zwickel and Bendell 1967) as described by Braun and Rogers (1971), and classified to age and gender and banded following Braun and Rogers (1971). Age of chicks was estimated following Giesen and Braun (1979). Numbered plastic bandettes were not used as in earlier years (Braun and Rogers 1971) as a color-code system using up to �50 4 different colored plastic bandettes was instituted in 1977-78. A check station was not operated on the Mt. Evans highway during the opening weekend of the ptarmigan season in that area as the area within 1/2 mile of the road was closed to all hunting. A volunteer wing collection station was available to hunters in the area until the season closed. RESULTS Breeding AND DISCUSSION Densities Mt. Evans. -- Ten pairs and 2 single males were recorded on the original study area in 1994 (Table 1). This represented an increase from the low breeding population in 1993 that resulted from the overharvest in 1992. Three of the 10 territorial males (with hens) were yearlings as were both single males in spring 1994 and 3 of the 10 hens. Rocky Mountain National Park. -- Timing of breeding events on the Trail Ridge study area was one week earlier than in 1993 and two weeks earlier than the 1966-93 average. Surveys of ptarmigan on breeding territories along Trail Ridge Road in May and June indicated a minimum population of 58 birds and included 21 pairs and 16 unpaired males. Densities on the original study area decreased slightly from 1993 (Table 1). The decrease in breeding density reflected high survival of banded adult males (47 of 64, 73.4%) but below average survival of banded adult females (12 of 25, 48.0%) from 1993. Four yearlings banded as chicks in 1993 (all males) recruited to the study area and yearlings comprised 21.1% (16 of 76) of all adult ptarmigan identified. Nesting Success and·Brood Size Mt. Evans. -- Timing of breeding in 1994 was early and most hens initiated egg laying in early June. Clutch size was small although clutches of 8 and 9 eggs were recorded (K. Martin, pers. commun.). Hatching success was poor in 1994 and only 2 of 10 hens seen in August-early September had broods (20%). Brood size was also poor as each hen had 1 chick. These chicks hatched on about 13 July and 4 August. This was the 4th consecutive year of poor nest success at Mt. Evans. However, the spring breeding population indicates good recruitment suggesting that nest success and production of young in areas adjacent to Mt. Evans were better than at the study area. Rocky Mountain National Park. -- Nest success was estimated from the proportion of hens with and without broods observed during July and August. Ten of 16 hens observed during summer surveys were with broods for an estimated nest success rate of 62.5%, up from 44% in 1993. The median hatch date calculated from wing molt of 20 juveniles was 1 July (range 26 Jun-22 Jul) and was two weeks earlier than the 1966-1993 average. Brood size in August averaged 3.2 chicks/hen (range 1-7). Harvest Mt. Evans. -- The ptarmigan hunting season was again delayed in the Mt. Evans area (east of the Guanella Pass Road and south of 1-70) from the statewide (1 Sep - 2 Oct) or Pikes Peak (10 Sep - 2 Oct) season dates and opened on 19 September and closed on 30 September. The bag/possession limit for Mt. Evans. was 1/2 and all hunters had to have a free, unlimited in number, permit. Further, only 2 ptarmigan were allowed for each permittee during the entire season and all birds harvested were to be tagged (not meaning banded). Also, hunting (all species) was not permitted within 1/2 mile of the Mt. Evans highway (Colorado 5). �51 Table 1. 1966-94. White-tailed ptarmigan breeding densities (birds/km2), Study area Rocky Mountain National Park Year (5.5 1966 1967 1968 1969 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 Colorado Mt. Evans (4.0 )on2) )on2) 11.3 9.8 11.5 12.0 9.6 9.1 8.7 7.8 8.0 11.1 13.5 12.9 10.7 8.7 8.4 8.2 7.8 6.7 5.8 6.0 4.5 6.0 5.4 6.2 7.6 6.7 7.6 6.0 5.8 3.0 2.7 2.7 2.2 2.0 4.2 7.5 6.2 6.2 6.2 6.7 > 6.0 7.5 10.3 9.5 9.0 6.5 6.5 8.0 8.0 6.5 5.0 7.5 8.0 8.0 8.2 6.5 4.0 5.5 A check station was not operated in 1994 but road checks were conducted. No hunters were found and no wings were deposited in the volunteer wing collection barrel. About 50 ± permits were issued in 1994 compared to 53 in 1993. Survey data for the permittees in 1993 or 1994 are not available. However, there is no evidence that ptarmigan were harvested within 1/2 mile of the Mt. Evans road in 1994. No bands were reported and the harvest appeared to be zero. LITERATURE Braun, C. E. 1969. Population dynamics, tailed ptarmigan in Colorado. Ph.D. Collins. 189pp. _____ CITED habitat, and movements Thesis, Colorado State 1971. Habitat requirements of Colorado white-tailed West. Assoc. State Game and Fish Comm. 51:284-292. _____ , and G. E. Rogers. Colorado Div. Game, of whiteUniv., Fort ptarmigan. 1971. The white-tailed ptarmigan in Colorado. Fish and Parks Tech. Publ. 27. 80pp. Proc. �52 _____ , R. K. Schmidt, tailed ptarmigan Jr., and G. E. Rogers. 1973. Census of Colorado whitewith tape recorded calls. J. Wildl. Manage. 37:90-93. Giesen, K. M. 1977. Mortality and dispersal of juvenile white-tailed ptarmigan. M.S. Thesis, Colorado State Univ., Fort Collins. 55pp. _____ , and C. E. Braun. 1979. A technique for age determination white-tailed ptarmigan. J. Wildl. Manage. 43:508-511. Zwickel, F. C., and J. F. Bendell. J. Wildl. Manage. 31:202-204. Prepared 1967. A snare for capturing of juvenile blue grouse. by Clait E. Braun Wildlife Research Leader Kenneth M. Giesen Wildlife Researcher C �53 Colorado Division Wildlife Research April 1995 of Wildlife Report JOB FINAL REPORT (Research) state Project: Work Plan: study Title: Colorado Upland W-167-R 21 Bird Research : Job __~8~ __ Increasing foods for wildlife eastern Colorado within the rangelands of Personnel: L. L. Bixler, L. K. Haynes, K. L. Martin, J. W. Moore, J. L. Mekelburg, and W. D. Snyder, Colorado Division of Wildlife ABSTRACT Evaluations were conducted to determine establishment, survival, growth, and seed production of selected herbaceous species as food sources for avifauna within the Tamarack Prairie in eastern Logan County, Colorado. Eight rangeland sites, each <1 ha in size, were tilled to destroy existing perennial vegetation in early spring 1992. Each site contained a randomly selected block of seeded ann~als, seeded perennials, disturbance tillage (on 5 of 8 sites in 1992), and an undisturbed control. Perennials were established in spring 1992. Annuals were planted during 1992, 1993, and 1994, but satisfactory results were obtained only during 1992. Annuals failed to attain satisfactory stands in 1993 and marginal results were obtained in 1994 because of severe drought. Disturbance tillage produced varied results among years but findings indicated late spring tillage is needed annually to thin stands and promote tall, sparse cover. Wild sunflowers (Helianthus annuus), millets, and sorghums were best adapted among seeded annuals and should dominate in mixtures with other tame species planted in late spring for upland birds. Alfalfa of falcata parentage should be the primary component within perennial mixtures planted in tracts large enough (1-3 hal to withstand grazing by wildlife. ��55 INCREASING FOODS FOR WILDLIFE WITHIN Warren THE RANGELANDS OF EASTERN COLORADO D. Snyder INTRODUCTION Lack of food is hypothesized as a primary limitation for greater prairiechickens (Tympanuchus cupido), lesser prairie-chickens (~. pallidicinctus), scaled quail (Callipepla squamata), northern bobwhite (Colinus virginianus), sharp-tailed grouse (~. phasianellus), and mourning doves (Zenaida macroura) in eastern Colorado sandhill rangelands. In most instances, densities of these species are relatively low and all prairie grouse are listed as threatened or endangered in Colorado. These species are primarily seed eaters as adults and prefer the energy-rich seeds of annuals. Prairie grouse also consume green vegetation including legumes, and green vegetation is used through fall and winter if available. Young of all species (except mourning doves) primarily consume insects during the initial 2-3 weeks after hatching. Perennial and annual forbs comprise <10% of vegetation species composition within the ungrazed Tamarack Prairie and many of these forbs do not yield seeds of value to upland birds. Seed-producing forbs are even less available in most grazed ranges. Prairie grouse have not used the Tamarack Prairie extensively even though livestock have been excluded for >15 years and heightdensity of residual vegetation has increased. Both numbers and distribution of greater prairie-chickens increased markedly in Yuma, Phillips, and eastern Washington counties following extensive development of corn and other irrigated crops using center-pivot irrigation systems in the early 1970's. Scaled quail and northern bobwhite reside in rangelands only if combinations of protective shrubby cover and abundant annuals are present, therefore, densities of these species are usually low. However, in one southeastern Colorado location, containing abundant wild sunflowers, tall sand sagebrush (Artemisia filifolia), and other shrubs, 4 or 5 coveys were found within a 68ha tract during December 1994. These instances and others provide strong evidence that food is limiting in most rangeland situations. There is need to determine what food species are best suited for use in dry rangeland situations in eastern Colorado. Both seed-producing species and those that will yield green vegetation, especially into early winter, need to be evaluated. P. N. OBJECTIVE Evaluate the establishment, survival, growth, and seed production of selected herbaceous species as food sources for avifauna within the Tamarack Prairie of eastern Logan County. Ascertain the presence/abundance of selected wildlife during brood rearing (summer) and fall-winter intervals. Assuming objectives 1 and 2 are attained, a potential 3rd objective will be to test their adapt ion to other rangeland sites in eastern Colorado for scaled quail, lesser prairiechickens, mourning doves, and other species. SEGMENT 1. Prepare sites 2. Monitor the relative establishment, qualities of tested species. 3. Monitor the presence/abundance 4. Monitor precipitation 5. Prepare a job final report. OBJECTIVES for planting. survival, of selected growth, wildlife and other environmental and food producing using factors. the test sites. �56 METHODS 1. Literature concerning avifauna food habits was reviewed and horticulturists and range specialists were contacted to develop a list of plant species and varieties potentially of high food value for wildlife and suited for rangelands in eastern Colorado. Seeds of several species and varieties were obtained using both acquisition apd hand collection. 2. Initially,S sites, each containing <1 ha were placed linear and perpendicular to prevailing winds among the Dailey and Valent loamy sandy soils (Amen et al. 1977) within the eastern part of the Tamarack Prairie (3 south of Highway 1-76 and 2 north of the highway (Fig. 1). A management directive precluded planting certain exotic species south of Highway 1-76. Therefore 3 additional plots were added north of the highway in April 1992. Five initial sites were mold-board plowed in March 1992 by J. L. Mekelburg and subsequently tilled with a cultipacker prior to planting. Three supplemental sites were double disced with a tandem disc in April 1992 to destroy existing vegetation. A tractor-mounted rototiller was used for tillage prior to planting in subsequent years. Each site contained 3 randomly selected plots: (1) seeded, (2) disturbance tillage, and (3) undisturbed control. The 3 sites established last did not contain disturbance tillage because of dry soils in spring 1992. Within seeded plots, species were separated for planting and evaluation into 2 groups: (1) seed producers (primarily annuals), and (2) species yielding green leafy vegetation (primarily perennials). Perennials were planted in 1992 and not in subsequent years. Selected species were planted within each site using randomized patterns. Planting was completed between late April and mid May 1992 and in early May 1993 and 1994. A push-type garden seeder, equipped with seeder plates for different seed sizes, was used for planting annuals in blocks containing 10 -12 rows, spaced 5 - 6 dm apart and 10 - 12 m long during 1992-93. Efforts were made, based on seed size, plant size, and past experience, to seed at proper rates and depths. In 1994, plantings were with a tractor-mounted 4-row surface planter (7.6 dm row spacings) and most plots were tilled with a tractor-mounted cultivator once in late June. Species blocks were marked with metal pins and labeled with metal tags for identification. Species planted for green, leafy vegetation were in linear strips approximately 6-m wide and as long as the tilled site was wide (approximately 40 m) on each site. A drill equipped with double-disk furrow openers (1.8 dm spacing between openers), 2.5-cm band attachments, packer wheels, and an alfalfa seed box was used. Within sites 6 - 8 (Fig. 1) all species were seeded by this method in 1992. A shop vacuum, powered by a portable generator, was used to remove surplus seed from the drill box before planting the next species. Disturbance tillage was established in plots at least 10 x 50 m (0.05 ha) placed perpendicular to prevailing winds at each site. Because of concern for wind erosion, wild sunflower was drilled into approximately one-half of each disturbance site in 1992 (100% of the disturbance tillage area within site 3). Transects to compare vegetation within controls were positioned about 30 m southwest of, and parallel to, each test site. 3. The relative establishment, survival, growth, and food producing qualities of species within seeded and disturbed sites were monitored periodically from June through October each year. Within plots planted for seed production, post-seeding inspections were conducted to determine status as (1) no establishment, (2) established but failed to survive, and (3) established and survived. �57 A 0.5 x 1.0-m Daubenmire frame was used to estimate canopy cover to measure establishment, survival, and growth (Daubenmire 1959). Sampling was replicated 12 times per plot at 1 to 2 pace intervals (depending on plot size) in a diagonal direction across each plot. sampling was conducted between 28 August and 3 September 1992 and in late September 1994. Vegetation was classified as the percentage of seeded species, competing annual forbs, annual grasses, perennial forbs, perennial grasses, bare ground, and dead vegetation. This procedure was used within seeded annuals, seeded perennials, disturbance tillage, and control plots on all sites. Height (dm) of seeded vegetation and total vegetation was sampled within the Daubenmire frames. Seed production within all plots was rated as: 0) - no production, 1) - low production, 2) - moderate production, and 3) high production, based on the expected potential of the species to yield seed under favorable growing conditions. 4. Wildlife use of seeded annuals, seeded perennials, disturbance tillage, and control plots was sampled during periodic visits to the 8 sites from mid-June 1992 through March 1993, and at infrequent visits during late summer, fall, and winter 1994-95. Wildlife presence or absence was determined, categorized by wildlife group: gallinaceous birds, passerines, mourning doves, and other avifauna (data were listed by species when possible). Abundance, when present, was delineated as ~ 5, and> 5/species or group. Narrow « 2 m), disturbed-soil buffers were maintained around seeded and disturbed tracts using a tractor~mounted rototiller in August 1992 but not in subsequent years. Observations of tracks within the buffer strips were recorded during periodic visits from mid-June through March to aid in ascertaining use by deer, rabbits (Lagomorpha), rodents (Rodentia), and gallinaceous birds. Establishment, survival, and food producing problems by species and variety, occurring because of depredation by deer, rodents, rabbits, or other wildlife, were documented. 5. Automatic precipitation recorders were used to monitor precipitation. Gauges were placed at sites 2 and 5 (Fig. 1). Precipitation during winter months was obtained using u.S. Weather Bureau records from nearby recording stations. �58 Fig. 1. Prairie, I I I i I , I I I .J Z- Q Z E ''"" ... "w - ••~ oJ a ,._ ~ -- __________ w a: > a: ~ Location and sequence of rangeland Logan County, Colorado, 1992-94. C") £!1l!!.0.2... 0 W Ict,) W a-: <t 0:: a. ~ 0 N '" , __ • ;::; 0 N e ..• •• ,.. w !:: 4 ct,)D ('II ~ iii _ ~ , I •, ,, 3: •• r ..• 3: <II ~a: ~ • ,! food test sites with the Tamarack �59 RESULTS Precipitation and Planting Conditions Although precipitation was deficient in April-May 1992, above average precipitation in late May and June stimulated establishment and growth of most annuals and perennials (Table 1). January through August precipitation averaged above the long-term mean in 1992 on the Tamarack Prairie. Rain gauges did not operate consistently during 1993, but data from 3 nearby U.s. Weather Bureau stations indicated January-August rainfall was about average within the area (Table 1). However, plantings in May 1993 failed to establish satisfactory stand~ of annuals and time constraints did not permit replanting. Precipitation was extremely deficient from April through June 1994 on the Tamarack Prairie. Only two-thirds the normal precipitation was received from January through August 1994 (Table 1). As a consequence, test plot vegetation was severely stunted and meaningful data during this study was primarily obtained in 1992 and reported earlier (Snyder 1993). Table 1. Precipitation (cm) in the vicinity of the Tamarack from 1992 to 1994 in relation to the long-term average. 1992 Month Jan Feb Mar Apr May Jun Jul Aug South 2.79b 1.25b 5.36b 0.56 2.44 14.58 3.35 6.53 Totals 8 b c 1993 2.79 1.25 5.36 0.46 1.70 14.58c 6.98 12.04 36.86 x8 1994 South North Prairie North 0.69 3.38 1.85 3.48 6.43 5.61 4.88 6.02 1. 75 1.19 1.07 2.74 2.41 2.24 8.43 1.52 1. 75 1.19 1.07 2.74 4.78 0.97 6.12 1.63 45.16 32.34 21.35 Long-term 0.81 0.89 1.98 5.13 6.48 6.40 5.31 5.18 20.2532.18 Based on U.S. Weather Bureau records during a 44-year interval. Data averaged between Sterling and Sedgwick stations. Rainguage destroyed by hail. Precipitation believed> than 14.58 cm. Rainfall averaged 21.41 cm at Sterling and Sedgwick weather stations. Plantings Completed During 1994 Three varieties of sorghums (1 grain sorghum plus 2 taller cane sorghums, [Sorghum vulgare]) and wild sunflowers were seeded in all sites in early May 1994. White wonder millet (Setaria italica), pearl millet (Pennisetum glaucum), Rocky Mountain beeplant (Cleome serrulata), safflower(Carthamus tinctorius), annual buckwheat (Polygonum sp.), and tame sunflowers were planted in single test plots within site 3. Characteristics of 1994 plots and limited comparisons with 1992 data varied (Table 2). Fair to good stands of sorghums were obtained on all sites, however, wild sunflower (planted at the same time and depth) did not establish on any site for unknown reasons. Due to dry weather in 1994, percent canopy cover approximated only 15-20% for planted sorghums and bare ground averaged >50%. In contrast, grain sorghum approximated 56% canopy cover and bare ground was 21% in 1992 plots (Table 2). Cane sorghums, which attain heights of 1.5 - 2 m under favorable growing conditions, were only slightly taller (about 0.6 m) than the grain sorghums. Cane sorghums did not stand well over winter to provide significant cover. �60 Table 2. Canopy cover (%) for tested species vegetation within controls, Tamarack Prairie, Vegetation Variable Plots (n) Bare grnd. Planted species and varieties, disturbance Colorado, 1992 and 1994. Canopy Cover Annual Peren. Peren. Annual forbs forbs grass grass tillage, and Height(dm) Seed Planted Total prod. species veget.ratea 1994 G Sorghum Occur. 8 1{ SE MorCane Occur. 8 1{ SE Frmt. Cane Occur 8 1{ SE 8 8 49.6 5.3 8 16.7 3.1 8 51.5 4.2 15.6 3.4 29.9 4.5 8 54.6 3.6 8 20.2 4.2 21.5 4.2 8 25.8 2.9 8 8 4 1.1 0.6 2 0.2 0.1 2 0.4 0.3 4.4 8 5.3 0.6 5.6 0.5 1.4 0.2 4 1.0 0.6 3 2.0 1.8 8 6.2 0.9 6.4 0.8 0.9 0.2 4 0.6 0.3 4 2.7 1.3 8 6.5 0.9 6.7 0.8 0.7 0.2 2 0.6 0.5 5 6.2 W.W. Millet 1 62.9 17.9 19.2 3.8 0.0 Pear!' Millet 1 60.4 29.6 10.0 4.1 0.5 8 8 39.7 3.7 5.2 0.4 1.3 0.2 2.4 0.2 0.2 0.1 6.2 0.6 2.3 0.2 11.0 2.0 2.5 0.3 3.3 0.1 0.3 Dis. till. Occur 1{ SE Rangeland (Control) Occur 8 1{ SE G sorghum Occur 8 1{ SE Dist. till. Occur. 5 SE Rangeland (control) a b c Occur. 1{ SE 8 42.8 2.5 8 20.6 3.3 5 12.4 1.6 1{ 8 8 53.7 3.9 8 24.7b 1.5 8 56.1 4.3 3 5 1.3 0.7 2.3 1.2 7 5 1.7 0.6 2.0 1.2 8 20.2 3.6 5 80.9 2.3 8 6.4 1.6 8 52.4 2.3 3 3.1 1.8 1 0.5 o 3 6 0.8 0.7 2.2 0.9 2 1 0.1 2.5 4 1.8 0.6 8 4.0 2.6 8 64.7c 4.0 1 0.2 4.4 5.8 0.7 Seed production was rated on an ascending scale from 1 to 3 based capability of the species to produce seed. Includes bare ground and dead vegetation. Includes sandsage. on the estimated �61 White wonder millet established 18% canopy cover (63% bare ground and 19% forbs), averaged 3.8 dm in height, and did not produce seed. In contrast, it had averaged almost 1 m tall and yielded excellent seed production (rated 2.9) in 1992 (canopy cover was 62%). Canopy cover of pearl millet was approximately 30%, (60% bare ground and 10% annual forbs), its height averaged 4 dm and its seed production rating was 0.5 or marginal within the one site. This species was not planted during 1992. Rocky Mountain beeplant was seeded into one plot within site 3 in May 1994 but, as in previous years, it failed to establish. Other species that failed to establish included safflower, annual buckwheat, and tame sunflower. Annual kali), cover, at 2.5 ground 1.3 in present forbs, primarily pigweed (Amaranthus sp.) and Russian thistle (Salsoli dominated in disturbance tillage plots during 1992 with 81% canopy 12% bare ground, a height exceeding 1 m, and good seed production rated (Table 2). Their percentage canopy cover (54%) was much lower, bare was 40%, height approximated 0.5 m, and seed production was rated at 1994. Few wild sunflowers, major contributors to height and food, were in disturbance tillage during 1994. Within untreated controls (native range), perennial grass continued to dominate at 52% canopy cover in 1994, whereas bare ground was higher than in 1992 (Table 2). Percentages of annual and perennial forbs were also less than during 1992. In combination, they equaled only 3.7% of the canopy cover within ungrazed controls during 1994. Perennials established during 1992 Dense stands of most perennials (and biennials) were attained within several sites during June 1992. Vegetation sampling of canopy cover, height, and seed production was conducted in late summer 1992 and results were generally positive (Snyder 1993). However, several factors resulted in poor survival and growth. Persistent heavy grazing by deer, rodents, and insects in combination with dry weather, competition with annual weeds, and poor soils did not permit most perennials to attain significant growth, become well established, or build energy reserves for survival. If the test plots had been much larger (1 to 2 ha), grazing intensity might have been markedly reduced. Several perennial legumes, including purple prairie clover (Petalostemon purpureum), ladino clover (Trifolium repens), perennial sweet pea (Lathyrus latifolius), and Illinois bundleflower (Desmanthus illinoiensis) showed marginal growth and survival during 1993, did not compete with annual weeds, and most failed by 1994. Perennial sweet pea was a tame "garden" variety, not the sweet pea native to the Tamarack Prairie. Cicer milkvetch and Sainfoin showed better survival and adaptation to rangeland sites but little growth and poor survival was evident during 1993 and 1994. Grazing, weed competition, and insects were believed factors in their suppression. Two varieties of alfalfa (Medicago sativa), spreador II and ladak, were each established in 4 sites in 1992. Stands, which initially were dense, thinned or died but alfalfa survived in several locations through 1993 and 1994. Observations in summer 1993 revealed intense grazing by deer and consumption by insects. Alfalfa was not able to make significant growth and build energy reserves for survival and competition with annuals because of high use. None of the biennials seeded in 1992, which included hairy vetch (Vicia villosa), common vetch (Vicia americana), and yellow sweet clover (Melilotus officinalis), persisted until 1994. Vetches are winter annuals and should be planted in late summer. Hairy vetch survived or reestablished on site 2 during 1993 but it and common vetch did not persist in other sites. Yellow sweet clover was too dense to establish tall cover in 1993 and was completely defoliated by grasshoppers in early summer. �62 Small burnet (Sanguisorba minor) grew well during 1992 but gradually died out in most locations during 1993 and 1994. Lewis blue flax (Linum perenne) established and retained near pure stands, approximately 0.3 m tall, that continued to thrive through 1993 and 1994. It out-competed annual forbs which did not persist within test plots. This species was best adapted to the site among all the perennials tested. Wildlife Occurrence Although considerable use by several species of wildlife was evident during 1992, wildlife use diminished drastically during 1993 and 1994. Deer grazing of alfalfa was noted during 1993 but was relatively minor during 1994 because of poor growth. Kangaroo rats (Dipodomys sp.) were the most common residents within all sites and created much disturbance with their burrowing and other activities. A greater prairie-chicken was flushed from site 3 in spring 1994, but other evidence of grouse activity there was not observed. Grouse use of the site had been noted in 1992. Seed production from seeded and wild annuals and growth of tall cover was not adequate to hold passerines or other upland birds in 1994. Observations of all plots during early January 1995 showed a small flock of American tree sparrows (Spizella arborea) at site 2 but no wildlife other than rodents at other plots. DISCUSSION This study was handicapped by time constraints in 1993 and 1994, and severe drought conditions through spring and early summer 1994. However, considerable information was obtained and supplemented by information obtained in previous studies (Snyder 1967, 1978) and from literature. High food production and wildlife use, as obtained during 1992, can not be expected every year within sandy rangelands in eastern Colorado. However, study findings indicate that with proper management, satisfactory stands of annuals can be obtained most years. Disturbance Tillage Plots Disturbance tillage, used to destroy native perennial grasses, was effective in promoting wild annuals which averaged less than 5% of the canopy cover within native rangelands. Plowing or discing to destroy nearly all perennial vegetation is usually needed in plot establishment. However, Noble blades or other sweep-plows that undercut while retaining surface residue to reduce erosion may be effective in subsequent years. Initial tillage of sod should occur in early spring (mid Mar - early Apr), but if plots are to be productive, tillage in subsequent years must be conducted in late spring (late May - early Jun). Early spring tillage usually allows high densities of forbs to reestablish resulting in short, stunted stands often dominated by Russian thistle. Sparse stands are essential to obtain satisfactory growth and seed production. Russian thistle, pigweed, lambsquarter (Chenopodium album), and kochia (Kochia scoparia), have some seed and cover values for wildlife, but, species such as wild sunflowers, Texas croton (Croton texensis), snow-on-the-mountain (Euphorbia marginata) which yield larger nutritious seeds are preferred. Observations indicate wild sunflower is one of the best adapted and most valuable species for both food and cover. Seeds of many annuals such as crotons and sunflowers are often present within rangeland soils but conditions for their germination and establishment are not favorable every year. Commercial seed sources are available for some, e. g., wild sunflowers, Rocky Mountain beeplant, which can be planted to augment wild populations. Commercial seeds of natives are usually expensive, sources are limited, and seeds do not always grow when planted due to soil moisture, soil temperature, planting depth, need for scarification, and other unknown factors. Spring plantings of Rocky Mountain beeplant were unsuccessful during this evaluation. This species possesses a �hard shell and should either drained, frozen, and planted (R. stevens, pers. commun.). similar manner. be fall seeded or soaked in water for a day, in early spring immediately after it is thawed Other hard-seeded natives should be treated in Planting may be the only way to get some species started in locations where they are absent or uncommon. Harvesting by hand or with machinery can be used to obtain seed from wild sources for native annuals. c~ammy weed (Polanisia dodecandra), toothed spurge (Euphorbia dentata), horse-mint (Monarda fistulosa), buffalo-bur (Solanum rostratum), and field pennycress (Thlapsi arvense) are suggested as possible species for collection and propagation in eastern Colorado rangelands. Field pennycress and other mustards must be planted in fall, late winter, or early spring or they will not produce seed. Tame Annual Plantings Individual varieties or species were planted in test plots during this study, however, mixtures should be used in most situations. At least some species or varieties will usually establish and grow even though weather conditions may not be suited for all species within a mixture. Individual species or varieties also have differing maturity dates, moisture requirements, and preferences by wildlife helping to increase the value of the plot. Numerous variations can be used in seed mixtures. Varieties of early and medium maturing grain sorghums, taller sorghums or sorghum/sudan grass hybrids, foxtail millet, pro so millet (hersey), safflower, wild sunflower, and small seeded tame sunflower are suggested components. Some species, such as proso millet, mature early to provide food for mourning doves and other wildlife in late summer and early fall. Sunflowers and sorghums provide seed into fall and winter. Safflower, in 1992, retained its seed until consumed by pheasants in mid winter when little other food remained. Hungerford (1948) reported excellent seed retention of safflower into winter and noted its drought tolerance after establishment. Tall sorghums usually are more important for the cover they provide to feeding wildlife than for the seed produced. Study findings indicate millets and sorghums were among the best adapted, most easily grown tame annuals, had good food producing potential for wildlife in eastern Colorado sandhills, and should be basic components in most seed mixtures. Foxtail and proso millets require less rainfall to produce seed than any other crop grown in eastern Colorado (Greb 1978). Farmers commonly grow sorghums in soils considered too sandy for winter wheat production in southeastern Colorado. Strips or plots to be planted should be tilled in early spring (Mar) to kill perennial grasses. Observations show that dense weed growth is seldom a problem during the first year that a site is broken from sod but it becomes an increasing concern in subsequent years. Shallow tillage should be replicated just before planting in late Mayor early June. Retention of surface residue by use of sweep tillage is recommended to reduce erosion. Use of strips up to 25 m wide, placed perpendicular to prevailing winds, or use of narrow undisturbed buffer strips in wider plots, will also help suppress erosion. A grain drill capable of planting different-size seed at once is needed. Tame annuals can be rotated biennially with disturbance tillage on two proximal strips to increase diversity and reduce effort. Perennial Species Establishment Alfalfa had the best survival among several perennial legumes planted on the Tamarack Prairie, but it was persistently suppressed by deer, rodents, and insects during 1992-93 and by drought in 1994. Alfalfa was the most promising among 14 legumes tested in sandy and loamy soils in northeastern Colorado in the early 1970's (Townsend et ale 1975). McGinnies and Townsend (1983) found that alfalfa lived longer (7 years) than sainfoin (5 years), but was impacted by pocket gophers (Geomys spp.). They planted it in mixtures with cool-season grasses. Sicklepod milkvetch (Astragalus falcatus) survived better than either �of the above but seed sources for this species remain unknown. Alfalfa was best among 7 legumes tested for rangeland seedling by Hewitt et ale (1982) who noted that it sustained a relatively low preference by insects. Varieties of alfalfa possessing falcata parentage characteristics such as spreading by root proliferation, broad crown development, dormancy during midsummer drought, and slow decumbent regrowth are recommended for semiarid rangelands in the northern Great Plains (Berdahl et ale 1989). Varieties ,with a sativa parentage, commonly used for hay, showed much lower survival. Spreador II, a falcata type, and ladak, a sativa type, were planted on the Tamarack Prairie but monitoring of comparative survival was not conducted. Townsend et ale (1975) noted that cicer milkvetch appeared to be adapted to the region and that sainfoin was able to grow only within sandy soils in northeastern Colorado. Cicer milkvetch, planted about 15 years ago at several ungrazed loam and sandy loam locations in northeastern Colorado, has persisted, sustained good annual growth, and continued to spread. This species establishes more slowly than alfalfa, grows more slowly in spring and summer, but is less subject to destruction by pocket gophers. The soaking and freezing technique previously described for Rocky Mountain beeplant should be used for establishing this hard-seeded specles. study findings in combination with results of previous data provide a general basis for recommendations, however, more testing is needed. Alfalfa should be the priority species in perennial legume plantings but should be planted in mixtures with other legumes and perennials. Cicer milkvetch and sainfoin should be included in lesser amounts. Townsend et ale (1975) noted that Illinois bundleflower, listed as a native of Colorado, did not survive the first winter after it was planted. Results were similar on the Tamarack Prairie indicating it should not be used in the future. Ladino clover, possessed a small growth form plus low survival and is also not recommended for use. Purple prairie clover is a native in eastern Colorado, however, if used, it should comprise only a small « 5) percentage of a seed mixture. Hairy vetch has been observed reestablishing itself in sandy sites in northeastern Colorado and along roadsides where planted but it gradually dies out if shallow tillage is not replicated annually to help it reseed. Yellow sweet clover also inconsistently reestablishes from seed. Shallow tillage in late winter may help sustain these species. Their use in seed mixtures is optional. If seed sources for native sweet pea can be found, small amounts of this species should be included in perennial mixtures. Testing is needed to determine if shallow « 7 cm) tillage in late winter would reduce competition of other vegetation with patches of native sweep pea. The same is true for western ragweed, a warm-season native that spreads by rhizomes and possesses moderately large seed. Western ragweed was reported to be of high value to bobwhite and scaled quail in the Rolling Plains of Texas and was increased by spring discing (Webb and Guthery 1983). Prairie coneflower (Rudbeckia occidentalis), prairie spiderwort (Tradescantia occidentalis), scarlet globemallow (Sphaeralcea coccinea) and other native perennials can be added to seed mixtures to increase diversity, but their establishment and value to wildlife is uncertain and they will rapidly increase seed costs if sources are available. Leadplant (Amorpha canescans) still occurs in remnant populations in ungrazed or lightly grazed rangeland in northeastern Colorado. It and sandcherry (Prunus pumila), another indigenous species subject to heavy browsing by livestock, are recommended for use in ungrazed sites primarily for their woody cover. Development of practical establishment techniques is needed as seed costs are high. Small burnet is not recommended for future planting in rangelands. Lewis blue flax has been seeded and has established in highway roadsides in sandy areas in northeastern Colorado. It is not native, apparently has little or no value to wildlife, and is not recommended in perennial seed mixtures. Dryland adapted mixtures of wild flowers are available for planting but most are not native, few will sustain themselves, and seed costs are high. �65 Much larger and wider tracts, at least 1 or 2 ha in size should be used when possible in planting perennial mixes. Portable solar-powered electric fences should be used to exclude deer until seedlings become well established. This study initiated what should be continued as long-term evaluations of species, varieties, and techniques to develop high quality foods for upland birds and other wildlife in eastern Colorado rangelands. Property management personnel have both the equipment and facilities, and are urged to continue these evaluations. Efforts need not be extensive or elaborate, but record keeping concerning what species will grow and are used by upland birds is needed. We are fortunate noxious weeds are not a concern when soils are disturbed within sandhill rangelands. This increases opportunities for habitat management in these areas. LITERATURE Amen, CITED A. E., D.L. Anderson, T. J. Hughes, and T. J. Weber. 1977. of Logan County, Colorado. U.s. Dep. Agric., Soil Conserve Washington. D.C. 252pp. Berdahl, S. D., A. C. Wilton, and A. B. Frank. 1989. performance of 25 alfalfa cultivars and strains rangeland. J. Range Manage. 42:312-316. Daubenmire, R.F. 1959. A canopy-coverage Northwest Sci. 33:43-64. method Greb, limited B. W. 1978. Millet production with Univ. Exp. Sta. Prog. Rep. 15. 3pp. Soil Survey Servo , Survival and agronomic interseeded into of vegetational water. analysis. Colorado state Hewitt, G. B., A. C; Wilton, and R. J. Lorenz. 1982. The suitability of legumes for rangeland interseeding and as grasshopper food plants. J. Range Manage. 35:653-656. Hungerford, K. E. 1948. Manage. 12:436-437. McGinnies, grown Safflower as a winter W. J., and C. E. Townsend. alone and in mixtures with game bird food. 1983. Yield of three range grasses legumes. J. Range Manage. 36:399-401. Snyder, W. D. 1967. Experimental habitat improvement for scaled Colorado Dep. Game, Fish and Parks. Tech. Bull. 19. 65pp. 1978. The bobwhite Publ. 32. 88pp. in eastern J. Wildl. Colorado. Colorado quail. Div. Wildl. Tech. 1993. Increasing foods for wildlife within the rangelands of eastern Colorado. Colorado. Div. Wildl., Wildl. Res. Rep., Fed. Aid Proj. W167-R. Apr.:101-118 Townsend, C. E., G. o. Hinze, W. D. Ackerman, and E. E. Remmenga. 1975. Evaluation of forage legumes for rangelands of the central Great Plains. Colorado State Univ. Exp. Sta. Gen. Series Rep. 942. 10pp. Webb, W. M., and F. S. Guthery. spring discing of mesquite Manage. 36:351-353. Prepared by WMMtJ ~ Warren D. Snyder 1983. Response of wildlife food plants rangeland in northwest Texas. J. Range to ��67 Colorado Division Wildlife Research April 1995 of Wildlife Report JOB PROGRESS State of: Colorado Project: W-167-R Work Plan: Upland Job Title: Period Author: Job 22 Covered: Upland Bird Research 1 Bird Research 01 January REPORT Publications through 31 December 1994 Clait E. Braun Personnel: Clait E. Braun, K. M. Giesen, R. W. Hoffman, W. D. Snyder, Colorado Division of Wildlife T. E. Remington, and ABSTRACT The following articles were published in 1994: Braun, C. E. 1994. Band-tailed pigeon. Pages 60-74 in T. C. Tacha and C. E. Braun, eds. Migratory shore and upland game bird management in North America. Int. Assoc. Fish and Wildl. Agencies, Washington, D.C. 1994. Colorado. Historic and present distribution and status J. Colorado-Wyoming Acad. Sci. 26:30. of sage grouse in ______ , K. Martin, T. E. Remington, and J. R. Young. 1994. North American grouse: issues and strategies for the 21st century. Trans. North Am. Wildl. and Nat. Resour. Conf. 59:428-438. ____ ~, K. M. Giesen, R. W. Hoffman, Upland bird management analysis Div. Rep •.19. 48pp. T. E. Remington, and W. D. Snyder. 1994. guide, 1994-1998. Colorado Div. Wildl., Giesen, K. M. 1994. Breeding range and population status chickens in Colorado. Prairie Nat. 26:175-182. 1994. Movements and nesting habitat in Colorado. Southwest. Nat. 39:96-98. of lesser of lesser prairie- prairie-chicken hens Joy, S. M., R. T. Reynolds, R. L. Knight, and R. W. Hoffman. 1994. Feeding ecology of sharp-skinned hawks in deciduous and coniferous forests in Colorado. Condor 96:455-467. Tacha, T. C., and C. E. Braun, editors. 1994. Migratory shore and upland game bird management in North America. Int. Assoc. Fish and Wildl. Agencies, Washington, D.C. 223pp. �68 _______, , and R. E. Tomlinson. 1994. Migratory shore and upland game bird resources - status and needs. Pages 219-213 in T. C. Tacha and C. E. Braun, eds. Migratory shore and upland game bird management in North America. Int. Assoc. Fish and Wildl. Agencies, Washington, D.C. Young, J. R., J. W. Hupp, J. W. Bradbury, and C. E. Braun. 1994. Phenotypic divergence of secondary sexual traits among sage grouse, Centrocercus urophasianus, populations. Anim. Behav. 47:1353-1362. Prepared .~.;__V._-__ 2 __ by __ --:---,-~.L.t:.... Clait E. Braun wildlife Research Leader _ �71 Colorado Division Wildlife Research April 1995 of Wildlife Report JOB PROGRESS state of: Colorado Project: W-167-R Work Plan: 27 Job Title: Period Author: Experimental Covered: Richard Personnel: REPORT Upland 1 Job Range 01 January Bird Research Expansion through of Ruffed 31 December Grouse in Colorado 1994 W. Hoffman Clait E. Braun, Richard W. Hoffman, Richard Olterman. Robin Olterman, Colorado Division Peckham, Mark Tucker, U. S. Forest Service. M. Lopez, Jim of Wildlife; Kathy A. ABSTRACT No ruffed grouse (Bonasa umbellus) were trapped and transplanted into Colorado during 1994. Efforts focused on conducting interagency and public meetings, gathering additional information, and preparing reports in accordance with the Memorandum of Understanding between the Colorado Division of Wildlife (CDOW) and U. S. Forest Service (USFS) regarding wildlife transplants and introductions. A revised schedule was prepared, with the first release planned for spring 1995. Site inspections of proposed release areas were conducted with personnel from the CDOW (Southwest Region) and USFS (San Juan National Forest). Stoner Mesa was identified as the preferred release area because of the diversity of aspen age classes and stand types occurring in association with coniferous and mountain shrub communities. Both Wyoming and Idaho have granted permission to trap and move birds to Colorado. Based on analysis of available information, it was concluded that the range expansion program would not adversely effect existing fauna and flora on the forest, conflict with present land uses, or preclude or limit other activities on the forest. Two meetings were held in September 1994 to provide information to the public about the program and to hear their concerns. The issues raised at these meetings are addressed in this report. �72 �73 EXPERIMENTAL RANGE EXPANSION Richard OF RUFFED GROUSE IN COLORADO W. Hoffman INTRODUCTION A small population of ruffed grouse (Bonasa umbellus) was first discovered in western Moffat County along the Colorado/Utah border in 1988. Prior to this discovery, it was believed that ruffed grouse did not occur in Colorado and that reported observations were of blue grouse (Dendragapus obscurus) or sharp-tailed grouse (Tympanuchus phasianellus). Other than observations and collections of birds from extreme northwestern Colorado, there is no evidence to suggest ruffed grouse occur elsewhere in the state. It also is unlikely this population will naturally expand into other suitable habitats because dispersal is effectively blocked by wide expanses of sagebrush (Artemisia spp.) and pinyon/juniper (Pinus edulis/Juniperus spp.) rangelands. Currently, the ruffed grouse is considered native to Colorado and is classified as a game species with full protective status (i.e., no open season). In July 1992, the Colorado Wildlife Commission passed a resolution supporting development of a program to expand the distribution of ruffed grouse in Colorado. P. N. OBJECTIVES To identify release sites, develop guidelines, conduct transplant, evaluate success or failure of ruffed grouse to establish a breeding population, refine guidelines, and prepare recommendations for possible future transplants. SEGMENT literature 1. Review 2. Evaluate 3. Send letter to U. S. Forest ruffed grouse. 4. Conduct joint evaluation of release area with U. S. Forest Colorado Division of Wildlife personnel. 5. Conduct 6. Locate 7. Trap and transplant up to 40 ruffed into southwest Colorado. 8. Radiomark at least 20 birds and monitor their movements, survival, habitat use, and reproductive performance. Conduct drumming survey. 9. Compile potential public release to the objectives of this study. sites. Service announcing intent to transplant Service and meetings. suitable data, pertinent OBJECTIVES trap sites analyze in nearby results, states. grouse and prepare from Wyoming, progress Idaho, or Utah report. STUDY AREA Personnel from the Colorado Division of Wildlife's (CDOW) Upland Bird Program conducted a reconnaissance of potential ruffed grouse habitats in western Colorado during September 1992. A proposal/analysis of the range expansion program was prepared by the CDOW following this investigation and submitted to the U. S. Forest Service (USFS) for consideration. This proposal identified the San Juan National Forest (SJNF) as the preferred release area. �74 Vegetative composition of the SJNF includes the following forested (72%) and non-forested (28%) types: spruce/fir (Picea/Abies) (28%), ponderosa pine (Pinus ponderosa) (18%), quaking aspen (Populus tremuloides) (16%), Douglasfir (Pseudotsuga menziesii)/mixed conifer (10%), pinyon/juniper «1%), meadows (9%), Gambel's oak (Quercus gambelii)/mountain shrub/sagebrush communities (7%), riparian (2%), and unvegetated areas (9%). The current condition of the SJNF reflects almost a century of human manipulation anq protection from natural disturbances. Timber harvesting has reduced the ponderosa pine type to less than 10% of its original distribution; conversely, logging has helped prevent the invasion of conifers into the aspen type and has created structural diversity within the aspen and spruce-fir types. However, extensive, homogeneous stands of mature to overmature aspen and spruce-fir still occur on the forest due to past control of fires. METHODS other than site inspections, no field work was conducted during 1994. Efforts focused on conducting interagency and public meetings and preparation of reports in accordance with Section C of the Supplemental Appendices to the Memorandum of Understanding between the Colorado Division of Wildlife and USFS (Region 2) regarding wildlife transplants and introductions. Trapping and transplanting, which was scheduled to begin during fall 1994, was postponed to allow time for additional public comment and to prepare ~. joint report outlining the expected effects of the range expansion program. A revised schedule was prepared, with the first release scheduled for spring 1995. This schedule along with a letter indicating the CDOW's intention to proceed with the release were sent to the USFS. RESULTS Site AND DISCUSSION Inspections Site inspections were conducted in June and August 1994, and included a meeting and tours of proposed release sites with personnel from the SJNF, CD OW (Southwest Region), and Ruffed Grouse Society (RGS). Specific sites inspected during these tours included Stoner Mesa (Deer Creek, Stoner Creek, and Sulphur Creek) and Haycamp Mesa (Lost Canyon Creek, Fish Creek, and Morgan Gulch). Both sites are within the Mancos/Dolores District of SJNF. Stoner Mesa, specifically T 39 N, R 11 W (sections 21,28) and R 12 W (sections 20,29) was identified as the preferred release area. The Mancos/Dolores District of the SJNF was identified as having high habitat suitability for ruffed grouse because of the diversity of aspen age classes and stand types that occur in association with coniferous and mountain shrub communities throughout the District. Approximately 121,720 ha of the SJNF is comprised of aspen. Over 20,000 ha of aspen have been harvested on the Forest since 1950. In the past 10 years, about 4,050 ha of aspen have been harvested. currently, about 400 ha are harvested annually. No other Forest in the state has harvested this much aspen. The end result has been the creation of a mosaic of aspen stands of varying ages and structural characteristics; conditions considered ideal for ruffed grouse. Expected Effects of Range Expansion Program Optimal habitats for ruffed grouse are expected to occur where 15-20 year old aspen stands are interspersed with mature (40+ years) stands, scattered clumps of conifers, smallorenings « 1 ha), and shrub thickets. Such areas should support 10+ males/km. Mesic sites are likely to be preferred because of the food and cover associated with these sites. Ruffed grouse also are expected to use mountain shrub/oak communities adjacent to the aspen type, heavily �75 vegetated riparian corridors, and Douglas-fir/lodgepole pine (~. contorta/~. lasiocarpa) communities with a dense understory of shrubs or conifer regeneration. Ruffed grouse will probably occur in a clumped distribution within these areas and densities are likely to be highly variable « 1 to >10 males/km2). Ruffed grouse may disperse through the ponderosa pine/oakbrush type, but are not expected to breed or winter in this type. Sagebrush and pinyon/juniper rangelands are unsuitable habitats for ruffed grouse; expanses greater than 10 km wide should effectively function as ecological barriers to dispersal. Principle winter and early spring foods should include flower buds and catkins of male aspen trees, with buds and twigs of serviceberry (Amelanchier spp.), hawthorn (Crataequs spp.), willow (Salix spp.), dogwood (Cornus stolonifera) , choke cherry (Prunus virqiniana), and Wood's rose (Rosa woodsii) serving as alternative food sources. Seeds, fruits, buds, leaves, and flower parts from forbs, grasses, and shrubs will constitute 90%+ of the summer and fall diets; about 10% of the summer diet will include insects and other small invertebrates. Insects will provide 90-95% of the chick's diet until about 1 month of age. By 2 months of age the chicks will be feeding almost exclusively on plant matter. Ruffed grouse are opportunistic feeders and tend to selectively feed on the most abundant foods available. For this reason, they are not likely to impact rare or endangered plants or insects. According to the SJNF Plan, there are no threatened or endangered plants on the Forest. Although ruffed grouse are not native to the SJNF, the species is native to the physiographic region that includes the SJNF. In Utah, Wyoming, and Idaho, ruffed grouse occur in similar habitats and coexist with the same animals that are found on the SJNF. There is no evidence to suggest ruffed grouse compete with other wildlife species, including blue grouse, in these habitats. There also is no evidence to suggest ruffed grouse will compete with any threatened, endangered, or sensitive wildlife species identified in the SJNF Plan. Of 9 T&E species listed in the plan, only the American peregrine falcon (Falco pereqrinus), bald eagle (Haliaeetus leucocephalus), and river otter (Lutra canadensis) actually exist on the Forest. The other 6 species, 4 of which are fish, are of questionable existence on the Forest. They include the grizzly bear (Ursus arctos), wolverine (Gulo qulo) , Colorado River cutthroat trout (Onchorynchus clarki pleuriticus), Colorado squawfish (Ptychocheilus lucius), humpback chub (Gila ~), and razorback sucker (Xyrauchen texanus). The introduction and establishment of ruffed grouse on the Forest will have no impact on aquatic environments and should not be an issue with respect to endangered fish species. The introduction of ruffed grouse should benefit the northern goshawk (ACcipiter qentilis), 1 of 3 sensitive species known to occur within the proposed release area, by providing additional prey. Ruffed grouse should have no impact on the purple martin (Proqne subis) or olive-sided flycatcher (Contopus borealis), the other 2 sensitive species, because they belong to different feeding guilds. The introduced population may eventually expand at the rate of 5+ km per year. The initial rate of expansion may be less as there should be an abundance of suitable habitat near the release sites. In general, ruffed grouse, especially males, are considered poor pioneers because of their short dispersal distances and sedentary nature. Males seldom disperse more than 3 km from their natal area and once established on a breeding area, most will remain within an area < 12 ha the remainder of their lives. Based on these attributes, one would expect a new population to expand slowly. However, the rate at which birds pioneer new habitats is also a function of their reproductive potential; i.e., the more birds produced each year, the greater the potential to expand into new areas. Most ruffed grouse hens attempt to nest, they lay on average 11 eggs per clutch, and they will renest if the first nest is destroyed. Thus, the species has a high reproductive potential that may partially compensate for its poor pioneering ability. �76 There are no ecological barriers to prevent the expansion of ruffed grouse into the Weminuche and Lizard Head Wilderness areas and onto the Rio Grande and Uncompahgre National Forests. This could occur within 3-10 years postrelease. Aspen stands within the Lizard Head Wilderness Area are not suitablebreeding habitat, but birds will disperse through these stands. Suitable breeding habitat does occur in the Weminuche Wilderness and on the Rio Grande and Uncompahgre National Forests. There is no ruffed grouse habitat in Mesa Verde National Park or on the Ute Mountain Ute and Southern Ute reservations. The vast expanses of sagebrush, ponderosa pine, and pinyon/juniper rangelands adjacent to these areas should function as ecological barriers to movements of ruffed grouse. Field investigations indicate suitable habitats for ruffed grouse are present on the SJNF. There is no evidence to suggest the range expansion program will adversely effect native wildlife and plant species, including T&E species. Furthermore, the program will not conflict with present land uses or ongoing programs on the SJNF, nor will it preclude or limit other management activities on the Forest by the USFS or CDOW; i.e., no closures nor special precautions will be necessary to protect ruffed grouse. Public Meetings Two public meetings were held in September 1994. One meeting was held in Durango and the other in Denver. Five people attended the Denver meeting and 20 attended the Durango Meeting. There were 5 major concerns raised at these meetings. 1. Those in favor of the program questioned why the CDOW was holding more public meetings when (1) the Colorado Wildlife Commission had already passed a resolution supporting the program, and (2) the USFS had already issued a letter to the CDOW stating that, although they had concerns about introducing a species that was not native to the SJNF, they could find no evidence that ruffed grouse would have any effects upon National Forest resources, programs or policies, and that consistent with the MOU, they found no need for the USFS to do any additional analysis. 2. Because of concerns about declining sage grouse (Centrocercus urophasianus) and sharp-tailed grouse populations, some individuals questioned whether implementing the ruffed grouse project was an appropriate use of Division resources. 3. Others expressed concern that the proposal was inconsistent with the principals of ecosystem management because no evidence was presented to indicate ruffed grouse historically occupied habitats in the SJNF. 4. Still others felt the USFS violated the National Environmental Policy Act by not preparing an environmental assessment before issuing a statement of no affect on National Forest programs, resources, or policies. 5. Some people contended that releasing ruffed grouse on the SJNF would be in violation of the Wilderness Act because the birds could potentially expand their range into the Weminuche and Lizard Head Wilderness areas. Trap sites Wyoming and Idaho have granted permission to trap and move birds to Colorado. The Portneuf Range, 20 km southeast of Pocatello, Idaho has been identified as the preferred trap site. Idaho Game and Fish personnel have trapped and moved birds from this site to other areas of the state. They also provided birds for release in Nevada. Their experience and knowledge of suitable trap sites will be valuable in successfully completing the trapping phase of this �77 project. Furthermore, southeastern Idaho is within the range of Bonasa umbellus incana, the subspecies of ruffed grouse that most likely occurs in northwest Colorado, and therefore, the subspecies best suited for release elsewhere in Colorado. Prepared � https://cpw.cvlcollections.org/files/original/213d58e4c80d489c22795e5225dddc43.pdf 2d9cf0940b8c468bffdd79313ad68e2b PDF Text Text Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS Colorado State of Project REPORT No. W-153-R-2 Mammals Research Work Plan No. Multispecies Job No. Consulting Services for Mark-Recapture Analysis Period Covered: Author: July Inyestigations 1, 1994 - June 30, 1995 G. C. White Personnel: R. M. Bartmann, R. B. Gill, T. D. I. Beck, D. J. Freddy ABSTRACT Progress towards the objectives of this job include: 1. A computer program (Program NOREMARK) developed to operate on personal computers for the computation of population estimates based on resightings of marked animals was accepted for publication by the Wildlife Society Bulletin. 2. Preliminary estimates of black bear numbers on the southwest study area were developed in cooperation with Tom Beck. 3. A study of compensatory effects of harvest on the Piceance Basin mule deer population was continued as part of Federal Aid Project W-153-R Work Plan 2 Job 15, entitled Compensatory Effects of Harvest in a Mule Deer Population. Experimental harvests have been conducted in December, 1989, 1990, and 1991. Radio collars to monitor over-winter survival of fawns wer,e.,placed on the animals dUI::ing,November, 1989, 1990, 1991, 1992, 1993, and 1994. 4. Preliminary cooperation 5. A draft of a paper providing estimates Dolores River has been prepared. analysis of elk sightability with David Freddy. models was performed of crayfish abundance Colorado in in the ��3 CONSULTING SERVICES FOR MARK-RECAPTURE ANALYSES G. C. White P. N. OBJECTIVES Evaluate density dependence in the Piceance SEGMENT 1. Basin mule deer populations. OBJECTIVES Evaluate methodology to detect density dependence in mule deer populations from data on the survival of mule deer fawns in the Piceance Basin. RESULTS AND DISCUSSION Introduction. Density dependence in a deer population has major implications for management because the concepts of maximum sustained yield, compensatory mortality, quality versus quantity assume density-dependent feedback in the population. Density dependence is generally defined as a negative relationship between population growth rate and population size (MCCullough 1990). As McCullough (1990) pointed out, growth rate is a misnomer in the sense that its value can be positive, zero, or negative; the latter resulting in a decreasing population. A positive growth rate is often associated with populations below carrying capacity, whereas a negative rate is more typical with populations above carrying capacity. A concept closely tied to density dependence is compensatory mortality, or a change in the rate of the remaining sources of mortality in response to a change in the rate of 1 mortality source. As Kautz (1990) stated, the only reasonable mechanism to explain compensatory mortality is density-dependent mortality. Objectives of this paper are 1) to present models of how density dependence can operate in a population's dynamics, 2) to review evidence in the literature of density dependence in deer populations, and 3) to discuss some potential problems in experimental design and statistical analysis procedures when investigating density dependence in a deer population. Acknowledgments. We thank the numerous personnel from the Colorado Division of Wildlife, Los Alamos National Laboratory, and Colorado State University, and numerous other volunteers that helped in conducting the 15 years of mule deer research in the Piceance Basin. Constructive comments provided by Tanya Shenk, Ron C. Kufeld, and Warren Snyder are greatly appreciated. Models Incorporating Population models Density Dependence can all be considered special cases of the model where Nt is population size at time t, Bt is number of new animals recruited to the population during the interval t to t+1 (births), Dt is number of deaths of animals alive at time t during the interval t to t+1, It is number of immigrants into the population during the interval t to t+1, and Et is number of emigrants leaving the population during the interval t to t+1. When the per capita rate of Bt' Dt, It' or Et changes with population size, then density-dependent population growth takes place (with population growth �4 meaning either increase their per capita rates, rewritten as B t.! Dt' It' and E t are replaced then the above equation can be or decrease). If i.e., bt BtlNt, = by When the per capita rates btl dtl itl and et are independent of population size, density-independent population dynamics result. That is these rates do not change regardless of population density, or they change randomly with no correlation to population density (see McCullough 1990:534-535 for definitions of density dependence, density independence, and inverse density dependence). = As a simplification of the above model, suppose rt bt - dt + it - et so that all changes in the population between time t and t+1 are modeled by 1 parameter. Here, rt corresponds to the population growth rate of MCCullough (1990). If r is modeled as a simple linear function of Nt as rt r 0 (1 K) , where the intercept is r 0 for Nt = 0 and the slope is rol K, the usual logistic growth equation results. When Nt = K, rt = 0 and growth ceases because the population has reached K carrying capacity (Macnab 1985) • = N/ Instead of modeling all density dependence in terms of rt' the relationship can be split between the 4 components of population change. For example, suppose a population is closed (~t = et = 0), all density dependence is in the mortality rate [dt = do (1 + N / K) ), and the per capita birth rate is constant across all population sizes (bt bo for all values of Nt). This simple model will also result in logistic growth because the per capita rate of change is still a linear function of population size. Thus, a densitydependent relationship between any 1 or combination of the 4 components results in density-dependent population growth. = Two extensions to this framework are often applied. First, multiple age classes are modeled with the requirement of per capita rates for each age class. However, if relationships between per capita rates and population size are assumed linear, growth equivalent to logistic growth still results. Second, rather than modeling per capita rates of change as linear functions, more complex functions can be substituted. An example is the Richard's curve (Fowler 1981) where per capita recruitment is with the exponent m changing the shape of the relationship from linear to either concave or convex (Fig. 1). For m = 10, density dependence is not invoked until the population approaches K. For m = 1, density dependence is invoked at a constant rate as the population grows. For m = 0.1, density dependence is invoked most strongly at low densities and relaxes as the population grows. Fowler (1981) argued that theory and empirical information support the conclusion that most density-dependent change occurs at high population levels (close to carrying capacity) for species with life history strategies typical of large mammals like deer (m > 1). The reverse is true for species with life history strategies typical of insects and some fishes (m < 1). McCullough (1990) also elaborated on this concept and suggested the �5 spatial scale of the population being measured and environmental heterogeneity affect the degree to which deer populations demonstrate density dependence near K carrying capacity. Evidence of Density Dependence in Deer populations The density dependent process has been shown or implied to operate within various aspects of deer biology, e.g., pregnancy rates, birth dates, etc. (Albon et al. 1983, Clutton-Brock et al. 1987). More important, however, are the cumulative and ultimate effects of these biological aspects reflected in fawn survival and recruitment. Therefore, these 2 parameters are considered in the following examples concerning density dependence in deer (including red deer and elk [Ceryus e1aphus) populations. Four studies incorporated 1 or more of the design requirements discussed by Kautz (1990): randomization, replication, and manipulation. Two other studies not meeting these requirements provided long-term data sets. Most studies designed to try and detect density dependence in deer populations have been conducted on small enclosed or well-defined free-ranging populations. The George Reserve in Michigan contained a white-tailed deer (Odocoi1eus yirginianus) population with a long and well-documented history (McCullough 1979). The population within the 4.64-km2 enclosure was manipulated through planned reductions to explore the relationship between density and recruitment. The size and age/sex compositions of the population during each year of study were reconstructed from age estimates for jaws collected from mortalities. Because nearly all mortalities of deer were from controlled shooting or could otherwise be accounted for, birth and death dates were available for virtually all deer ~6-months-01d. During the 19-year period, pre-hunt populations varied from 16.1-34.1 deer/km2 and post-hunt populations from 5.0-11.9 deer/km2. A significant negative relationship was found when recruitment rate (fawns 6 months old) was regressed on total post-hunt females (r2 = 0.501, P < 0.01). A similar regression replacing recruitment rate with total fawns at 6 months of age yielded a weaker (r2 = 0.317), but still significant (P < 0.05), relationship. Application of this regression equation supported a declining recruitment rate as post-hunt females increased to further corroborate a density-dependent relationship. As discussed below, the first regression has an increased Type I error rate because of the induced correlation caused by including total females on both sides of the equation. The second regression should be significant for both density-dependent and density-independent populations, because, even with density dependence, the number of young produced is a function of the number of adults present to produce them. Bartmann et al. (1992) demonstrated density dependence in the winter survival of mule deer (~ hemionus) fawns stocked in large fenced pastures on a pinyon (~ edu1is)-juniper (Juniperus osteosperma) winter range in northwest Colorado. During each of 3 winters, 3 pastures of 1.69, 1.01, and 0.69 km2 were each stocked with 50 radio-collared fawns and enough adults to achieve densities of 44, 89, and 133 deer/km2, respectively. These stocking rates simulated hunting removals of 67, 33, and 0%, respectively. An inverse relationship between fawn survival rates and density (P < 0.001) indicated a strong compensatory mortality process was operating in the population. In a companion study with free-ranging mule deer fawns on an approximate 50km2 winter range, predation accounted for 49-77% of radio-collared fawn mortality over 4 winters (Bartmann et al. 1992). During the next 3 winters, coyotes were intensively removed. Predation rates decreased (P = 0.004) and starvation rates increased (P = 0.042) compared to the pre-removal period while survival rates were unchanged (P = 0.842). This study provided evidence of compensatory mortality supporting results from the pastures. However, as �6 discussed below, they did not correct for the year-to-year variation. Reanalysis of this data suggests that they concluded a compensatory mortality effect existed because of this temporal variation. Predation rate was likely not reduced (P = 0.126), and starvation rate was not shown to increase (P = 0.552). Red deer on the 12-km2 north block of the Island of Rhum, Scotland (CluttonBropk et ale 1985) comprised a small free-ranging population that simulated confinement because of negligible immigration/emigration. Annual culling that occurred prior to the study was discontinued and the population of hinds increased from 57 to 166. Because all deer could be individually identified and the population was intensively monitored during the 13-year study, fairly rigorous testing for density-dependent effects was possible. As hind density increased, proportions of hinds calving as 3-year-olds and of milk hinds calving decreased, while winter calf survival increased (P < 0.01). Consequently, calf/hind ratios in spring also decreased (P < 0.01). Again, these authors did not correct for temporal variation in their analysis, so may have concluded that density dependence exists when it does not. However, year-to-year variation in the maritime environment of the Island of Rhum is probably small compared to the temporal variation observed in a continental climate. Hamlin and Mackie (1989) evaluated a 28-year data set for a non-migratory mule deer population on a 275-km2 portion of the Missouri River Breaks in Montana. Fall population estimates ranged from 565 to 1,720, or 2.1 to 6.3 deer/km2. Although hunting occurred, it was management-oriented rather than a planned perturbation. Integrity of the data set was maintained over the 28-year period, but data collection procedures varied. Per capita fawn recruitment to 1 year of age (fawns/female ~2 years old) was not significantly related to total adults the previous spring (r = -0.052, P > 0.05). A stronger, but still non-significant, relationship resulted when total adults were replaced with breeding-age females (r = -0.246, P > 0.05). The authors concluded that any density dependence in the recruitment process was obscured by extreme environmental variation. However, such variation could not be separated from sampling variation that, although not quantified, was probably quite large. The northern Yellowstone elk population has been extensively monitored for many years (Houston 1982). During all but the last 11 years in this report, the elk herd was subjected to varying levels of removals by shooting and trapping within the national park to curb population growth. Public hunting adjacent to the park occurred all years. Calf recruitment to 6-9 months (proportion of cows with calves at heel) was inversely related to population size the previous year (~ = 0.62, P < 0.001). This density-dependent relationship was mostly attributed to calf mortality rather than to reproductive changes. Although the author acknowledged the questionable quality of data collected some years, the large span of years (24) and wide range in estimates of population size (3,000-13,000) were probably the main reasons density dependence was still detectable. Testing for Density Dependence Experimental Design The need for adequate experimental design to detect density dependence operating in a species' population dynamics has been reviewed by Kautz (1990). As with inferences from any experiment, strength of the design directly affects the strength of inferences resulting from the experiment. We agree with Kautz (1990) that many tests of density dependence are flawed because design principles are ignored. �7 Probably the most commonly violated principle of experimental design is lack of replication. Most experiments to detect density dependence are too costly and time consuming to be able to replicate the experiment to allow inferences across an entire species. consequently, results of most experimental tests of density dependence cannot be extrapolated beyond the specific population being studied. None of the studies reported above had adequate replication to make their inferences widely applicable. Realistically, however, we cannot expect massive experiments to be replicated by the same investigators, so replication must occur by other investigators and reported in the literature. Also, we believe management by experimentation (Macnab 1983) and adaptive management (Walters 1986) provide state agencies the opportunity to test for density dependence by invoking different management strategies on different management units. Unfortunately, we perceive that most state agencies succumb to political pressure and invoke new management practices state-wide rather than by an experimental approach to actively test them. Statistical Tests Two general methods of testing for density dependence have been developed. For a time series of population sizes, tests of a relationship between change in population size over an interval and population size at the start of the interval are commonly used. The second approach is to regress independently measured population rates, such as birth, death, etc., rates, against population size. For both types of tests, the null hypothesis is density independence and the alternative hypothesis is density dependence. Failure to reject the null hypothesis, however, does not constitute evidence of density independence in these cases. Instead, the evidence may suggest the test lacked sample size or the experimental variance was too high to reject the null hypothesis. In cases where the null hypothesis of density independence is not rejected, the investigator should report the confidence interval on the parameter being tested. This confidence interval will include the parameter value that suggests density independence because the test failed to reject this hypothesis. However, a narrow confidence interval is evidence the true parameter value may not differ much from density independence. In contrast, a wide interval suggests the test lacked power to reject a false null hypothesis and little information is contained in the data relative to the alternative hypothesis of density dependence. A procedures for testing density dependence in a time series of population sizes was developed for the Hudson Bay trapping data on lynx and snowshoe hares by Bulmer (1975). Pollard et al. (1987) extended the procedure and Dennis and Taper (1994) developed it further. These procedures have not been particularly useful in deer research because the long time-series of population sizes needed to achieve reasonable power by these tests have not been available. Typically, >10 years of data are needed to achieve power >0.5 for populations fluctuating around K carrying capacity. Further, all procedures suffer increased Type I errors when population sizes are estimated and include 10gnorma11y distributed sampling error (T. Shenk, Colorado State Univ., Pers. Comm.). Considerable controversy has developed over the usefulness of these tests (Wolda and Dennis 1993, Holyoak and Lawton 1993, Hanski et al. 1993, and Wolda et al. 1994), so we will not consider them further. Procedures to test for a relationship between recruitment (including per capita birth or death rates) and population size have been extensively used to test for density dependence (Tanner 1966, McCullough 1979). The approach is to regress population growth rate against population size. If population growth is density independent, the expected slope of the regression is o. If density dependent growth is occurring, the slope and correlation of the regression should be negative. However, as first pointed out by Eberhardt (1970), population growth rate (recruitment) must be estimated independently �8 of population size. series of population When population sizes as growth rate is estimated from the time- , and i: t is regressed against Nt' a correlation is induced because Nt occurs on both sides of the regression. Eberhardt (1970) pointed out that correlations of about -0.7 are expected for sequences of random numbers when tested with the regression procedure used by Tanner (1966). The induced correlation problem also occurs when the number of new recruits (fawns, Ft) in the population is divided by Nt to obtain the per capita recruitment rate and then this rate is regressed against Nt. Nt is in the denominator of the dependent variable (Ft/Nt) and is the independent variable. Hence, this simple linear regression is likely to suggest density dependence more often than it should. The appropriate analysis is to regress fawns against Nt and N; without an intercept, i.e., r, = ~lNt + ~2N; and then test the null hypothesis of ~2 = O. If the test rejects then fawn recruitment has been shown density dependent. and ~2 < 0, If the estimate of reproductive rate is independent of the population size, then similar procedures can be used to assess density dependence in reproduction. The trap to avoid is a regression with the same variable on both sides of the equation, i.e. an induced correlation is created. A similar problem would occur if the number of animals dying in the population were divided by the population size and then regressed against population size. However, a more common procedure is to estimate the mortality rate in the population from radio tracking. A logistic regression procedure can then be used to regress the mortality rate (or equivalently, survival rate) against population size, i.e., where ntd and nt1 are the number of radio marked animals that died and lived, respectively. Note that the radio-marked sample and estimate of mortality rate are independent of population size so no induced correlation is present in this logistic regression. A significant positive value of ~1 indicates density-dependent mortality whereas, if the survival rate is regressed against Nt' density dependence is indicated with a negative value of ~1. Improving Power of Tests for Density Dependence Power of a statistical test is defined as the probability the test will reject the null hypothesis given that the null hypothesis is false (or conversely, that the alternative hypothesis is true). Three factors are usually under control of the experimenter and can be manipulated to increase power of statistical tests for density dependence: increasing the size of the densitydependent response, decreasing sampling variation, and removing environmental variance. �9 To maximize power of the tests of the slope of recruitment rate, birth rate, or mortality rate against population size, the population should be manipulated over a range of values. For example, if the population is at K, the slope of the regression is expected to be zero and failure to reject the null hypothesis that the coefficient is zero does not constitute evidence of density independence. To see if density dependence operates in the population, the population must be observed at densities less than K. The larger the difference between low and high densities where observations are made, the greater the probability of detecting density dependence. McCullough (1982) provided an excellent example of manipulating a population to increase power of the experiment when he reduced the George Reserve whitetailed deer population to about 10 animals and then allowed them to increase unhindered. Survival monitoring with small sample sizes of radio-tracked animals will result in low power because of a large sampling variance. For survival rates, the variance of the estimate is inversely proportional to sample size. Thus, for S = 0.4, a sample of size 10 has a variance of 0.4(1 - 0.4)/10 = 0.024 while a sample of size 100 has a variance of 0.0024. Bartmann et ale (1992) used 50 radio-marked mule deer fawns for each of 3 treatments in each of 3 years in their pasture study. Still, they had power <80% to detect a difference in survival of <0.1. To overcome the limitations of sample size, they attempted to maximize the treatment effect to increase the power of the experiment. Another factor that influences power because of increased sampling variation is precision of the estimate of population size. Most studies do not have a census of the population, so population size is estimated and includes sampling variation. The effect of sampling variation is to lower the power of the statistical test of density dependence. Another source of variation that will lower the power of regression tests is temporal and spatial (environmental) variation in population growth rates, i.e., the beta noise of MCCullough (1990). For example, we monitored mule deer fawn survival on 3 study areas and obtained estimates of 30 annual survival rates in the Piceance Basin in northwest Colorado (see Bartmann et ale 1992 for a description of study areas and methods). The mean survival rate was 0.357 (SE = 0.038). However, the year*area survival rates varied from S = 0.03 to S = 0.81. Using the technique suggested by Burnham et ale (1987) to estimate variation of the survival process separately from sampling variation caused by a finite sample of radio-marked animals, we find a process variance of 0.040 (95% CI 0.024 - 0.076). If the true survival rate for each year is drawn from a normal distribution with a mean 0.357 and a variance 0.040, then 95% of true survival rates would occur in the interval -0.035 to 0.749. The following example demonstrates this calculation. Suppose survival rates of 0.3, 0.4, and 0.5 are estimated from 50 radio-marked fawns during 3 winters. The estimated total variance for these 3 estimates is n - 1 = 0.01 However, this estimate includes both sampling and temporal variation. Sampling variation is a funct~on of flample ~ize and can be estimated for each estimate by the formula Var (S .) S. (1 - S.) / n . . For n. = 50 each year, ~ ~ ~.J ~ estimates of sampling variance are 0.0042, 0.004H, and 0.0050. Variation due to time, although unknown, is the variation in the true parameter across the 3 years. Variation in the process is not a function of sample size, but a = �10 result of year-to-year differences in the true survival rate because of weather or other random effects. To estimate this process variation, a numerical optimization is required (Burnham et al. 1987). For this example, the estimate of process variation is 0.0054. To overcome the loss of power because of high temporal (process) variation, a control area is needed to remove the effects of time. Thus, Bartmann et al. (1992) stocked deer in pastures at 3 different densities to test for density effects. Even though there were differences in survival across years, the pattern of density effects between pastures was consistent across years and allowed detecting a response in survival as a function of density. Example of Power for a Test of Density Dependence Although tests for density dependence based on regression analysis seem simple and easy to apply, little evidence of density dependence has been detected in deer populations. Here, we demonstrate that even strong density-dependent effects will not be detected without experimental designs that remove environmental variation. Typically, data used in regression analyses are over some time period without a spatial control to account for environmental variation. Suppose fawn survival in a deer population is monitored for 5 years with 50 radio-marked fawns each year. Density is then reduced and the population monitored for another 5 years, again with 50 radio-marked fawns each year. A test of density dependence would be whether the mean survival rate for the second 5-year period (low density) is greater than the first 5year period (high density). To evaluate the power of this approach, assume annual survival for the first 5 years is 0.357, a value based on survival for mule deer fawns in the Piceance Basin of northwest Colorado. Further assume the true survival for each year in the first 5 years is selected from a normal distribution with mean 0.357 and variance 0.040. Let mean annual survival in the second 5 years increase over the first 5 years by the amount 8. The larger the value of 8, the greater the mean annual survival and the stronger the density-dependent response. The null hypothesis is that mean survival for the first 5-year period equals mean survival for the second 5-year period which can be tested with logistic regression. The appropriate sampling unit for this study would be a year, so sample size is 10 years. However, most logistic regression procedures do not have methods to treat year as a random effect to allow constructing the proper test of the null hypothesis stated above. Typically, users might specify the model P.~ where P. is a dummy variable specifying the 5-year period effect, i.e., ~ period 1 or period 2. However, this analysis treats the sample size as the total number of radio-marked fawns, or 500. The appropriate model is logi t( n:, n" n,J = Pi + Yi (P;I where Y. (P .) is the year effect nested within the period effect. The ~ ~ appropriate test is then constructed as the ratio of the chi-squared values divided by their respective degrees of freedom to form an F ratio, i.e., �11 = X~1/1 X~8/8 Power of this test for the null hypothesis is shown in Fig. 2 for values of a from 0 to 0.6, and values of the process variation of survival from 0 to 0.04. Each estimate of power is based on 1,000 simulations. The power of the test to detect density dependence decreases as the process variation increases. Even with no process variation, the probability of detecting density dependence with an increase in survival of 0.1 is only 0.53. For a = 0.1, any process variation lowers the power substantially, down to 0.11 for a process variance of 0.04. The problem with using a simple chi-squared test is shown in Fig. 3, where the rejection rate (power) of the chi-squared test is shown. For a = 0, the rejection rate exceeds a = 0.05 for values of process variance >0. Particularly note that the Type I error (probability of rejecting a true null hypothesis) rate reaches 0.54 for a process variance of 0.04. Thus, if process variance exists during the study, the chi-square test will reject a true null hypothesis more often than it should. Investigators would conclude that density dependence exists when it does not. When density dependence does exist, i.e., a > 0, -this test suggests better power than the F test. However, the Type I error rate >0.05 invalidates this test. As this power calculation exercise demonstrates, even with a large number of radio-marked animals to estimate survival, a planned experiment that lacks a spatial control to remove environmental variation has little chance of detecting a reasonable (0.1 - 0.2) increase in survival. We doubt that increasing survival beyond 0.2 is biologically feasible. Therefore, we are not surprised density dependence is seldom detected by merely observing a population, i.e., no manipulation or temporal and spatial controls, when the correct statistical test is performed. Lack of a treatment to increase power of the statistical test, lack of spatial controls to remove temporal variation in the population process, and the usual lack of an exceptionally large sample size all preclude detecting density dependence even when a strong effect may be operating in the population. Conclusions Detection of density dependence in deer populations is complicated by the presence of environmental variation (process variation). Proper analysis methods correct for process variation, but none of the published tests for density dependence have used these methods. Other studies have concluded that density dependence exists in a population, but have likely induced a correlation by including the same variable on both sides of the regression equation, and hence may have incorrectly concluded that density dependence exists in a population. Finally, studies that conclude density dependence does not exist in a deer populations generally fail to reject the null hypothesis of density independence and then conclude that the null hypothesis is true. Given the difficulty of conducting large scale, long term studies to detect density dependence, it is not surprising that researchers are unable to provide conclusive evidence that density dependence commonly exists in deer populations. �12 Literature Cited Albon, S. D., B. Mitchell, and B. W. staines. 1983. Fertility and body weight in female red deer: a density dependent relationship. J. Anim. Ecol. 52:969-980. Bartmann, R. M., G. C. White, and L. H. Carpenter. 1992. Compensatory mortality in a Colorado mule deer population. Wildl. Monogr. 121. 39pp. Bulmer, M. G. 1975. The statistical analysis of density dependence. Biometrika 31:901-911. Burnham, K. P., D. R. Anderson, G. C. White, C. Brownie, and K. H. Pollock. 1987. Design and analysis methods for fish survival experiments based on release-recapture. Am. Fisheries Soc. Monogr. 5. 437pp. Clutton-Brock, T. H., M. Major, and F. E. Guinness. 1985. Population regulation in male and female red deer. J. Anim. Ecol. 54:831-846. ________, , S. D. Albon, and F. E. Guinness. 1987. Early development and population dynamics in red deer. I. Density-dependent effects on juvenile survival. J. Anim. Ecol. 56:53-67. Dennis, B., and M. Taper. 1994. Density dependence in time series observations of natural populations: detecting stability in stochastic systems. Ecol. Monogr. 64:205-224. Eberhardt, L. L. 1970. Correlation, regression, and density dependence. Ecology 51:306-310. Fowler, C. W. 1981. Density dependence as related to life history strategy. Ecology 62:602-610. Hamlin, K. L., and R. J. Mackie. 1989. Mule deer in the Missouri River Breaks, Montana: a study of population dynamics in a fluctuating environment. Montana Dep. Fish, Wildl., and Parks, Fed. Aid in Wildl. Restor. Final Rep., Proj. W-120-R-7-18. 401pp. Hanski, I., I. Woiwod, and J. Perry. 1993. Density dependence, population persistence, and largely futile arguments. Oecologia 95:595-598. Holyoak, M., and J. H. Lawton. 1993. Comment arising from a paper by Wolda and Dennis: using and interpreting the results of tests for density dependence. Oecologia 95:592-594. Houston, D. B. 1982. The northern Yellowstone elk: ecology and management. Macmillan Publ. Co., Inc., New York. 474pp. Kautz, J. E. 1990. Testing for compensatory responses to removals from wildlife populations. Trans. North Am. Wildl. and Nat. Res. Conf. 55:527533. Macnab, J. 1985. Carrying capacity and related slippery shibboleths. Wildl. Soc. Bull. 13:403-410. McCullough, D. R. 1979. The George Reserve deer herd: population ecology of a K-selected species. Univ. Michigan Press, Ann Arbor. 271pp. 1982. Population growth rate of the George Reserve deer herd. J. Wildl. Manage. 46:1079-1083. -1990. Detecting density dependence: filtering the baby from the bathwater. Trans. North Am. Wildl. and Nat. Res. Conf. 55:534-543. Pollard, E., K. H. Lakhani, and P. Rothery. 1987. The detection of density dependence from a series of annual censuses. Ecology 68:2046-2055. Tanner, J. T. 1966. Effects of population density on growth rates of animal populations. Ecology 47:733-745. Walters, C. J. 1986. Adaptive management of renewable resources. Macmillan Publ. Co., Inc., New York. 374pp. Wolda, H., and B. Dennis. 1993. Density dependence tests, are they? Oecologia 95:581-591. _______ , , and M. Taper. 1993. Density dependence tests, and largely futile comments: Answers to Holyoak and Lawton (1993) and Hanski, Woiwod and Perry (1993). Oecologia 98:229-234. �13 +J ~ 2 a +J .~ e 1.5 o Q) ~1 ItS +J ·~o. 5 ItS U ~ 0 o Q) AI 40 20 60 80 Population Size Figure 1. Formof density dependence as modeled by the Richards' curve from Fowler (1981). For m = 1, logistic growth results. For m > 1, density dependence is strongest close to K, and vice versa for m < 1. 10080 Of> 60 $.I Q) 3: 0 ~ 40 20 0 0.1 ____0 -e-- 0.2 0.3 0.4 Effect Size (f1) 0.01-e- 0.02-9- 0.03~ 0.5 0.6 0.04 Process Variation Figure 2. Power (probability of rejecting the null hypothesis) of F test for fawn survival for 7 effect sizes (f1) and 5 values of process variation. Each plotted value is based on 1,000 simulations. �14 , 80 , , , 60 , .............. f...I cv l 40 o Il< 20 - • -- ••• ---- •• ( ••.• ---.--- , dI> , I •••• y ••••••••••••• _--- ---···················,········------········· , •.••••••••••• , 1···········-----········1························ , , , , , , , , , , , , ..r····················--··1··········----··········1···-······ , , , , , I I . , I • , , , , • I , , , I I , I , , _.. I _ ..•. ··-···-----~-·······-··----·····-·--T----·--·--·-············r············----·-·-···,························r····_···········_--·_I I I , •• I , I • , I , I I I I , I , I I , o , .5 Effect ---0 --0.01 -£:3-0.02 b.6 Size ---'<7-0.03 ____"'_0.04 Process Variation Figure 3. Power (probability of rejecting the null hypothesis) of X2 test for fawn survival for 7 effect sizes (~) and 5 values of process variation. Each plotted value is based on 1,000 simulations. �15 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS state of Project REPORT Colorado No. Mammals W-153-R-8 Research Work Plan No. 1 Multispecies Job No. 7 Mammals Library Period Author: Covered: July Jacqueline Personnel: Investigations Publication, Services Editing, and 1, 1994 - June 30, 1995 A. Boss Jacqueline A. Boss and Nancy W. McEwen ABSTRACT During the segment the following were accomplished: * 10 publications were purchased at the request of Mammals Researcher personnel and placed into the Colorado Division of Wildlife Research Center Library collection. * 17 free reports and short publications from state or federal agencies from private sources were located, ordered, and obtained for use by mammals Research personnel. * 30 theses or books were obtained by Mammals Research personnel. * 641 individual articles were Mammals Research personnel. * 25 manuscripts by Mammals for <,publication •.. * 0 manuscripts were prepared for peer review. on Interlibrary located Research Loan or as gifts and delivered personnel and submitted on request were published by Mammals or for use for use by or accepted Research personnel ��17 MAMMALS PUBLICATION, EDITING Jacqueline AND LIBRARY SERVICES A. Boss P. N. OBJECTIVE To provide a centralized support program for manuscript editing services to facilitate publishing results of research conducted Federal Aid Project W-153-R. SEGMENT and library by staff of OBJECTIVE To provide a centralized support program for Mammals Research editing, library, and publishing services so that Mammals Research personnel can be most efficient in publishing results of their research. SUMMARY Publications Purchased Center Library with Mammals Caughley, G. 1994. Wildlife Scientific Publications. 1994. Island Research ecology 334pp. Fair, J. 1990. The great American 192pp. Grumbine, R. E., ed. Washington, D.C. OF SERVICES Funds and Placed and management. bear. Minocqua, Environmental policy Press. 415pp. Knight, R. L., and K. J. Gutzwiller, eds. 1995. : coexistence through management and research. Press. 372pp. Lee, K. N. 1993. Compass and gyroscope: for the environment. Washington, D.C. in the Research Boston WI Blackwell NorthWord Press. and biodiversity. Wildlife and recreationists Washington, D.C. : Island integrating science and politics : Island Press. 243pp. MacDonnell, L. J., and S. F. Bates, eds. 1993. Natural resources law : trends and directions. Washington, D.C. : Island Press. policy and 241pp. Moore, J. A. biology. of modern 1993. Science as a way of knowing: the foundations Cambridge, Mass. : Harvard University Press. 530pp. Western, D., R. M. Wright, and S. C. Strum, eds. perspectives in community-based conservation. Press. 581pp. 1994. Natural connections Washington, D.C. : Island Yaffee, S. L. 1994. The wisdom of the spotted owl : policy century. Washington, D.C. : Island Press. 430pp. lessons Zaslowsky, D., and T. H. Watkins. 1994. These American lands: wilderness, and the public lands. Washington, D.C. : Island 398pp. for a new parks, Press. In addition to books purchased with Federal Aid Funds, about 17 free reports and short publications from state or federal agencies or from private sources were located, ordered, and obtained for use by Mammals Research personnel. �18 Theses and Books Obtained Researchers on Interlibrary Loan or as Gifts for Use by Anderson, R. M., and R. M. May. 1982. Population biology of infectious diseases : report of the Dahlem Workshop on Population Biology of Infectious Disease Agents : Berlin 1982, March 14-19. New York: Springer-Verlag. 314pp. Au Yeung, W. M. 1993. An assessment of trade in bear parts between North America and Hong Kong. M.S. Thesis, University of Manitoba, Winnipeg, Manitoba, Canada. 104pp. Bahre, C. J. 1991. A legacy of change in the Arizona borderlands. Tucson historic human impact on vegetation University of Arizona Press. 231pp. Berwick, S. H. 1968. Observations on the decline of the Rock Creek, population of bighorn sheep. M.S. Thesis, University of Montana, Missoula, MT. 245pp. Montana, Comly, L. M. 1993. Survival, reproduction, and movements of translocated nuisance black bears in Virginia. M.S. Thesis, Virginia Polytechnic Institute and State University, Blacksburg, VA. 113pp. Cartmill, M. 1993. through history. Covell, D. F. Colorado. 111pp. A view to death in the morning : hunting and nature Cambridge, MS : Harvard University Press. 331pp. 1992. Ecology of the swift fox (VULPES VELOX) in southeastern M.S. Thesis, University of Wisconsin - Madison, Madison, WI. Dietz, D. R. 1967. Chemical composition and digestibility by mule deer of selected forage species, Cache la Poudre Range, Colorado. Ph.D. Dissertation, Colorado State University, Fort Collins, co. 162pp. Evernden, N. University The natural alien of Toronto Press. humankind 160pp. and environment. Ginsberg, J. R., and D. W. Macdonald. 1990. Foxes, : an action plan for the conservation of canids. IUNC. 116pp. Glenn, E. M. 1993. Use of wetlands Thesis, University of Vermont,. by black 96pp. bears Toronto wolves, jackals, and dogs Gland, Switzerland: in southern Vermont. M.S. Hanson, P. P. 1986. Environmental ethics: philosophical and policy perspectives. Burnaby, B.C. : Simon Fraser University. 199pp. Herscovici, A. Enterprises 1985. Second nature: : Toronto. 254pp. the animal-rights Hines, T. D. 1980. An ecological study of Vulpes velox Thesis, University of Nebraska, Lincoln, NE. 103pp. controversy. CBC in Nebraska. M.S. Horejsi, B. L. 1976. Suckling and feeding behavior in relation to lamb survival in bighorn sheep (Ovis canadensis Shaw). Ph.D. Dissertation, University of Calgary, Calgary, Alberta. 265pp. International Air Transport Montreal : International Kingsland, S. E. 1985. population ecology. Association. 1993. Live animals Air Transport Assoc. 252pp. regulations. Modeling nature : episodes in the history of Chicago : University of Chicago Press. 267pp. �19 Kucera, T. E. 1988. Ecology and population dynamics of mule deer in the eastern Sierra Nevada, California. Ph.D Dissertation, University of California-Berkeley, Berkeley, CA. 207pp. Lindberg, M. 1986. Swift fox distribution in Wyoming : a biogeographical study. M.A. Thesis, University of Wyoming, Laramie, WY. 64pp. Logan, K. A. 1983. Mountain lion population and habitat characteristics in the Big Horn Mountains of Wyoming. M.S. Thesis, University of Wyoming, Laramie, WY. 101pp. MacIntyre, A. A. 1982. The politics of nonincremental domestic change: major reform in federal pesticide and predator control policy. Ph.D Dissertation, University of california-Davis, Davis, CA. 876pp. (microfiche) Mayr, E. 1969. McGraw-Hill. Principles 428pp. of systematic Mayr, E., and P. D. Ashlock. 1991. edition. New York : MCGraw-Hill, zoology. 1st edition. principles of systematic Inc. 475pp. New York zoology. 2nd Rollin, B. E. 1989. The unheeded cry: animal consciousness, science. New York : Oxford University Press. 308pp. animal Shepard, P. 1973. The tender carnivore Charles Scribner's Sons. 302pp. New York: and the sacred game. pain and Shorrocks, B., ed. 1984. Evolutionary ecology: the 23rd symposium of the British Ecological Society: Leeds 1982. Boston: Blackwell Scientific Publications. 418pp. Sweanor, L. L. environment. 1990. Mountain lion social organization in a desert M.S. Thesis, University of Idaho, Moscow, 10. 172pp. Tresner, C. 1980. History of Larimer county, Colorado : by Ansel 1911. Fort Collins, CO : Fort Collins Public Library. 74pp. Watrous U.S. Fish and Wildlife Service. 1987. Florida panther (Felis concolor coryi) recovery plan. Prepared by the Florida Panther Interagency Committee for the u.S. Fish and Wildlife Service, Atlanta, Georgia. 75pp. Wishart, W. D. 1958. Thesis, University Reference Document The bighorn sheep of the Sheep of Alberta, Edmonton, Alberta. Location River Valley. 66pp. and Delivery The Research Center Library staff also located and delivered individual articles or free documents on request for Mammals during this segment. Manuscripts Job Progress Published Reports; M.A. approximately Researchers 641 FY 1994-95 Federal Aid. All studies. Baker, D. L., M. W. Miller, and T. M. Nett. 1995. Gonadotropin-releasing hormone analog-induced patterns of luteinizing hormone secretion in female wapiti (Cervus elaphus nelsoni) during the breeding season, anestrus, and pregnancy. Biology of Reproduction 52:1193-1997. Beck, T. D. I., and R~ B. Gill. 1995. Colorado Western Black Bear Workshop 5:119-131. status report. Proc. �20 Beck, T. D. I., D. S. Moody, D. B. Koch, J. J. Beecham, G. R. Olson, and T. Burton. 1995. Sociological and ethical considerations of black bear hunting. Proc. Western Black Bear Workshop 5:119-131. Bowden, D. C., and R. C. Kufeld. 1995. Generalized mark-sight population size estimation applied to Colorado moose. J. Wildl. Manage. 59:(in press). Carpenter, L. H. 1995. Integrating research and management - how do we develop reliable knowledge? Proc., Western States and Provinces Joint Deer and Elk Workshop. (abstract). Freddy, D. J., G. C. White, and D. C. Bowden. 1995. Survival and sightability of adult and calf elk in Colorado: a progress report. Western states and Provinces Joint Deer and Elk Workshop. p.139 (abstract). Proc., Gerhardt, T. D., J. K. Detling, and N. T. Hobbs. 1994. Interactive effects of mowing, fire and primary production on patch selection by large herbivores. Bull. Ecol. Soc. Am. (Suppl.) 75(2):75 (abstract). Gill, R. B., and M. W. Miller. 1995. Thunder in the distance: the emerging debate over wildlife contraception. Proceedings of the Symposium on Contraception in Wildlife Management. (in press). Hobbs, N. T., D. L. Baker, G. D. Bear, and D. C. Bowden. grazing in sagebrush steppe I: mechanisms of resource Ecolog. Appl. (in press). 1995. Ungulate competition. Hobbs, N. T., D. L. Baker, G. D. Bear, and D. C. Bowden. 1995. Ungulate grazing in sagebrush steppe II: effects of resource competition on secondary production. Ecolog. Appl. (in press). Hobbs, N. T., and J. E. Gross. 1994. Predicting impacts of landscape change on biotic diversity : dynamic multispecies models. Bull. Ecol. Soc. Am. (Suppl.) 75(2):95 (abstract). Kufeld, R. C. Antler point restrictions. 1994. In: Deer. eds. D. Gerlach, S. Atwater, and J. Schnell. Mechanicsburg, Penn: Stackpole Books p.341. Kufeld, R. C. The human hunter. 1994. In: Deer. eds. D. Gerlach, Atwater, and J. Schnell. Stackpole Books: Mechanicsburg, Penn. 340pp. Kufeld, R. C. [1995J. Neck circumference calves during winter. Alces 30:63-64. Kufeld, R. C. 30:41-44. [1995J. Status of shiras moose and management of moose S. 337- (Alces a. shirasi) in Colorado. Alces Kufeld, R. C., and D. C. Bowden. 1995. Mule deer and white-tailed deer inhabiting eastern Colorado plains river bottoms. Colo. Div. of Wildl. Technical Publication No. 41. 58pp. Miller, J. R., T. T. Schulz, N. T. Hobbs, K. R. Wilson, D. L. Schrupp, and W. L. Baker. 1995. Changes in the landscape structure of a southeastern Wyoming riparian zone following shifts in stream dynamics. Biological conservation 75(1995):371-379. Miller, M. W., M. A. Wild, and W. R. Lance. naltrexone hydrochloride in antagonizing immobilization in captive Rocky Mountain Wildl. Dis. (in press) 1995. Efficacy and safety of carfentanil citrate elk (Cervus elaphus nelsoni). J. �21 Olterman, J. H., D. W. Kenvin, and R. C. Kufeld. southwestern Colorado. Alces 30:1-8. [1995]. Moose transplant Pojar, T. M., D. C. Bowden, and R. B. Gill. 1995. Aerial counting experiments to estimate pronghorn density and herd structure. J. Manage. 59(1):117-128. to Wildl. Popel, A. S., P. C. Johnson, M. V. Kameneva, and M. A. Wild. 1994. Capacity for red blood cell aggregation is higher in athletic mammalian species than in sedentary species. J. of Applied Physiology 77(4):1790-1794. Reed, D. F., and K. A. Green. [1995]. Mountain goats on Mount Evans, Colorado - conflicts and the importance of accurate population estimates. Bien. Symp. North. Wild Sheep and Goat Council. 9: (in press) Reed, D. F., J. Vayhinger, S. R. Ogilvie, E. B. Brekke, and T. P. Huber. 1994. Mountain sheep habitat use in the Arkansas River Canyon, Colorado. Fort Collins, Colo. : Colo. Div. of Wildl. 38pp. Wagner, F. H., R. Foresta, R. B. Gill, D. R. McCullough, M. R. Pelton, W. F. Porter, and H. Salwasser. 1995. Wildlife policies in the u.S. National Parks. Washington, D.C. : Island Press. 242pp. White, G. C., and R. M. Bartmann. 1995. Detecting density dependence in mule deer populations. Proc., Western States and Provinces Joint Deer and Elk Workshop. p. 110 (abstract). Manuscripts in Review FY 1994-95 At the end of FY 1994-95 all manuscripts were either 'in preparation' press.' None were in the stage of 'in review' by periodicals. Prepared by Jacqueline Librarian A. Boss or 'in ��23 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS State of Project REPORT Colorado No. Mammals W-153-R-8 Research Work Plan No. 1 Multispecies Job No. 9 Mammals Period Covered: Author: Personnel: R. Bruce July Investigations 1 Research Administration 1, 1994 - June 30, 1995 Gill R. Bruce Gill and Diane K. Hall ABSTRACT Human and fiscal resources were allocated among 7 Mammals 1 research jobs. Highlights of work progress on each study are summarized. All research objectives were accomplished within the resources allocated to each job. Thirteen scientific manuscripts publication during the segment. or abstracts were published or accepted for ��25 MAMMALS JOB PROGRESS REPORT 1 RESEARCH ADMINISTRATION R. Bruce Gill P. N. OBJECTIVE Administer research studies within productivity at the lowest cost. the Mammals SEGMENT 1. Supervise and administer Research Section. research 1 Research Unit for the highest OBJECTIVES on deer, elk, and moose in the Mammals INTRODUCTION Seven projects were active during the segment, funded with Pittmann-Robertson grants and 1 of originally allocated to the Watchable Wildlife objectives were completed successfully for all include: 6 of which were partially which was funded from money Project budget. Segment 7 projects. Highlights • Acquisition of 10 publications requested by Mammals Researchers and included in the Research Library collection; acquisition of 17 free reports and short publications for use by research staff; requests for 30 graduate theses which either were included in the Research Library collection or were obtained on loan for temporary use by research staff; acquisition of 641 published scientific articles for use by research staff. • Publication of 13 scientific articles in professional journals or in-house technical publications including 4 dealing with wapiti, 4 with moose, and 3 with deer. • Heart rate telemetry equipment was purchased and successfully implanted first into domestic goats (to assure that neither the surgical procedures nor the equipment was inimical to health or behavior of the recipient) and second into captive bighorn sheep. Equipment was evaluated for signal strength and clarity and for longevity. If equipment tests are successful, hear-rate telemetry will be tested experimentally as way of measuring stress in bighorn sheep ultimately to evaluate and mitigate stresses imposed by wildlife viewers. • Computer programs were developed to generate preliminary mark-resight estimates for a black bear population occupying a 465-km2 study area on the Uncompahgre Plateau in southwestern Colorado. Preliminary tests of computer models estimating elk sightability functions were tested successfully with real life data. • A draft study plan was developed for a management experiment to test the hypothesis that early-season hunting disturbance of elk results in movement of elk from public to private land. • Preliminary tests indicate that helicopter counts of elk groups detect average of 82% of the groups present. However, differences between observers were significant. The best sightability model out of 2,048 candidate models was a complex 12 parameter model. an �26 • Results of the North Park moose study have been published in 4 separate scientific articles. Data describing movements, home range size, and habitat use are being collected continuously and will be analyzed at the completion of the study. • Experiments with conjugates of gonadotropin releasing hormone (GnRH) and phytotoxins to assist in controlling population growth of mule deer at the Rocky Mountain Arsenal were negative because the entire conjugate was not successfully transported across the pituitary cell walls. Preliminary tests of remote delivery of contraceptives via biobullet injection were successful suggesting that once an efficacious fertility control agent is developed, it should be possible to remotely deliver effective doses in a biobullet fired from an air powered rifle. �27 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS state of Project Colorado No. W-153-R-8 Work Plan No. Job No. Period REPORT Mammals 2 Deer 15 Covered: Author: Personnel: Research Investigations Compensatory Effects of Harvest in a Mule Deer Population July 1, 1994 - June 30, 1995. R. M. Bartmann, G. C. White. J. Frothingham, T. Lytle, D. G. Saltz, C. L. Vardaman, E. White. ABSTRACT Aerial line transects were flown on the Ridge study area 4-6 January 1995. Estimated deer densities on the control and treatment units were nearly the same (23.1 and 24.3 deer/km2, respectively). Neither fawn nor doe survival rates over the past few years provide a rational explanation for the declining density on the control unit. Helicopter netgunning was used to capture 140 fawns between 19 November and 1 December 1994. Another 26 were captured with dropnets from 15-22 November. Estimated fawn survival on the treatment unit (0.825, SE 0.042) was not significantly higher (~ = 0.079) than on the control unit (0.701, SE 0.051). The extremely mild winter may have tempered any effects of deer density on fawn survival. As of 30 June 1995, 6 does died on the control unit and 1 on the treatment unit for preliminary survival rate estimates of 0.869 (SE 0.050) and 0.977 (SE 0.022), respectively. One fawn each from the control and treatment units died off the Ridge study area. Essentially all remaining fawns and does were on the units where captured during most of the winter. There were no significant differences in weight, left hind foot length, or weight:length ratio of fawns from the 2 units (~~ 0.247), but control fawns were 2.0 cm longer than treatment fawns (~ = 0.023). There were also no differences in the same body size indicators for adult does from the 2 units (~~ 0.127). ��29 COMPENSATORY EFFECTS OF HARVEST IN A MULE DEER POPULATION Richard M. Bartmann and Gary C. White P. N. OBJECTIVES 1. Increase the winter survival rate of mule deer fawns by lowering total deer density to reduce competition for forage during winter. 2. Increase the harvest rate of deer through increased productivity of adult does and decreased natural mortality of fawns resulting from closer alignment of population size with carrying capacity. SEGMENT OBJECTIVES 1. Maintain the winter population of mule deer on the Ridge treatment a density <40/km2 for 5 years. 2. Estimate winter 3. Estimate units. annual survival rates of adult females on control and treatment 5. Estimate condition of fawns on control and treatment 6. Estimate condition of adult females on control and treatment survival rates of fawns on control and treatment unit at units. units. units. METHODS Except for deer trapping, methods remained the same as previously reported by Bartmann (1990) and Bartmann and White (1991) with modifications by Bartmann and White (1992). Most deer were captured with helicopter net guns by Helicopter Wildlife Management, Inc., and the rest captured with dropnets. RESULTS AND DISCUSSION Maintain Population Line transects were flown on the Ridge study area 4-6 January 1995. Estimated deer density on the treatment unit has remained at a low level (24.3/kmt) while density on the control unit has unexplainably continued to decline (23.1/km2) (Fig. 1). Although far from significant, the density estimate for the control unit was marginally lower than on the treatment unit which was contrary to expectations. Estimates of adult and fawn survival do not provide a rational explanation for declining density estimates on the control unit. We originally suspected the change in capture method to helicopter netgunning may have had some affect on subsequent deer response to helicopters. However, we would have expected the affect to be similar, i.e., proportional, on the 2 units rather than the greater negative impact realized on the control unit. Therefore, we conclude some other problem exists with the line transect methodology and a new approach to density estimation is needed. Fawn Survival In 1994, 26 fawns were captured with dropnets and 140 by helicopter netgunning. Dropnetting occurred 15-22 November to capture deer in locations �30 not conducive to netgunning. Netgunning was done 19 November-1 December with effort alternated daily between control and treatment units. A hand-held Global Positioning System unit was used in the helicopter to get capture locations for each deer. We radiocollared 84 fawns on the control unit and 82 on the treatment unit. One fawn death on each unit was considered capture-related. In addition, 1 fawn from each unit died off the study area and was deleted from survival analyses. -+- CONTROL ... ~.. mEATMENT 1~r---~----------------------~ YEAR Fig. 1. Deer density estimates (w/95% confidence intervals) from aerial line transects on control and treatment units during early to mid-winter. The 1994-95 winter was even milder than in 1993-94 and fawn survival on the treatment unit (0.825, SE 0.042) was the highest ever recorded (Table 1). On the control unit, the survival rate of 0.701 (SE 0.051) was surpassed only during the 1989-90 winter. Unlike in 1993-94, fawn survival rates between the 2 units did not differ (~= 0.079). The winter may have been sufficiently mild to negate any effects of reduced deer density on the treatment unit. " ) for radio-collared mule estimates of survival rates (.§. Table 1. Kaplan-Meier deer fawns on control and treatment units of the Ridge study area in Piceance Basin, Colorado, from time of collaring in November and December until the following 15 June 1982-83 through 1994-95. Hunting mortalities are censored. Winter 1982-83 1983-84 1984-85 1985-86 1986-87 1987-88 1988-89 1989-90 1990-91 1991-92 1992-93 1993-94 1994-95 n 28 28 34 59 60 32 34 38 34 28 80 80 82 Control unit SE(.§.) .§. 0.321 0.071 0.196 0.537 0.431 0.241 0.270 0.779 0.320 0.456 0.112 0.550 0.701 0.088 0.049 0.078 0.070 0.064 0.077 0.083 0.078 0.090 0.107 0.036 0.056 0.051 n Treatment .§. 31 32 26 58 58 28 28 44 36 42 74 80 80 0.387 0.033 0.431 0.439 0.471 0.107 0.445 0.745 0.339 0.548 0.148 0.767 0.825 unit SE(.§.) 0.087 0.033 0.105 0.070 0.067 0.058 0.096 0.070 0.106 0.098 0.043 0.048 0.042 P of eqUal .§. (:t.) 0.578 0.774 0.075 0.157 0.565 0.006 0.509 0.659 0.909 0.481 0.102 0.006 0.079 Predation continued to be the leading mortality cause for the few fawns that died during 1994-95 (Table 2). In addition, all fawn deaths in the "other" category were suspected predation losses. starvation was attributed as the cause of death for only 1 fawn on the treatment unit. Location checks during mid-month in January, February, and March again indicated most deer, except for the 2 that died off the study area, remained on the units where they were captured. As in previous years, the few that were off their units usually had been captured near the boundary and were close to it during subsequent locations. �31 Adult Doe Survival The difference in adult doe survival on the 2 units approached significance (~ = 0.058) for the first time during the study (Table 3). Six does died on the control unit and 1 died on the treatment unit for preliminary survival rate estimates of 0.869 (SE 0.050) and 0.977 (SE 0.022), respectively. Table 2. Cause of mortality for radio-collared mule deer fawns on control and treatment units of the Ridge study area in Piceance Basin, Colorado, from time of collaring in November and December until the following 15 June 1982-83 through 1994-95. percentages are of total uncensored8 fawns. Winter n 1982-83 1983-84 1984-85 1985-86 1986-87 1987-88 1988-89 1989-90 1990-91 1991-92 1992-93 1993-94 1994-95 29 28 34 59 60 32 34 38 34 28 80 80 82 1982-83 1983-84 1984-85 1985-86 1986-87 1987-88 1988-89 1989-90 1990-91 1991-92 1992-93 1993-94 1994-95 31 32 26 58 58 28 28 44 36 42 74 80 80 Censored Hunting other 1 7 11 6 2 5 14 9 7 1 4 5 2 1 1 1 4 9 5 5 9 9 6 3 9 7 9 1 10 3 Starvation No. % Control unit 15 54 22 79 16 59 21 10 14 26 22 73 10 34 3 14 6 24 7 33 15 12 3 4 Treatment 15 27 8 17 16 19 9 6 8 7 23 3 1 Mortality: cause Predation No. % 4 4 5 7 17 14 14 19 15 31 5 17 10 3 50 22 15 40 14 64 29 19 2 3 2 11 13 1 1 7 10 9 22 25 4 4 3 2 26 8 9 15 8 39 11 11 other No. % 1 7 3 2 7 3 3 3 7 11 9 4 15 6 7 24 14 12 14 9 14 12 2 7 3 1 1 5 5 4 4 4 7 8 4 14 2 2 18 20 13 20 17 10 11 5 unit 50 87 36 35 30 68 36 20 40 29 34 4 1 8 Uncensored fawns are those that were not killed by hunters, that had nonfailing radios, or that had collars that did not drop off prematurely. Condition of Does Only 11 adult does were captured on the control unit and 10 on the treatment· unit. These small sample sizes together with high variability resulted in no significant differences for any of the 4 condition indices (~~ 0.127) (Table 5). Two yearling does were captured on the control unit and only 1 on the treatment unit. �32 A Table 3. Kaplan-Meier estimates of annual (1 Dec-30 Nov) survival rates (~) for radio-collared adult female mule deer on control and treatment units of the Ridge study area in Piceance Basin, Colorado, 1982-83 through 1994-95. Hunting mortalities are censored. n Winter 1982-83 1983-84 1984-85 1985-86 1986-87 1987-88 1988-89 1989-90 1990-91 1991-92 1992-93 1993-94 1994-958 8 10 15 9 25 27 14 7 23 39 41 46 49 47 Survival Condition Control unit Hunting ~ 1 3 2 1 0.800 0.779 1.000 0.917 0.756 0.818 0.857 1.000 0.969 0.758 0.716 0.939 0.869 rate estimates SE(~) a 0.126 0.113 11 15 10 21 18 10 5 28 41 42 52 46 44 0.056 0.087 0.116 0.132 0.031 0.071 0.067 0.034 0.050 for 1994-95 Treatment Hunting ~ 1 9 12 6 7 0.909 0.929 1.000 0.900 0.878 1.000 0.800 1.000 0.906 0.806 0.719 0.935 0.977 are only through unit SE(~) P of eqUal § 0.087 0.069 0.067 0.081 0.179 0.065 0.072 0.066 0.036 0.022 30 June 0.448 0.271 1.000 0.821 0.432 0.329 0.854 1.000 0.474 0.641 0.910 0.933 0.058 1995. of Fawns Weight and left hind foot length of fawns did not differ between the control and treatment units (~~ 0.909) (Table 4). But the total body length of fawns on the control unit averaged 2.0 cm longer than on the treatment unit (~ = 0.023). Weight/length ratios (treatment 0.253, SD 0.021; control 0.258, SD 0.022) also did not differ between units (~ = 0.247). LITERATURE CITED Bartmann, R. M. 1990. Compensatory effects population. colo. Div. Wildl., Wildl. of harvest Res. Rep. _____ , and G. C. White. 1991. Compensatory population. Colo. Div. Wildl., Wildl. effects of harvest in a mule Res. Rep. July:27-40. _____ , and G. C. White. 1992. Compensatory population. colo. Div. Wildl., Wildl. effects of harvest in a mule deer Res. Rep. July:27-37. Prepared by Richard M. Bartmann LSSR III Dr. Gary C. White Professor in a mule deer July:187-196. deer �33 Table 4. Weights (kg) and body measurements (cm) of mule deer fawns trapped on control and treatment units of the Ridge study area in Piceance Basin, Colorado, 1982-94. Weight so Year n 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 28 28 34 60 60 33 34 40 35 28 82 85 83 34.6 31.7 32.2 32.6 31.9a 29.9 29.5 32.7 30.8 30.7 30.4 30.7 33.6 Control 3.10 4.40 4.65 4.02 3.89 3.60 3.10 3.31 4.29 3.58 3.80 3.91 3.59 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 31 32 26 60 61 28 30 47 36 43 83 80 82 32.8c 32.3 32.3 32.3 31. 7 30.2 28.8 30.6 30.7 32.6 30.3 32.9 33.9 Treatment 4.18 3.12 5.07 4.62 4.13 5.34 4.13 3.33 4.42 3.54 4.68 3.96 3.90 a b nn == c !! 58 27 30 R Total body length so R Left hind foot length so R unit 124.0 124.2 123.9 124.4 128.1 127.3 123.8 131.0 126.5 128.2 126.8 129.1 132.9 4.64 5.65 7.25 6.26 6.53 6.12 7.83 5.81 9.02 5.47 6.67 6.17 7.13 41.1 40.6 40.8 41.1 41.0 40.8 41.0 41.8 40.8 40.6b 40.6 40.5 41.7 1.08 1. 73 1.53 1.48 1.95 1. 72 1.37 2.16 1. 75 1.32 1.58 1.68 1.41 unit 121. 7 123.6 124.7 124.4 125.9 127.5 124.9 126.6 127.8 129.8 126.7 131.6 131.0 5.45 5.53 7.25 6.28 6.58 8.86 7.07 5.33 6.54 6.11 7.94 5.76 6.26 41.1 40.6 40.8 40.8 41.0 41.2 40.6 40.7 41.1 40.8 40.5 41.3 41.9 1.65 1.34 1.89 1.77 2.11 1.66 1.88 1.41 1.69 1.29 1.77 1.43 1.29 �34 Table 5. Weights (kg) and body measurements (cm) of yearling and adult female mule deer trapped on control and treatment units of the Ridge study area in Piceance Basin, Colorado, 1988-94. Weight ~ Total body: length so ~ Left hind foot len9!;:h SO ~ Unit Year a Control 1988 1989 1990 1991 1992 1993 1994 2 2 10 8 4 46.2 46.0 49.48 50.9 50.6 Yearlings 1.91 6.43 2.38 2.50 4.85 146.1 148.4 147.9 149.2 145.6 10.75 2.26 2.49 3.38 4.19 45.8 47.8 45.8 45.7 45.5 0.99 3.04 1.08 1.02 1.59 2 49.1 5.51 152.0 8.49 46.5 3.25 Treatment 1988 1989 1990 1991 1992 1993 1994 2 7 8 10 10 1 1 50.2 49.3 50.8 49.4 50.3 48.7 49.8 0.35 3.58 3.38 3.45 5.38 151.2 154.3b 150.1 149.8 149.8 150.0 144.5 3.18 6.20 4.94 5.31 5.35 46.0 46.6 45.6 45.3 47.2 45.1 45.1 0.00 0.69 1.26 1.51 6.37 Control 1989 1990 1991 1992 1993 1994 19 21 30 31 14 11 67.7 66.9 62.9 63.0 65.3 65.0 Adults 5.22 5.51 5.18 5.67 8.32 8.04 168.6 166.7 162.3 164.3 169.3 169.2 5.20 6.98 6.40 7.89 7.70 8.49 48.1 47.3 47.3 47.5 47.3 48.2 1.10 1.09 2.19 1.29 1.32 1.66 Treatment 1989 1990 1991 1992 1993 1994 39 25 32 36 9 10 65.8 67.6 62.8 62.4 61.9 64.5 5.04 5.14 6.24 6.91 2.58 7.74 166.6 166.8 162.2 162.4 163.8 164.5 6.57 5.81 7.07 7.32 7.08 5.94 47.8 48.1 47.0 47.2 47.4 47.7 1.32 1.44 1.06 1.47 1.04 0.97 8 b n = n= 9. 6. SO �Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS State of Project REPORT Colorado No. Mammals W-153-R-8 Research Work Plan No. 3 Elk Investigations Job No. 8 Effects of Early Hunting on Elk Distribution Period Author: Covered: July Seasons 1, 1994 - June 30, 1995 R. Bruce Gill Personnel: R. Bruce Gill, Gary C. White, Jeff Madison, and George D. Bear Mary M. Conner, John Ellenberger, ABSTRACT George D. Bear, the Principal Investigator on this job, retired during the segment. He and Dr. Gary C. White are preparing a manuscript describing the results of the pre-experimental phase of this study. Dr. White and Mary M. Conner, a PhD candidate from Colorado State University, have prepared a draft study plan for the experimental phase. A copy of that draft study plan is appended. ��37 EFFECTS JOB PROGRESS REPORT OF EARLY HUNTING SEASONS ON ELK DISTRIBUTION R. Bruce Gill P. N. OBJECTIVE Evaluate the effects of early big game hunting seasons muzzleloading deer and elk seasons) on the distribution River Data Analysis unit. SEGMENT 1. (archery and of elk in the White OBJECTIVES Prepare a manuscript or Job Final alternative management strategies Report and make recommendations and for additional research. for INTRODUCTION During the Segment, George D. Bear, the Principal Investigator on the project retired. Several meetings and discussions were held with interested parties and those who had a stake in the outcome of this study to plan a future course of action. originally this project was conceived as a 2-phase investigation. Phase 1 gathered and analyzed preliminary data to describe patterns of elk distribution in relation to archery and muzzleloading hunting activity in August and September. Conventional wisdom held that early hunting seasons caused elk to migrate prematurely to privately owned ranches at lower elevations. Preliminary analyses of Phase 1 data suggest that the majority (>85%) of radio-collared cow elk on the study area redistributed themselves from public lands with unrestricted public to private lands where access was limited within the first week of archery season in mid-August. These results suggested that early hunting season could be a primary causative agent effecting elk redistribution. Currently, we are developing a study plan which will include treatment areas and controls to test this hypothesis. RESULTS A draft of of Phase 1 experiment results in a scientific manuscript has (the descriptive phase). A to test the hypothesis that movement of elk from public been prepared summarizing the results draft study plan describing a proposed early-season hunting disturbance of elk to private land (Appendix A). ��39 ELK MOVEMENTS IN RESPONSE TO EARLY-SEASON HUNTING IN THE WHITE RIVER AREA DRAFT STUDY PROPOSAL Mary Conner Department of Fishery and Wildlife Biology, Colorado State University. Fort Collins, Colorado 80523 Fall 1995 EXECUTIVE SUMMARY The White River elk herd (Cervus e/sphus) has been growing since its re-establishment to the area in 1929 and its numbers are now at the upper bounds of the desired management objectives. The number of hunters using the area has grown along with the elk herd, with especially significant increases in the number of early-season hunters during the past ten years. Also over the past 10 years, there have been increasing observations and complaints that elk have been moving off easily accessible public lands to lower elevation private lands or to remote and inaccessible areas during the early-season hunting period. The increasing disturbance of early-season hunting may be causing the elk to move off their summer ranges before fall migration. Early movement has lead to complaints by local landowners onto whose land the elk are moving, by resource managers, and by early-season hunters. All parties indicate that it is not the number of elk in the area causing the problem, but the distribution of the elk. Documented responses of elk to hunting from previous studies includes increased movement, movements away from hunted areas, movement into inaccessible areas, and movement into no-hunting areas. There is some evidence that elk response may be dependent on the hunter density and the amount of escape cover in the viCinity. Further, there is an indication that the responses of elk to hunters is of short duration. All the previous elk studies with respect to hunting have been observational in nature and have not tested for a cause and effect relationship between hunting activity and elk movement. A preliminary study on the proposed study area was conducted by George Bear and Jeff Madison from the Colorado Division of Wildlife. They trapped and radio-collared 20 elk cows in 1992. From 1992 to 1995 the radio-collared elk were intensively monitored during August and September, approximately one month before and one month after the opening day of early-season hunting. Corresponding to the opening of early-season hunting, all elk located on non-refuge areas (areas easily accessible to hunters) moved to refuge areas (private land or wildemess area not easily accessible to hunters), while all elk located in refuge areas stayed in refuge areas. The mean date of movement was not significantly different from the opening date of early-season hunting. Although the proportion of elk moving from non-refuge to refuge areas and the date of movement indicated that elk movement correlates with the opening of early-season hunting, there are several altemative hypotheses that could also explain these movements. To infer a casual relationship between elk movement and early-season hunting, the study needs to be continued with a manipulative experiment. The objective of the proposed study is to evaluate the impacts of early-season hunters on the movement and distribution of elk with the basic premise or null hypothesis that early-season hunting does not effect elk movements. A manipulative experiment will be conducted to determine if the presence of hunters is contributing to the movement of elk off of public land, to private land or wilde mess areas. The principle objectives to answer this question are: 1. 2. 3. Test the hypothesis that early-season hunting disturbance of elk results in movement of elk from public to private land. Evaluate altemative hypotheses about causes of elk movement such as sheep grazing, woodcutting, recreationalists, weather, or forage quality. Compare the movements of this heavily hunted elk population with movements of elk from concurrent studies in other parts of Colorado. Elk will be captured and radio-collared on Game Management Units (GMUs) 12,23,24, and 33, and treatment and control groups established. A total sample size of 80 radio-collared elk are recommended based on power analyses. Telemetered elk will be located at least twice a week for the month preceding �40 and the month following the opening date of early-season hunting. The locations will be labeled as either refuge or non-refuge for the analysis. Three experiments are presented in this proposal. The preferred altemative consists of a crossover design with the study area split into two experimental units of the GMU areas north and south of the White River. In the first year of the study, one area would be treated with hunting moved forward one week, while the remaining area would be treated with hunting delayed two weeks from traditional opening day, yielding a three week difference in opening dates. The treatments would be reversed in the second year of the study. Data from the United States Forest Service on sheep grazing areas, woodcutting permits, and recreational use will be used to evaluate the effects of these disturbances on elk movements during the study period. Analysis of the elevation shifts and movements of the control and treatment elk will be used to evaluate the effects of weather and forage quality on elk movements. Results from this study will provide managers with scientifically defensible and publicly credible information that can be used in to define management strategies that will decrease the early-season elk distribution problems. The conclusions will have direct applicability to the White River area and will also contribute to the body of scientific knowledge on elk in the he westem United States. Outputs from this research will include written reports, scientific publications and spatial use maps. �41 TABLE OF CONTENTS INTRODUCTION ...............................................................•..... BACKGROUND ............................................................•........... " 43 44 ELK MOVEMENT IN RESPONSE TO HUMAN ACTIVITIES ••••••••••••••••••••••••••••••••••••••• " 44 ELK MOVEMENT IN RESPONSE TO HUNTING •••••••••••••••••••••••••••••••••••••••••••••••• 45 CONCLUSIONS • • • • • • • • • • • • • • • • • . • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 46 PROJECT OBJECTIVES ................•............................................... 46 PROPOSED STUDY AREA ..................•....................................•...... 47 PILOT STUDY ...•........................•........•................................... 47 PROPOSED METHODS .............................•.....................••..••...•.••. 50 Common Study Design Parameters ..........................•........................ 51 Study Design ............................•........................•............ 51 Sample Sizes ..............................•.................................•. 52 Capture and Constraint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • • . . . . . . . . . . .. 52 Radio-Telemetry Methods ...........................................•.......... " 52 Comparative Studies ..............................................•............. 52 Alternative Study Designs ..................................•.....................•.. 53 Study Design Under Ideal Conditions ..............•.....................••..•......•.. 58 Application of Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 58 LITERATURE CITED 59 BUDGET -1996 AND 1997 61 Appendix A: Logistic regression analysis for 1992 and 1993 . FIGURES Rgure 1. Number of early-season hunters for DAU E-6 in the White River area, Colorado, 1984 -1994 .................................................................•. Rgure 2. White River study area ......................•................................... Figure 3. Example of a logistic regression curve to estimate the date of movement for an elk moving from a non-refuge area to a refuge area (A), and elk staying in a refuge area (B). ..•....... Figure 4. Proportion of elk on refuge areas for elk in non-refuge and refuge areas before opening of early-sea son hunting in 1992 and 1993. ...•.•.....•........•.......•.••..•.••.•. Rgure 5. Study design layout showing nested levels of experimental units. ...•............•...... 43 47 49 50 54 TABLES Table 1. Table 2. Table 3. Table 4. Table 5. Table 6. Table 7. Locations of radio-collared elk before and after opening day of early-season hunting, White River elk herd, 1992 - 1994. .....................................•.......... Mean dates of movement of elk from non-refuge to refuge areas, opening date of early-season hunting, and p-values from a t-test for differences for the White River elk herd, 1992 - 1994. . Example of crossover design blocking out effects of sheep grazing activity with a hunting and no-hunting treatment. .......•..................................•..•......... Layout of Latin square crossover design with corresponding model for alternative one. . .... Sample size (number of collared elk) per haH for treatment groups based on estimates of effect size and 90% power for primary response variables. Layout of Latin square crossover design with corresponding model for alternative two Layout of Latin square crossover design for alternative three ..............•............ 48 49 51 54 56 57 57 ��43 INTRODUCTION The White River elk (CelVUS elaphus) herd of Data Analysis Unit (DAU) E-6 has been one of the most heavily hunted, managed, and documented elk herds in Colorado (Boyd 1970, Freddy 1987, Gray et al. 1994). The herd has been growing in number since its re-establishment to the area in 1929 and is now at the upper bounds for the desired management objectives (Gray et al. 1994). The number of hunters using the area has grown along with the elk herd, with especially significant increases in the number of early-season hunters between 1985 and 1992 (Figure 1). l:! ~ 3,000 • ::t: ill fi 2,600 ••• tI • (II ~ 2,000 OJ t 1,500 ::J :z 1,000 +---+---+---+---+--I---+---+---+--~ 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 Year Figure 1. Number of early-season hunters for DAU E-6 in the White River area, Colorado, 1984 - 1994. Along with the increase in early-season hunters has been an increase in the use of four-wheel drive (4 WD) and all-terrain vehicles (ATV's) (Grayet al. 1994). The disturbance caused by an increasing number of early-season hunters and hunters using 4WD vehicles and ATV's may be causing elk to move off summer range areas early. Historically, the main migration from summer to winter ranges took place between late November and early January (Boyd 1970). However, Freddy (1987) noted that elk were commonly found in lower elevations, typical winter range, during the September and October hunting seasons, which was uncommon during the 1960's. Now there is evidence that elk are moving during August and early September during early-season hunting (Gray et al. 1994), from areas easily accessed by hunters to areas not easily accessed. The easily accessed areas are primarily public forest lands with abundant road access, and the more difficult access areas are either private lands or wilderness areas with sparse or no public road access. These movements and resulting redistribution of elk has lead to complaints by local private landholders, resource managers, and hunters (Grayet al. 1994). As the number of early-season hunters continues to increase, early-season hunter induced elk movements could lead to elk redistribution with corresponding overpopulation problems in localized areas. Also, early fall movements may leave elk summer range under used at the cost of over using winter range. In the White River area, there have been complaints by private landholders bordering public areas of range damage and loss of hunting income, complaints by resource managers that riparian areas are being damaged by this redistribution, and complaints by early-season hunters of lower success rates in the public areas they hunt. All parties indicate that it is not the number of elk in the area causing the problem, but the distribution of the elk (Gray et al. 1994). Although the phenomena of elk movements from public to private lands has been discussed, it is not well documented. A three year pilot study from 1992 and 1994 was conducted on 14 to 20 radio-collared White River elk from Game Management Units (GMU) 12,23,24, and 33. During the study, all radio-collared elk located on nonrefuge areas moved to refuge areas. For all years combined, the mean day of movement was not different than opening day of early-season hunting. Although a causal relationship was not established by this study, it provided strong evidence of a correlation between elk movements and the opening of early-season hunting. �44 To more thoroughly understand elk response to hunting this study proposes a manipulative experiment to evaluate the cause and effect relationship between the movement of elk and the opening of early-season hunting in the White River DAU. The null hypothesis is that elk movements are the same on hunted and non-hunted areas, or that early-season hunting does not cause elk to move. Hypotheses on the timing and extent of other factors that may explain the elk movements will also be examined. BACKGROUND Elk Movement In Response to Human Activities Elk response to recreational activities such as hiking, cross-country skiing and driving seems to vary depending on how habituated the elk are and whether the population is hunted. In the Rocky Mountain National Park, where elk were accustomed to people and were not hunted, elk showed little response to approaches of automobiles and people on foot, day or night (Schultz and Baily 1978). Another study of an unhunted elk population in Yellowstone National Park found variation in elk response to cross country skiers (Cassirer et al. 1992) depending on how accustomed the elk were to people. In areas of typically low human activity during the winter, elk responded with flight distances between 125 1,700 m, while in an area of high human activity, the flight distance was much less, ranging between 0 - 300 m. Interestingly, elk in the area of high human activity showed a three-fold increase in flight distance when they were disturbed just outside the IOO-ha area where people were present 24 hours each day (Cassirer et al. 1992). Thus, even habituated elk appear to be influenced by changes in human space use. Acceptance of human activity may be a learned response ofunhunted elk. Movements with respect to roads seems to be related to the habituation of elk to vehicles and to the level of road use. In Rocky Mountain National Park, where the elk were not hunted and there was high traffic volume, Schultz and Bailey (1978) found that none of their 14 delineated elk behaviors changed with traffic volume and there was little to no avoidance of the roads in the winter. In contrast, on the Roosevelt National Forest, which is adjacent to the Rocky Mountain National Park, Rost (1975) found that a population of hunted elk avoided roads in winter ranges. Wright (1983) found a variable response; mean distance to jeep trails more than doubled (800 m to 2,100 m) during fall from just before and during hunting season. Elk may learn to accept human activity that poses no threat. However, behavior of elk in areas with respect to recreational activity has not been examined in hunted areas for comparison. Elk may also respond to the level of activity on a road. Wright (1983) found that elk had double the average distance to heavily traveled paved roads than to lightly traveled jeep trails. Similarly, Hershey and Legee (1982) found elk crossed secondary roads more frequently than primary roads. Czech (1991) found a significant increase in mean elk distances to a road after it was opened to the general public with a corresponding increase in traffic levels. If elk respond to different levels of activity on a road they likely also respond to different levels of hunting activity. Effects oflogging and mining disturbances on elk movements depend on the cover available near the disturbances. Edge and Marcum (1985) found normal elk movements were changed by logging disturbances; elk moved significantly longer distances away from logging areas than towards them. Typically, a buffer zone of between 500 1,000 m separated areas of high elk use from areas of disturbance depending on the cover in the area (Edge and Marcum 1985). Czech (1991) found that elk tolerance of logging operations was correlated positively with proximity to hiding cover. In one of the few manipulative experiments on elk movements, elk calves were subjected to simulated surfacemining activities (Kuck et al, 1985). Compared to undisturbed calves, disturbed calves moved greater distances, used larger areas, showed greater use of coniferous forest, and lacked selection for favorable physiographic elements (Kuck et al. 1985). A strong case for a cause and effect relationship between elk movement and mining disturbance was built with this manipulative experiment. All studies that evaluated elk movements after the disturbance ended found that displacement appears to be temporary in most situations. The Yellowstone elk displaced by cross-country skiers typically returned close to their original locations after people left the area (Cassirer et al. 1992). Road proximities returned to pre-hunting distances after hunting season closed (Wright 1983) and mean distance to roads decreased soon after the roads were closed for the season (Hershey and Legee 1982). During the weekends, and immediately following the end oflogging activities, elk moved back into logged areas (Ward 1976, Edge and Marcum 1985). �45 Elk Movement In Response to Hunting How and to what extent hooting affects elk is not clearly understood (Adams 1982). Elk response to hooting pressure has been examined in several studies (Martinka 1969, Knight 1970, Craighead et al. 1972, Lemke 1975, Morgantini and Hudson 1979, Wright 1983) and two theories about elk response have been tendered. The first theory is that elk in recent years have learned to move to unhunted refuges and choose migration routes through lightly hooted areas (Altmann 1956, Martinka 1969) (learned behavior theory), while the second theory is that elk populations were decimated if they migrated through or lived in heavily hooted areas whereas elk living in lightly hooted areas persisted (Rudd et al. 1983, Boyce 1991) (population redistribution by differential hooting pressure theory). Neither theory has been experimentally tested. Documented responses of elk to hooting includes movement away from hooters or heavily hooted areas, increased movement, movement to thick cover, shifts in circadian patterns, and elevation shifts in migration patterns. Most responses seem to occur before winter migration, but some studies have noted changes in migration patterns that were attributed to hooting pressures. Responses of the elk may depend on density of hooters, roughness of terrain, or density of cover. Altmann (1956) described an evasive "migration" of elk hooted adjacent to Yellowstone National Park. With opening of hooting season, movements of these elk became relatively long (5 - 13 km), and they stopped moving only when they reached the sanctuary of the Park. Similarly, Martinka (1969) found that all of 12 marked elk located in the National Elk Refuge just prior to hunting season moved between 8 and 22 km to areas closed to hooting in Grand Teton National Park. Other studies have found less dramatic movement distances to refuge areas. With opening of hooting season, elk moved to densely forested (Irwin and Peek 1979) or shrubby (Morgantini and Hudson 1979) areas adjacent to their typical areas of activity, which provided refuge from hooting. In a study of elk responses to hooting pressure, elk moved toward winter range earlier than normal fall migration time, and the number of locations on private land increased during rifle season (Wright 1983). Wright (1983) also found that mean straight-line distances traveled by radio-collared elk tripled during hooting season versus the months before hooting season. Hooting disturbances appeared to affect the timing of cropland use; a hooted elk population limited open cropland grazing to early morning hours (roughly 0200-0400), while an unhunted population would enter cropland during daylight hours (Strohmeyer and Peek in press). A reversal in downward elevation migratory movement of elk, which coincided with the opening of the hooting season, was found by Knight (1970) and Morgantini and Hudson (1979). While some elk appear to have learned to move to refuge areas, elk in refuge areas may learn to stay there in response to hooting pressure. In Jackson Hole, both resident and migratory elk use the National Elk Refuge for their winter range (Martinka 1969). Of the 183 marked elk in the study, resident elk were defmed as elk located July through August within 15 km of the winter range, while migratory elk were not located in the area. In the fall, both resident and migratory elk had to traverse areas where hooting was permitted while moving to the National Elk Refuge winter range. Resident elk were conditioned to hooters presence and tended to remain on areas closed to hooting while migratory elk moved through them to the winter range. These studies of elk moving to, or staying in, refuge areas support the theory that elk are responding, as possibly a learned behavior, to hooting pressure. Other studies found evidence to indicate elk are not responding to hooting pressure, but are being differentially removed from heavily hooted areas. In Wyoming, a resident population of elk just east of Yellowstone National Park were being detrimentally reduced by differential hooting pressure on them (Rudd et al, 1983). Like the Jackson Hole herd, migratory elk that summered in protected areas of Yellowstone wintered with resident elk in the unprotected range just east of the park. Hooting seasons occurred before normal elk migrations from the park, placing heavy hooting pressure on the resident elk. The proportion of migratory elk from refuge areas had increased proportionally to the resident elk from no apparent behavior shifts due to hooting. Boyce (1991) noted that migration routes of elk wintering on the National Elk Preserve in Jackson Hole have changed dramatically between 1950-1954 and 1980-1984. The primary migration route shifted from being predominately on open hooting national forest lands to routes through Grand Teton National Park, where hooting is more restricted. In the 1950-1954 period, 22.1 % of the migrating elk were estimated to have moved through Grand Teton National Park, growing to 51.4% by 1980-1984. Boyce hypothesized that this may be to differential removal of the migrating animals from the population by hooting and not due to individuals shifting to alternative routes. However, the mechanism of these movement shifts in response to hooting has not been identified or tested. Elk responses to hooting pressure may depend on density of hooters. Wright (1983) examined elk response to different hooting seasons: archery/muzzleloading, rifle special elk, rifle special deer, and deer and elk rifle. The greatest density of elk hooters was during the rifle special elk, with distances traveled by elk the greatest during that season. Elk traveled distances three to four times greater during this season than during early-season hooting and muzzleloading season. Wright (1983) concluded that radio-collared cow elk were not adversely affected by hooters or muzzleloaders. �46 However, it is possible that the lack of effect by hunters and muzzleloaders could be due to their low densities (0.13 hunters/knf ) compared to the higher densities of rifle hunters (1.42 rifle hunters/km'), In a previous study on the proposed study area in 1985, Consolidation Coal Company had 23 elk collared to evaluate the perceived movement of elk onto their protected land during early-season hunting. In that study, 87% of the elk were still found on National Forest lands mid-way through early-season hunting and muzzleloading seasons (Camp, Dresser and McKee Inc. 1986). However, due to a change in hunting regulations in 1985, only 37% as many hunters were afield during the study as the previous year (Gray et al. 1994) perhaps explaining the lack of movement that year. Also, Zahn (1974), Lemke (1975), and Hershey and Legee (1982) noticed an increase in movements during the first 10-12 days of hunting season which decreased to normal during the remaining of the hunting season. These authors noted that hunting pressure was heavy during that time and relatively light for the remainder of the season. These studies suggest that there is perhaps a critical density of hunters that must be reached before elk begin to move in response to the hunting disturbance. An interesting observation from the Zahn (1974), Lemke (1975) and Hershey and Legee (1982) studies is that elk response to hunter disturbance was short lived. All studies noted that elk movements returned to normal after the initial 10-12 days of opening day of hunting, and all studies noted that the initial 10-12 days was when hunting activity was the greatest. Elk not only discontinued their erratic and increased movements but returned to their pre-season activity areas, indicating that hunting activity was a temporary disturbance. Elk movements during hunting may depend on the amount, location and quality of hiding cover, as well as the topography of the area. Elk either greatly increased their daily movements (Altmann 1956, Martinka 1969) or were found in thick vegetation inaccessible to most hunters (Lemke 1975, Irwin and Peek 1979, Morgantini and Hudson 1979). Also, preliminary data from George Bear's study (unpubl. data) on the elk on the proposed study area found that elk located in rugged and relatively inaccessible areas = 9) moved short distances (mean = 2.5 /an?) to dark timber and rough terrain when early-season hunting opened (Gray et al. 1994). In contrast, elk located in accessible areas = II) moved an average of 13 km to private land. There were no areas of dense timber or rough terrain for elk on the accessible areas; therefore they needed to move to private land for a refuge. Areas of rough topography and dense timber may serve as an refuge; if it is available, elk may not move far even in the face of high hunting pressure. en en Conclusions Elk are extremely plastic in response to human activity, habituating to people in areas where there is no hunting, and actively avoiding hunters in areas where they are hunted. Elk response to human activities appears to depend on the amount of contact and the danger of contact with humans. Avoidance of roads changes with the amount of use and the danger level. The presence of dense cover for hiding seems to attenuate movement responses to human disturbance. Finally, elk responses to human disturbances do not seem to persist after the disturbance is removed. Whether hunting is causing learned responses of avoidance and refuge seeking, or whether populations with certain behaviors are differentially removed are two theories of elk responses to hunting. Most hunter induced movements appear to occur before migration. As with other human activities, the level of activity, hunter density, and the amount of cover in the vicinity seem to affect elk responses to hunters. The literature indicates that elk responses to disturbance is short-lived. All of the studies of elk responses to hunting have been observational and it remains untested whether there is a causal relationship between hunting and elk movements. This study will examine this issue with an experiment designed to determine if elk move from non-refuge to refuge areas in response to early-season hunting. PROJECT OBJECTIVES The objective of this study is to evaluate the impacts of hunters on the movement and distribution of elk with the null hypothesis that elk movement is not affected by early-season hunting. The study will focus around a manipulative experiment to determine if the presence of the hunters is contributing to the movement of elk off of public land (easily accessible to hunting), to private land or wilderness areas (not readily accessible to hunting), but will also examine alternative hypotheses. Basic study objectives are: I. Test the hypothesis that early-season hunting disturbance of elk results in movement of elk from nonrefuge to refuge areas. 2. Test hypotheses about other possible causes of elk movement such as sheep grazing, woodcutting, recreationalists, weather, or forage quality. �47 3. Compare the movements of this heavily hunted elk population with movements of elk from concurrent studies in other parts of Colorado. Early-season hunters are the focus of the study because they are the first to begin hunting and they are alleged to be causing the premature movement of elk off summer ranges. Although later-season rifle hunters may also cause movement of elk to refuge areas or keep elk in refuge areas, timing of the rifle season movements are more acceptable to resource managers and private landholders. To eliminate confounding factors from rifle hunting pressure, the manipulations will be limited to the early-season hunting. PROPOSED STUDY AREA The study area consists of the GMUs 12,23,24, and 33, a subset ofDAU-E6, located in and adjacent to the White River and Routt National Forests (Figure 2). The area is approximately 4,540 km2 and is bounded on the north by the William's Fork of the Yampa River, on the west by state highway 13 (west side of the Great Hogback), on the south by 170 between Rifle and Canyon Creek (12 km east of New Castle), and on the east by Canyon Creek in the south, through the eastern side of the Flattops, joining up with the East Fork of the William's Fork in the north. Topography, climate, and vegetation vary widely throughout the study area. Elevation ranges from 1,629 m along the Colorado River to 3,700 m on the Flattops. Higher elevations have severe winters with heavy snowfall, while the lower elevations have comparatively mild winters. Mean annual precipitation at 3,000 m in the Routt National Forest is about 100 em, while at Rifle (1,629 m) and Craig (1,856 m) mean annual precipitation is about 30 em, Vegetation types range from the montane/subalpine zone in the higher elevations (>2,600 m), to the transitional zone in the middle elevations, to the Great Basin zone at the lower elevations «1,980 m) in the southern and northern parts of the study area. The area is described in detail by Gray et al. (1994). Higher elevations of the Flattops, Sleepy Cat Ridge and the White River/Colorado River divide areas provide good summer and fall forage for elk. Lower elevations are typically used as winter range by elk. Hunting pressure is relatively light on the Flattops, due to limited road access, and relatively heavy on the Sleepy Cat Ridge and the White/Colorado River divide areas due to abundant road access. The study area is 34% private land and 66% public land, with most of the public land being USFS (54%). Unit 24 is mostly (92%) public land, while the other three units average 57% public land. Private land is comprised mainly of ranches and coal mines. Much of the public land is grazed in the summer by sheep and cattle. Hunting for large and small game is economically important to local residents in the study area (Gray et aI. 1994). Guides, outfitters, local service sectors, and private landowners (who sell trespass permits) make a large proportion of their annual income during the elk and deer hunting seasons, with most of the revenues coming during rifle hunting season. Figure 2. White River study area. PILOT STUDY George Bear and Jeff Madison from the Colorado Division of Wildlife (CDOW) trapped and radio-collared 20 elk cows in the proposed study area in 1992. In 1995, an additional 11 cows were radio-collared in the study area, and the number of early-season hunter permits was reduced by 65%. From 1992 to 1995 the radio-collared elk were intensively monitored during August and September, from approximately one month before the opening day of earlyseason hunting to one month after opening day. There were 20, 16, 14, and 13 elk tracked in 1992, 1993, 1994, and 1995 respectively. A minimum of 18 locations was collected for each animal used in the analysis. Each location was classified as either being in a "refuge" or "non-refuge" area. Refuge areas were defined as having limited or no vehicle access, typically privately owned land or wilderness areas. Non-refuge areas were accessible by motor vehicles, typically USFS or BLM land. For each location of a telemetered cow, a zero was assigned to non-refuge locations, and a one was assigned to all refuge locations. All elk located on refuge areas remained on refuge areas throughout hunting season, while elk located in non-refuge areas moved to refuge areas with the advent of hunting season (Table 1). �48 Table I. Locations of radio-collared elk before and after opening day of early-season hunting, White River elk herd, 1992 - 1994. Movement Combinations: YEAR pre-opening -> post-opening 1992 1993 refuge -> refuge 10 10 1994 Total 0 0 refuge -> non-refuge 0 0 non-refuge -> refuge 10 6 non-refuge -> non-refuge 0 0 0 0 sample size 20 16 14 40 From these data, a logistic regression was done for each animal to estimate the date of its movement from nonrefuge to refuge area (Appendix A). The date of movement was defined as the date at which the logistic curve crossed 0.5 on the y-axis, indicating an equal probability of being on a non-refuge or refuge area (Figure 3). From this analysis, a mean date of movement for all animals was computed. A two-tailed t-test was computed to determine if the mean date of movement was different from the opening date of early-season hunting for each year (Table 2). The analysis was not applied to the refuge elk since none of them moved to a non-refuge area during the study period. These data were also used to evaluate the proportion of elk began on a non-refuge area and moved to a refuge area, before and after opening of early-season hunting (Figure 4). Because only the elk in non-refuge areas moved, the proportion of elk beginning in refuge areas remained constant (upper line in Figure 4) in the analysis. These data were appropriate for a Before-After-Control-Impact-Paired-Sampling (BACIPS) analysis with a ttest for the difference in proportions (Stewert-Oaten and Murdoch 1986, Osenberg et a1. 1994). Instead of comparing the mean difference in a response, such as proportion moved from non-refuge to refuge areas between a control and impact site, the BACIPS approach compares the mean difference (control- impact) in the before and after periods, where samples are temporally paired, thereby controlling for naturally occurring spatial and temporal variation (Osenberg et al. 1992, 1994). The average of the proportional differences Before and After the opening of early-season hunting was tested with the null hypothesis; Ho: = J_, and Ha: Jb < J.. The null hypothesis was rejected for 1992 (p < 0.0002) and 1993 (p < 0.0001), which supports the alternative hypothesis. The difference before was less than the difference after, that is, the proportion of elk that moving to refuge areas increased for the non-refuge group after the opening of early-season hunting. This suggests that early-season hunting opening had an influence on elk locations, specifically, elk moved to refuge areas after the opening of early-season hunting. Although mean date of movement, and difference in proportion of elk found on refuge or non-refuge areas before and after the opening of early-season hunting indicates that elk movement correlates with the opening of earlyseason hunting, several alternative hypotheses, such as woodcutting activity or weather, could also explain the movements. There are three main criteria required to strongly infer a cause-and-effect relationship between two factors: I) there is a correlation between factor A and factor B, 2) the presence of factor A by itself causes factor B, and 3) factor B does not occur when factor A is removed. Criteria one has been satisfied by the pilot study. Criteria two, everything else being the same, a change in early-season hunting causes a change in elk movement, and criteria 3, there is no elk movement when there is no early-season hunting, can only be tested by a manipulative experiment. An effect consistently seen in a replication of a well-designed experiment can only reasonably be explained as being caused by the manipulation of the experimental variables (Manly 1992). With an observational study the same consistency of results may occur because all of the data are affected in the same way by some unknown and unmeasured variable (Manly 1992). Thus, a manipulative experiment, that isolates the effect of early-season hunting, and controls for, or blocks for, other potentially confounding variables (weather, woodcutting activity etc.) is required to infer that early-season hunting is causing elk to move from refuge to non-refuge areas. a;, �49 (A) 1.5 Refuge ( Estimated date of movemlt • 0 = non-refuge, 1 = refuge at 0.5 probability 0( being on refuge -Filled logistic curve 0.5 Non-refuge 0 7f26193 ) 8ISI93 8/25/93 = estimated date at movement 8115193 8125193 914193 9114193 004/93 (8) 1.5.,--------------------, Refuge 1 No movement • 0 1 = non-refuge, = refuge -Filled logistic curve 0.5 Non-refugeo i---t---t----t----t,......--t----+-.,--+---\ 7120193 7J3O$3 8/9/93 8119/93 8/29/93 9/8/93 9'18193 912819310/8193 Figure 3. Example of a logistic regression curve to estimate the date of movement for an elk moving from a non-refuge area to a refuge area (A), and elk staying in a refuge area (B). Table 2. Mean dates of movement of elk from non-refuge to refuge areas, opening date of early-season hunting, and pvalues from a t-test for differences for the White River elk herd , 1992 - 1994 1992 1993 28Aug,n=6 1994 Mean date of movement 27 Aug, n= 10 95% Confidence Interval 24 Aug - 31 Aug 23 Aug-2 Sep Opening date of early-season hunting (da) 29 Aug 28 Aug 26 Aug Ho: d,. = opening date p=O.28 p=l.OO p= Combined Years n= p= �50 1992 1.2 1.0 lll. III III ~ CD IJ..e. 0.8 E d~ 0 ~ Vt- '0 0.4 C 0 a after, d, / CD a::: 0.6 c iii "~~. -Begin in non-reluge ~Beginin refuge V e a.. 0.2 / 0.0 -17 -12 -4 0 7 12 Archery Season Opening Date Days before opening day 18 22 28 Days after opening day 1993 r 1.2 ~ 1.0 8, ~ 0.8 ·~-rrrr-r-r~rrl-ft[~~\~-~~'-~-~~ d ifferences before, d, QI a::: 0 6 5 . V r- ~ iii '0 0.4 f- c .2 a. £. 0.2 V r- V -16 Days before opening day I I 1/"- 'd",ere"~ s after, d. Ir-------~ ~ ~inin nonrefuge / I -,,- Begin in refuge I 9 -9 0 Archery Season Opening Date 18 26 Days after opening day Figure 4. Proportion of elk on refuge areas for elk in non-refuge and refuge areas before opening of early-season hunting in 1992 and 1993. PROPOSED METHODS There are several alternative methods that could answer the question of whether the elk movement patterns in late August are related to the presence of hunters. The common factors to all alternatives will be discussed first, then the alternative study designs will be presented with a preferred design. Finally, an optimal study design, one with no limitations on money, resources, or time is presented for a comparison with proposed alternatives. �51 Common Study Design Parameters Study Design Although elk may respond in many ways to disturbance, effects of early-season hunter-induced movement will be tested by four response variables: mean date of movement, proportion of elk that move from one classification to the other, mean elevation changes, and mean distance moved between successive locations. The primary hypothesis tested by this study is: Ho: Elk movement from non-refuge to refuge areas is not different between hunted and non-hunted groups. Ha: Elk movement from non-refuge to refuge areas is greater for hunted groups. To build a strong case inferring a causal relationship between elk movement and the early-season hunting pressure we need to: 1) devise and test alternative hypotheses, and nearly as possible, exclude one or more of the hypotheses (platt 1964). Elk have been shown to move in response to recreational activity, logging activity, and road activity, so other activities that occur in the area should be considered. The following alternative hypothesis could also explain elk non-migratory movements in the White River area: Hal: Ha2: Ha3: Ha4: HaS: Elk Elk Elk Elk Elk are moving in response to sheep grazing activity/forage quality are moving in response to presence or activity of woodcutters are moving in response to recreationalists are moving in response to weather/forage quality have a learned, for an unknown reason, to move at this time of year. To assess the distribution of resource activity on the study area, data from the forest service on sheep grazing allotments, woodcutting permits, and recreational use will be evaluated. If the distribution of these activities remains uniform from year to year, then the crossover designs presented in this proposal will block out for these effects and only hunting will vary. What makes the crossover designs good for isolating for hunter effects alone is that each hunting treatment is used on each area. As a result, the effects of resource use averages out for each treatment. For example, say there were 10 bands of sheep grazing on one half of the study area, and only 5 bands grazing on the other half As long as this was consistent from year to year each treatment would experience each level of grazing, averaging them out when comparing between treatment (Table 3). Table 3. Example of crossover design blocking out effects of sheep grazing activity with a hunting and no-hunting treatment. North half of study area South half of study area Year 1 no-hunting treatment 5 bands of sheep hunting treatment 10 bands of sheep Year 2 hunting treatment 5 bands of sheep no-hunting treatment 10 bands of sheep To some extent HaS, that elk have a learned behavior to move at this time of year, will be evaluated by the 65% reduction in early-season hunter permits in 1995. If the movement is a persistent behavior, no reduction in the movements is expected. That is, if the same proportion of non-refuge elk move to refuge areas, even with the reduced density of hunters, then there may be a learned component to this action. If a lower proportion of elk move in response to the lower density of hunters, then it would seem that elk behavior is flexible to meet the changing conditions or level of danger or disturbance and may inferred to have little persistent learned effect. Any persistent learned behavior will cause a bias toward acceptance of the null hypothesis, that elk movement is not different between hunted and non-hunted groups. For example, if elk are hunted one year, and the next year still move as a learned behavior even when they are not hunted, there will be less difference between the movement patterns of hunted and unhunted groups the second year. Thus, the learned behavior could confound hunting induced �52 movements. Because the literature indicates that elk responses to disturbance are short lived (Zahn 1974, Lemke 1975, Ward 1976, Hershey and Legee 1982, Edge and Marcum 1985, Cassirer et al. 1992), the study is designed on the assumption that there will be little persistent behavior from year to year. Sample Sizes The number of animals needed for the study will depend on study design, estimation of variance for the response variable, and effect size. A power calculation has been done for primary response variables, and the larger required sample used, providing enough power for all tests. Tests will be done at ~90% power, to ensure powerful tests for a statistically significant conclusion that elk did not move in response to early-season hunting pressure. Sample size calculations are discussed with each alternative because they depend on study design. Capture and Constraint Helicopter netgunning will be the primary method of elk capture and will be contracted out to Helicopter Wildlife Management from Salt Lake City, Utah. Once a group of elk are located and an individual is "randomly" selected from the group, it is perused (typically < 30 sec) until the netgunner can fire a net over the elk. Once the elk becomes entangled in the net, it is blindfolded, hobbled and the net is removed to allow for installation of the telemetry collar (phillips 1994). Collars will be in the 148 - 151 MHz transmitting range with a mortality sensor. Because only cows will be collared, the collars will not be of an expanding design. Preferably, elk will be captured on their summer range in early July. Of the seven elk radio-collared by netgunning in August of 1995, the average distance between their capture sites and location four days after capture was 6.4 Ian. The elk moved < 2.5 Ian between their first second location after capture (3 days between first and second relocation). Therefore, capturing elk in early July will be early enough to avoid confounding capture movements with early-season hunter effects, but late enough so that the elk will be in their summer ranges to help ensure the desired distribution for the samples. Once the number of cows from each GMU is decided, capture sites will be randomly selected until an appropriate number of sites are located within the refuge or non-refuge areas. The pilot will go to the randomly selected location, then capture the first cow found from that location. In 1997, cows will be captured to replace losses from the sample due to mortality and transmitter failure. The cows will be replaced to balance the number of elk in the GMUs and classifications (refuge or non-refuge) to compensate for elk shifting location as well as lost elk. Radlo- Telemetry Methods All elk locations will be collected using aerial telemetry. An H-antenna will be mounted to each strut of an airplane, and the signal received using a four-band scanning receiver. Location will be determined using a Global Positioning System (GPS). Telemetry system error is a combination of observer error and GPS error. The error will be tested by selecting a minimum of 40 locations from a variety of topographical and vegetative types represented on the study area (stratified random sample). These selected sites will be located to ±50 m using the GPS and a beacon will be left at the site to be located by air. To ensure the error contains the component of observer error as well as location error, this will be a blind test with the selected locations unknown to the observer. Elk locations will be collected at least three times a week at regular intervals of one and two days between collection. The main hypothesis that this study addresses is the relationship between elk movements and early-season hunting, especially around the opening day of early-season hunting. Therefore, the time frame of analysis is defined as one month before to one month after opening day of early-season hunting. Elk locations before and after this time interval will be collected and may yield information about elk responses to other variables (sheep grazing, woodcutting etc.), but confining the intensive data collection to around the opening of early-season hunting will eliminate confounding factors from the analysis and focus on the disturbance of interest. I will attempt to collect a minimum of 15 locations on each animal the month before and after the opening date of early-season hunting. A Graphical Information System (GIS) map will be used to record all spatial information, such as refuge and non-refuge polygons, and elk locations. The map will be a 1:100,000 scale with 30 m elevation intervals, and will contain hydrography, property ownership, boundaries, and roads. Universal Transverse Mercator (UIM) 1983 will be the coordinate system used for all spatial analysis. Comparative Studies Data from at least two ongoing CDOW elk studies will provide additional data for comparison to data collected from this study. One study, conducted south of New Castle and Rifle involves 75 radio-collared cow elk. The other �53 study, which is of elk dispersal conducted around Dinosaur National Monwnent, is addressing similar questions to this study. Some of those collared cow elk are presently located north of Hayden in the Bears Ears area of Routt National Forest providing a potential comparison area close to the study area with heavy early-season hunting pressure and similar weather conditions. Alternative Study Designs Alternative one - open hunting season one week earlier on one area (chosen randomly) and two weeks later on the remaining area; Reverse treatments for year two. Alternative one is the preferred study design because it allows for blocking of confounding effects of space (area effects) and time (year effects), and is relatively easy to implement. The study area to the north or south of the White River would be chosen for application of treatment one, opening hunting season one week earlier than the typical opening date of the last weekend in August. Treatment two would consist of opening hunting season two weeks later than opening date on the remaining half of the study area. Hunters would choose and hunt on the study area GMUs to the north or south of the White River. Hunters choosing the early opening area would have an extra week of hunting. Hunters choosing the later opening season area would loose only one week of hunting because the season will be extended one week later than usual and, hence, will have more time during the elk rut period. The study area will be split roughly east-west by the White River and North Fork of the White River into two halves for application of early or late opening treatments. GMU 12 and part of GMU s 23 and 24 will comprise the north half, while GMU 33 and part ofGMUs 23 and 24 will comprise the south half. The primary response variables will be mean date of movement and proportion of elk that change classification (refuge -> non-refuge or non-refuge-e-refuge). Elk movement responses of elevation changes and distance between successive locations will also be analyzed to examine effects of hunting activity, as well as the effects of sheep grazing, woodcutting, and recreational activity. This is a nested type of design, with two levels of analysis. The lowest level of analysis is within half, where samples will be selected by location in refuge or non-refuge areas. This is a stratified random sample, with refuge or non-refuge area as the experimental unit, elk as the observational unit (Figure 5), and hunting as the treatment. If hunting has no effect on elk movements, then date of movement to refuge areas should not be the same as opening date, date of movement should not differ between non-refuge or refuge areas, and proportion of elk on refuge and non-refuge areas should not change with the opening of hunting. The hypothesis tested would be: Ho: date of movement = opening date Ha: date of movement * opening date Ho: date of movement of elk on non-refuge areas = date of movement of elk on refuge areas Ha: date of movement of elk on non-refuge areas * date of movement of elk on refuge areas Ho: proportional difference of elk on non-refuge and refuge areas difference of elk on non-refuge and refuge areas after hunting Ha: proportional difference of elk on non-refuge and refuge areas difference of elk on non-refuge and refuge areas after hunting before hunting opens = proportional opens before hunting opens * proportional opens Differences in time of movement for the fist two hypotheses will be tested with a one sample t-test, and the proportion of elk moving will be tested as done for the 1992-1993 pilot data. There would be two spatial replicates and two temporal replicates of each test. �54 Figure 5. Study design layout showing nested levels of experimental units. On the next level, this is a two-period crossover design (Manly 1992, Ott 1993) which is a essentially a 2 x 2 Latin squares design (Table 4). In this design, the experimental unit is north of south half of the study area, the observational unit is individual elk (Figure 5), and the treatment is early or late opening of early-season hunting. Here, the responses between the differently treated halves would be compared. A key issue in this wider analysis is that only elk on non-refuge areas will be compared between halves in statistical tests since response in movement of hunted elk is the primary concern of this elk study. This is not to say that elk on refuge areas are not important - they serve as a crucial behavioral control in the within-half analysis. If hunting has no effect on elk movements, then there should be no difference between the date of movement to refuge areas or the proprotion of elk on non-refuge areas for either treatment. The null hypothesis tested at this level would be: Ho: mean date of elk movement on early hunting GMU = mean date of elk movement on late hunting GMU Ha: mean date of elk movement on early hunting GMU '" mean date of elk movement on late hunting GMU On Early and/or Late Opening Date Ho: proportion of elk on non-refuge areas in early hunted GMUs = proportion of elk on non-refuge areas in late huntedGMUs Ha: proportion of elk on non-refuge areas before hunting opens '" proportion of elk on non-refuge areas after hunting opens Table 4. Layout of Latin square crossover design with corresponding model for alternative one. Factor B (periods) Sequence North half of study area South half of study area Collared Elk 1996 n Early opening Late opening n 1997 Late opening Early opening �55 This analysis is based on an assumption of minimal interaction between the blocked sources of variation (GMU and year). The corresponding analysis of variance for the model is: where: Ok 7t1(k) a, = = = p 9' = fixed effects for kth sequence random effects for individual animal fixed effects due to treatment fixed effects due to period (year) A power analysis to determine the number of cows needed from each half of the study area was done for: I. date of movement compared to opening date within half; 2. proportion of elk that move from non-refuge to refuge between halves on each opening date; 3. date of movement between halves. Power of at least 90% was specified for strong confidence that elk movement was not attributable to early-season hunting. Variance was estimated from the pilot study data of 1992 and 1993 as 4 day' for mean date of movement tests. Previous proportions of elk on refuge and non-refuge areas before and after hunting season from pilot data were used in the estimation of variance and effect sizes for change in proportion tests. Because the hunted elk, or elk on non-refuge areas, are the key issue in this study, emphasis in sample size calculations was given to the comparisons between halves. Sample size calculations (Chapman 1995) were run for different effect sizes for all three variables (Table 5). Between half analysis showed response variable 2 to require the largest sample size (Table 5). Based on an expected effect size of ~0.5 (i.e., at least 50% difference in proportion of elk on non-refuge area between treatments, at an opening date) a sample size of 22 cows per half are required for non-refuge areas. For refuge areas, the behavioral control, collaring of 16 cows per half would allow for comparison of movement dates at 9 days. This also allows for testing the difference of proportion of elk on refuge and non-refuge areas for likely scenarios (slightly more extreme differences), although this leaves slightly less power to detect more subtle differences. Because some cows may move from area to area, or move from non-refuge to refuge before the study period, some extra cows should be collered. If two extra cows are collared on non-refuge areas for contingency during the study, this totals to 40 cows per half (22 nonrefuge + 16 refuge + 2 contingency). Therefore, 80 total collars are required for the study. Testing of alternative hypotheses would be straightforward under this alternative. The forest service data of sheep grazing allotments, woodcutting permits, and recreational use will be evaluated for each half of the study area. If these activities are relatively constant from year to year they will be averaged out. Weather will be assumed to be relatively constant by elevation band between the halves. If there is a marked difference in activity on the two halves between year one and year two, then there a slightly different analysis would be performed. Only elk at distance greater than 1 km from a disturbance would be used in the analysis. A distance of 1 km was chosen since most studies found that elk farther away from disturbances ranging from hikers to logging did not respond to the disturbance (Ward 1976, Hershey and Legee 1982, Wright 1983, Edge and Marcum 1985, Cassirer et a1. 1992). The strength of a crossover design that it blocks for two sources of external variation (GMU and year), allowing for much stronger inference of cause and effect than the same design without a strict crossover (Ratti and Garton 1994), and is more efficient (smaller sample sizes needed) than a randomized design (Ott 1993). Further, the second two factors required to establish a causal relationship of factor A on B would be established with this design. That is, if elk move to refuge areas in the GMUs where hunting is moved forward one week, while remaining in non-refuge areas on the other GMUs, the presence offactor A (hunting) by itself can be inferred to be causing factor B (elk moment to refuge areas). Conversely, if the elk on the late opening GMUs do not move to refuge areas before hunting begins, then it can be inferred that factor B (elk movement to refuge areas) did not occur in the absence of factor A (hunting). �56 Table 5. Sample size (number of collared elk) per half for treatment groups based on estimates of effect size and 9()01o power for primary response variables. Prim nse variable 1- Nwnber of da s between date of movement and opening date within half Effect Size Number of days between 7 16 mean date of movement 10 10 and 14 8 0 date Primary response variable 2 - difference in proportion of elk on non-refuge areas before and after an opening date (early or late) between early and late hunting treatments. Also can be used to evaluate proportion of elk on refuge and nonrefuge (within halves) areas before and after an opening date (early or late). Proportion of elk on non-refuge areas for late hunting treatment area Proportion of elk on non-refuge areas for early hunting treatment area 0.1 0.2 0.3 0.7 0.8 0.9 15 II 8 22 16 11 36 22 15 Primary response variable 3 - date of movement between non-refuge and refuge. Sample Size Effect Size Number of days between 7 22 mean date of movement 10 12 for early and late hunting treatments 14 8 Two weaknesses of this design apply to all alternatives. There may be a learned effect between years that the design cannot determine. For example, elk that were on non-refuge areas and were hunted early the first year, may move early the second year, well before late hunting begins, as a random effect or a learned response. However, because all studies indicate that disturbances have a short term effect on elk responses, this error is assumed to be minimal. The second weakness is the lack of spatial replication. There is only one spatial replicate, and it borders on a pseudoreplicate because it is not randomly chosen. Thus, the results will only be applicable for the GMUs in the study. Alternative two - Close hunting one year on two GMUs(chosen randomly) and issue the fuU number of permits for the other two GMU's. Reverse the treatments the following year. This alternative is essentially identical to alternative one with different treatments on the north and south halves of the study area. Treatment one would be business as usual, with early-season hunting beginning on the last weekend in August on one half. Treatment two would consist of no early-season hunting permits (deer or elk) issued on the other half. There would be no limitation on the number of early-season hunting permits issued, thus the treatment one area could have high hunter densities during the study. GMUs12 and 23, and 24 and 33 would be paired as in alternative one, with the same response variables and testing of alternative hypothesis. Like alternative one, this would be a two-period crossover design (Manly 1992, Ott 1993) with different treatments (Table 6). �57 model for alternative two. Factor B ( Collared Elk iods) 1996 1997 n Early-season hunting No early-season hunting n No earl -season huntin Statistical analysis, sample size calculation, and testing of alternative hypothesis would be the same as for alternative one. The weakness of alternative one are applicable to this alternative, but there are additional logistical and practical problems. Hunters may boycott the White River area because they can't hunt in their traditional areas, or because they fear the crowds on the two areas left open to hunting. There would be numerous hunter complaints about this alternative. Second, the guides and outfitters have their traditional hunting areas defined. Closing of GMUs could cause them time and money in the establishment of new camps. Further, guides, outfitters, and local businesses in general, may have a decrease in their overall early-season revenues, ifhunters boycott the area. Alternative three - Move the early-season hunting forward for two weeks on non-refuge areas only, then close hunting on public land and open itfor the following two weeks on refuge areas onlJ!. Alternative three is a crossover design similar to alternative one (Table 7), but with the experimental units being refuge and non-refuge areas, the observational units being elk, and treatment being hunting or no hunting. However, the number of collared elk would change between the first and second periods, some to all of the elk in the non-refuge areas may move to refuge areas during the second period seriously unbalancing the sample sizes. Analysis of this alternative would be the same as for alternative one for both the main and alternative hypothesis. Table 7. Layout of Latin square crossover design for alternative three. Factor B (periods) Collared Elk First 2 weeks of season Last 2 weeks of season Non-refuge n Hunting No hunting Refuge n No Hunting Hunting Sequence The recommended sample size calculation is the same as for alternative one except area is refuge or non-refuge area instead of each half of study area. Therefore, 22 cows would be collared on refuge areas, and 22 cows would be collared on non-refuge areas (44 total collars). Alternative three suffers from all the drawbacks of alternative one and two, except for hunter displeasure. Additionally, alternative three would be more logistically difficult than either alternative one or two, because the opening and closing of areas would have to be coordinated with public and private landowners. Timing and coordination of closing and opening of areas to hunting is likely to be a logistic nightmare requiring many personnel out in the field coordinating the activity. Almost daily aerial radio tracking of the elk would be required to ensure enough samples for reaction to two perturbations so close in time. Finally, the analysis and inferences may be weakened by unbalanced sample sizes. �58 Study Design Under Ideal Conditions Observational studies and small manipulative experiments cannot predict responses of elk to hunter pressure. This information is only attainable through large-scale field experiments, which due to economic, administrative, and biological factors, often are not adequately replicated. Meta-analysis is one alternative to assist in prediction of widespread or typical responses to hunting (Gurevitch et al. 1992), but the best alternative is to have adequate spatial and temporal controls. Before the experiment is designed, much thought should be given to defining the important responses of interest, that is, responses that managers would like to be able to predict for management of an elk herd. To evaluate movement of elk to refuge areas in response to the opening of early-season hunting, I would use a spatially replicated crossover design, with four treatments on four adjacent areas for each study area. The four treatments would be: Area Area Area Area I 2 3 4 - no hunting, - full hunting pressure at normal time, - full hunting pressure two weeks early, - full hunting pressure two weeks late. This design would take four years, as each treatment would be administered each season. The spatial replicate would be the study area (each with four sub-areas for treatment). The study areas would be chosen by random stratified sampling throughout the state where there is a reasonable amount of elk hunting. I would recommend at least six study areas, this way all possible treatment sequences could be randomly assigned (4! possible sequences of treatments = 24 sequences, or 6 study areas x 4 sequences per study area). The sequences would have to be related in each study area so that each row and column received each treatment as required by the Latin Block design, which the crossover design is based on. However, each study area could have a different sequence to cover every possible sequence and a sequence effect could be eliminated from the analysis. A weakness of this design is that it cannot separate out persistent learned behaviors from the error term. Allowing several years for each treatment or adding a treatment including the import of elk naive to hunting would allow measurement of the learning effect of elk. However, because the literature indicates that disturbance effects tend to be short lived, I would begin with this experiment on the assumption that there will be little persistent behavior year to year. This experiment would not address the issues of effect of density of hunters, cover etc., it would be specifically designed to test for a cause and effect relationship between opening of early-season hunting and movement of elk to refuge areas. Once the relationship between opening of early-season hunting and movement of elk was established, an experiment could be designed to evaluate the effect of other factors on elk movement during early-season hunting. Application of Results The results from this study will falsify or establish a causal relationship between elk movements and earlyseason hunting activity. These results will provide managers with scientifically defensible and publicly credible information that can be used to decrease the early-season elk distribution problems. The conclusions will have direct applicability to the White River area and will also contribute to the body of scientific knowledge on elk in the western United States. Outputs from this research will include written progress reports, scientific publications and spatial use maps. The primary scientific publication would be an article in the Journal of Wildlife Management on 'Elk movements in response to early-season hunting in the White River area'. Offshoots from this project may include an article in the Wildlife Society Bulletin on 'Video methods for viewing animal movement' , or an article about large-scale field experimentation. �59 LITERATURE CITED Adams, A. W. 1982. Migration. Pages 301-322 in 1 W. Thomas and D.E. Toweill, eds., Elk of North America: ecology and management. Stackpole Books, Hanisburg, PA. Altmann, M. 1956. Patterns of herd behavior in free-ranging elk of Wyoming, Cervus canadensis nelsoni. Zoologica. 41:65-71. Boyce, M.S. 1991. Migratory behavior and management of elk (Cervus elaphus). App. Animal Beh. Sc. 29:239-250. Boyd, RJ. 1970. Elk of the White River Plateau, Colorado. Colorado Div. of Game, Fish and Parks. Tech. Publ. No. 25. 126pp. Camp, Dresser and McKee Inc. 1986. Meeker PRLA elk mitigation study: monitoring report Volume 2. Prepared for Consolidation Coal Company. Camp Dresser and McKee Inc., Denver, Colorado. Cassirer, E.F., D.J. Freddy, and E.D. Ables. 1992. Elk responses to disturbances by cross-country skiers in Yellowstone National Park. Wildl. Soc. Bull. 20:375-381. Chapman, P. 1995. SAS power calculation programs. Statistics Department, Colorado State University, Ft. Collins, co. Craighead, 11,G. Atwell, and B.W. O'Gara. 1972. Elk migrations in and near Yellowstone National Park. Wildl. Monogr. 29. 49pp. Czech, B. 1991. Elk behavior in response to human disturbance at Mount St. Helens National Volcanic Monument. App. Animal Beh. Sc. 29:269-277. Doncaster, C. D. 1990. Non-parametric estimates of interaction from radio-tracking data. 1Theor. BioI. 143 :431443. Edge, W.D., and C.L. Marcum. 1985. Movements of elk in reaction to logging disturbances. 49:926-930. 1Wildl. Manage. Freddy, D.l 1987. The White River elk herd: A perspective, 1960-85. Colorado Div. ofWildl. 64pp. Tech. Publ. No. 37. Gray, J. P., G. Byrne, and J. Madison. 1994. White River elk data analysis unit plan: game management units: 11,211,12,13,131,231,23,24,25,26,33. Colorado Div. of Wild I. 45pp. Gurevitch, 1,L. L. Morrow, A. Wallace, and 1 S. Walsh. 1992. A meta-analysis of competition in field experiments. Am. Nat. 140:539-572. Hershey, T.l, and T. A. Leege. 1982. Elk movements and habitat use on a managed forest in north-central Idaho. Idaho Dept. ofFish and Game. Wildl. Bull. No. 10. 23pp. Irwin, L. L., and 1M. Peek. 1979. Relationships between road closures and elk behavior in northern Idaho. Pages 199204 in M. S. Boyce and L. D. Hayden-Wing eds., North American elk: ecology, behavior, and management. University of Wyoming, Laramie. Knight, RR 1970. The Sun River elk herd. Wildl. Monogr. 23. 66pp. Kuck, L., G. L. Hompland, and E.H. Merrill. 1985. Elk calf response to simulated disturbance in southeast Idaho. J. Wildl. Manage. 49:751-757. �60 Lemke, T.O. 1975. Movement and seasonal ranges of the Burdette Creek elk herd, and an investigation of sport hunting. Montana Fish and Game Dept. Job Final Rept., Study No. 32.01, Job No. BG-3.1S. 127pp. Manly, B. F. J. 1992. The design and analysis of research studies. Cambridge University Press, New York. 353pp. Martinka, lC. 1969. Population ecology of summer resident elk in Jackson Hole, Wyoming. J. Wi1dl. Manage. 33:465-481. Minta, S. C. 1992. Tests of spatial and temporal interaction among animals. Ecol. App. 2: 178-188. Morgantini, L. E., and R J. Hudson. 1979. Human distribution and habitat selection by elk. Pages 132-139 in M. S. Boyce and L. D. Hayden-Wing eds., North American elk: ecology, behavior, and management. University of Wyoming, Laramie. Osenberg, C. W., S.J. Holbrook, and R 1Schmitt. 1992. Implications for the design of environmental assessment studies. Pages 75-89 in P. M. Grifinan and S. E. Yoder eds., Perspectives on the Marine Environment. Proc. on the Marine Env. of Southern Calif., Los Angeles, CA. Osenberg, C.W., R J. Schmitt, S. J. Holdbrook, K. E. Abu-Saba, and A. R Flegal. 1994. Detection of environmental impacts: natural variability, effect size, and power analysis. Ecol App. 4: 16-30. Ott, R 1. 1993. An introduction to statistical methods and data analysis. Fourth ed. Wadsworth, Inc., Belmont, CA. 1051pp. Philips, G.E. 1994. Upper Eagle River Valley elk study: draft study plan. Dept. Fishery and Wildl. Bio. Colorado State Univ., Ft. Collins. 26pp. Platt, lR 1964. Strong inference. Science. 146: 347-353. Ratti, 1T., and E. O. Garton. 1994. Research and experimental design. Pages 1-23 in S. Hieb, ed., Research and management techniques for wildlife and habitats, Fifth ed. The Wildl. Soc., Bethesda, Maryland. Rost, G. R 1975. Responses of deer and elk to roads. M.S. Thesis, Colorado State Univ., Ft. Collins. Rudd, W. J., A. L. Ward, andL. L.Irwin. 1983. Do split hunting seasons influence elk migrations from Yellowstone National Park? Wildl. Soc. Bull. 11:328-331. Schultz, RD., and J. A. Bailey. 1978. Responses of national park elk to human activity. J. Wildl. Manage. 42:91-100. Stewart-Oaten, A., and W. W. Murdoch. 1986. Environmental impact assessment: "pseudoreplication" in time? Ecology. 67:929-940. Strohmeyer, D.C., and J. M. Peek. in press. Wapiti home range and movement patterns in a sagebrush desert. Northwest Sci. Ward, A. L. 1976. Elk behavior in relation to timber harvest operations and traffic on Medicine Bow Range in southcentral Wyoming. Pages 32-43 in S. Hieb, ed., Proc. of the Elk--Logging--Roads Symposium. Univ. ofIdaho, Moscow .. Wright, K.1. 1983. Elk movements, habitat use, and the effects of hunting activity on elk behavior near Gunnison, Colorado. M.S. Thesis. Colorado State Univ., Ft. Collins. 200pp. Zahn, H. M. 1974. Seasonal movements of the Burdette Creek elk herd. Montana Fish and Game Dept. Job Final Rept., Study No. 32.01, Job No. BG-3.13. 68pp. �61 Budget - 1996 Item Costs SUbTotals 18kCollars 1m collars Im~ x $200 lcollar $300 Jelk captured $16,000 $24,000 ~ 2 Aighttime Aug-Sept July, Oct-Nov Dec-May ~ davslweek x 1 davslweek x 1 daY/month 12 weeks x 10 weeks x § monthsx ~ hrs/dayx ~ hrs/day x ~ hrs/day x $185 Ihr ~ Ihr ~ Ihr $26,640 ~ ~ ~ ~ Salary Ever-faithful graduate research assistant salary and tuition Volunteer for 2 months intensive data collection Equipment maintenance and office supplies Trawl money for Presentations $103A80 Budget - 1997 18kColiars 10 collars x 10 elk x Item Costs $200 lcollar $300 Jelk captured SubTotals $2,000 $3,000 $5,000 2 Righttime Aug-Sept July, Oct-Nov Dec-May ~ davslweek x 1 davslweek x 1 davtmonth x 12 weeks x 10 weeks x § months x ~ hrs/dayx ~ hrs/dayx ~ hrs/dayx $185 Ihr $185 Ihr $185 Ihr $26,640 ~ ~ ~ ~ Salary Ever-faithful graduate research assistant salary and tuition Volunteer for 2 months intensive data collection Equipment maintenance and office supplies Trawl money for Presentations ��63 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS Colorado State of Project Work No. Plan No. Job No. Period Author: REPORT Covered: Mammals W-153-R-8 Research 3 Elk Investigations 9 Estimating Survival Rates and Developing Techniques Estimate Population Size July of Elk to 1, 1994 - June 30, 1995 D. J. Freddy Personnel: F. Barnes, J. Broderick, G. Byrne, A. Coriell, D. Crane, J. Ellenberger, J. Frothingham, V. Graham, J. Gray, R. Hays, G. Loucks, P. Will, R. Witt CDOW; D. Bowden, C. Vardeman, G. White, CSU; K. Crane, C. McCarty, D. Ouren, volunteers; USFS Rifle, BLM Glenwood Springs, cooperating. ABSTRACT We radio-collared 69 calf elk (Cervus elaphus nelsoni) (6 months old) and an additional 14 adult female elk (~ 1 year old) in December 1994 to estimate survival rates during winter and used these same elk in 103 aerial sighting bias trials to develop models for estimating degree of negative sighting bias when counting elk with a helicopter on sample quadrats. Elk were captured using portable corral traps and helicopter net-gunning. Survival rates (± 95% CI) from December 1994 to June 1995 were 0.90 ± 0.07 for calves and 0.96 ± 0.04 for adult females. Suspected causes of death were malnutrition and predation for calves and shooting and predation for adult females. Calves suspected of dying from malnutrition had marrow fat values <13% at death and body weights <102 kg at capture while calves dying from predation had marrow fat values of 3.7-35.6% and body weights of 86-140kg. For adults ~1 year-old, hunting accounted for 88% of 24 deaths. Average sighting probability of elk groups was 82.3% which equates to a 17.7% negative sighting bias. There were no differences in sighting bias between years (P > 0.70) but there were differences Ln si,g.btingbias among observers (P = 0.06) which ranged from 9.3'% to 22.4.%. Univariate tests indicated·elk age (calf or adult), elk sex (calves only), initial group size, log(ln)' initial group size, total group size, log(ln) total group size, activity. behavior, vegetation type, percent vegetation occlusion cover, observer, navigator, pilot, and trapping method affected the probability of sighting elk (P ~ 06). Multivariate analyses incorporated elk age, In total group size, activity behavior, vegetation type, percent vegetation occlusion cover, percent snow cover, navigator, observer, and year. The best sightability model out of 2,048 possible models was a complex 12 parameter model involving nearly all major variables. We are currently assessing the effects on sighting probability of using less complex models. ��65 ESTIMATING SURVIVAL JOB PROGRESS REPORT RATES OF ELK AND DEVELOPING ESTIMATE POPULATION SIZE David TECHNIQUES TO J. Freddy P. N. OBJECTIVE Estimate estimate survival rates of adult population size. female SEGMENT and calf elk and develop techniques to OBJECTIVES 1. Radio-collar Management 75 calf and 15 adult female elk during Unit 42 south of Rifle, Colorado. 2. Estimate winter and annual survival from known fates of radioed elk. 3. Estimate probability of sighting and counting elk during surveys using radioed elk as a known population. 4. Estimate density system. 5. Analyze survival and sightability Aid Job Progress reports of elk in a portion rates December of calf and adult of GMU-42 using data and summarize 1994 in Game female elk helicopter a quadrat annually sampling in Federal INTRODUCTION Our objectives are to provide reliable estimates of survival rates for calves and adult females during winter and for adult females throughout the year for the period 1993-94 through 1997-98. Additionally, we will develop and test a system for estimating population size that will incorporate estimates of sighting bias in conjunction with a random sampling system using search quadrats as sample units. Our winter study area encompasses about 839 km2 2 (324 mi ) in the eastern half of Game Management Unit 42 south and east of Rifle, Colorado. Elk winter range vegetation types include juniper-pinyon woodland (Juniperus osteosperma-Pinus edulis), oakbrush-mountain shrub (Quercus gambelii-Amelanchier alnifolia), aspen (Populus tremuloides), sagebrush (Artemisia tridentata), and agricultural fields (Freddy 1993, 1994). METHODS Marking We placed radio collars (172-176MRz) having mortality sensors on 69 calves (6 months old), of which 33 were males and 36 were females, and 14 adult females (~ 1 year old). Of these, 65 calves and 2 adults were trapped from 7-11 December 1994 using helicopter net-gunning and 4 calves and 6 adult females were trapped from 13-21 December using portable corral traps. Six adult females were captured in corral traps 2 March 1995 to elucidate movements of elk associated with a specific area of private land. Helicopter capture occurred at 11 remote sites located primarily on public lands while corraltrapping occurred at 3 sites, 2 on private and 1 on public lands. Trapping effort was allocated among 8 geographic trap zones to assure that radioed elk were representative of most if not all segments of the population (Table 1). Radio collars were of the same type used in 1993 (Freddy 1994). �Calves captured by net-gunning were ferried by helicopter to processing points usually within 1.6 km of capture sites. At processing points, body weight, total body length, hind foot length, and rectal body temperature (F) were measured and calves were then radio-collared and released. Similar measurements were also made on calves that were corral-trapped and then released at the trap site. Body measurements for calves were compared between sexes using Proc FREQ, GLM, and REG (SAS 1988). Survival We monitored life or death status of radioed elk during daily ground surveys and aerial surveys conducted at 2-4 week intervals from December 1994 through April 1995 and via monthly aerial surveys from May to November 1994 and May to June 1995. Survival rates (S) of radioed elk were calculated using the binomial estimator with a variance, VAR(S) = S(l-S)/n (White and Garrott 1990) (Proc FREQ, SAS 1988). We chose not to use the staggered entry approach and did not use a Kaplan-Meier approach because few animals were censored (White and Garrott 1990). Life or death status of all calves radioed in December 1994 (68 with functional collars) was known for the period 7 December 1994 through 15 June 1995 (1 male calf collar, 173.949/94, failed 2 weeks post-capture and although the calf was seen alive through 30 January 1995 it was excluded from estimates of survival rates). On 15 June, calves become yearlings for purposes of calculating rates of calf survival. Life or death status for adult females, yearling females, and yearling males collared in December 1993 or 1994 was known for 117 of 118 animals through 15 June 1995. All 6 adult females collared 2 March 1995 were alive as of 15 June 1995 but were not used in calculations of survival for time periods prior to 14 June 1995. Fat content (percent dental cementum were Collins). Sighting dry matter) of femur marrow and estimates of age based on obtained for dead radioed elk (Colo. Div. Wildl. Lab, Ft. Bias Procedures for conducting aerial sighting trials were identical in 1994 and 1995 (Freddy 1994). In 1995, we conducted 103 sighting bias trials that targeted radiocollared elk on 23, 24, 26, 28, and 30 January and 6-8 and 21 February. We again used a Bell-Soloy helicopter and the same observers and navigators as in 1994. The pilot which flew all trials in 1994 flew 77 (75%) of the trials in 1995 with 2 additional pilots flying the remaining 26 trials. Observers indicated that all pilots provided comparable and acceptable service. Sighting bias models were developed using logistic regression (Proc GENMOD, SAS). The dichotomous classification of groups seen or missed was the dependent variable. Initial group size (In), total group size (In), activity behavior, vegetation type, percent occlusion cover, percent snow cover, snow type, year, observer, navigator, trap method, temperature, wind, lighting conditions, pilot, time interval, and time of day were independent variables. Observer, navigator, pilot, trap method, elk sex, elk age, activity, vegetation type, snow type, time interval, time of day, light conditions, and wind were treated as class variables and group size, percent occlusion cover, and percent snow cover were treated as continuous variables. Significance level for independent variables was P ~ 0.06 for univariate tests. Relative efficiency of models using all possible combinations of significant variables was assessed using AIC values (Akaike Information Criterion = -2[-10g likelihood value] + 2[No. model parameters]). Population Estimates Elk population size in GMU 42 from East Alkali Creek to Beaver Creek was estimated during January and February 1995. Three independent estimates of size or density were obtained. On 19 January, a nonrandom elk sex/age �67 classification flight using a helicopter was flown during which numbers of marked (radiocollared) and unmarked elk were counted. From 22-24 February, 40 randomly selected quadrats approximately 1 mi2 (2.59 km2) in size were flown using a helicopter and during these flights numbers of marked and unmarked elk were counted. For these 2 flights, we used a simple Lincoln-Peterson estimator based on ratios of marked and unmarked elk (White and Garrott 1990) to estimate total numbers of elk. The third estimate of population size resulted from projecting the average number of elk counted per sample quadrat to the entire segment of winter range sampled by the quadrat system. The number of elk counted by direct observation at the time of the flights that were on private land areas not surveyed by the 3 flights were added to each of the 3 estimates of population size to arrive at an estimate of total elk in GMU 42 east of Beaver Creek. At this time, no adjustments have been make for sighting bias for counts of elk on quadrats. Movements We continued to locate 37 radioed elk at least once per month since capture to document seasonal movements via telemetry using a Cessna 185. These elk were selected at random from within trap zones and equalized by age class. As of 14 June 1995 these elk were classified as 15 adult females, 7 yearling females, 4 female calves, 7 yearling males, and 4 male calves. During 199394, 6 of 34 elk monitored for locations died. These 6 were replaced at random primarily with 6 month-old calves captured in December 1994 in the same trap zone(s) as those elk that died. During June 1995, we selected an additional 28 adult females at random to document locations during the calving period. As needed, we located other elk to document unusual movements. RESULTS AND DISCUSSION Survival Between 1 December 1993 and 14 June 1995, 37 of 190 radiocollared elk died (Appendix 1). Of these 37, hunting was apparently involved in 57% of the deaths. For adults ~1 year-old, hunting accounted for 88% of 24 deaths. There were 2 periods of mortality during the year. Calves died from February to May while adults died during fall and early winter when hunting seasons occurred (Fig. 1). survival rates (± 95% CI) for calves during winter, 1 December - 14 June, were 0.92 ± 0.06 in 1993-94 and 0.90 ± 0.07 in 1994-95 (Table 3). Sex of dead calves was 4 male and 2 female in 1993-94 and 4 females and 3 males in 1994-95 (Table 2). These survival rates were associated with winters considered mild in temperature and having low or moderate snow depths. In 1994-95, considerable snow fell during March and April but usually melted rapidly at lower elevations. Suspected causes of death for calves during winter were, mountain lion predation (31%), malnutrition (23%), and unknown (46%) (Table 4). For the unknown category, bear predation was suspected once (3/1/95) as was lion predation (3/18/94) (Table 2). Those calves dying from malnutrition had marrow fat values <13% at death and body weights <102 kg at capture while calves likely dying from predation had marrow fat values 3.7-35.6% and body weights 86-140 kg (Table 2). There were no calf mortalities during capture and no evidence that capture procedures directly induced mortality in either year. In 1993-94 the first mortality involved an 82 kg female suspected of dying from malnutrition at 59 days post-capture while in 1994-95 the first mortality was an 86 kg female suspected of dying from mountain lion predation at 72 days post-capture. At capture, this last female had an abnormally articulating front shoulder which may have been injured during or prior to capture but the injury appeared to minimally affect her mobility upon release at capture. �Survival rates (± 95% CI) for adult females during winter, 1 December - 14 June, were 0.96 ± 0.05.in 1993-94 and 0.96 ± 0.04 in 1994-95 (Table 3). Of 7 winter deaths during both years, 5 (71%) were due to legal (1) and illegal (1) hunting or wounding loss (3) during late rifle seasons (Table 4). Natural deaths (2) were attributed to mountain lion predation and unknown but possible lion predation. Of the 3 elk lost to wounding, 2 were suspicious kills. Both of these elk died from 1 rifle shot to the upper neck or head in relatively plain sight near or on an agricultural field where retrieval of the carcass was not difficult. These animals may have been accidentally shot, maliciously shot, or abandoned because of the radiocollar. Survival rates during winter 1994-95 were 1.00 for juvenile males and females age 18-23 months and radiocollared as calves (Table 3). Annual survival rate for adult females, 1 December 1993 - 30 November 1994, was 0.77 ± 0.10 (Table 3). Of the 15 deaths involving adult females marked in December 1993, 13 (87%) were associated with hunting. The 2 natural deaths were due to lion predation during winter and an unusual breech birth that occurred about 1 October 1994. Hunting deaths were attributed to legal hunting (6, fall seasons), illegal hunting (1, late season), wounding loss (5, 4 fall seasons, 1 late season), and presumed hunting (1, fall seasons). We examined all 5 carcasses attributed to wounding loss. Based on locations and physical position of the carcasses, we found no compelling evidence that hunters had likely found the animal and then left the carcass because it was radiocollared. Thus, we do not believe at this time that wounding loss during fall seasons is resulting from hunters being afraid to claim a radiocollared animal. Depending on the type of hunting season and GMU, yearling males with spike antlers are not legal quarry. In general, yearling males are not legal quarry in any part of the study area until the third combined rifle season in late October. In August 1994, there were 32 yearling males alive that were radiocollared as calves. Life/death status of all 32 was determined through 14 June 1995. Of these 32 yearlings, 4 (12.5%) were known or presumed to be illegally taken during hunting seasons. Three carcasses were recovered in the following general condition: shot, eviscerated, and left in the field (1st rifle season); shot, not eviscerated, and left in the field (2nd rifle season); shot, eviscerated, head removed, partially packed-out, and radiocollar buried under debris (3rd rifle season). The telemetry signal of the fourth animal, presumed shot and removed, could not be heard after 4 October 1994. Calf Body Size Male calves had larger body weights (P = 0.001), longer total body length (P = 0.06), longer hind leg lengths (P = 0.06), and higher condition indexes (P = 0.06) than female calves (Tables 5, 6). Body weights and measurements were not different between years for male or female calves (P > 0.30). Predicting body weight from body measurements does not look promising at this time, although all regressions were significant (P < 0.001). The multiple regression using body length and hindfoot length as independent variables provided the best correlation coefficients (~) of 0.59-0.67. Sighting Bias In 1994 and 1995, we conducted 213 sighting trials of which 192 (90%) were successful, and each observer was involved in 70-72 sighting trials during both years (Table 7). Successful trials occurred when the targeted elk was on the assigned flight quadrat whether or not that elk was seen by observers. Unsuccessful trials occurred when the target elk was found off the assigned quadrat immediately subsequent to the time the quadrat was flown by the helicopter team. Elk were off quadrats primarily due to their movements between the time elk were located with the Cessna 185 and the time the helicopter team flew the quadrat. Usually only 1 elk was targeted per quadrat even though several radiocollared elk may have been present on a quadrat to insure that targeted elk were in separate and independent groups. �During the 2 years, 143 different elk were used in 192 successful trials (Table 8). Most elk were targeted only once during the 2 years. Targeted elk included adult females, yearling females, female calves, yearling males (spike-antlered), and male calves. Calves comprised 53-54% of the targeted elk within each year. Yearling males were only available as targets during 1995 (Table 9). We used as many different radiocollared elk as possible to avoid any positive effects on sightability that might have occurred if an observer or navigator had been involved in locating a target elk more than once. Observers were not assigned the same target· elk within the same year, except in 1995 an observer was given the same elk twice but the animal was on 2 different search quadrats. During both years, there were 11 cases where a target elk was given to the same observer for 2 trials. In 7 of these cases, the targeted elk was a calf the first year and a yearling the second year, while in the 4 remaining cases, the target elk was an adult in both years. In all 11 instances, elk were on different search quadrats in both years. During both years, there were 24 cases where a navigator was involved with the same target elk during 2 trials. Only 3 of these cases occurred in the same year and the targeted elk was on a different search quadrat on each occasion. Average sighting probability was 82.3% which equates to a 17.7% negative sighting bias (Table 7). There were no differences in sighting bias between years (P > 0.70) but there were differences in sighting bias among observers (P = 0.06) which ranged from 9.3% to 22.4% (Table 7). Univariate tests indicated elk age (calf or adult), elk sex (calves only) initial group size, log(ln) initial group size, total group size, log(ln) total group size, activity behavior, vegetation type, percent occlusion cover, observer, navigator, pilot, and trapping method affected the probability of sighting elk (P ~ 0.06, Tables 11, 12). Adult females were more sightable than calves (P = 0.03) as were male calves compared to female calves (P = 0.06). Elk caught in corral traps were more sightable than those caught with a helicopter and this effect was primarily associated with adult females (P 0.05), not calves. Multivariate analyses incorporated elk age, ln total group size, activity behavior, vegetation type, percent vegetation occlusion cover, percent snow cover, navigator, observer, and year. According to AIC criteria, the best sightability model out of 2,048 possible models was a complex 12 parameter model involving nearly all major variables. Ln group size, percent vegetation cover, and observer persisted as significant variables as number of parameters was reduced (Table 13). Models incorporating elk age, observer, and navigator are likely not practical for field application. We are currently comparing these 12 best fitting models to assess what effects each model has on predicting sightability of our known population of elk groups. This process will hopefully produce a less complex sighting model. When the same variables are used to build sighting models, results from Idaho (Samuel et ale 1987) and Colorado strongly suggest there are common factors affecting sightability of elk in broadly different landscapes. For coniferous landscapes in Idaho the best elk sightability model was: Y = 1.22 + 1.55 (LnGroupSize) - 0.05(% vegetation cover). For oakbrush landscapes in Colorado, elk sightability was: Y = 1.23 + 1.08 (LnGroupSize) - 0.03(% vegetation cover). Effects on sightability of group size and vegetation cover are shown in Fig. 2. Unfortunately at this time, this 3 parameter model ranked 1,165 out of 2,048 Colorado models. The most efficient 3 parameter model incorporated elk activity instead of vegetation cover (Table 13). Group size is a variable that must be incorporated into any sighting model to correct for the propensity to miss groups containing <7 elk (Fig. 2). Our data indicates that groups of elk observed but not targeted for sighting trials were smaller than our target groups (P =0.02). This indicates our radiocollared elk groups biasly represented all possible groups of elk which �70 likely reflects a trapping bias. This data however, also strongly suggests that the need to correct for group size will be greater in actual surveys. Population Estimates Initial estimates of elk population size ranged from 2,049 to 4,339 and were generally imprecise (Table 10). For mark-resight estimators, the nonrandom flight provided the most precise estimate (± 27%) because more unmarked and marked elk were observed compared to the quadrat flight (± 47% precision). However, both mark-resight flights provided estimates of similar magnitude: 3,810 and 4,339 elk. The lower quadrat estimate of 2,049 elk probably reflects a need to restratify and reallocate sample units. Precision of the quadrat sample estimate can likely be improved by altering strata, increasing sample size, and increasing the quadrats allocated to high density strata. The imprecision of both the quadrat and quadrat mark-resight estimates (+ 47%) indicate more quadrats need to be flown along with refining strata boundaries. count data indicates that major changes are needed in delineating the Garfield High and East Divide Creek low strata. Of the 15 (38%) quadrats where no elk were counted, 8 occurred in these 2 strata. Consideration will be given to assigning each individual quadrat to high and low strata even if quadrats are adjacent to each other. Elk Movements As of 15 April 1995, 22 radioed elk had dispersed to a winter range outside the winter range in GMU 42 where they had originally been trapped in December 1993 (Table 14). Of 35 females collared as calves in December 1993 and recruited into the population as yearlings in June 1994, 8 (23%) dispersed during summer-fall 1994 to another winter range. Likewise, of 32 male calves recruited as yearlings in June 1994, 6 (19%) dispersed during summer-fall to another winter range. Eight (12%) of 65 adult females alive in June 1994 dispersed during summer-fall to another winter range. Seven (32%) of the dispersing elk moved to areas outside of DAU E-14, with 6 of these elk moving into GMU 43 immediately east of DAU E-14 (Table 14). The longest dispersal movement involved an adult female elk (172.480/93) that moved to Beaver Creek west of Gunnison, a straight line distance of about 80 miles. This elk had been neckbanded as a young adult in Beaver Creek in 1991 (GMU 54), then radiocollared in GMU 42 in 1993, and subsequently returned to the drainage where it was originally marked in January 1994 (visual and telemetry location). As of June 1995, this elk had returned to GMU 42. We are currently creating a GIS land database for the area frequented by radiocollared elk. This is a volunteer cooperative effort through the National Biological Service. We believe plots of elk locations and movements using this database will be forthcoming in 1996. CONCLUSIONS We obtained acceptably precise estimates of calf and adult survival rates during winter and recommend continuing our current sampling effort of monitoring 75 radioed calves and >75 radioed adult females in 1995-96. Efforts to measure and develop criteria to adjust for negative bias in counts of elk were promising and we are currently assessing the relative efficiency of several sighting models. We recommend conducting the planned replicate surveys using sample quadrats to estimate elk density from mark-resight estimators and sighting bias correction models in 1995-96. �71 LITERATURE CITED Freddy, D. J. 1993. Estimating survival rates of elk and developing techniques to estimate population size. Colo. Div. Wildl. Game Rep. July: 83-117. Res. Freddy, D. J. 1994. Estimating survival rates of elk and developing techniques to estimate population size. Colo. Div. Wildl. Game Res. Rep. July: 27-42. Samuel, M. D., E. O. Garton, M. W. Schlegel, and R. G. Carson. 1987. Visibility bias during aerial surveys of elk in northcentral Idaho. Wildl. Manage. 51:622-630. SAS Institute Inc. 1988. SAS/STAT Cary, NC. 1028pp. White, User's Guide, 6.03. SAS Institute, G. C., and R. A. Garrott. 1990. Analysis of wildlife data. Academic Press, Inc., San Diego. 383pp. Prepared by ~~~-= __=-~~ David J. Freddy Life/Science Researcher _ Inc., radio-tracking J. �Table 1. capture objectives and numbers of elk radioed in 8 trapzones, December 1993 and December 1994. GMU 42. __Obtec~iYe_s_andelk captured for 1994 are in parentheses. capture Elk Collareda Calves Adult Trap Obiective Total Helio Corral Males Females Females Zone Name 4( 0) Garfield 8( 5) 8( 6) 8( 6) O( 0) 3( 3) 1( 3) A 12 ( 6) 20( 8) 25(17) 19( 7) 6{10) 5( 5) 8( 6) Gibson B 12( 0) Uncle Bob 24(13) 29(13) 29(13) O( 0) 10( 7) 7( 6) C 26(13) 21(11) 21(11) O( 0) 5( 1) 6( 8) 10( 2) West Divide D Hightower 10(10) 17(17) 17(17) O( 0) 4( 9) 3( 8) 10( 0) E O( 0) Middle Mamm 10( 8) 0(13) 0(13) O( 0) O( 8) O( 5) F 8( 8) 6( 0) 6( 0) O( 0) 2( 0) 3( 0) 1 ( 0) West Mamm G 44(21) 35( 6)b O( 0) 35( 6) 7( 0) 9( 0) 19 ( 6) Dry Hollow H 150(86) 141(83) 100(67) 41(16) 36(33) 37(36) 68(14) All Helio = Helicopter net-gunning, Corral = corral-trap. bAll 6 adult females captured 2 March 1995 a Table 2. Causes of mortality and body condition for radioed calf elk in GMU 42, 1 December 1993 - 14 June 1995. Radiocollar Date Estimated Body Marrow Fat Frequency Sex Agea Dead Cause of Death Wt.Ckglb Percent Dry Matter 172.899/93 173.000/93 173.262/93 173.289/93 173.461/93 173.469/93 173.870/94 174.119/94 174.140/94 173.789/94 173.640/94 174.170/94 173.589/95 F F M M M M F M M F F M F 9 10 9 9 8 10 8 9 9 9 9 10 12 mos mos mos mos mos mos mos mos mos mos mos mos mos 3/18/94 4/25/94 3/18/94 3/22/94 2/07/94 4/25/94 2/21/95 3/01/95 3/14/95 3/20/95 3/30/95 4/25/95 6/01/95 Mt.Lion kill Malnutrition Unknown, Predation? Unknown Malnutrition Malnutrition Mt.Lion kill Unknown, Predation? Mt.Lion kill Unknown Unknown Mt.Lion kill Unknown 86 102 108 111 82 77 86 140 112 100 89 140 117 a Approximate age at death, assume 15 June birthdate. b Whole body weight at capture in December 1993 or 1994. c Fat content of either the right or left femur bone marrow at death. d Fat content of lower jaw bone marrow at death. 28.5d 12.6c 28.9d not available 2.1c 2.7c not available 3.7c 35.6c 76.2c not available 17.4c not available ~ �Table 3. Survival rates of radiocollared elk in GMU 42 for different age and sex classes during 4 time periods from 1 December 1993 through 14 June 1995. Survival rates calculated as a simple binomial of elk alive at end of time period divided by elk alive at beginning of time period with a variance of S(1-S)/n. Calves were 6-12 months old and collared at 6 months of age. Yearlings were 12 - 18 months old and juveniles were 18 - 23 months old and both were collared as 6 month old calves in December 1993. Adult females were ~1 year-old when captured or recruited in December. Time Period Age/Sex Class 1 DEC 93 - 14 JUN 94 A8 Db Surv. RateC Calves + F 73 6 15 JUN 94 - 30 NOV 94 A D Surv. Rate 1 DEC 93 - NOV 30 94 A D Surv. Rate 0.92 ± 0.06 1 Dec 94 - 14 JUN 95 A D Surv. Rate 68 7 0.90 + 0.07 28 0 34f 0 1.00 1.00 94 4 0.96 ± 0.04 190 11 M Yearlings Male Female 32 35 4d 1e 0.88 ± 0.12 0.97 ± 0.06 Juveniles Male Female Adult Females 68 3 Totals 141 9 0.96 + 0.05 64g 12h 131 17 0.81 ± 0.15 67 15 67 15 0.77±0.10 8 Number of radiocollared elk alive at beginning of time period. b Number of radiocollared elk dying during time period. C Survival rate + 95% confidence interval. d One elk (173.309/93) disappeared during October hunting seasons and presumed dead. e One elk (172.800/93) disappeared during November hunting seasons and presumed dead. f Yearling females are included in adult female survival rates during this time period. g One elk (172.011/93) censored from calculations because animal was missing. h One elk (172.649/93) disappeared during October hunting seasons and presumed dead. (j �74 Table 4. Causes of deaths in radiocollared elk from GMU 42 between 1 December 1993 and 14 June 1995. Calves were 6-12 months old and collared at 6 months of age. Yearling males and females were 12-18 months old and collared as 6 month old calves. Juvenile males and females were 18-23 months old and collared as 6 month old calves. Adult females were ~1 year-old when collared or recruited in December. Cause of Death Calves Yearl. Males 0 0 6 3 18 0 0 1b 0 0 0 0 0 0 0 1 1c 1 4 3 7 Total 13 4 1 0 0 19 37 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 7 Total 3 4 0 0 0 0 0 0 0 Adult Females Malnutrition Predation-Lion Accident Legal Hunting Archery/Muzzle Rifle Rifle/Late Wounding Loss Archery/Muzzle Rifle Rifle/Late Illegal Hunting Presumed Hunting Unknown Elk 0 0 0 0 Elk AgeLSex Class Yearl. Juv. Juv. Females Males Females 0 0 0 0 0 0 0 3 5 1 7 2 4 1 7 0 0 0 7 1 3 3 was a spike-antlered male that disappeared during October (173.309/93) hunting seasons and presumed dead and illegally shot. b Elk (172.800/93) disappeared during November hunting season and is a presumed hunting mortality. c Elk (172.649/93) disappeared during October hunting seasons and is a presumed hunting mortality. a �Table 5 • Frequency distribution of whole body weights for male and female elk calves trapped in Game .Manaqement Unit 42, December, 1993 and 1994. Percentage per weight class shown in parentheses. Year Sex 70-79 80-89 90-99 Body Weight Class 100-109 110-119 1993 1994 M M 1(2.9) 1(3.0) 1(2.9) 1(3.0) 3(8.6) 2(6.0) 6(17.1) 6(18.2) 12(34.3) 13(39.4) 1993 1994 F F 1(2.9) 2(5.7) 4(11.4) 4(11.4) 8(22.9) 7(20.0) 12(34.3) 10(28.6) 4(11.4) 10(28.6) Body measurements for elk calves 130-139 140-149 Total 10(28.6) 6(18.2) 1(2.9) 2(6.0) 1(2.9) 2(6.0) 35(100) 33(100) 6(17.1) 2( 5.7) 0(0.0) 0(0.0) 0(0.0) 0(0.0) 35(100) 35(100) Table 6. Year Measurement 1993 Body Weight (kg) 112.7 Body Length (cm) 191.2 Hindfoot Length (cm) 56.3 Condition Index 0.59 (Wt./Body Length) 13.7 10.2 2.2 0.05 77.0 164.0 52.0 0.43 141. 0 210.0 61.0 0.67 35 36 36 35 103.8 188.9 54.8 0.55 12.2 9.8 2.1 0.05 76.0 167.0 51.0 0.46 123.0 207.0 59.0 0.64 35 35 34 34 1994 Body Weight (kg) 113.5 Body Length (cm) 190.3 Hindfoot Length (cm) 56.9 Condition Index 0.59 (Wt./Body Length) 14.2 9.2 1.8 0.05 71.0 168.0 50.0 0.42 140.0 204.0 59.0 0.69 33 32 32 32 103.0 186.3 55.1 0.55 13.3 8.8 2.1 0.06 70.0 166.0 50.0 0.42 128.0 207.0 59.0 0.65 35 36 36 35 Mean trapped (kg} 120-129 in Game Management Male Calves SO Min Max n Unit 42£ December £ 1993£ 1994. Mean Female Calves SO Min Max n 01 �76 Table 7. Summary and 1995. Observer Yr Broderick 94 95 94-95 94 95 94-95 94 95 94-95 94 95 94-95 Ellenberger Masden Totals of elk sighting Trials AttemEted bias trials Trials Not Useable 29 42 71 36 36 72 45 25 70 110 103 213 Trials Useable 3 7 10 4 1 5 4 2 6 11 10 21 in GMU 42 during conducted 26(90%) 35(90%) 61(90%) 32(89%) 35(97%) 67(93%) 41(91%) 23(92%) 64(91%) 99(90%) 93(90%) 192(90%) Target Elk Detected Target Elk Not Detected 22 27 49 25 27 49 36 22 58 83 76 159 4(15.4%) 8(22.9%) 12(19.&%) 7(21.9%) 8(22.9%) 12(19.7%) 5(12.2%) 1( 4.3%) 6( 9.3%) 17(17.2%) 17(18.3%) 34(17.7%) Table 8. Numbers and frequency of individual elk targeted during sighting bias trials in GMU 42, 1994 and 1995. No. Elk No. Elk No. Elk No. Used in Total Used in Used in Elk Trials 1 Trial 2 Trials 3 Trials Year Used 1994 1995 1994-95 77 90 143 67 87 101 0 0 7 16 3 35 Table 9. Age and sex of elk targeted GMU 42 1994 and 1995. Female Yearling Calf Year Adult 1994 39 (39%) 1995 20(21%) 1994-95 59 (31%) 6( 6%) 28(28%) 10(11%) 26(28%) 16( 8%) 54(28%) Table 10. Summary February, 1995. of population during Male Yearling estimates 1/19/95 2/24/95 2/24/95 Method Mark-Resight NonRandom Flight Mark-Resight Random Quadrats Flight Random Quadrats Flight sighting Calf bias trials Total 26(26%) 22(24%) 48(25%) 99(100%) 93(100%) 192(100%) for elk in GMU 42, January and Private Land Elkb Total Estimate +399 3,810 Densit Elk/mi1 C 14 (267 mi2) 3,744±47% +565 4,339 19 (234 mi2) 1,484±47% +565 2,049 9 (234 mi2) Estimate Date successful 99 93 192 successful o ( 0%) 15(16%) 15( 8%) 1994 + 95% cr3,411±27% a Number of elk estimated in survey area actually flown. b Elk directly counted on private land area not included in survey flight. C Density for area represented by survey flight area plus private land area. in �Table 11. Elk sightability survey results 1994 and 1995. January-March, Variable Group Size 1 2 3 4 5 6 7+ Vege. Type Clearing/Ag Riparian Sagebrush Oakbrush Pinyon-Juniper Aspen Tall Conifer occl. Cover(%) 0-19 20-39 40-59 60-79 80-100 a V-visibility No. Grou:Qs Seen Missed 12 9 2 5 0 1 4 0 0 1 14 9 7 2 2 11 10 6 4 13 15 21 14 15 12 69 13 1 6 89 42 7 1 35 60 43 18 3 (groups seen 7 Va 0.52 0.63 0.91 0.74 1.00 0.92 0.94 1.00 1.00 0.86 0.86 0.82 0.50 0.33 0.95 0.85 0.81 0.75 0.43 groups by major independent Variable Behavior Bedded Standing Moving Navigator GB VG FB Observer JB JE DM Elk Age Adult Female Calf (M + F) Yearling Calf Sex Male Female Trap (All elk) Corral Trap Helicopter Trap (calves) Corral Trap Helicopter variables No. Grou:Qs Missed Seen va from Game Management Variable 9 4 20 6 41 112 0.40 0.91 0.85 Snow Cover 0-19 20-49 50-99 100 22 11 0 68 81 10 0.76 0.88 1.00 12 15 6 49 52 58 4 21 8 6 15 Missed Unit 42, Seen va (%) 3 5 5 20 9 9 47 94 0.75 0.64 0.90 0.82 Snow Type Fresh Old 5 28 35 124 0.88 0.82 0.80 0.78 0.91 Wind Light Mode.rate Strong 32 1 0 137 20 2 0.81 0.95 1.00 55 81 23 0.93 0.79 0.74 Light Bright Dull Hazy 22 3 8 97 24 38 0.82 0.89 0.83 42 39 0.88 0.72 Pilot ET BB PP 32 0 1 139 10 10 0.81 1.00 0.91 Time AM PM 24 61 98 0.87 0.8.0 4 29 45 114 0.92 0.80 3 18 14 67 0.82 0.78 9 seen plus groups missed). ....• ....• �78 Table 12. Variables tested in univariate tests using logistic regression for elk sightability trials, Game Management Unit 42, January-March, 1994 and 1995. Significance level at P < 0.06. Significant Variable Date of Trial Sex of Elk Calves Age of Elk Initial Group Size Log(ln) Initial Group Size Total Group Size Log(ln) Total Group Size Elk Activity Vegetation Type Occlusion Cover % Observer. Navigator Pilot Snow Cover % Snow Type Temperature Wind Conditions Light Conditions Time Interval Time of Day Trap Type No Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes No No No No No No No Yes Sign. Year Effect Sign. Year Interaction No No No No No No No No No No No No Yes No No No No No No No No Yes No No No No No No No No No No Yes No No No No No No Yes Table 13. The best fitting sightability model at each level of 1 to 12 parameters according to Akaike Information Criterion (AIC). Rank is the relative rank of each model among 2,048 models which used variables in all possible combinations. "Yes" denotes variables used in models. variables In Models Ln No. Group Elk Vege. %Vege. %Snow Elk ParamAIC Size Act. Type Cover Cover Nav. Obs. Age Year eters Rank 1 2 7 4 48 61 152 463 869 1019 1403 1890 2028 12 11 10 9 8 7 6 5 4 3 2 2 18 Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes The one parameter model includes only the intercept average sightability of elk which was 0.823. 8 Yes which equates to the �79 Table 14. Radiocollared elk captured in GMU 42 during Oecenber 1993 and thought to have dispersed out of GMU 42 to a new winter range as of 15 Aeril 1995. Locations ~GMUs2 determined by aerial telemetr~. Trap New AgeA Sex GMU Zone Location Oescrietion Fr!19uenc~ 3+ F 43 Fourmile Ck (Loc.Elk) 1n.207/93 0 3+ F 421,42 Beaver-Buzzard Cks, Alkali Ck. (Loc.Elk) 1n.219/93 0 5+ East Muddy Ck 1n.249/93 C F 521 2+ Middle Thompson Ck 1n.308/93 0 F 43 3+ 1n.480/93 H F 54 Beaver Ck, Gunnisonb 5+ 1n.459/93 F 521 N.Fk.Gunnison, Near Paonia E 2+ 1n.509/93 H F 43 Roaring Fk R., Glenwood Sprgs. (Loc.Elk) 1+ 1n.830/93 B F 421 Hawxhurst Ck 1+ 1n.849/93 A F 43 Threemile Ck, Roaring Fk. River 1+ 1n.921/93 C F 521 East Muddy Ck, Paonia Reservoir 1+ 1n.929/93 C F 43 Redstone; Porcupine Ck 1+ 1n.961/93 F 521 East Muddy Ck, Paonia Reservoir 0 1+ 173.010/93 G F 421 Hawxhurst Ck, Brush Ck 1+ 173.020/93 F 421 Hawxhurst Ck, Brush CK G 2+ 173.070/93 E F 42,West Parachute;Monument Gulch 1+ 173.090/93 F H 43 Roaring Fk River, Crystal Ranch 1+ 173.279/93 C M 521 East Muddy Ck, Paonia Reservoir 1+ 173.300/93 C M 521 East Muddy Ck 1+ 173.332/93 0 M 421 Clover Gulch, Hawxhurst Ck 1+ 173.370/93 E M 421 Plateau Ck/Salt Ck 1+ 173.390/93 0 M 521 West Muddy Ck, Pilot Knob 1+ 173.450/93 421 Kimball Ck, Wallace Ck H M b Age of elk in years as of April 1995 This elk returned to GMU 42 during June 1995, a movement distance of about 80 airline miles. Appendix t , Mortalities of radiocollared elk from 1 December 1993 - 14 June 1995. Age is approximate age in years of elk at death; C=calf 6-12 months old, Y=yearling 12-18 months old. Body weight measured in December when elk caetured as calves. Frequency 10/ Body Date Trae Site Zone Sex Age Year Caetured Heard Dead Cause of Death Wt.{kg2 1n.039/93 GR B 06/14/95 Undetermined mortality F 2+ Legal harvest rifle season 1n.080/93 GM A F 5+ 11/05/94 Wounding loss late rifle season 1n.090/93 GR B F 5+ 01/16/94 Legal harvest rifle season 1n.160/93 CC C F 2+ 10/23/94 Wounding loss rifle season MC F 2+ 11/03/94 1n.181/93 C Wounding loss rifle season 1n.201/93 GS C F 2+ 11/15/94 Legal harvest archery/muzzle season 10/04/94 1n.258/93 HY C F 2+ Wounding loss archery/muzzle season 1n.277/93 GS C F 2+ 10/04/94 Legal harvest rifle season 1n.290/93 SG D F 5+ 10/23/94 Legal harvest archery/muzzle season 1n.369/93 AC E F 2+ 09/18/94 Wounding loss late rifle season 172.409/93 AC E F 5+ 12/29/94 Mountain lion predation 1n.542/93 FS H F 9+ 02/01/94 Wounding loss rifle season 1n.570/93 PG H F 9+ 11/03/94 Legal harvest rifle season 1n.581/93 PG H F 2+ 10/17/94 H Accident, breech birth of calf 1n.610/93 PG F 2+ 10/04/94 1n.649/93 11/15/94 Disappear rifle season PG H F Y+ 01/16/95 Legal harvest late rifle season 1n.670/93 PG H F 2+ Wounding loss late rifle season 1n.678/93 PG H F 9+ 12/22/94 Illegal harvest, poached 1n.690/93 FM B F 5+ 01/24/94 Disappear rifle season 1n.800/93 SR B F Y+ 11/15/94 F C 86.0 03/18/94 Mountain lion predation 1n.899/93 BC C WM 04/25/94 Malnutrition 173.000/93 G F C 102.0 111.0 Illegal harvest rifle season 173.190/93 BC C M Y 10/29/94 Illegal harvest rifle season M Y 105.0 11/15/94 173.232/93 BR A Undetermined mortality 173.262/93 BC C M C 108.0 03/18/94 111.0 Undetermined mortality 173.289/93 MC C M C 03/22/94 HY C M Y 124.0 11/15/94 Disappear rifle season 173.309/93 Illegal harvest rifle season 173.320/93 HY C M Y 118.0 10/20/94 173.461/93 PG H M C 82.0 02/07/94 Malnutrition 173.469193 FS H M C 77.0 04/25/94 Malnutrition 06/01/95 Undetermined mortality 173.589/94 OG B F C 117.0 173.640/94 Undetermined mortality MC C F C 89.0 03/30/95 03/20/95 Undetermined mortality 173.789/94 MR E F C 100.0 86.0 Mountain lion predation 173.870/94 F C 02/21/95 SM 0 M 140.0 03/01/95 Undetermined mortality 174.119/94 MM F C MM F M C 112.0 03/14/95 Mountain lion predation 174.140/94 Mountain lion predation M C 140.0 04/25/95 174.170/94 FM B ��81 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS Colorado State of project REPORT No. ~W~-~1~5~3_-~R~-~8~ _ Mammals Research Work Plan No 4 Moose Job No. 1 Development of census methods and determination of movements, habitat selection, and degree of calf mortality of moose in North Central Colorado. Period Author: Covered: July Investigations 1, 1994 - June 30, 1995 R. C. Kufeld Personnel: D. Bowden, D. Younkin. ABSTRACT Instrumented moose captured during 1991, 1992, 1993, and 1994 were located at approximately 2-week intervals from January 1992 through June 1994. Four papers concerning moose were published during the past year. Three were in Alces Vol. 30, and one is in press in the Journal of Wildife Management. ��83 DEVELOPMENT OF CENSUS METHODS AND DETERMINATION OF MOVEMENTS, HABITAT SELECTION, AND DEGREE OF CALF MORTALITY OF MOOSE IN NORTH CENTRAL COLORADO Roland P. N. C. Kufeld OBJECTIVES 1. To determine the proportion of moose counting moose in North Park. 2. To determine 3. To determine the degree of dispersal of young animals, and seasonal movements, home range size, and habitat selection of North Park moose. the extent of moose actually observed calf mortality when aerially in late winter. SEGMENT OBJECTIVES 1. To determine the extent of moose calf mortality in late winter. 2. To determine the degree of dispersal of young animals, and seasonal movements, home range size, and habitat selection of North Park moose. STUDY AREA The study area was described by Kufeld (1992). METHODS AND MATERIALS Instrumented moose captured during 1991, 1992, 1993, and 1994 (Kufeld 1992, 93, 94) were located at approximately 2-week intervals from January, 1992, through June, 1995, and plans call for such monitoring to continue for at least 0.6 more years. Most locations were made by aerial telemetry using a Cessna 185 aircraft with a 2 element, "H" configuration receiving antenna mounted on each strut. A switchbox permitted the telemetry operator, a passenger in the aircraft, to operate antennas jointly or separately. Some locations were made by tracking on the ground until the animal was observed when the airplane was not available. Moose locations were plotted on USGS 1:50,000 scale maps and recorded by UTM coordinates. Vegetation type was also recorded for each moose location. RESULTS Moose monitoring Analysis of data for movements, home range tagged moose will be presented in a future moose is completed. size, and habitat use for all report when periodic monitoring of �The following year. four papers concerning moose were published during the past Bowden, D. C., and R. C. Kufeld. 1995. Generalized mark-sight population size estimation applied to Colorado moose. J. Wildl. Manage. In Press. Kufeld, R. C. 1994. Neck circumference of Shiras calves during winter. Alces 30:63-64. Kufeld, R. C. 1994. 30:41-44. status and management of moose Olterman, J. H., D. W. Kenvin, and R. C. Kufeld. southwestern Colorado. Alces 30:1-8. LITERATURE moose (Alces alces in Colorado. 1994. Moose shirasi) Alces transplant to CITED Kufeld, R. C. 1992. Development of census methods and determination of movements, habitat selection, and degree of calf mortality of moose in North Central Colorado. Colo. Div. Wildl. Wildl. Res. Rep. July:95108. Kufeld, R. C. 1993. Development of census methods and determination of movements, habitat selection, and degree of calf mortality of moose in North Central Colorado. Colo. Div. Wildl. Wildl. Res. Rep. July: (119124). Kufeld, R. C. 1994. Development of census methods and determination of movements, habitat selection, and degree of calf mortality of moose in North Central Colorado. Colo. Div. Wi1dl. Wildl. Res. Rep. July: (43- 47). Prepared by __~~~ ~~~ Roland C. Kufeld LS Res/Sci III. _ �Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS State of Project REPORT Colorado No. Mammals W-153-R-7 Research Work Plan No. SP1 Deer Job No. 1 Regulation of Mule Deer Population Growth by Fertility Control: Laboratory, Field, and Simulation Experiments Period Covered: Authors: Personnel: Investigations July 1, 1994 - June 30, 1995 Dan L. Baker and M. W. Miller M. A. Wild, T. M. Nett, D. J. Kessler, McCarty, J. Griess, M. Lockhart P. E. Bleicher, C. W. ABSTRACT The Rocky Mountain Arsenal has the potential to become one of the premier wildlife viewing areas in the Nation. However, realizing that potential depends on wise management. In particular, the mule deer population contained within the boundary fence must be regulated in balance with the resources the area offers. Contraception offers a potential alternative to recreational hunting and professional culling to achieve this objective. Of the techniques potentially available for reducing reproduction in wild deer, conjugates of gonadotropin releasing hormone (GnRH) analog and cytotoxins appear to be the most promising noninvasive method for providing lasting infertility after a single treatment. Research is currently underway to evaluate the efficacy of a GnRH-cytotoxin conjugate in reducing fertility in mule deer and other wild ungulates. In previous experiments, we determined the minimum effective dose of GnRH analog required for contraception in mule deer. During 1994-1995, we attempted to treat. female mule deer with GnRH-toxin conjugate but initial in vitro laboratory experiments using various cytotoxins were unsuccessful. Thus, in vivo experiments with mule deer are delayed until further evaluation and testing of other toxins is completed. For some management applications, remote delivery of contraceptives via projectile syringe or ballistic implant may be required. We conducted pilot experiments to evaluate intramuscular delivery and ballistic implant. Initial results are promising but many unanswered questions must be addressed before this technology is an acceptable method of administering contraceptives. Before contraceptive treatments can be allocated to wild deer at the Rocky Mountain Arsenal, basic knowledge of the reproductive and genetic characteristics of this deer population are needed. These studies are currently in progress. Initial results of reproductive studies estimate average conception date to be about November 23 with 95% of adult females bred between November 19 and November 27. Pregnancy rate across all females sampled was 93.6%; fetal rate 1.75 fetuses/doe. Finally, we attempted to evaluate fecal progesterone concentrations as a method for diagnosing pregnancy. Although there was considerable individual variation in fecal progesterone levels, this technique appears to be a potentially reliable method of pregnancy determination for free-ranging mule deer. ��87 REGULATION OF MULE DEER POPULATION GROWTH BY FERTILITY LABORATORY, FIELD, AND SIMULATION EXPERIMENTS CONTROL: Dan L. Baker and M. W. Miller P. N. OBJECTIVES 1. To develop a practical and acceptable method populations using GnRH-toxin conjugates. 2. To demonstrate the feasibility the Rocky Mountain Arsenal. 3. To predict population simulation modeling. for controlling mule deer of such control in a field application impacts of alternative contraceptive at regimes using SEGMENT OBJECTIVES 1. Evaluate the effectiveness and duration of a single dose application GnRH-toxin conjugate to prevent normal production of reproductive hormones in captive mule deer. 2. Evaluate a remote delivery system for administering GnRH-toxin to captive mule deer. 3. Describe Mountain the reproductive and genetic characteristics Arsenal mule deer population. an effective of dose of. of the Rocky INTRODUCTION The Rocky Mountain Arsenal, located in close proximity to Denver, Colorado, offers an exceptional opportunity for an urban population to view and enjoy Colorado's wildlife. However, the Arsenal presents unique problems as well as unique opportunities. For reasons of security, the perimeter of the area was fenced in 1990. While this fence does not impede the usual movements of birds and small mammals, it does create an unnatural barrier to the movements of ungulates, particularly mule deer (Odocoileus hemionus). Before the Arsenal was fenced, mule deer numbers were held at a relatively constant equilibrium of about 300 animals. As a result of preventing those movements, deer numbers within the Arsenal will certainly rise exponentially during the next decade. Experience with enclosed deer populations elsewhere has shown that such increases will lead to degradation of habitat, widespread starvation, and eventually to catastrophic declines in animal numbers. The only way to prevent this outcome is to control the abundance of the enclosed deer population. This is usually done by public hunting or by professional culling of animals. However, public hunting cannot be allowed on the Arsenal because of concerns for security. Moreover, although professional culling would eliminate excess animals, it would also reduce the value of the deer population as a watchable resource. In this situation, alternatives to hunting as a means of regulating ungulate numbers are needed. Fertility control offers a viable alternative to hunting as a means of population control when hunting is infeasible. However, current fertility control technology does not provide a means of controlling ungulate numbers practically and economically. Here, we propose to develop a practical and economical method of fertility control in mammalian wildlife that overcomes many of the shortcomings of current technology, particularly problems of treatment duration and environmental safety. We propose to use conjugates of gonadotropin-releasing hormone (GnRH) and cellular toxins to selectively �destroy gonadotropin-producing cells in the anterior pituitary preventing gamete production by the ovaries and testes. gland, thereby This research project consists of laboratory, field, and modeling phases, each of which is designed to address questions that must be answered before widespread application of hormonal-toxin conjugates is possible. Details of the experiments to be conducted in each of these phases are described by Baker (1992). In this report, we discuss a) results of laboratory studies with captive mule deer to determine the most effective dose of GnRH-toxin conjugate, b) a pilot experiment to evaluate the effectiveness of a delivery system for remotely administering contraceptives, and c) reproductive and genetic characteristics of the Rocky Mountain Arsenal mule deer population. A. EVALUATION OF SINGLE DOSE APPLICATION OF GNRH-TOXIN CONJUGATE One of the most promising new approaches to contraception involves linking synthetic analogs of gonadotropin-releasing hormone (GnRH) to cytotoxins. GnRH is a molecule produced in the hypothalamus of the brain. It directs specific cells in the pituitary gland to synthesize and secrete two important reproductive hormones; follicle stimulating hormone (FSH) and luteinizing hormone (LH). These latter two hormones, known as gonadotropins, control proper functioning of the ovaries in the female and testes in the male. By coupling a superactive analog of GnRH to a cytotoxin, it should be possible to specifically target that toxin to LH and FSH-secreting cells in the anterior pituitary gland. Of the techniques potentially available for reducing reproduction in wild deer, conjugates of gonadotropin releasing hormone (GnRH) analog and cytotoxins appear to be the most likely noninvasive method for providing lasting infertility after a single treatment (Baker et ale 1993, Nett et ale 1993). Consequently, research is underway to evaluate the efficacy of a GnRHcytotoxin conjugate in reducing fertility in mule deer (Baker 1994). During 1994-1995, in vi~ro laboratory experiments using conjugates of GnRH analogs and either a diphtheria or plant toxin were moderately successful in reducing LH secretion from the pituitary but neither conjugate maintained its effectiveness over time. Therefore, in vivo experiments using mice, domestic sheep and mule deer are pending until laboratory evaluation and testing of other toxins has been completed. B. EVALUATION CONJUGATES OF A REMOTE DELIVERY SYSTEM FOR ADMINISTERING TO CAPTIVE MULE DEER GNRH-TOXIN INTRODUCTION GnRH-cytotoxin conjugates disrupt reproduction in a variety of mammalian species by binding to and destroying pituitary gonadotrophs; as a result, luteinizing hormone (LH) secretion ceases and ovulation cannot occur (in males, lack of LH suppresses testosterone secretion) (T.M. Nett, unpublished data). Because virtually all pituitary gonadotrophs must be destroyed to effect infertility using this approach, determining the amount of GnRH analog needed to bind and stimulate at least one receptor on each pituitary gonadotroph is prerequisite to estimating an effective GnRH-cytotoxin conjugate dosage in mule deer (Nett et ale 1993, Baker 1994). Experiments to determine a minimum effective GnRH analog dosage that elicits maximum serum LH concentrations (presumably by stimulating virtually all pituitary gonadotrophs) revealed that ~ 2 ~g GnRH analog/50 kg body mass (BM), delivered intravenously (IV), induced equivalent maximum serum LH responses in mule deer does; however, the magnitude of those responses varied widely among �89 individuals 1994). and across different reproductive states (Nett et ale 1993, Baker Based on these experiments, GnRH-cytotoxin conjugate doses ~ 2 ~g/50 kg BM IV should be sufficient to cause infertility in mule deer does. However, the need for IV administration will significantly limit the practicality of using GnRH-cytotoxin conjugates in managing a free-ranging deer population because animals must be captured, handled, and treated individually. Although GnRH analog-induced LH secretion curves are essentially the same regardless of whether analog is delivered IV or intramuscularly (1M), reliance on 1M hand injections will place equally severe limits on using GnRH-cytotoxin conjugates in free-ranging settings. It follows that the ability to remotely deliver GnRH-cytotoxin conjugates to wild deer will be necessary before long-term application to population management is practical, and that estimates of effective dosage should anticipate and accommodate potential influences of delivery via projectile syringe or ballistic implant on the pharmacokinetics of GnRH-cytotoxin conjugates. Here, we describe a pilot experiment comparing LH secretion patterns in mule deer does stimulated by GnRH analog delivered intramuscularly via syringe injection and ballistic implant. METHODS AND MATERIALS We used captive hand-raised pregnant adult (~ 2.5 yrs old) mule deer does (n = 6) in this pilot experiment. All does were housed at the CDOW's Foothills Wildlife Research Facility (FWRF) throughout the study; they resided together in a 7 ha pasture before and between treatment/sampling periods, and were held in pairs in 50 ref isolation pens during the two 24 hr treatment/sampling periods. Alfalfa hay and a pelleted high-energy supplement was provided as prescribed under established feeding protocols for mule deer throughout the study; fresh water and mineralized salt were provided ad libi~um. Health of each doe was evaluated daily throughout the study, and does were weighed immediately prior to each treatment/sampling period. We compared LH secretion patterns stimulated by GnRH analog delivered intramuscularly via syringe and ballistic implant in a blocked, crossover experiment with a repeated measures structure. Does were randomly assigned to syringe or ballistic implant treatment groups (n = 3 does/treatment) for the initial 24 hr treatment/sampling period; we then switched treatment assignments for the second period conducted 6 days later. For each treatment/sampling period, we sedated does with 40-80 mg xylazine HCL injected 1M. Once sedated, each doe was fitted nonsurgically with an indwelling jugular catheter. After all 6 does were catheterized, we collected 5 Ml of blood from each (pretreatment). We then partially reversed sedation by administering 10 mg yohimbine HCL IV. Once standing, the upper hind leg of each doe targeted for ballistic implantation was covered with a transparent, adhesive-backed film. Each doe then received 12 ~g GnRH analog (about 8.5 ~g/50 kg BM), delivered in one of two forms: 1) via 1 Ml of a 12 ~g GnRH analog/Ml solution, injected 1M by hand, or 2) via a soluble ballistic implant ("biobullet") carrying 12 ~g GnRH analog and about 100 mg lact~se powder, implanted 1M using an air-powered delivery system (BallistiVet Implant System). Analog from a single batch was used in making both solutions and ballistic implants. All does received GnRH in the right hind leg during the first treatment/sampling period and in the left hind leg during the second. Immediately after delivery and periodically thereafter, we examined and palpated ballistic implant entry sites to ensure delivery was achieved and that bleeding or injury at entry sites was not excessive. After all sampling had been completed, we administered 3 X 106 U penicillin G benzathine/penicillin G procaine injected subcutaneously, removed catheters, and returned does to their 7 ha pasture. �In addition to pretreatment (time 0) samples, we collected about 5 Hl of blood from each doe at 1.5, 3, 4.5, 6, 7.5, 9, 10.5, 12, 15, 18, 21, and 24 hrs after administering GnRH. All blood samples were held for 6-12 hrs at 4 C, then centrifuged. Serum was collected and stored at -20 C until analyzed. Serum concentrations of LH fng/Ml) was measured using an ovine LH radioimmunoassay (Niswender et al. 1969) previously validated for use in mule deer. To date, data have not been analyzed. In doing so, however, we plan to compare 1) maximum LH responses (highest LH concentration achieved after treatment, minus pretreatment concentration), 2) time intervals for reaching maximum serum LH concentrations, and 3) total 24 hr LH secretion (estimated by calculating the area under the LH curve) (Abramowitz and Stegun 1968) stimulated by GnRH analog delivered intramuscularly via syringe injection and ballistic implant. Data will be analyzed using least squares ANOVA for General Linear Models (Freund et al. 1986) and the SAS Interactive Matrix Language. Responses to treatments will be analyzed with two-way factorial analysis of variance for a randomized complete block design with a repeated measures structure. We will use delivery approaches as treatments and individual animals as blocks; factors in the analysis will be treatment and time. The animal within treatment variance will be used as the error term in testing for treatment effects. Time will be treated as a within subject effect using a multivariate approach to repeated measures (Morrison 1976). In addition, we will use a priori orthogonal contrasts to test for differences among individuals (Miller 1966). RESULTS AND DISCUSSION Because data have not been analyzed, results discussed here are preliminary and largely descriptive. Neither ballistic delivery nor hand-injection of GnRH had any apparent acute or chronic effect on health of does used in this pilot study. In general, ballistic implant sites were minimally traumatized and, in 5 of 6 cases, were difficult to detect without extensive examination. Does reacted minimally to being shot with implants and mild, transient «15 sec) lameness was the only observable clinical effect. All implants appeared to penetrate the haircoat and skin and embed >1 cm deep in the semimembranosus or semitendinosus muscles. In one doe (E91), the implant apparently struck the caudal margin of the semitendinosus muscle and passed through the muscle mass, lodging in subcutaneous tissues at the lateral margin of the perineum; otherwise, implants were not seen or palpated. In addition, we observed no residue on the adhesive-backed overlay that might indicate contents had been released from the rear of implants on impact. Serum LH responses stimulated by l2 ~g GnRH delivered via ballistic implants appeared to vary widely among individual does as compared to responses stimulated by the same dose delivered via syringe injection (Fig. 1). In 2 does (D92, W92), we observed no measurable response to implant-delivered GnRH; a delayed and somewhat protracted response was observed in a third doe (E91) whose implant lodged in subcutaneous tissue. For the 3 other does (A91, S90, Y92), LH responses were comparable between delivery systems, although the onsets of their responses to implant-delivered GnRH were delayed about 1.5-6 hrs compared to responses to hand-injected GnRH. In 3 of the 4 does responding to both treatments, maximum serum LH concentrations stimulated by hand-injected GnRH exceeded maximum responses stimulated by implant delivered GnRH by ~50%. Based on our observations during the study, we cannot explain the failure of 2 does to respond to GnRH delivered via ballistic implant. It is possible implants never entered muscle on these 2 does or were extruded before GnRH was released; however, implants penetrated the skin of both does and we did not see or feel casings or residue in their hair, under the adhesive film, or on the ground. Alternatively, GnRH could have inadvertently been omitted or removed from implants during their assembly, or release could have been • �91 delayed or precluded by impeded breakdown of implants and/or local binding, destruction, or malabsorption of GnRH analog. Regardless of cause, additional evaluation and refinement will clearly be needed before ballistic implants can be substituted for syringe injections in delivering GnRH analogs or conjugates in future fertility control research or management applications. C. REPRODUCTIVE AND GENETIC CHARACTERISTICS MOUNTAIN ARSENAL OF FEMALE MULE DEER AT THE ROCKY INTRODUCTION Applying GnRH-toxin conjugates to control the growth of deer populations at the Rocky Mountain Arsenal will require that wildlife managers choose specific tactics for treating animals. Choices must be made on the number and age to treat, the frequency of treatment, timing of treatments and so on. We will provide support for these decisions by developing an interactive model of mule deer population dynamics. This model will combine knowledge of deer biology with an understanding of the constraints intrinsic in the GnRH-toxin conjugate technique. Fundamental to the insights offered by this model is knowledge of the reproductive and genetic characteristics of the Rocky Mountain Arsenal mule deer population. Here, we present results of reproductive stUdies conducted during 1994. The objectives of this study were 1) estimate conception dates and breeding season of female mule deer in order to determine optimum time of delivery of contraceptives 2) estimate fetal rates of female mule deer in order to increase reliability of model predictions of population growth rates 3) evaluate fecal progesterone levels in female mule deer as a potential noninvasive technique for determination of pregnancy following contraceptive treatments 4) collect the anterior pituitary gland and hypothalamus from pregnant females in order to more precisely estimate dosage requirements for contraception 5) collect tissue samples to evaluate genetic diversity of male and female mule deer populations. METHODS AND MATERIALS Collection of Deer Approximately 10 female mule deer were collected each month at 2-3 week intervals from November ~, 1993 to April 19, 1994. Specific collection dates and sample sizes were: November 10 (n = 10), November 19 (n = 10), December 8 (n 10), December 29 (n 11), January 25 (n 10), February 16 (n 10), March 9 (n = 9), April 19 (n = 9). A total of 80 females were examined during this period. Animals were shot through the spine in the cervical region; all died within 5 minutes; no wounding loss occurred. = Estimation = = = of Age The left incisor tooth was collected from each deer. Deer with deciduous dentition were assigned an age using the tooth replacement chronology of Robinette et al. (1957). Deer with permanent dentition were assigned an age from counts of dental cementum in the first incisor (Erickson and Seliger 1969) • Pregnancy Rates Reproductive tracts of 80 females were examined for pregnancy. Does lacking identifiable embryos or fetuses after December 29 were regarded as current breeding failures. If corpora lutea were present, we used only those does in which there was gross evidence, through presence of embryos or pigmented degenerating corpora lutea, that the doe either had been or was currently pregnant. Reproductive data was summarized by the following age classes: �92 yearling, 2-year old, prime, old, and unclassified. Yearlings examined for pregnancy were 18-23 months of age; 2-year olds were 30-35 months of age; prime does were those estimated to be 3-7 years old, inclusive, and old does were 8 years and older. Unclassified does were those known only to have been older than fawns. Breeding Season The female reproductive tract including the ovaries were removed in the field and transported to the laboratory for examination. The primary purpose of measuring the embryos and fetuses was to estimate probable age at the time of death; hence conception dates and the breeding season for this population. Fetal forehead-rump measurements were made as described by Armstrong (1950). Measurements of twins and triplets were averaged to provide a single value. We estimated the age of each fetus by using the linear regression of foreheadrump length (mm) (Y) and age (days) (X) established from known-age mule deer fetuses between the ages of 48-174 days old (Hudson and Browman (1959). Fetal ages in this study were estimated to be within this age range, and Short (1970) indicated that a linear relationship adequately reflected this period of fetal growth. Pregnancy Determination Using Fecal Progesterone Progesterone concentration have been used to diagnose pregnancy in a variety of wildlife species. Monitoring fecal progesterone concentrations may provide a means to remotely diagnose pregnancy without risk, stress, or cost associated with capturing animals. If fecal progesterone were an accurate predictor of pregnancy, then it would be a useful tool in evaluating contraceptive treatments. To evaluate this technique, we collected 5 ml of blood and 10 g of fresh feces from each female mule deer. Samples were placed in a cooler with dry ice and transported to the laboratory for later analysis (Miller et ale 1991). Serum and fecal progesterone concentrations were quantified using radioimmunoassay (Niswender et ale 1969). We compared known reproductive status of females with measured levels of progesterone in serum and feces. Anterior Pituitary/Hypothalamus The brain was completely removed from the cranium by first making three-four cuts on the skinned skull with a bone saw or cleaver, two along the edge of the frontal and parietal bones, and one perpendicular to the longitudinal axis of the skull, and at or just above, the postorbital process of the frontal bone. Following removal of the brain, the anterior pituitary gland was removed from the sella turcica by cutting the diaphragm sellae at the rim of the sella turcica. The pituitary and hypothalamus were wrapped in aluminum foil to facilitate fast-freezing, labelled and placed on dry ice. Genetic Evaluation Muscle, liver, and blood samples were collected from all females. Tissue samples were placed in plastic bags and kept on dry ice during collections, then stored at -70C until analyzed. Samples will be subjected to horizontal starch gel electrophoresis and mtDNA restriction enzyme analyses. RESULTS AND DISCUSSION Age Characteristics of Females Age distribution of females collected during 1994 is summarized in Fig 2. Approximately 53% of the females collected were of prime breeding age (3-7 years); 7% yearlings, 25% were 2 years old, and 15% were 8 years and older. The oldest female collected was estimated to be 14+ years of age. Most (> 90%) females examined were judged to be in good to excellent condition. This �93 estimate was based on condition. It should of the entire breeding selected over smaller be underepresented in Pregnancy visual assessment of body fat reserves and overall body be noted that these collections were not a random sample population since larger females were intentionally animals. Thus, the proportion of yearling females may these collections. Rates Reproductive data from 48 does collected after December 29 are shown in Table 1. Examinations of these does were made sufficiently long after conception that fetal counts could be made and breeding success of failure determined. Before this date, pregnancy could not be determined by gross examination of the reproductive tract, thus 32 females were excluded from this data set. Table 1. Pregnancy records Rocky Mountain Arsenal. for female mule deer collected Age class Total does examined Percent does pregnant Yearlings 2-years Prime Old Total 2 14 28 4 48 100.0 92.8 96.4 75.0 93.0 aFetal Rate = total Erenatal vouna total females of breeding during 1994 at the Fetuses per pregnant doe 2.00 1.84 1.78 2.00 1.86 Fetal Ratea 2.00 1. 75 1. 72 1.50 1. 75 age Pregnancy rate averaged across all age classes was estimated to be 93.6%; fetal rate for all does was 1.75 fetuses per pregnant doe. In summary: Yearlings: Only two yearling females were examined; both were pregnant and both carried two fetuses. Two-year-olds: Fetal rate for this age class was similar to that observed for both prime and old age groups. Twins outnumbered singles by almost 6:1; no triplets were observed for this age class. Prime (3-7 years): Approximately 96% of these females were pregnant but the rate per pregnant doe was lower than that for 2 year-olds and old does. Two sets of triplets were found in this group. Old (8+ years): Pregnancy and fetal rates for this group was substantially lower than that for other age classes. Although the sample size here is small, it does not appear that productivity of does at the RMA is significantly impaired by senescence. Two females, aged 13 and 14 were pregnant with two fetuses each. Breeding Season We examined and measured 53 fetuses during the period of January 25 to April 19, 1994. The sex ratio of fetuses was 1.5 males: 1.0 females. The growth rate for fetuses in this study was similar to the rate for known-age mule deer fetuses (Hudson and Browman 1959) and to the growth rate reported for freeranging mule deer in the Piceance Basin in western Colorado (Bartmann 1986) (Fig 3). We used the above fetal growth rates to estimate conception date. Linear regressions of estimated breeding date on collection date were then calculated (Fig 4). The slope of the regression line was not significantly (~ = 0.21) different from O. Thus, estimated conception dates were similar regardless of collection date. We estimate that peak of conception occurs about November 23 and 95% of the adult females conceive between November 19 and November 27. �Pregnancy De~ermina~ion Serum and fecal progesterone concentrations followed similar patterns during gestation (Fig. 5 and 6). Concentrations of fecal progesterone gradually increased from conception to a peak of 175 ng/g at 64 days of pregnancy, then gradually declined to the end of the collection period. Although there was substantial individual variation in fecal progesterone concentrations, after 48 days of gestation, levels for non-pregnant females were noticeably lower than those for pregnant females (Fig. 6). Serum progesterone levels rose to a peak of 3.76 ng/ml by day 35, then showed little change between days 36 to 156. It appears from these data that both serum and fecal progesterone can be used to diagnose pregnancy in RMA mule deer. Anterior Pituitary/Hypothalamus Analysis of these samples is currently Physiology, Colorado State University, Genetic in progress at the Department Ft. Collins, Colorado. of Evaluation Analysis of these samples is currently in progress Ecology Laboratory, Aiken, South Carolina. LITERATURE at the Savannah CITED Armstrong, R. A. 1950. Fetal development of the northern (Odocoileus virginianus borealis). Amer. Midl. Nat. Abramowitz, M., and I. A. stegun. 1968. Dover Publications Inc., New York, River white-tailed 43:650-666. Handbook of mathematical NY, pp. 645-652. deer functions. Baker, D. L., N. T. Hobbs, T. M. Nett, M. W. Miller, and R. B. Gill. 1993. Regulation of mule deer (Odocoileus hemionus) population growth by fertility control: laboratory, field, and simulation experiments. Contraception in Wildlife Management symposium, Oct. 26-28, Denver, CO, (Abstract). Baker, D. L. 1994. Regulation of mule deer population growth by fertility control: laboratory, field, and simulation experiments. Quarterly Report, July-Oct 1994, Colo. Div. Wildl., Ft. Collins, CO. 6pp. Bartmann, R. M. 1989. Growth rates of mule deer fetuses conditions. Great Basin Nat. 46:245-248. Freund, R. J., R. C. Littell, models. SAS Institute, Hobbs, and P. C. Spector. Cary, NC, 187-201. N. T. 1994. Regulating stage structured models. under 1986. different SAS system for linear populations by controlling fertility: Ecological Applications: in review. Hudson, P., and L. G. Browman. 1959. Embryonic mule deer. J. Wildl. Manage. 23:295-304. winter and fetal development general, of the Miller, M. W., N. T. Hobbs, and M. C. Sousa. 1991. Detecting stress responses in Rocky Mountain bighorn sheep (Ovis canadensis): reliability of cortisol concentrations in urine and feces. Can. J. Zool. 69:15-24. Miller R. G. 1966. Simultaneous New York, NY, pp. 152-168. statistical Morrison, D. F. 1976. Multivariate Co, New York, NY, pp. 145-194. inference. statistical methods. McGraw-Hill McGraw-Hill Book Co, Book �95 Nett, T. M., D. L. Baker, M. W. Miller, and A. L. Case. 1993. GnRH induced patterns of LH secretions in mule deer (Odocoileus hemionus) during the breeding season, anestrous, and pregnancy. contraception in Wildlife Management Symposium, October 26-28, 1993, Denver, CO, (Abstract). Niswender G. D., L. E. Reichert Jr., A. R. Midgley 1969. Radioimmunoassay for bovine and ovine Endocrinology 84:1166-1173. Robinette, W. L., and J. S. Gashwiler. J. Wild. Manage. 19:115-136. Short, C. 1970. tailed deer 1955. Jr., and A. V. Nalbandov. luteinizing hormone. Fertility of mule deer Morphological development and aging of mule deer fetuses. J. Wildl. Manage. 34:383-388. in Utah. and white- �96 Intramuscular Injection vs. 8iobuilet Delivery ot GnAH Analog in Mule Deer on A91 .................... 10 --0- '""eellon ~- 8100uUeI ;- ::::a 8IOou ••• r e Ci I s ;; J """'A' ! ... 14 :. . ''" 21/ ... .. _ ..•/<::::::~····::·::.·>·~·S:~~ o _ •• '. ", ".~ " o lj·=-=.•. ·_.=--__.•· .. 2 ". . ';' ;O-~Q--">- o 9 3 15 12 nm. 21 18 o 24 ~_ ~ 3 12 tnrs, Time 61 10 I 15 21 ••••••.• "24 Ihrs, 590 10 . ~ ~ 9 I .....•.•...•....•.••.... _ .."..·-==r"""'-.= •••.=-=-""" ="""'·_ ••.•• •••• . ......•..... " . Infection ~~ ~!~o.u"~.. i:; '''::::::.::..::::::::::::..::::::::::: . ~~I·::::~:::··· :'.:::.::::.:::::::::::::::::::::::' . . . . ., 2 /. ... . . . . . .. . .. . o ..,/"'~~"""'l A- t6 o 3 IC • .......... 10 . .:...------" .•.. 9 :.,. 12 15 18 21 o 24 3 6 12 9 nmeOus) Timel,I'lrSI Y92 W92 .. ·············~i,;,~;o~· 10 :::: 3 E -9 - a l: ...• -0 ... j ... , .. '" 2 !i • 4 \ ."\ .: \. " ... -r-' <II 24 "0' ····f··· ." .. L ; 21 '- .. ,.. Ci c _ E 18 :::::<)\: E ~e IS 2 ··4·· o ~J .... \. .. .. \ '" ~ ..... 12 1S 0 0 3 6 9 12 IS 18 21 24 o 3 6 18 21 24 TIm. thrs, Figure 1. Serum lli responses stimul,ated by 12 I-lgGnRH delivered via ballistic implant and via syringe inj ec t Lon, Paired responses of' individual does, identified by alphanumeric ear tags , are shown; solid' lines are responses during the first trial, dashed lines &re responses during the second trial. �97 60 50 •...... ~ 40 0 IZ w o a: w a.. 30 20 10 0 1 (yearling) 2 3-7 (prime) 8+ (old) AGE (years) Figure c.- Age distribution of female mule deer collected at the Rocky Mountain Arsenal during 1994. �98 - E E ..•.... - + 300 ..c: 0> c: Q,) ...J C. E 200 :::I r2 - 0.98 a: I "'C ell Q,) ..c: ...0 Q,) Y 100 - -14.2 +.3.4x LL o o 25 Collection Figure 3. Forehead-rump length as,a Rocky Mountain Arsenal during 1994. 75 50 fdnction of collection 100 125 Date (Julian Day) date for mule deer. fetuses collected at the 350 % o 340 I- + c: ell + :::I :::2. 330 I- Q,) C\l o OJ c: 320 I- "'C Q,) Q,) + :j: + + + + + + + + + •.. rn iii + 310 r2 o 0.018 y - 326.8-0.02 I- UJ 300 = + + + + + + I I I I 25 50 75 100 Collection Figure 4. Estimated breeding date ~f'female calculated from data of HUdson and Browman 125 Date (Julian Day) mule deer at. the Rocky (1959), Mountain Arsenal using fetal growth rates �99 SERUM -II20 ---E - PREG (CAP) PROGESTERONE - •.• _. NON (CAP) -..... PREG (RMA) r-------------------------------------------~------------------_, 16 - Ol c: ..._.. 12 "'=t a. :2 8 :::> a: w CIJ 4 ~~~~~_.--I._~~ Ok-~~----J---~~~~~==~~~~~ o -22 48 22 64 __ 86 106 GESTATION 132 156 186 216 204 (DAYS) Figure 5. Serum progesterone concentrations (rig/mll of pregnant Arsenal during 1994, and from captive mule deer at the Foothills mule deer collected at the Rocky Mountain Wildlife Research Facility. FECAL PROGESTERONE -II- - •.• _. NON (RMA) PREG (CAP) -..... PREG (RMA) 350 r-----------------------------------------------~I ----I 1 ...Ol --Ol 300 1 1 250 I I 200 I I c: o:::t 0... _j < o w u.. I 150 1 100 TI ..•... 50 ......... O*-~~ -22 o L_ __ _L __ ~L_ 22 48 64 __ ~ __ ~ 86 GESTATION 1 .---------~ 106 __ ~ 132 I ~ 156 .186 __ ~ __ ~ 204 (DAYS) Figure 6. Fecal progesterone concentrations (ng/mll of pregnant mule deer collected Arsenal during 1994, and from the Foothills Wildlife Research Facility. at the Rocky Mountain 216 ��101 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS REPORT Title: Heart Rate as a Potential Indicator of Stress: Sheep Exposed to Human Disturbance Period Covered: Author: Application to Bighorn July 1, 1994 - June 30, 1995. M. A. Wild and D. L. Baker. Personnel: P. E. Bleicher, R. B. Heath, D. L. Piermattei, J. L. Schaefer. ABSTRACT Introduced disturbances may adversely affect bighorn sheep in various manners, but the most widely hypothesized detriment is that of stress leading to excessive stimulation of the endocrine system with associated immunocompromise. To better understand the effects of disturbances on bighorn sheep, a reliable, longterm, noninvasive, quantitative indicator of stress is required. We propose using remote monitoring of heart rate to meet this need. Various stimuli, including disturbance by humans, elicit heart rate changes in wildlife, even in the absence of overt behavioral responses. These changes likely correlate with pituitary-adrenal axis and metabolic responses. Remote monitoring of heart rate might provide a quantitative measure of stress level in free-ranging bighorn sheep if such a correlate could be demonstrated. We planned experiments to investigate this correlation of heart rate and serum cortisol in captive bighorn sheep. We developed a successful technique to remotely monitor heart rate in bighorn sheep that combined new technology and a new surgical approach. Three domestic goats and then 5 bighorn sheep received Telonics model HR400 heart rate transmitters placed subcutaneously on the dorsolateral thorax using aseptic technique. Of the 3 transmitters placed in goats, one functioned well, one required a minor procedure for correction, and one produced a spurious signal that interfered with data collection and could not be corrected. Minor modifications were made and the surgical technique was applied to bighorn sheep. Currently, all transmitters in bighorn sheep are functioning properly and no adverse reactions have been obl3erv.ed. The Lotek SRX 400 telemetry receiverjdatalogger .Witll, ~vent Log W9 wae{ determined to be us.eful for collect'ion of h~a~t rate datia, .. - ��100 HEART RATE AS A POTENTIAL INDICATOR OF STRESS: APPLICATION TO BIGHORN SHEEP EXPOSED TO HUMAN DISTURBANCE Margaret A. Wild and Dan L. Baker P. N. Objectives Develop a technique to monitor disturbance wildlife viewing areas. to bighorn sheep from humans in SEGMENT OBJECTIVES 1. Prepare a study plan for the evaluation of heart rate collected telemetry as an indicator of stress in bighorn sheep. via 2. Develop a safe, reliable, and unobtrusive system to remotely monitor rate in bighorn sheep over an extended period (~ 1 year). heart METHODS AND MATERIALS We prepared a detailed study plan for the evaluation of heart rate collected via telemetry as an indicator of stress in bighorn sheep (Appendix A). Because we found no documentation of a safe, reliable, unobtrusive system to remotely monitor heart rate in ungulates, we proposed a new approach which required evaluation with 2 pilot studies followed by limited application in bighorn sheep. Determination of optimal electrode placement: In the first pilot study, we determined the optimal location for electrode placement. We sedated each of 3 adult, female bighorn sheep held at Foothills Wildlife Research Facility (FWRF) using about 90 mg ketamine + 9 mg xylazine administered intravenously (IV). We clipped ECG leads to various locations on the body of the sedated sheep and monitored real-time ECG tracings. ECG tracings were subjectively evaluated based on height of positive and negative waves, particularly, Rand T waves. Implantation of heart rate transmitters in goats: In the second pilot study, we used domestic goats as a model for bighorn sheep in the initial evaluation of our surgical technique. We obtained 3, l-year-old domestic goats (mean weight about 50 kg) from a local producer. After a period of acclimation to the facility, goats were implanted in November 1994. Goats were fasted for about 48 hr prior to surgery. Anesthesia was induced with 200 mg ketamine + 7.5-10 mg valium administered IV and was maintained using isoflurane. Vital signs and aesthetic depth were monitored regularly through the anesthetic period. Intermittent positive pressure ventilation was used as necessary. Using aseptic surgical technique, an approximately 15 cm skin incision was made on the left lateral thorax just caudal to the scapula. We evaluated 3 different surgical approaches: vertical, curved, and horizontal (paramedian) skin incision. Fat was elevated and the latissimus dorsi muscle incised. Deep fascia was incised parallel to the ventral lateral border of the longissimus dorsi muscle to allow for its elevation. The sterilized Telonics model TR400 heart rate transmitter was placed in the naturally occurring shelf between the ribs and longissimus dorsi muscle. The transmitter was secured into place by reattaching the split fascial plane over the transmitter using a far-far near-near pattern tied in a cruciate. The positive lead was looped, laid just ventral to the transmitter, and the electrode sutured to muscle fascia using a simple interrupted suture. The negative lead was passed through a subcutaneously placed trocar inserted from the primary incision site to a secondary incision site located about 5-10 cm dorsad from the xiphoid. With the lead in place, the trocar was removed, excess lead laid in a loop, and the electrode secured in place. The transmitter was checked for proper �function by listening to the signal using a telemetry receiver. The incisions were then closed. Muscle, subcutaneous, and skin layers were closed using simple interrupted, cruciate, or simple continuous patterns. Maxon suture, size a or 2-0, was used through out the procedure. About 1,800,000 U of procaine G and benzathine penicillin was administered subcutaneously and 200 mg phenylbutazone administered orally (PO) on recovery from anesthesia. Implantation of heart rate transmitters in bighorn sheep: We used the technique developed in goats, with minor modifications, to implant heart rate transmitters into 5 captive bighorn sheep at FWRF in April-May 1995. Based on experience from surgery on goats, we incorporated several minor improvements to our technique. These included: using a 15 cm paramedian skin incision, incising the latissimus dorsi muscle parallel to the muscle fibers, placing the transmitter several centimeters more caudad, placing the positive electrode ventral-caudal to the transmitter, placing the negative electrode just caudal to the antebrachiohumoral joint, securing both the leads and electrodes with staple sutures, and administering ceftiofur (about 2 mg/kg, intravenously) intraoperatively. We monitored a Lotek model to our novel receiver and function of heart rate transmitters at specified intervals using SRX 400 telemetry receiver with Event Log version 2.5x W9. Due applIcation of the software, we subjectively investigated ideal datalogger function settings. RESULTS AND DISCUSSION Determination of optimal electrode placement: We considered the optimal electrode location to be not only where R waves were high, but as importantly, where T waves were markedly smaller than the R waves. This difference in electrical potential of R waves as compared to other waves in the complex is important in avoiding multiple counts of each heartbeat when using the telemetry equipment. Based on these data, we determined that the optimal electrode location on bighorn sheep was placement of the positive electrode on the cranial-dorsal thorax (at the proposed site of transmitter implantation) and of the negative lead at the xiphoid or directly dorsal to the xiphoid approximately 6-8 cm. Average required lead length for transmitters was determined to be 15 cm for the positive lead and 60 cm for the negative lead. Implantation of heart rate transmitters in goats: Goats were ambulatory about 1 hr post-surgery. Two were clinically normal by 36 hr post-surgery. The third was in moderate pain and required further analgesia (200 mg phenylbutazone PO) 48 hr post-surgery but was clinically normal by 72 hr postsurgery. All transmitters remained functional; however, spurious signals (extra beats) were a problem with 2 transmitters. In goat BK93 a spurious signal was apparent from soon after surgery. We determined that spurious signals were likely due to the more dorsal location of the negative electrode as compared to the other goats. To correct the problem, we sedated the goat (BK93) with 80 mg ketamine + 8 mg xylazine IV (with an additional 27 mg ketamine + 3 mg xylazine administered during the procedure). Using aseptic technique, we made an about 4 cm incision and located the negative electrode. When the electrode was located, we observed that the lead was looped so that the electrode was "upside down" and also turned on its side. This apparently resulted in inappropriate contact with the muscle. We made a second incision about 6 cm directly ventral to initial incision and pulled the electrode subcutaneously to the new site using tissue forceps. The electrode was secured with 2 simple interrupted sutures of 2-0 Maxon. Muscle, subcutaneous, and skin layers of each incision were closed using cruciate or simple continuous suture patterns of 2-0 Maxon. Yohimbine (10 mg IV) and penicillin (1,800,000 SQ) were administered postoperatively. The goat recovered well and the problem of spurious signals was resolved. This problem may be avoided in the future by �100 assuring that the electrode is placed no more than about 7 cm from the sternum and by more tightly securing the electrode to the underlying tissue. A staple suture may increase the amount of contact between the electrode and muscle. In a second goat, BR93, the signal became worse with time (beginning about 2 wk post-surgery). In this case, we believed that the positive electrode was the cause of the spurious signals. We performed a second surgery on 30 January. Anesthesia and surgical approach were as described above. The transmitter was observed to have migrated cranially about 5 cm and was partially beneath the scapular cartilage. Consequently, the positive electrode was in close apposition to the scapular cartilage. We hypothesized that in this location the electrode transmitted spurious signals when the scapular cartilage moved. The transmitter was now well affixed to the body wall by fibrous tissue and so was left in place. The lead to the positive electrode was dissected from fibrous tissue and the electrode reattached to muscle caudal and dorsal to the transmitter. In this position, the electrode was <5 cm from the scapular cartilage. Surgical closure and post-operative care were as described above; however, a body wrap of elasticon was applied as well and removed about 48 hr post-op. Although the signal was accurate for about 2 days post-op, reliability quickly deteriorated. starting at 3 days post-op the signal was interpreted to be normal at times but at other times appeared to be the default mortality signal plus occasional beats. Ten days post-op the skin incision began to dehisce and a purulent exudate was noted. Antibiotic therapy (Naxcel, 75 mg daily, SQ) cleared the infection and the skin defect was able to heal. Unfortunately, the transmitter continued to function on the mortality signal with only occasional beats. This indicated that the acute inflammatory response was not the cause of the errant signal. Because the transmitter was not properly functioning within the time for normal healing after the corrective surgery, the goat was euthanatized. Postmortem examination of the goat was unremarkable except for the surgical site. A large accumulation of fibrous tissue was found'over the site of the second surgery, especially at the attachment site of the positive electrode. The positive electrode was isolated in the fibrous tissue and was not in direct contact with the muscle. This isolation likely precluded transmission of the electrical impulse to the electrode, therefore, the mortality signal occurred. The transmitter appeared in good condition with no sign of leakage; however, the wax coating on the transmitter had started to breakdown. All wax was missing from an about 1 cm strip along the back of the transmitter where it had laid against the body. The wax was apparently removed from the site by the body. Telonics was contacted about the loss of wax but did not know of this problem occurring before or why it had happened. The transmitter will be submitted for a failure analysis to determine if damage to the transmitter occurred. Although the transmitter did not provide a useful signal in this goat, we learned several important points from the failure. First, the size of the goat may have predisposed the system to difficulty. The goat was likely about 5-10 kg smaller than the 2 successful goats. Slight difficultly was encountered on the initial surgery because the "pocket" made by the epaxial muscles was smaller than expected and the transmitter did not fit nicely into place. This may have resulted in the migration of the transmitter craniad under the scapular cartilage. Second, placement of the positive electrode caudal to the transmitter (instead of over it) to fully avoid the scapular cartilage would likely have avoided the problem of interference if the transmitter migrated for any reason. Third, although a more severe inflammatory response is anticipated after a repeat surgery, response was greater than expected. These observations lead to 3 conclusions: 1) there may be a lower limit on body size for successful implantation, 2) the positive electrode should be placed caudal to the transmitter, and 3) the prognosis for a functional transmitter after a second surgery (involving the site of the transmitter) is questionable, if an animal requires follow-up surgery, the electrode should be placed in a "new" location that has not been previously manipulated and antibiotics should be continued for 1 week post-op if possible. �100 Based on results from this pilot study, we believed that this surgical approach was highly likely to be successful in bighorn sheep. Further, it should provide a relatively simple, safe, and humane approach to implant transmitters for collection of reliable heart rate data. Implantation of heart rate transmitters in bighorn sheep: Surgical implantation of transmitters into bighorn sheep was successful. We devoted more attention to assuring optimal placement of electrodes intraoperatively to avoid need for correction postoperatively. No adverse reactions were noted. All transmitters are currently functioning properly. The receiver/datalogger appeared acceptable for collection of heart rate data. For experimental use at FWRF, ideal settings are: noise index = 20; noise blank level = 48; gain = 92; scan time = 6 sec; continuous record timeout = 0; collision threshold = 250 ms; and group size = 8. These settings appeared to optimize data screening (filtering spurious signals) and maximize data collection. �107 APPENDIX A STUDY PLAN HEART RATE AS A POTENTIAL INDICATOR OF STRESS: APPLICATION TO BIGHORN SHEEP EXPOSED TO HUMAN DISTURBANCE I. NEED As human populations increase and disperse in Colorado, and throughout the west, human interactions with wild species increase. This increased interaction occurs unintentionally when urban sprawl expands into wildlife habitats. Moreover, humans increasingly venture into wildlife habitats intentionally with the purpose of encountering, viewing, and enjoying freeranging wildlife. Wildlife agencies have recently increased efforts to provide such wildlife viewing experiences. Management programs for wildlife viewing opportunities strive to optimize human satisfaction while protecting the wildlife resource. Accomplishing this dual goal requires knowledge both of human desire for and evaluation of recreational activities and the impacts of these activities on wildlife populations. A survey of residents of Denver, Colorado (Manfredo and Larson 1993) indicated that bighorn sheep (Ovis canadensis) were among the most important species for wildlife viewing recreation. Opportunities to view bighorn sheep in Colorado are numerous, and several designated viewing areas have been established. The impacts of viewing areas on bighorn sheep remain unclear. Bighorn sheep may habituate to predictable patterns of human activity and be unaffected by viewing. However, due to potential deleterious effects of human disturbance on bighorn sheep, restrictive limitations are placed on humans using some areas (e.g. Rocky Mountain National Park). These restrictions may limit the satisfaction of some groups of wildlife viewers, but are deemed necessary for the welfare of the bighorn sheep. This reasoning is based on the assumption that bighorn sheep exhibit fidelity to traditional ranges, regardless of changes in suitability of those ranges (Geist 1971), and may endure increased levels of stress from disturbance rather than expand to new ranges to avoid it. Introduced disturbances may adversely affect bighorn sheep in various manners, but the most widely hypothesized detriment is that of chronic stress (Selye 1950) leading to excessive stimulation of the endocrine system with associated immunocompromise (Hudson 1973, Spraker et al. 1984). Increased levels of glucocorticoids, as occurs under stress, have been shown to adversely impact several aspects of host defense in domestic bovids (reviewed by stephens 1980, Roth 1985). Immunosuppressive effects of stress have been implicated in the bovine shipping fever complex (reviewed by Confer et al. 1988), and may be an important component of the bighorn sheep pneumonia complex (Hudson 1973, Spraker et al. 1984). The pneumonia complex certainly impairs bighorn sheep population performance (Buechner 1960, Spraker and Hibler 1982, Thorne 1982, Onderka and Wishart 1984). Given the potential deleterious effects of stress on bighorn sheep, conservative management practices seem prudent. To better understand the effects of disturbances on bighorn sheep, a reliable, longterm, noninvasive, quantitative indicator of stress is required. Serum cortisol levels are commonly measured, but reflect only the status of an animal at a point in time (which is commonly influenced by the invasive sampling technique itself). Fecal and urine cortisol (Miller et al. 1991) provide a reliable method for noninvasive monitoring; however, this technique can be difficult to apply to monitoring of individual free-ranging bighorn sheep, especially in their response to specific events. Behavioral responses alone are inadequate indicators of excitation in bighorn sheep (MacArthur et a1. 1982, Stemp 1982) and other species (Kreeger et al. 1989, Baldock and Sibly 1990) because various stimuli, including disturbance by humans, elicit heart rate changes even in the absence of overt behavioral responses. Heart rate changes likely correlate with metabolic and pituitary-adrenal axis responses, including serum cortisol levels (Harlow et al. 1987a, Harlow et al. �100 1987b, Chabot et al. 1991). Remote monitoring of heart rate might provide longterm, noninvasive (after initial placement of telemetry device), quantitative index of cortisol level in free-ranging bighorn sheep if a correlation could be demonstrated. a Harlow et al. (1987a) suggest that heart rate has a good potential of reliably predicting cortisol levels. This conclusion was based on finding a strong correlation between heart rate and plasma cortisol levels in a group of domestic sheep (Harlow et al. 1987a) and in 2 individual bighorn sheep (Harlow et al. 1987b). Several questions remain to be answered before this method can be applied to free-ranging bighorn sheep. First, the correlation between heart rate and serum cortisol levels in a more representative number of bighorn sheep must be determined using graded disturbances as stressors. If a strong correlation is demonstrated, the relationship could then be used to predict cortisol responses by other bighorn sheep based on their heart rate. However, factors other than the disturbance will likely influence heart rate and cortisol levels and must be considered. These factors may include sex, age, physical condition, and reproductive status of the individual, it's activity status, and influences of circadian, prandial, and seasonal cycles. Therefore, the second set of questions to be answered will involve the independent influences of some of these factors on heart rate and cortisol levels. Understanding the effects of these factors will provide insight into possible correction rates that should be applied to the correlation. Finally, a safe and reliable field system must be established for monitoring heart rate in free-ranging bighorn sheep for extended periods (~ 1 year). Because application of the system may be in bighorn sheep located near human viewing areas, the system should also be inconspicuous and not detract from the aesthetic value of the animal. Although numerous attempts have been made (MacArthur et al. 1979, Stemp 1982, Bunch et al. 1989, Coates et al. 1990, Wallace et al. 1992), no system reported for use in bighorn sheep fully meets these criteria. Therefore, modification of the most promising technique needs to be attempted and evaluated. A remote technique, such as heart rate monitoring, that accurately predicts cortisol response of bighorn sheep to various disturbances could substantially increase our ability to wisely manage bighorn sheep. The technology would serve as a basis for further research to determine, based on physiologic response, what does and does not disturb bighorn sheep. This information would allow us to encourage human activity that maximizes satisfaction without adversely affecting the bighorn sheep in wildlife viewing areas. II. OBJECTIVES Use of heart rate as an indicator of stress in bighorn sheep shows promise. We propose research to investigate the utility of heart rate data collected via telemetry systems to predict cortisol levels in bighorn sheep. In a future study, we will apply this technology to free-ranging bighorn sheep exposed to human disturbance. The objectives of this research project are: 1) To develop a safe, reliable, and unobtrusive system to remotely monitor heart rate in bighorn sheep over an extended period (~ 1 year). 2) To determine the correlation between heart rate and serum cortisol levels in bighorn sheep and to understand the effects of some other physiologic parameters on this correlation. III. EXPECTED RESULTS AND BENEFITS We believe that the proposed research will provide a novel and effective means for monitoring stress in bighorn sheep, and other species given establishment of species-specific regressions of serum cortisol to heart rate. We anticipate that the degree of disturbance (by humans, or other environmental, �109 social, or physiological stressors) will be predicted using heart rate as an indicator of serum cortisol level in bighorn sheep. Thus, a means will be available to identify activities that do, and do not, disturb bighorn sheep. These findings will be applied in future research on free-ranging bighorn sheep and should ultimately guide management decisions that will maximize human satisfaction without endangering the well-being of bighorn sheep. IV. APPROACH We will conduct technique development and controlled experiments using tractable, penned bighorn sheep held at Foothills Wildlife Research Facility (FWRF), Fort Collins, Colorado. Between treatment periods bighorn sheep will be housed in groups in about 3 ha pastures. Grass/alfalfa mix hay, water, and mineral block will be provided ad libitum in addition to limited amounts of pelleted supplement (Baker and Hobbs 1985). We will develop and refine surgical techniques and data monitoring and collection methods using these captive bighorn sheep. Bighorn sheep equipped with heart rate telemetry devices will be used to measure heart rate and serum cortisol responses to graded stressors under controlled conditions. We will also investigate effects of other physiologic states on heart rate and serum cortisol levels. The work will be organized into a technique development stage and 2 exper iments • Technique Development Rationale: A safe and unobtrusive remote monitoring system that functions reliably for extended periods is prerequisite to successful field monitoring of heart rate in bighorn sheep. Currently used systems lack one or more of these necessary criteria. Objectives: 1) To develop/refine an anaesthetic and surgical protocol for implantation of heart rate transmitters in bighorn sheep that emphasizes animal welfare and maximal lifespan of telemetry signal. 2) To identify applicability, including accuracy, range, and ease of use, of a automated data acquisition system for collecting heart rate data. Methods: Transmitter Implantation Based on preliminary work using domestic goats as a model for bighorn sheep (Wild, unpub. data), we have developed a safe and reliable method to remotely monitor heart rate. We will apply this newly developed technique to surgically implant transmitters in bighorn sheep. Surgery will be performed in the designated surgical area at FWRF. Adult, female bighorn sheep (n=15) will be fasted for 48 hours then anesthetized. Anaesthesia will be induced with intravenous (IV) administration of ketamine (1-2 mg/kg) plus either xylazine (0.1-0.2 mg/kg) or diazepam (0.2 mg/kg). Alternately, anesthesia may be induced with mask administration of isoflurane. Anesthesia will be maintained using isoflurane. Vital signs and anesthetic depth will be monitored regularly throughout the anaesthetic period. Bighorn sheep will be placed in right lateral recumbency and an aseptic site prepared for surgery. Following aseptic technique, we will make an approximately 15 cm skin incision on the left lateral thorax just caudal to the scapula. We will use sharp and blunt dissection through superficial fat and muscle tissue to reach the epaxial muscles. The sterilized Telonics model HR400 heart rate transmitter will be inserted just ventral to and parallel to the epaxial muscles with the lead exits pointing caudad. The transmitter will be secured into place by suturing previously transected muscle over the transmitter using a cruciate pattern of 0 or 2-0 Maxon. Alternately, the transmitter may be placed into SupraMesh polyamide mesh sheets which will be sutured into place as well. Placement of electrodes will be based on previous investigations using clip-on ECG leads on bighorn sheep (Wild, unpub. data). The positive lead will be • �110 looped and laid just ventral to the transmitter. The negative lead will be passed through a trocar inserted from the primary incision site directly ventral to a secondary incision (about 3 cm) located about 5 cm dorsad from the xiphoid. With the lead in place, the trocar will be removed, excess lead laid in a loop, and the electrode secured in place using 0 or 2-0 Maxon. The transmitter signal will then be checked for proper function. Muscle, subcutaneous, and skin layers will be closed using 0 or 2-0 Maxon in cruciate or simple continuous patterns. We will administer about 3,000,000 U procaine G and benzathine penicillin subcutaneously (SQ) and 300 mg phenylbutazone orally (PO). Bighorn sheep will be monitored until recovery from anesthesia is complete. All stages of the protocol will be performed by, or under the direct supervision of, a veterinarian. Health status of bighorn sheep will be subjectively evaluated by trained caretakers at least once daily for the remainder of the study. Transmitter function will be checked once daily for 4 weeks, then at least once weekly for the remainder of the study. Medical or surgical intervention will be provided by a veterinarian if required to assure health of research animals and for proper functioning the transmitters. If transmitters regularly produce a signal that is not accurate (e.g., spurious signals) beyond 2 weeks post-implantation, we will intervene surgically. We will use the approach described above; however, only specific portions of the anesthetic and surgical approach will be required based on the site of failure. For example, a loose negative lead may require only induction with IV anesthetics, a small incision, securing of the electrode, and closure. In all cases aseptic and proper technique will be followed and post-operative antibiotics and analgesics will be administered. Yohimbine (10 mg) will be administered IV if required for recovery after short procedures. Transmitters will remain in place for the life of the bighorn sheep unless adverse reactions are observed. If any chronic problems occur due to the transmitter, it will be surgically removed using the procedure described above for placement. Data Collection System Evaluation Transmitters will be checked for function using an appropriate telemetry receiver. For data collection, the receiver will be interfaced with a scanner/programmer, digital data processor, and data logger. Data will be transferred from the data logger to an IBM compatible personal computer using an appropriate interface and software. We will also assess accuracy of heart rate determination by the data collection system through comparison with the "standard" of a hard-wired electrocardiograph (ECG) output. We will hold individual bighorn sheep (n=5) in a 0.5 by 1.5 m enclosure, attach clip-on ECG leads, and record heart rate using the data collection system and ECG strip output simultaneously for 3 independent 1-min sampling periods. This protocol will be repeated under periods of low, moderate, and high heart rate for each bighorn sheep. Heart rate may need to be manipulated by administration of xylazine (0.1-0.2 mg/kg IV) or atropine (0.2 mg/kg IV) or by exercise. We will determine line of sight range of the signal from transmitters to the data acquisition system. We will place bighorn sheep (n=5) in a fixed location in a pasture and determine mean signal strength over 1 min sampling periods from each of 6 experimenter distances (20, 100, 400, 700, 1,000, and 1,300 m). Signals will be evaluated with bighorn sheep facing toward, away from, at a right angles to (left and right) the experimenter. This protocol will be repeated with each bighorn sheep at 2 mo intervals for the lifespan of the transmitters. Analysis: Transmitter Implantation Evaluation of anaesthetic clinical assessment. and surgical techniques will be based on subjective �111 Data Collection System Evaluation Agreement between methods for heart rate determination will be evaluated by preparing a calibration line. We will compute the regression line for heart rate determined by the data collection system versus heart rate from the ECG (the "true" value) using the method of least squares and determine prediction intervals. Analyses will be performed using SAS (SAS Institute Inc. 1988). We will use the general linear model to determine the function that describes the relationship between signal strength and distance for each of the 4 body positions. Analyses will be performed using SAS (SAS Institute Inc. 1988). Because many factors, especially terrain, will influence range of the data collection system, we will not make inference to other locations using these data. Instead, these data will provide general information on possible utility of the system in the field. Experiment 1: Heart Rate-Serum Cortisol Correlation Rationale: According to the General Adaptive Syndrome (GAS) (Selye 1950), when a stress-inducing stimulus is perceived by the cerebral cortex, a cascade of physiological events occur in an animal (Reviewed by Stephens 1980). The autonomic nervous system is stimulated via the hypothalamus. Adrenaline and noradrenaline output from the adrenal medulla is increased and leads to the "fight or flight" reaction. Increased heart rate is one of these responses. The hypothalamus also secretes corticotrophin-releasing factor (CRF) which, in addition to adrenaline, stimulates the anterior pituitary gland to secrete adrenocorticotrophic hormone (ACTH). In turn, ACTH stimulates the adrenal cortex to secrete corticosteroid hormones, including cortisol. Because increased heart rate and cortisol levels are both components of this response to a stressor, we hypothesize that measurement of heart rate will serve as a predictor of serum cortisol. In fact, heart rate changes have been shown to correlate with plasma cortisol levels in domestic sheep and in 2 individual bighorn sheep under controlled situations with graded stressors (Harlow et al. 1987a, Harlowet al. 1987b). If a strong correlation between heart rate and serum cortisol level was determined for a specific class of bighorn sheep (e.g. adult, nonpregnant ewes), predictions of serum cortisol for other individuals in that class could be made by remotely monitoring heart rate of those individuals. Objective: To determine the correlation between heart rate and serum cortisol level in adult, female bighorn sheep under controlled situations with graded stressors. Design: Each of 15 bighorn sheep will be exposed to graded stressors as treatment stimuli. Because accurate data cannot be collected from >3 bighorn sheep that are simultaneously exposed to a stressor, we will randomly assign bighorn sheep to 5 sets of 3 animals each for exposure to stressors. After each set of bighorn sheep has been exposed to a given stressor, we will rerandomize sets for exposure to the next stressor. We will expose each of the 5 sets to the mild stressor, then to the moderate stressor, and then to the marked stressor. We will also collect data when the set is not subjected to a stressor. These control sessions will occur prior to initiation of exposure to stressors and after exposure to the range of stressors. Bighorn sheep will be exposed to no more than 1 treatment (stressor or control) per 5 days. Methods: Tractable, adult, female bighorn sheep (n=15) equipped with heart rate transmitters will be trained (retrained) to reside in isolation pens (50 m2) and also to stalls measuring 2.3 x 1.1 m for periods of 6 hr. All measurements for heart rate-cortisol correlation will be made while bighorn sheep are in these stalls. Acclimation to stalls and to presence of monitoring equipment will be determined based on heart rate and/or serum cortisol values of confined bighorn sheep during training sessions. Approximately 12-18 hr prior to treatment initiation, we will sedate bighorn �112 sheep with ketamine (1.0 mgjkg) and xylazine (0.1 mgjkg) administered IV. Each bighorn sheep will be fitted nonsurgically with an indwelling jugular catheter using aseptic technique. Yohimbine (10 mg) will be administered IV to reverse the effects of xylazine. With catheters taped securely in place, bighorn sheep will be placed in isolation pens for the night. The following morning, bighorn sheep will be placed in stalls. Treatments will be initiated 1 hr later, or when the heart rate of each bighorn sheep has returned to baseline for >0.5 hr. If heart rate of an individual does not return to baseline within 2.5 hr, she will be released from the trial. The bighorn sheep that are released from a trial will be grouped into a new set to be retested. Retesting will be performed after all other bighorn sheep have been exposed to the stressor, but before application of the next level stressor. Treatments will consist of graded stressors presented in the following sequence: no stimulus (control), a technician appearing suddenly and pacing in front of the stalls for 1 min, a technician appearing suddenly shouting, running, and banging on stalls for 1 min, a technician with a dog on lead appearing suddenly and running in front of the stalls for 1 min, no stimulus (control). Blood samples will be obtained by technicians positioned near, but visually isolated from, individual bighorn sheep. Blood collections will be made using the catheters attached to about 2 m of sterile tubing. Care will be taken to flush catheters and tubing to maintain patency. Heart rate will be recorded using the data acquisition system previously described. Heart rate measurements and blood collections will be made 5 min before, and 5, 10, 15, 20, 30, 40, 60, 80, 100, 120, and 180 min after application of treatments. Additional heart rate measurements will be made at about 30 sec intervals from 0-10 minutes post-treatment. Serum cortisol will be determined using the radioimmunoassay procedure described by Miller et ale (1991). After the final sample collection, we will remove catheters, administer 1,800,000 U procaine G and benzathine penicillin SQ, and return bighorn sheep to pastures. To determine if a state of chronic stress has occurred during the trials, we will collect feces at the time of the first and final control trials. Fecal cortisol will be determined using the technique of Miller et ale (1991). Analysis: Regression lines of peak serum cortisol level versus peak heart rate will be calculated using the method of least squares. Additionally, regression lines based on the area under the cortisol curve versus the heart rate curve will be calculated. Prediction intervals for cortisol values predicted from new heart rate values will be obtained. Fecal cortisol levels (pre- and post-trial) will be compared using a paired t-test. All statistical analyses will be made using SAS (SAS Institute Inc. 1988). Experiment 2: Influence of Various Physiological States Rationale: Several variables may effect heart rate or serum cortisol level independently, i.e. through pathways other than the GAS response. These effectors likely include: circadian, prandial, and seasonal cycles, extreme temperature ranges, reproductive status, and activity level. Although we can attempt to control these variables in heart rate-cortisol correlation experiments, they are uncontrolled in field situations. Therefore, understanding the influence of these effectors is required to accurately predict serum cortisol levels based on heart rate. A circadian cycle for plasma cortisol level has been demonstrated in cattle (Fulkerson et ale 1980), domestic sheep (McNatty et ale 1972), and desert bighorn sheep (0. canadensis cremnobates) (Turner 1984). This cycle is suggested to be due to an endogenous adrenocortical rhythm (Demura et ale 1966, Turner 1984); however, magnitude of the cycle may be intensified due to confounding effects of feeding (Holley et ale 1975) or changes in visibility and mobility associated with light and dark periods (Turner 1984). Other species such as the dog (Thun et ale 1990), white-tailed deer (Odocoileus virginianus) (Bubenik et ale 1983), and eld's deer (Cervus eldi thamin) (Monfort et ale 1993) apparently lack a diurnal rhythm of cortisol secretion. Circadian cycles for heart rate have not been described. Kreeger et ale �113 (1989) observed no diurnal changes in heart rate of foxes (Vulpes vulpes). In this experiment, we will describe the daily rhythms of heart rate and cortisol secretion in bighorn sheep. We predict that similar to previous studies of domestic and wild sheep, serum cortisol will exhibit circadian cycles and that this may not be correlated to changes in heart rate. A seasonal cycle, with summer peak and winter decline, in heart rate has been demonstrated in many ruminant species and parallels the cycle of basal metabolic rate that is determined by day length (Moen 1978, Baldock et ale 1988, Fran90ise Domingue et ale 1992). Reports describing seasonal cortisol secretory rhythms are limited and equivocal. No distinct seasonal differences in cortisol secretion have been observed in domestic rams (Kennaway et ale 1981), white-tailed deer (Bubenik et ale 1975), or eld's deer (Monfort et ale 1993). However, results from previous studies with wild ruminants were based on serum cortisol levels collected from restrained and/or anesthetized animals which may have acutely influenced cortisol secretion. We hypothesize that cortisol levels are cyclic in phase with basal metabolic rate, being greater in summer during periods of increased feed intake and activity and lower in winter. However, if higher cortisol levels are associated'with the dark phase of the circadian cycle, daily mean cortisol levels may be higher than expected in winter due to short daylength. Feeding may influence heart rate and cortisol levels. In cattle, a decrease in ruminoreticular fill results in a reflex slowing of heart rate due primarily to increased parasympathetic tone (Clabough and Swanson 1989). Meal feeding was suggested to be a significant signal in the circumdiel release of cortisol in domestic sheep (Holley et ale 1975); however, Turner (1984) observed that although bighorn sheep fed throughout the daylight hours, cortisol levels were at their nadir during this period. We hypothesize that the pattern of feeding will determine the response of heart rate and cortisol levels. Fasting will likely decrease heart rate and, at least initially, cortisol levels. Bolus feeding will likely increase heart rate and cortisol level, while consumption of frequent small meals will likely have minimal impact. Exercise increases the physiological demands on an individual. Heart rate increased linearly with running speed in deer (Mautz and Fair 1980) and reindeer (Nilssen et ale 1984). Alexander et ale (1991) demonstrated that exercise resulted in a rise in plasma cortisol level but not CRF in horses. This suggests that increased cortisol during exercise is not directly related to the GAS pathway but to an alternate pathway, likely via arginine vasopressin (AVP) (Alexander et ale 1991). We hypothesize that both heart rate and serum cortisol levels will increase in association with exercise, but that their correlation will be different than when the response is elicited by psychological stressors. Reproductive processes of wild ruminants include growth of body tissue, growth of the fetus, and the production of milk. The metabolic cost of these processes has been studied in both domestic animals and wild ruminants and have been shown to increase significantly during the last trimester of gestation and early lactation (Brockway et ale 1963, Robbins 1983). We hypothesize that a change in baseline heart rate and serum cortisol will occur in late pregnancy in response to these increased metabolic demands. This change in baseline will likely influence magnitude of response to graded stressors and may change the observed correlation between heart rate and serum cortisol level. Objective: To determine the heart rate and serum cortisol response of bighorn sheep to 5 potential effectors: circadian cycle, seasonal cycle, prandial cycle, exercise, and reproductive status. Design: Five adult, female, non-pregnant bighorn sheep will be used in Experiments A, B, and C. Experiment B will be performed in a cross-over �114 design using 2 blocks of 3 bighorn sheep. For Experiment D, 12 adult females will be randomly assigned equally to bred and unbred groups. Methods: Bighorn sheep will be housed in stalls for experiments A, B, and D, and maintained on a halter for experiment C. Jugular catheters will be placed about 12-18 hr prior to treatment initiation as previously described. Heart rate and blood collections for experiment D will be performed as in Experiment 1. Data collection for other experiments in this section will be similar, but a roving technician that is not visually isolated from the bighorn sheep will collect blood samples. Bighorn sheep will be trained to each collection protocol prior to data collection. Experiment A: Effects of Circadian and Seasonal Cycles Circadian and seasonal effects will be monitored by measuring changes in parameters over time of day and time of year. We will sample bighorn sheep every 2 hr for 24 hr periods to determine circadian rhythms at the summer and winter solstice and the vernal and autumnal equinox. Bighorn sheep will be fasted for 12 hr before beginning collections, and will receive 200 g pelleted supplement (Baker and Hobbs 1985) after each sampling. Experiment B: Effects of Feeding Feeding patterns will be manipulated to study effects of prandial cycles on heart rate and serum cortisol levels. Bighorn sheep will be fasted for 24 hr prior application of treatment I or 2. We will provide bighorn sheep in treatment 1 with ad libitum quantities of high quality grass/alfalfa mix hay and pelleted supplement for 1 hr, then remove all feed. Bighorn sheep in treatment 2 will receive 200 g pelleted supplement every hour. We will collect heart rate and blood samples 5 min before initial feeding and at hourly intervals for 12 hr after feeding. Treatment assignments will be switched and the experiment repeated about 1 week later. Experiment C: Effects of Exercise Activity of bighorn sheep will be manipulated to study effects of exercise on heart rate and serum cortisol levels. We will train bighorn sheep to stand and to run slowly while on lead. We will collect heart rate and blood samples 5 min before and for 1 hr after (at 1, 5, 15, 30, 45, 60 min) 1 min of forced exercise. Additionally, heart rate will be monitored continuously during and at 1 min intervals for 15 min following exercise. Experiment D: Effects of Reproductive Status We will investigate effects of reproductive status on heart rate and serum cortisol levels. We will repeat the protocol described in Experiment 1 using 6 pregnant and 6 nonpregnant adult ewes. Pregnancy will be predicted using serum assay for pregnancy specific protein B (G. Sasser, pers. comm.) and confirmed at parturition. One repetition of the trial will be conducted during each of the 3 trimesters of pregnancy. Analvsis: We will conduct all statistical analyses using SAS (SAS Institute Inc. 1988). Heart rate versus serum cortisol values from each experiment will be compared to prediction intervals from the heart rate versus cortisol regression line identified in Experiment 1. Experiment A: Effects of Circadian and Seasonal Cycles We will use the general linear model to determine the function that describes the relationship between serum cortisol values and heart rate over the 24 hr sampling period and during the daylight hours. To detect possible seasonal effects on heart rate and serum cortisol levels, we will repeat Experiment A during each season. We will use the general linear model to determine and �115 compare the functions that describe level and heart rate versus time. Experiment B: Effects the relationship between serum cortisol of Feeding To detect possible effects of the prandial cycle, we will compare data from trials where bighorn sheep were fasted and then bolus-fed to results when the same animals were fed small, frequent quantities. We will use the general linear model to determine and compare the functions that describe the relationship between serum cortisol level and heart rate versus time for the 2 feeding regiments. Experiment C: Effects of Exercise We will compute the regression lines for serum cortisol level versus heart rate over the range of collected data at various stages of exercise. We will compare these regression lines to those obtained in Experiment 1 to determine the effects of activity on the correlation. Experiment D: Effects of Reproduction We will compute and compare regression lines of serum cortisol level versus heart rate for pregnant and nonpregnant ewes, and determine the effects of time (stage of gestation) on these regressions. V. SCHEDULE Technique Development Initial Transmitter implantation Calibration, evaluation Experiment 1. Heart Rate-Cortisol Correlation Additional transmitter implantation Training, pilot experiments Graded stressors Experiment Experiment Experiment Experiment Experiment 2. Physiological (10) LITERATURE oct - Nov 1995 Nov - Dec 1995 Jan - Mar 1996 Effectors Mar, Aug Aug Dec A B C D Final data analysis VI. Apr-May 1995 Jul 1995 (5) and manuscript prep Jun, Sep, Dec 1996 1996 1996 1996; Feb, Apr 1997 Apr - Dec 1997 CITED Alexander, S. L., C. H. G. Irvine, M. J. Ellis, and R. A. 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Physiol. and Pharm. 53:787792. Bubenik, G. A., A. B. Bubenik, D. Schams, and J. F. Leatherland. 1983. Circadian and circannual rhythms of LH, FSH, testosterone (T), prolactin, cortisol, T3 and T4 in plasma of mature male white-tailed deer. Compo Bio. Physiol. 76A:37-45. Buechner, H. K. 1960. The bighorn sheep in the United present, and future. Wildl. Monogr. 4:1-74. Bunch, T. D., G. W. Workman, and R. J. Callan. and heart rate monitoring in desert bighorn Trans. 33:1-5. 1989. sheep. states, its past, Remote body temperature Desert Bighorn Counc. Chabot, D., M. Bayer, and A. de Roos. 1991. Instantaneous heart rates and other techniques introducing errors in the calculation of heart rate. Can. J. Zool. 69:1117-1120. Clabough, D. L., and C. R. Swanson. 1989. fasting-induced bradycardia of cattle. Heart rate spectral analysis of Am. J. Physiol. 257:R1303-1306. Coates, K. P., J. C. Undem, B. C. Weitz, J. T. Peters, and S. D. Schemnitz. 1990. A technique for implanting heart-rate transmitters in bighorn sheep. Bienn. Symposium North. Wild Sheep Goat Counc. 7:143-148. Confer, A. W., R. J. Panciera, and D. A. Mosier. 1988. pasteurellosis: Immunity to Pasteurella haemolytica. Assoc. 193:1308-1316. Bovine pneumonic J. Am. Vet. Med. Demura, H., C. D. West, C. A. Nugent, K. Nakagawa, and F. H. Taylor. 1966. sensitive radioimmunoassay for plasma ACTH levels. J. Clin. Endocrinol. Metabol. 26:1297-1302. A Fran~oise Domingue, B. M., P. R. Wilson, D. W. Dellow, and T. N. Barry. 1992. Effects of subcutaneous melatonin implants during long day length on voluntary feed intake, rumen capacity and heart rate of red deer (Cervus elaphus) fed on a forage diet. Br. J. Nutr. 68:77-88. Fulkerson, W. J., G. J. Sawyer, and C. B. Gow. 1980. Investigation of ultradian and circadian rhythms in the concentrations of cortisol and prolactin in the plasma of dairy cattle. Aust. J. Bio. Sci. 33:557-561. Geist, V. 1971. Mountain University of Chicago sheep a study in behavior and evolution. Press, Chicago, Ill. 383pp. The Harlow, H. J., E. T. Thorne, E. S. Williams, E. L. Belden, and W. A. Gern. 1987a. Adrenal responsiveness in domestic sheep (Ovis aries) to acute and chronic stressors as predicted by remote monitoring of cardiac frequency. Can. J. Zool. 65:2021-2027. Harlow, H. J., E. T. Thorne, E. S. Williams, E. L. Belden, and W. A. Gern. 1987b. Cardiac frequency: a potential predictor of blood cortisol levels during acute and chronic stress exposure in Rocky Mountain bighorn sheep. Can. J. Zool. 65:2028-2034. �117 Holley, D. C., D. A. Beckman, and J. W. Evans. on the circadian rhythm of ovine cortisol. Hudson, R. J. 1973. phytohemagglutinin 51:479-482. 1975. Effect of confinement J. Endocrinol. 65:147-148. stress and in vitro lymphocyte stimulation by in rocky mountain bighorn sheep. Can. J. Zool. Kreeger, T, J., D. Monson, V. B. Kuechle, U. S. seal, and J. R. Tester. 1989. Monitoring heart rate and body temperature in red foxes (Vulpes vulpes). Can. J. Zool. 67:2455-2458. Kennaway, D. J., J. M. Obst, E. A. Dunstan, and H. G. Friesen. 1981. Ultradian and seasonal rhythms in plasma gonadotropins, prolactin, cortisol, and testosterone in pinealectomized rams. Endocrinology 108:639-646. MacArthur, R. A., V. Geist, and R. H. Johnston. 1982. Cardiac and behavioral J. Wildl. Manage. responses of mountain sheep to human disturbance. 46:351-358. MacArthur, R. A., R. H. Johnston, and V. Geist. 1979. Factors influencing heart rate in free-ranging bighorn sheep: a physiological approach to the study of wildlife harassment. Can. J. Zool. 57:2010-2021. Manfredo, M. J. and R. A. Larson. 1993. Managing for wildlife recreation experiences: an application in Colorado. Wildl. 21:226-236. viewing Soc. Bull. Mautz, W. W., and J. Fair. 1980. Energy expenditure and heart rate activities of white-tailed deer. J. Wildl. Manage. 42:715-738. for McNatty, K. P., M. 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York. 343pp. Wildlife feeding and nutrition. Academic Press, Roth, J. A. 1985. Cortisol as mediator of stress-associated immunosuppression in cattle. Pages 225-242 in G.P. Moberg, ed. Stress. American physiological Society, Bethesda, Maryland. SAS Institute, Inc. Cary, Inc. N.C. 1988. SAS/STAT 1028pp. user's guide, release New Animal 6.03 ed. SAS Inst., �118 Selye, H. 1950. Montreal. The physiology and pathology of exposure to stress. Acta, Spraker, T. R., and C. P. Hibler. 1982. An overview of the clinical signs, gross and histological lesions of the pneumonia complex of bighorn sheep. Bienn. Symposium North. Wild Sheep and Goat Counc. 3:163-172. Spraker, T. R., C. P. Hibler, G. G. Schoonveld, and W. S. Adney. 1984. pathologic changes and microorganisms found in bighorn sheep during a stress-related die-off. J. Wildl. Dis. 20:319-327. Stemp, R. 1982. Heart rate responses of bighorn sheep to some environmental factors. Bienn. Symp. North. Wild Sheep and Goat Counc. 3:314-319. Stephens, D. B. 1980. Stress and its measurement in domestic animals: a review of behavioral and physiological studies under field and laboratory situations. Pages 179-210 in Advances in veterinary science and comparative medicine. Academic Press, Inc. Cambridge, England. Thorne, E. T., 1982. Pasteurellosis. Pages 72-77 in E. T. Thorne, N. Kingston, W. R. Jolley, and R. C. Bergstrom, eds. Diseases of wildlife Wyoming. Wyoming Game and Fish Dep., Cheyenne. in Thun, R., E. Eggenberger, and K. Zerobin. 1990. 24-hour profiles of plasma cortisol and testosterone in the male dog: absence of circadian rhythmicity, seasonal influence and hormonal interrelationships. Reprod. Dom. Anim. 25:68-77. Turner, J. C. 1984. Diurnal periodicity of plasma cortisol and corticosterone in desert bighorn sheep demonstrated by radioimmunoassay. Can. J. Zool. 62:2659-2665. Wallace, M. C., P. R. Krausman, D. W. DeYoung, and M. E. Weisenburger. 1992. Problems associated with heart rate telemetry implants. Desert Bighorn Council Trans. 36:51-53. �119 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS State of Project Work Colorado No. Plan W-153-R-8 No. Job No. Period Author: REPORT Covered: Mammals Research lA Multispecies 1 Animal and Pen Support Facilities for Mammals July Investigations Research I, 1994 - June 30, 1995. M. A. Wild. Personnel: D. L. Baker, P. E. Bleicher, A. L. Case, Johnson, D. W. Mock, and J. L. Schaefer. W. S. Graffam, L. ABSTRACT The Colorado Division of Wildlife's Foothills Wildlife Research Facility (FWRF) maintained captive animals (up to 136 wild ungulates of 5 species, 3 domestic ungulates, and 50 migratory game birds of 4 species) and facilities supporting 11 major research projects. In fall 1994, we recruited 12 individuals into our captive herd. In addition, 3 domestic goats were added to the research herd and 5 adult pronghorn were returned to the herd from the Sybille Wildlife Research Unit. In FY1995, reductions in animal numbers were due to mortalities (5 adults and 7 neonates) and release of the captive waterfowl. The most common cause of death in adults was Chronic Wasting Disease (CWO) (n=2) and in neonates hypothermia due to a late spring snow (n=2) and elective euthanasia (n=2). FWRF was inspected by USDA APHIS and found to be in compliance with federal animal welfare standards. A formal Preventive Medicine protocol was established. This protocol and the CWO protocol constitute the basis for animal health care at FWRF. Routine animal care and facility maintenance programs were conducted as previously described with an emphasis on quality and conservation. Volunteers contributed 392 man hours. Numerous improvements were made to facilities at FWRF to increase usefulness, efficiency, quality, and/or safety. The single largest project was the addition of a lab and surgery suite to the west side of the facility. We performed 3 experiments concerning animal maintenance. First, we completed a .2 year study investigating the effects of oral vitamin E supplementation on bighorn sheep •. '~esults indicated that serum vitamin E levels could be increased by oral supplementation via the pelleted ration~ Based on these results and the fact that vitamin E is a safe and inexpensive feed additive, the'vitamin E level in pelleted supplement for all species at FWRF was increased initially from 0 to 70 IU/kg, then to 500 IU/kg. In the second experiment, we examined the reproductive tracts of 6 pregnant pronghorn does using ultrasonography between about 2-10 weeks of gestation. Embryos were observed in 5 of 6 does by 3 weeks of gestation (the sixth was not observed until 5 weeks of gestation). A reduction in litter size was observed in 5 of 6 does between 7-9 weeks of gestation. Finally, in the third experiment we determined a safe and effective technique to induce abortion in mule deer in the first and early second trimester of pregnancy. Fenprostalene (0.5 mg) , divided into 2 subcutaneously administered doses 4 hr apart combined with a single 14 mg dose of dexamethasone produced abortion in 2 of 4 does; however, when the dose of fenprostalene was increased to a total 1 mg, 4 of 5 aborted after 1 treatment and the fifth doe aborted after 2 treatments. No adverse reactions were observed. ��121 ANIMAL AND SUPPORT FACILITIES Margaret FOR MAMMALS RESEARCH A. Wild P. N. OBJECTIVES 1. To provide and maintain captive wildlife populations and facilities supporting CDOW's Terrestrial Wildlife Research Program, as well as programs of CDOW cooperators. 2. To develop improved animal and facility management practices provide maximum research opportunities at minimum cost. 3. To enhance needs. facilities to serve a growing diversity that will of anticipated research SEGMENT OBJECTIVES 1. Maintain and improve animal research 2. Coordinate 3. Maintain up to 30 elk, 35 mountain sheep, 30 pronghorn, white-tailed deer, and 80 ducks according to prescribed animal welfare for the use in research experiments. 4. Conduct management experiments to increase efficiency and efficacy feeding, husbandry, and maintenance of research animals. 5. Follow a conservation-oriented approach for providing services to operate research facilities. 6. Follow Standard Operating facility records. all rearing, training, and holding maintenance, Procedures facilities. and research in maintaining activities. 45 mule deer, 15 standards of utilities detailed of and animal and METHODS AND MATERIALS Routine animal care and facility maintenance programs supporting new and ongoing Terrestrial Wildlife Research Program projects were conducted as previously described. We emphasized a quality and conservation-oriented approach in this work by striving to increase efficiency and longterm benefit from the programs and projects undertaken. Specifically, we performed the following tasks: ANIMAL MAINTENANCE General: Again this year, routine feeding and caretaking of research animals, including health observations, training, weighing, and clean-up, was performed primarily by well trained work-study and temporary employees, as well as volunteers. FWRF was inspected by USDA APHIS for compliance with federal animal welfare regulations on 15 March 1995. In 1994, two pronghorn fawns were hand-raised at FWRF. Three additional mule deer fawns that were hand-raised by rehabilitators were received by FWRF after weaning. Eight bighorn sheep lambs were dam-raised by tame ewes. In 1995, 10 pronghorn fawns were born to captive does. Fawns were removed from does at about 24 hours of age for bottle-raising. Twelve lambs were born to, and raised by, captive bighorn ewes. �122 NUTRITIONAL MAINTENANCE Feeding protocols: Feeding protocols were as previously described (Wild and Graffam 1994). In FY1995, all bighorn sheep received a special formulation of the pelleted supplement (Baker and Hobbs 1985) with lower copper. The special low copper pellet contains 10 ppm copper in contrast to the 66 ppm contained in the basic formulation provided to other species. Bottle-raising neonates: In 1994, pronghorn fawns received ad libitum evaporated milk until about 70 days of age. At that time, milk offered to fawns was limited to 660 ml/day. Thereafter, the amount of milk offered was reduced by 80-100 ml/day each week until fawns received 230 ml/day. Fawns were weaned at about 102 days of age. Ten pronghorn fawns are being handraised in 1995. Thus far, fawns have received ad libitum evaporated milk. Response of captive Rocky Mountain bighorn sheep to oral vitamin E supplementation: Phase II of the experiment was completed in fall 1994 following the protocol reported by Wild and Graffam (1994). Briefly, ewes received pelleted supplement with projected vitamin E levels of 80 IU/kg for the low level supplementation group (LOWE) and 500 IU/kg for the high level supplementation group (HIGHE). We sampled all available feeds (pellets, hay, and pasture) at monthly intervals to estimate daily vitamin E intake/kg feed by bighorn sheep in each treatment group. Blood samples were collected at monthly intervals (13 December 1993 - 12 September 1994) from ewes and every 2 weeks from lambs (24 hr of age - 12 september 1994). We compared plasma levels of vitamin E between treatment groups and also between Phase I and Phase II of the study using least square means and analysis of variance for repeated measures using the general linear models procedure of SAS (SAS 1988). HEALTH MAINTENANCE General: Over the past several years a comprehensive preventive medicine program has been established for animals at FWRF. The goals of the program are to: 1) provide captive research animals that are physiologically representative of their free-ranging counterparts and 2) to minimize disease, injury, and stress, to optimize production, and to maximize animal welfare. A few aspects of the program have been reported in detail previously. A summary of general components of the animal care program at FWRF that relate to preventive medicine are included in Addendum A. Surveillance for chronic wasting disease (CWO) is an important aspect of the program in cervids. Early embryonic loss in pronghorn: In 1968, O'Gara reported a unique aspect of the reproductive strategy of pronghorn. From postmortem observations.in does, he observed that the number of implanted embryos was generally 2 in each uterine horn (total 4) in does collected at about 4 weeks of gestation but only 1 in each horn (total 2) in does at about 8 weeks of gestation. The mechanism suggested for this reduction in litter size was the killing of one embryo by its larger sibling in the same uterine horn. Prenatal fatal sibling competition has not been documented in any other mammalian species. The evolutionary explanation of this event may be 1)that the initial overproduction is adaptive and the dam avoids missing a reproductive opportunity or 2) the mother overproduces for quality control, i.e., as an early means of identifying genetically superior offspring. Conversely, the siblicide may be counter to maternal fitness but is a selfish action to improve an embryo's personal fitness. To better understand this strategy, we must first document reduction of litter size using repeated observations in individual pronghorn. Further, we must identify when the embryonic loss occurs. Here, we began collection of these data. We used real time B mode ultrasonography to examine the reproductive tracts of 6 pregnant does at about weekly intervals between 20 October and 2 December 1994. Does were immobilized with about 170 mg Telazol administered intravenously, placed in sternal recumbency, and a intrarectal probe inserted �123 for examination. We collected descriptive data on characteristics of the pronghorn reproductive tract during early pregnancy. We estimated the number of embryos/fetuses present using a conservative count of individuals, hence, we were more likely to underestimate than to overestimate litter size. Size of structures were measured using electronic calipers. Pharmaceutical-induced abortion in mule deer: The ability to safely and reliably induce abortion in captive wild ungulates is useful for husbandry as well as research purposes. At FWRF, we require a means for pharmaceuticalinduced abortion in mule deer; however, no published reports are available for this species. Prostaglandin F~ (PGF~) is a safe and effective abortifacient in cattle when given as a single 25 mg dose (Hudson 1986). A preliminary study in white-tailed deer (Becker and Katz 1994) suggested that PGF~ alone was not effective but that 50 mg PGF~ combined with 15 mg betamethasone may be an effective abortifacient. Although PGF~ (as Lutalyse, Upjohn Co., Kalamazoo, MI 49001) has been used as an abortifacient in llamas, adverse reactions have been observed (L. Johnson and M. Fowler, pers. corom.). Fenprostalene (Bovilene, Syntex Animal Health, West Des Moines, IA 50226), a synthetic analogue of PGF~, is believed to be an effective, yet safer, alternative to PGF~ for use in llamas (L. Johnson and M. Fowler, pers. corom.). We evaluated the efficacy and safety of fenprostalene for inducing abortion in 7 captive mule deer between about 65-113 days of gestation. Prior to treatment, all does were sedated with 60-90 mg xylazine administered intramuscularly and confirmed pregnant using real-time B mode ultrasonography with the probe placed intrarectally. Daily direct observation and ultrasonography about 7 days post-treatment were used for evaluation of treatments. Does were held in 50 m2 isolation pens for 7 days following their initial treatment. Thereafter, does were housed together in an about 5-ha pasture. Initially, 4 does each received 0.25 mg fenprostalene and 14 mg dexamethasone (Vedco, Inc., St. Joseph, MO 64504» followed by an additional 0.25 mg fenprostalene 4 hr later, all administered subcutaneously (SQ). Due to equivocal results, subsequent treatment dosages were increased to 0.5 mg fenprostalene with 14 mg dexamethasone followed by an additional 0.5 mg fenprostalene 4 hr later. This high dose protocol was administered as the initial treatment to 3 does and as a repeat treatment after failure of the low dose in 2 does. FACILITY MAINTENANCE/REPAIRS/IMPROVEMENTS A variety of scheduled and unscheduled maintenance and repair activities were necessary to support facility operation and ongoing research programs. We worked toward a conservation-oriented approach for facility care by undertaking preventive maintenance projects, and performing high-quality new construction and repairs to existing facilities. RESEARCH PROJECTS Facility operations offered support for pilot studies, student special studies, and CDOW and cooperative research experiments that were initiated, conducted, or continued using ~~F animals and facilities throughout the year. EDUCATIONAL CONTRIBUTIONS Facility tours and educational lectures were provided to school, university, and professional groups visiting FWRF. We emphasized the importance of maintaining captive wildlife for performing controlled experiments and the contributions made by research projects performed at FWRF. FWRF animals and facilities were also used occasionally for hands-on training for professional groups. �124 RESULTS AND DISCUSSION ANIMAL MAINTENANCE General: When volunteers were carefully selected and trained similarly to paid employees, their contribution was remarkable. Eight volunteers contributed a total of 392 hours of work at FWRF during FY1995. These volunteers performed primarily caretaker tasks and also assisted in weighing and collecting samples from animals. In addition, volunteers were recruited for a one day painting project that totalled 36 man hours. Contributions by volunteers represented a savings of about $4015 to FWRF (vs. cost of temporary employees). The animal welfare inspection by USDA APHIS revealed one minor infraction (out dated drugs in an emergency kit) that was corrected immediately. At the close of FY1995, FWRF maintained 19 elk, 39 bighorn sheep, 11 whitetailed deer, 44 mule deer, and 21 pronghorn. During FY1995, we recruited 7 bighorn sheep, 3 mule deer, and 2 pronghorn into our captive herd. Five of the 6 adult female pronghorn previously transferred to the Wyoming Game and Fish Sybille Wildlife Research Unit were returned to FWRF (one pronghorn, SL87, died at Sybille). In addition, 3 domestic goats were added to the research herd. In ungulate species, 12 mortalities occurred. All captive waterfowl were released to the wild. The US Fish and Wildlife Service provided financial support to maintain 37 mule deer at FWRF. The USDA Animal Damage Control provided financial support for 11 white-tailed deer at FWRF. The watchable wildlife program financially supported 3 domestic goats and 1 bighorn sheep equivalent. NUTRITIONAL MAINTENANCE Feeding protocols: All species maintained reasonable body condition on available diets. Mule deer continue to maintain generally fair body condition; however, we have begun to investigate alternate feedstuffs that more closely resemble native diets in an attempt to improve nutritional condition of mule deer. At the start of FY1995 we will begin feeding a browser maintenance diet (PMI Feeds, Inc.) in addition to our basic diet in 2 groups of mule deer. The browser diet will be evaluated by following clinical health and weight change in these deer vs. deer on the traditional diet. Bottle-raising neonates: Weaning weights of 2 pronghorn fawns were 23.2 and 21.9 kg. Weights at about 104 days did not differ (P > 0.05) from pronghorn bottle-raised in the 2 previous years. The gradual weaning method is more similar to natural weaning and appears to stimulate fawns to consume more dry feeds prior to weaning. Response of captive Rocky Mountain bighorn sheep to oral vitamin E supplementation: Despite attempts to standardize vitamin E supplementation, intake of vitamin E differed from predicted treatment values. Based on analyses of the complete diet, bighorn sheep in the LOWE group received about 197 IU vitamin Ejheadjday while bighorn sheep in the HIGHE group received about 375 IU vitamin Ejheadjday. Variation in the levels of vitamin E fed occurred due to inaccuracy in the milling process for pellets, loss of vitamin E from pellets over time, and changes in pasture vegetation. Mean plasma vitamin E levels did not differ (P > 0.05) between the LOWE and HIGHE group. Least square means were 108 ugjdl (SE=6.7) for LOWE and 123 ugjdl (SE=7.2) for the HIGHE group. These levels at least met the minimum normal value reported for domestic sheep; PuIs (1994) reported vitamin E values <200 ug marginal and <100 ugjdl deficient in domestic sheep. This is in contrast to results from Phase I of the experiment. In Phase I, bighorn sheep received an estimated 75 IU vitamin Ejheadjday for the control, low vitamin E group, and about 107 IU vitamin Ejheadjday in the high vitamin E treatment group at maintenance. This level of treatment resulted in least square means of 62 ugjdl (SE=5.7) for the low E group and 71 ugjdl (SE=6.2) for the high E group. No difference (P > �125 0.05) was observed between treatment groups in Phase I and neither treatment level resulted in plasma vitamin E values out of the range considered deficient in domestic sheep. When treatments are combined, plasma vitamin E levels were higher (P=0.0003) for bighorn sheep in Phase II than in Phase I of the study. This increase can also be appreciated in plots of plasma vitamin E levels (Fig. 1). This difference may be because levels of vitamin E supplemented in both treatment groups of Phase II were higher than the high vitamin E treatment in Phase I. Conversely, the difference may have been due to the decrease in level of copper supplemented in feed (from 66 to 11 ppm) between Phase I and Phase II or due to an unknown time effect. The effects on lambs from vitamin E supplementation of their dams was difficult to determine due to the small number of lambs born to treatment ewes. In Phase I, 3 lambs were available and in phase II, 8 lambs were available. Vitamin E levels were more consistent over time in Phase II versus Phase I (Fig. 2). No respiratory disease was observed during the treatment periods. Although beneficial results of vitamin E supplementation in this study were equivocal, it appeared that high levels of vitamin E supplemented in the pelleted feed increased plasma vitamin E values. Based on these results, and the fact that vitamin E is a safe and inexpensive additive, the vitamin E level in pelleted supplement for all species at FWRF was increased from about 70 to 500 IU/kg. HEALTH MAINTENANCE General: Overall, captive wildlife maintained at FWRF remained healthy throughout the year. During FY1995, 5 mortalities occurred in adult ungulates and 7 in neonates at FWRF (Table 1). The most common cause of death in adults was CWO (n=2). In neonates, elective euthanasia was performed in 2 intractable male pronghorn fawns. Hypothermia associated with late spring snows was the likely cause of death of 2 bighorn lambs. In contrast to many recent years, no deaths in bighorn sheep were attributable to respiratory disease; however, 3 lambs did develop mild pneumonia after weaning. Lambs were treated with oxytetracycline (LA200, 20 mg/kg SQ) as needed. We followed newly implemented protocols for the preventive medicine program and management of CWO (Wild and Graffam 1994). However, we did make one exception to the CWO protocol by introducing domestic goats to FWRF. The threat posed by goats was determined to be negligible, especially if goats were isolated from other species at FWRF. In spite of the strict CWO protocol, 2 cases occurred in FY1995. An a-year-old cow elk was diagnosed with CWO in February 1995. The cow had shown subtle behavioral changes (reduction in socialization with cohort and reduced tractability) for about 6 months and a marked stepwise weight loss for 3 months. This was the fourth case of CWO in elk since the depopulation in 1985. The first case of Cwo diagnosed in a deer at FWRF since the depopulation in 1985 occurred in October 1994. The 3-year-old mule deer doe was born at FWRF and never left the facility. She exhibited an inability to regain weight that was lost over the previous winter and behavioral changes. Behavioral changes were not entirely typical of CWO (Williams and Young 1992) and were characterized simply by lack of tractability and slight hyperexcitability. The pen where the doe was housed was cleaned and disinfected. Although CWO has been diagnosed in only 1 captive mule deer since the depopulation, another chronic progressive disease has been responsible for the euthanasia of 6 young adult mule deer since 1991, including 1 in March 1995. In addition, 32-year-old does currently show clinical signs of the disease. Cause of the disease is undetermined, but clinical signs include fluid bloat, esophageal reflux, failure to gain weight, and depression. These signs somewhat resemble CWO, however, on histopathology characteristic spongiform changes are not observed. As more sophisticated diagnostic tests for CWO • �126 become available, samples from these mule deer will be further examined. Future management experiments at FWRF could also be used to investigate if the disease may have a nutritional basis. Early embryonic loss in pronghorn: Using date of parturition, we backcalculated 252 days (Pojar and Miller 1984) to determine conception date. Using this information, we were able to estimate the stage of gestation when specific events occurred. We examined pronghorn weekly between about 2 and 10 weeks of gestation. At least 1 embryo was observed in 5 of 6 does by 3 weeks gestation using ultrasound (the sixth was not observed until 5 weeks of gestation). We documented embryonic loss in 5 of 6 pronghorn (Table 2). We observed embryos that appeared "unhealthy" (solid echoic areas on ultrasound) in 3 of the pronghorn. In each case, one less embryo was observed on subsequent examinations suggesting loss of the "unhealthy" embryo (Table 2). We estimated that embryos were lost at 7-9 weeks of gestation. This coincides with O'Gara's (1968) observations in postmortem examinations. Although we did not identify the mechanism of loss, we did collect valuable data on the pronghorn reproductive tract during early pregnancy, document a reduction in litter size in vivo, and confirm the stage of gestation when the loss occurs. This information will aid in further study of pronghorn early embryonic loss and reproductive strategies. Pharmaceutical-induced abortion in mule deer: No adverse reactions to the abortifacient were observed. Abortion occurred in 5 of 7 does after 1 dosage protocol (Table 3). Efficacy increased with the high dose protocol (Fig. 3). Two does (animal ID W92 and Y92) did not abort using the initial low dose protocol. These does were re-examined using ultrasound 27 and 22 days, respectively, post-initial-treatment. Although the fetus of Y92 appeared healthy, no heart beat or movement were observed in the fetus of W92 suggesting that it had died in utero at some point between 7 and 27 days posttreatment. Both does were treated with the higher dosage protocol and returned to their pasture. Doe W92 aborted 5 days after the second treatment. Although doe Y92 did not abort following the second treatment, mucoid and serosanguinous vulvar discharges were noted and the fetus appeared "unhealthy" (slow heart rate, decreased volume of amniotic fluid) on ultrasound examination 9 days post-treatment. Y92 was treated with the high dosage protocol again 44 days post-initial-treatment. Abortion occurred 3 days posttreatment. Of the 5 does that aborted after their initial treatment, time from treatment to abortion was 5 days in 2 does and 10 days in 3 does (Table 3). In the latter group, all fetuses were alive 7 days post-treatment based on ultrasound findings. Abortion generally occurs in 4-5 days in domestic ruminants treated with chemical abortifacients. The discrepancy in time to abortion in our mule deer remains unclear, but may be related to "protection" of the pregnancy by an adrenal source of progesterone during periods of stress (e.g., while housed in isolation pens). To investigate this hypothesis, we will measure cortisol and progesterone levels in fecal samples collected from mule deer during this trial. Results will be presented in subsequent reports. Does showed mucoid or serosanguinous vulvar discharge and abdominal pressing near the time of abortion. Fetuses were recovered from 5 of the 7 does. The remaining does likely consumed the expelled fetuses. Of the 8 fetuses recovered, crown-rump length ranged from 87-240 rom and estimated age of fetuses based on a growth curve (Hudson and Browman 1959) ranged from 65-113 days and increased with date of abortion (Table 3). Based on these measurements, conception by does likely peaked during the last 1-2 weeks in November. This time of peak conception corresponds with data from previous years for does at FWRF. Based on these results, a 1 mg dose of fenprostalene divided into 2 SQ injections 4 hr apart combined with a single 14 mg dose of dexamethasone appears to be a safe and effective abortifacient. Follow-up ultrasound examination revealed no apparent abnormalities in the reproductive tracts of �127 treated does. impaired. FACILITY This suggests that future reproductive capability should not be MAINTENANCE/REPAIRS/IMPROVEMENTS In addition to numerous daily repairs and maintenance projects, we performed several major improvements. Significant maintenance/repair/improvement projects completed at FWRF this year included: Construction of facility. Installation of of the facility. - Construction of Construction of a new lab and surgical a 1000 gallon septic suite tank on the west and leach side of the field on west side feed storage area and covered feed area in pen W3. cut-off fences to provide catch pens in pastures A and W. - - Construction of a holding pen for bighorn sheep trials. Preparation of a goat pen in location of old cow pasture. Repair and painting of shelters and feed sheds on west side of facility using a volunteer day. Revegetation of pen E4. Replacement of old, dangerous pen fencing in pens E1, E2, W4, and W7. Replacement of older section of hay shed. Repair and preventive maintenance to west side alleyway and isolation pens. Replacement of visual barriers in east isolation pens. Replacement of vehicle gate in pen W4. Landscaping for erosion control in pen W2. Replacement of a corroded water pipe that burst in the isolation pens. Addition of road base to roughest/muddiest portions of facility roads. Repairs to old roofs after wind damage. RESEARCH PROJECTS In addition to ongoing facility management experiments and improvements described above, the following pilot studies, special studies, and research experiments were initiated, conducted, or continued using FWRF animals and facilities this year: - Response of captive supplementation--W. Rocky Mountain bighorn sheep to oral vitamin Graffam, M. Wild, and N. Irlbeck. - Effect of stress on Pasteurella haemolytica cytotoxin dependent of neutrophils from bighorn sheep--B. Kraabel and M. Miller. Feasibility of using carriers--L. Miller, Early embryonic liposomes B. Johns, as deer immunocontraceptive R. D. Thompson (ADC). loss in pronghorn--D. Mock, M. Wild, E killing oral vaccine and L. Johnson. - Comparative effects of Saccharomyces cerevisiae and a commercial supplement on voluntary feed intake and digestibility of low quality grass hay fed to captive wapiti (Cervus elaphus)--D. Murphy, N. Irlbeck, and D. Baker. Surgical implantation of heart rate transmitters model for bighorn sheep--M. Wild, D. Piermattei, - Determination of an effective HCl immobilization in elk--T. in domestic goats as a D. Baker, and B. Heath. dose of naltrexone to reverse Holz, M. Miller, W. Lance. etorphine �128 - Experimental evaluation of a multivalent Pasteurella haemolytica toxoidbacterin (AI, A2, T10) in captive bighorn sheep (Ovis canadensis)--M. Miller, J. Conlon, and A. Ward. - Mule deer pituitary stimulation by a gonadotropin releasing hormone analog: comparison of intramuscular delivery via injection and ballistic implant--M. Miller, D. Baker, M. Wild, and T. Nett. - Effects of condensed tannins and curl leaf mountain-mahogany on digestibility and salivary tannin-binding protein concentration mountain sheep--J. Peterson Alldredge and D. Baker. - Heart rate as a potential indicator of stress D. Baker, D. Piermattei, and B. Heath. EDUCATIONAL in bighorn in sheep--M. Wild, CONTRIBUTIONS FWRF provided formal educational instruction for 2 grade school classes, 3 high school classes and 1 high school career day, 2 university classes, 1 Pueblo Youth Naturally group, and 2 Youth in Natural Resources groups. Animals and facilities were used for hands-on training with 1 professional group. Numerous other informal tours were provided individually to visiting professionals. LITERATURE CITED Baker, D. L. and N. T. Hobbs. 1985. Emergency feeding of mule deer during winter: tests of a supplemental ration. J. Wildl. Manage. 49:934-942. Becker, S. E., and L. S. Katz. (PGF~) on pregnancy status 1994. Effects in white-tailed of exogenous prostaglandin-F~ deer. Zoo BioI. 13:315-323. Hudson, P., and L. G. Browman. 1959. Embryonic mule deer. J. Wildl. Manage. 23:295-304. and fetal development Hudson, R. S. 1986. Diseases of the reproductive and urinary 765-818 in J. L. Howard, ed. Current Veterinary Therapy: Practice 2. W. B. Saunders Co, Philadelphia, Pa. of the systems. Pages Food Animal O'Gara, B. W. 1968. A study of the reproductive cycle of the female pronghorn (Antilocapra americana Ord.). PhD Thesis, University of Montana, Missoula. 161 pp. pojar, T. M. and L. L. W. Miller. pronghorn. J. Wi1d1. Manage. PuIs, R. 1994. Mineral levels International, Clearbrook, 1984. Recurrent 48:973-979. in animal B. C. health. SAS Institute, Inc. 1988. SAS/STAT user's Inst., Inc. Cary, N. C. 1028pp. Wild, guide, estrus Second and cycle ed. releases length in Sherpa 6.03 ed. SAS M. A, and W. S. Graffam. 1994. Animal and pen support facilities for mammals research. Colorado Div. Wildl. Res. Rep., WP1a, J1, Jul 1993 Jun 1994, Fort Collins. Williams, E. S., and S. Young. 1992. Spongiform encephalopathies Cervidae. Rev. Sci. Tech. Off. Int. Epiz. 11:551-567. in �129 Table 1. Summary Species of mortalities Animal in hoofstock Age (yrs) ID at FWRF during Cause FY 1995. of Death C995 E894 o o L795 M395 QG95 o o Elk C8G 8 Chronic Pronghorn BN91 Ya95 Yb95 3 o o Perforated Population Population Qb91 R91 3 3 Chronic Chronic Br93 1 study protocol-transmitter Bighorn sheep Mule deer Domestic goat Stillborn Ruptured urethra secondary to naval ilIa Hypothermiab Hypothermiab Malnutrition associated with mammary abnormality in ewe o Wasting Diseasea large colon control/intractable control/intractable fawna fawn8 bloat, failure to thrive8 Wasting Diseasea failure8 Euthanatl.zed bunconfirmed diagnosis Table 2. Numbers of embryos observed 20 October-02 December 1994. Animal BL91 ID Week of Gestation8 5 7 G in pronghorn 3 4 0 2 3 3 2 1 1b 1 2 0 2 2 2 2 0 2 2 3 2 2 2 2 2 3c 2 2 1 2 2 3 3c 3 1 3 0 2 3c 2 DD91 0 SE91 Y091 ultrasonography 2 BN91 NK91 using 1 8 9 2 8Week of gestation estimated based on conception date 252 days prior to f,arturition (Pojar and Miller 1984). Subsequent ultrasound examination revealed 2 fetuses. COne embryo appeared "unhealthy" (bright echoic area on ultrasound) • from 10 2 �130 Table 3. Results Animal 10 of treatment with abortifacient Treat date Abort date No. fawnsb E91 10 2-02-95 2-07-95 2 092 10 2-07-95 2-17-95 2 R86 hi 2-14-95 2-19-95 A91 hi 2-14-95 2-24-95 S90 hi 2-17-95 2-27-95 W92 Y92 10 hi 2-02-95 3-01-95 3-06-95 10 hi hi 2-07-95 3-01-95 3-23-95 3-26-95 in captive Sex mule deer at FWRF. Length C-R (rom) Est. age (days)c 87 nd 65 65 f 145 83 nd nd nd 1 f 155 86 2 m m 180 175 100 100 nd nd nd m 240 113 1 ~ne dose of 14 mg dexamethasone combined with either 0.5 mg (10) or 1.0 mg (hi) fenprostalene divided into 2 doses 4 hr apart administered SQ. bNumber determined by ultrasound and confirmed by recovery of aborted fetuses except where noted. Coetermined from growth curve (Hudson and Browman 1959). dNot determined. ~umber determined by ultrasound only; fetuses not recovered. �131 A 200~------------------------------------------------~ 150 o L Q) .c Q. o o o Lambing 100 l- I o ..c Q. « 50 o E (f) o 0... O+-~-+~~~~~~+-~-+~~~-+~--+-~-+~ o 28 56 84 112 140 168 196 224 252 280 308 336 364 .-·LOWE 0·······0 HIGHE Day of Study B oI..... 150 Q) ..c Q. o o o 100 I t._.I o ..c u Q. « • 50 1 o E 0 1 (f) o [L Lambing O+-~-+~~r-+-+-~-+~--r-+-;-~-+~~r-+--r-+_'--~+-~ 390 418 446 474 502 530 558 Day of Study 586 614 642 670 698 .-. 0·· ····0 LOWE HIGHE Fig. 1. Mean plasma vitamin E (alpha-tocopherol) levels of bighorn ewes at FWRF during vitamin E supplementation study A) Phase I and B) Phase II. Day 0 represents 1 October 1992 (Phase I initiation) and Day 439 represents 13 December 1993 (Phase II initiation). Error bars indicate ±1 SE. �132 A 400 ------"0 <, 350 0' ::J • 300 0 I.... OJ 250 ..c a. 0 0 200 0 l- I 150 0 ..c a. <i: 100 0 E 50 (/) 0 o, 0 200 228 256 284 312 340 Date of Blood Sampling 368 396 .-. LOWE 0······· 0 HIGHE (n=1 ) (n=2) B 450 --. 400 "0 <, 0' ::J ..__.. 350 0 I.... OJ 300 ..c a. 0 u 0 I- I 250 200 0 ..c a. <i: 0 E (/) 0 0: 150 100 50 0 550 578 606 634 662 Date of Blood Sampling 690 718 746 .-. LOWE 0········0 HIGHE (n=3) (n=S) Fig. 2. Mean plasma vitamin E (alpha-tocopherol) levels of bighorn lambs at FWRF during vitamin E supplementation study A) Phase I and B) Phase II. Day 0 represents 1 october 1992. Error bars indicate ±1 SE. �133 _ Aborted ~ Non-aborted 5 4 ~ o 3 -•... "C o CD .c E :;, z 2 1 o low High Fenprostalene dose Fig. 3. Number of abortions occurring in captive mule deer at FWRF following initial treatment with fenprostalene at low (total 0.5 mg) and high (total 1.0 mg) doses in combination with 14 mg dexamethasone. �134 ADDENDUM A PREVENTIVE MEDICINE PROGRAM FOR RESEARCH ANIMALS FOOTHILLS WILDLIFE RESEARCH FACILITY Care of Adult AT Animals Each animal is checked at least once daily by a trained caretaker (See FWRF Animal and Facility Protocols for information provided in training). During periods of special concern, e.g., during the period of parturition, animals are monitored more frequently. Any abnormalities are recorded and a veterinarian is contacted immediately with concerns related to animal health. Further, all animals are checked by a veterinarian at least once every 2 weeks. Veterinary care is provided promptly. Diagnostic tests and treatments are provided as needed for the benefit of individuals and to minimize risk to the captive population. Every animal that dies at FWRF receives a complete postmortem examination. Postmortem examination provides information not only on the cause of death in an individual, but insights into its prevention and treatment in other animals (for both infectious and "husbandry-induced" mortalities) and screening for presence of other diseases. vaccination and anthelmintic treatment are preformed at least annually, usually in the spring. Standard procedure is vaccination is with a 7-way Clostridium toxoid and treatment with Ivermectin. other treatments are provided as needed based on imminent threats to animal health. Hooves are trimmed as needed. In general, hooves are trimmed 2-3 times per year for bighorn sheep, 1-2 time per year for pronghorn, and once a year for deer. Housing and Feeding of Adult Animals Adult research animals are housed with regard to species, sex, and physical status. Proper pen assignment is important to minimize intraspecific aggression and as a means to provide appropriate diets. Animals brought into FWRF are screened for diseases of concern and held in isolation (generally for ~2 months) prior to mixing with residents. Intact males are generally housed separately from females except when breeding is desired. Antlers are removed from male cervids as soon as possible after the velvet is shed. Potentially bred females are checked for pregnancy status or are assumed to be pregnant for management purposes. Holding and handling facilities are designed and maintained with attention to animal and human safety. Animals are trained to handling facilities to minimize risk of injury. Sanitation is a high priority, especially in feed areas. Animal enclosures are cleaned at weekly intervals. A rodent control program is followed. Special procedures are followed to minimize the spread of Chronic wasting Disease (CWO) (See Wild and Graffam 1994). Type and amount of feed provided are customized based on species and physical status of individuals. Currently, pronghorn and deer receive alfalfa hay and pelleted supplement (Baker and Hobbs 1985), bighorn sheep receive grass hay and pelleted supplement, and elk receive hay cubes, grass hay, and pelleted supplement. Two blends of pelleted supplement are used (Wild and Graffam 1994). The custom blend is provided to all species except bighorn sheep. Bighorn sheep receive a low copper (11 ppm) blend of the same pelleted feed. As a basis, feed amounts follow estimates made by Baker (Miller 1990); however, amounts of pelleted supplement provided to deer and pronghorn are consistently higher than predictions. Feeding programs are evaluated and adjusted based on condition of animals. Condition of animals is determined by subjective assessment and using body mass data collected from most animals at 2 week intervals. This data is also useful in surveillance for CWO. • �Care of Neonatal Animals With a few exceptions, care of neonatal animals follows that described for adults. Neonates are generally handled when about 24 hrs old. At that time a brief physical examination is performed, the neonate is weighed, and iodine is placed on the umbilicus. If the neonate was received from outside FWRF, it is vaccinated with 7-way Clostridium toxoid and treated with Ivermectin. If it is to be dam-raised it is then returned to its mother. Hand-raised neonates are handled as described by Wild and Miller (1991). Strict sanitation procedures are an especially important aspect of neonatal care. Clostridium vaccination is generally repeated for all neonates at about 1 and 4 months of age. Literature Cited Baker, D. L. and N. T. Hobbs. 1985. Emergency feeding of mule deer during winter: tests of a supplemental ration. J. Wildl. Manage. 49:934-942. Miller, M. W. 1990. Animal and pen support facilities for mammals research. Colorado Div. Wildl. Res. Rep., WP1a, Jl, Jul 1989 - Jun 1990, Fort Collins. Wild, M. A., and M. W. Miller. 1991. Bottle-raising wild captivity. Colorado Div. Wildl. outdoor Facts. ruminants in 114. 6pp. Wild, M. A, and W. S. Graffam. 1994. Animal and pen support facilities for mammals research. Colorado Div. Wildl. Res. Rep., WP1a, J1, Jul 1993 - Jun 1994, Fort Collins. ��137 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS State of Project Work Colorado No. Plan No. Job No. Period REPORT Covered: Mammals W-153-R-B Research lA Multispecies 3 Mammals July Investigations 2 Research Administration 1, 1994 - June 30, 1995 Author: R. Bruce Gill Personnel: R. Bruce Gill and Diane K. Hall ABSTRACT Human and fiscal resources were allocated among 13 Mammals 2 research jobs. Highlights of work progress on each study are summarized. All research objectives were accomplished, but expenditures exceeded resources allocated to the Mammals 2 Research Program by approximately 7% ($11,000). Eleven during scientific manuscripts the segment. were published or accepted for publication ��MAMMALS 1 RESEARCH ADMINISTRATION R. Bruce Gill P.N. Objective Administer research studies within the Mammals productivity at the lowest cost. 1 Research Unit for the highest Segment Objectives 1. Lead and administer research on mammalian species in the Mammals Research Program other than deer, elk, and moose. 2 RESULTS Eleven projects were active during the segment and segment objectives successfully completed for all 11. Highlights include: were • Extensive repairs and improvements have been made to the Colorado Division of Wildlife's Foothills Research Facility. It is now one of the premier captive wildlife research facilities in the U.S. currently the facilities support the research of six Mammals research jobs. The facility also is being used to augment research of researchers from the Denver Wildlife Research Center aimed at developing immunocontraceptives for white-tailed deer. • Statewide monitoring of diseases among wildlife populations continues. The distribution of incidents of chronic wasting disease among deer and elk has been analyzed and a preliminary report has been prepared. Seventeen cases of chronic wasting disease (CWO) were confirmed in free-ranging deer and elk in Larimer County during FY 1994-95. Fortynine cases of CWO in free-ranging deer and elk have been confirmed in Colorado since 1981. To date, all but 2 of these cases have been from Larimer County. Statewide samples for the occurrence of brucellosis among elk populations were negative for antibodies to Brucella spp. on the standard card test. • Preliminary tests to compare Pasturella haemolytica isolates from 8 indigenous Rocky Mountain bighorn herds in Colorado suggest that strain of P. haemolytica are carried by healthy populations and may vary within and among wildlife populations. Preliminary results also suggest the combination of nenomic RNA fingerprinting and cytoxicity determination may offer a useful approach for studying the epizootiology of pasteurellosis within and among bighorn herds and may provide insights into strategies for effectively preventing or managing pneumonia epizootics in bighorn populations. • Measures of the rate of growth of a naturally reintroduced pronghorn population in Middle Park, Colorado suggest the population is approaching the natural carrying capacity over its current area of winter distribution. As habitat saturation approaches, there are signs that pioneering of unoccupied winter habitats may be occurring. • Mark-resight estimates were generated for black bears occupying a 465 km2 study area on the Uncompahgre Plateau. Resighting of marked bears was accomplished with cameras activated by active infra-red sensors. Six resighting replications of approximately 14 days produced 564 photographs of black bears. A total of 119 of those photos contained marked bears. A Lincoln-Peterson estimator indicated the total study area population averaged 168 bears (± 20%). Black bear density was estimated to be 36 bears/100 km2• �• In conjunction with ongoing activities in the spatial analysis of elk survival project, a landscape simulator was developed to evaluate a proportional hazard rate model and logistic regression was used to detect annual differences in animal survival in relation to habitat use. • studies continue on the status of kit fox in western Colorado. Eighteen additional foxes were captured and individually marked with radiocollars and/or eartags. To date 31 individual foxes have been captured. Of 37 collared or eartagged foxes caught since 1992, the fate of we is unknown, 7 are known to be dead, and 7 are still alive. Reproductive success of the population has been low. Of 14 adult females capture since 1992, only 6 litters of pups have been produced. • The System for Conservation Planning Project (SCoP) has initiated pilot programs in 3 Colorado counties: Larimer, Summit, and Boulder. In Larimer and Summit Counties, project staff are working with local citizens and planners to design information systems that support land use decisions which preserve and protect wildlife habitats. In Boulder County, studies have been initiated to evaluate the effects of residential development on avian communities in riparian zones. The project staff has been expanded with the recruitment of Dr. Tanya Shenk, a recent graduate of Colorado State University. Prepared by: R. Bruce Wildlife Gill Research Leader �141 Colorado Division Wildlife Research July 1995 of wildlife Report JOB PROGRESS State of Project Work Colorado No. Mammals W-153-R-8 Plan No. Job No. Period REPORT Covered: July Research 1A Multispecies Investigations 6 Monitoring and Managing Health in Colorado Wildlife 1, 1994 - June 30, 1995 Authors: M. W. Miller, Williams C. W. McCarty, C. A. Mehaffy, R. Ford, and E. S. Personnel: W. J. Adrian, J. Bredehoft, G. Byrne, A. Case, D. Clarkson, M. cousins, B. Davies, H. Dietrick, L. Evans, R. Forde, D. Freddy, T. Fulk, D. M. Getzy, K. Green, J. Jackson, K. Kinney, S. Kolus, M. Lamb, C. Leonard, K. Madriaga, B. Olmstead, J. Ritchie, G. Schoonveld, H. Spear, M. L. Stevens, R. Spowart, and T. R. Spraker. ABSTRACT Wildlife populations throughout Colorado were monitored for occurrence of disease using a combination of extensive and intensive approaches. We continued to develop and modify a statewide surveillance program for acquiring, examining, reporting on, and summarizing sporadic wildlife disease cases occurring throughout Colorado. At least 60 carcasses and/or tissue samples representing 55 wildlife cases were submitted for diagnostic examination during July 1994-June 1995. Bronchopneumonia and chronic wasting disease were the most common diagnoses in wild cervids submitted (although case submissions were undoubtedly biased by intensive monitoring for both diseases); all bighorn sheep submitted showed gross and/or histologic lesions of bronchopneumonia. Aside from pneumonia epizootics in bighorn herds near Loveland and Granite and salmonellosis in songbirds, all cases completed to date appear-ed to... represent. isolated cases of trauma or disease. Among carnivore·cases, trauma, bronchopneumonia; and malnutrition 'were were diagnosed. Salmonellosis cases in passerine birds continued through July 1994, and cases were submitted from 10 locations throughout Colorado during April 1994-June 1995. Other mammalian and avian cases appeared to represent isolated incidents of unusual maladies (e.g., hairball in a mountain lion, carcinoma in an elk). For 15 cases, cause of death could not be determined. We continued the annual statewide survey of deer and elk hunters to collect sera for brucellosis screening, and also continued modifying and evaluating our survey program. Of 9,825 elk hunters surveyed, 1,337 (14%) returned blood samples for brucellosis screening from animals harvested throughout Colorado during October 1994-January 1995. Of samples returned, 662 (49%) were usable; marked hemolysis and/or contamination precluded evaluation of the remaining samples. All elk sera tested were negative for antibodies to Brucella spp. on the standard card test. Overall, about 7% of the survey kits distributed to deer or elk hunters in 1993-1994 provided usable samples, as compared to 7% in 1993-1994, 7% in 1992-1993 and 5% in 1991-1992. �142 Seventeen cases of chronic wasting disease (CWO), a spongiform encephalopathy, were confirmed in free-ranging deer and elk in Larimer County during FY 19941995. Forty-nine free-ranging CWO cases have been confirmed in Colorado since 1981; to date, all but 2 of these cases have been from Larimer County (GMUs 9, 191, 19, or 20). We prepared a manuscript summarizing these cases. To obtain reliable estimates for distribution and prevalence of CWO in wild cervids, we continued to survey for CWO in select deer and elk populations throughout Colorado. Brains from about 150 mule deer and 65 elk harvested in GMUs 19 and 20 (DAUs D4, D10/E4, E9), were collected for examination for CWO. Histologic evaluation of all samples has not been completed, but of 343 deer and 212 elk brains from hunter-killed animals collected during 1991-1994 and examined to date, 3 deer were spongiform encephalopathy suspects; ancillary tests for confirmation are in progress. Based on survey data collected to date (and assuming all 3 suspects are confirmed positive), estimated prevalence of CWO in mule deer in DAUs D4/D10 combined is about 0.009 (95% CI 0.002-0.025); 95% CI for prevalence in DAU E4/E9 elk is 0-0.01 based on survey data (0/212) collected to tate. Retropharyngeal and other cranial lymph nodes and tonsils were collected from deer and elk heads through game processing establishments in Durango and Hesperus for gross and histologic examination and culture of cranial lymph nodes; examination of 36 elk and 49 mule deer heads revealed no gross evidence of bovine tuberculosis infection, but abscessed tonsils and/or lymph nodes from 2 elk and 1 deer have been submitted for further microscopic evaluation; no microscopic lesions compatible with bovine tuberculosis have been observed in similar samples from previous years examined to date. Although no evidence ot tuberculosis has been detected in this survey, present sample sizes are insufficient to rule out the possibility of infection; for the number of negative samples examined to date, upper 95% confidence limits for prevalence estimates are about 0.08 for elk, about 0.06 for deer, and about 0.03 for deer and elk combined. Plans for additional surveillance are uncertain. We continued developing a generalized, stochastic, individual-based simulation model of infectious disease in wild ungulate populations. We incorporated parameters to simulate introduction of bovine tuberculosis into a wild elk population and conducted sensitivity analyses of 50-year simulations (n = 500) where an infected elk was introduced into a population of 200 wild elk under assumptions of varying conservancy. As with previous analyses, transmission coefficient (tc) assumptions markedly influenced outcomes: assuming tc = 0.3 new infections/infected individual/year, the probability that tuberculosis became established in simulated populations ranged from 0 to 0.4, depending on survivorship of infectious indivisuals and likelihood of cow-calf transmission. Our preliminary results suggest introduction of bovine tuberculosis into wild elk populations could represent a significant obstacle to nat.LonaI. eradication .goals. �MONITORING M. W. Miller, AND MANAGING C. W. McCarty, WILDLIFE C. A. Mehaffy, HEALTH IN COLORADO R. Ford, and E. S. Williams P. N. OBJECTIVES Develop and implement a program for enhancing statewide efforts to monitor manage health of Colorado's terrestrial wildlife populations. AGREEMENT and OBJECTIVES 1. Modify and improve systems for submitting, diagnosing and reporting sporadic disease cases in wild animals throughout Colorado. 2. Develop and use databases for assimilating and analyzing problems identified through surveillance and surveys. on data on disease 3. Design, conduct, and report results of surveys for brucellosis, tuberculosis, and chronic wasting disease in specific deer and/or elk populations. 4. Provide assistance in Colorado. in investigating and managing wildlife disease outbreaks 5. Design experiments to develop and/or improve techniques for investigating wildlife diseases; begin conducting approved and funded research. Maintaining healthy wildlife populations is a fundamental component of sound wildlife management practices. Habitat degradation, high animal density, extreme weather, and disease can act singly or in combination to compromise the overall health of a wildlife population. As Colorado's wildlife managers, we have developed a variety of tools for monitoring and assessing the effects of habitat loss, animal numbers, and weather on wildlife populations. We have also invested considerably in developing tools to manage these factors to optimize performance of the wildlife populations in our stewardship. In contrast, monitoring and managing the effects of disease on wildlife population performance have received relatively little attention (with a few notable exceptions). This lack of attention may be rooted to some extent in a widely-held belief that wildlife diseases are symptoms of larger underlying population problems that will be resolved if those larger problems are managed properly. Despite this belief, disease can be a significant obstacle to effective and efficient wildlife management in Colorado. Disease outbreaks account for substantial mortality in some wildlife populations. Introduced pathogens have potential to decimate local wildlife popUlations. Some diseases depress wildlife population performance to levels below resource-based carrying capacity. Many wildlife diseases are shared with domestic animals and/or humans, and in some cases wildlife populations serve as reservoirs for these agents. Disease also detracts from the aesthetic value of wild animals, and may convey a perception of mismanagement to uninformed publics. For these reasons, diseases should be regarded as an integral part of wildlife population dynamics and wildlife management. Select wildlife health problems have been monitored in Colorado for more than 30 years. These longstanding efforts have provided useful information on the diseases studied. However, because these efforts have not always been coordinated on a statewide basis, and because some findings have not been widely available to managers and policy makers, applications to overall �144 management programs have been limited. In order to improve our collective ability to manage wildlife health in Colorado, we need a more coordinated and systematic approach for monitoring, investigating, and reporting on health problems in free-ranging wildlife. A more complete understanding of wildlife diseases and their effects on population performance is fundamental to comprehensive wildlife management. Enhanced surveillance efforts will provide a mechanism for detecting health problems throughout the state before serious impacts to wildlife populations occur. Assimilating diagnostic data will aid in assessing trends suggestive of population-level disease problems. Programs for conducting extensive and intensive surveys for potential and realized wildlife diseases will provide reliable prevalence and distribution data for managers and administrators to use in decision making. Expertise in investigating and managing epizootics and epornitics will ameliorate efficacy and efficiency of efforts to control outbreaks. Improved techniques for diagnosing and studying wildlife diseases will provide a firm foundation for health management programs designed to enhance the quality of Colorado's wildlife populations. MATERIALS Disease AND METHODS Surveillance We monitored wildlife populations throughout Colorado for occurrence of disease using a combination of extensive and intensive approaches. These were organized and conducted as follows: Statewide Surveillance We continued to develop and modify a program for acquiring, examining, reporting on, and summarizing sporadic wildlife disease cases occurring throughout Colorado. All carcass submissions were subjected to necropsy. Ancillary diagnostics, including histopathology, bacteriology, virology, serology, parasitology, and toxicology were performed at the discretion of CDOW personnel and/or the attending pathologist. Preliminary examination and/or test results were telephoned to CDOW's Wildlife Research Center Laboratory, usually within 3-5 days of completion, and a final report were usually provided within 15 business days of submission. Pertinent data from preliminary and final reports were entered into a permanent database (described below), and copies of reports were filed as well as sent to appropriate field personnel. Pertinent data, including species, age, sex, location, number affected, diagnosis, and other information (as available) were entered into a computerized database. This database was used to generate quarterly and annual wildlife morbidity and mortality reports. In addition, data are available for analysis of long-term trends in select wildlife disease problems. Surveys Brucellosis Survev: We continued the statewide survey of deer and elk hunters to collect sera for brucellosis screening. Over the next several years, however, we plan to continue developing and implementing strategies for expanding utility and improving efficacy and efficiency of this survey. In particular, we will focus on improving return rates on sampling kits, quality of samples returned, and ability to target specific areas or populations for surveillance. We continued examining performance of the existing survey to determine average return rates and sample usability by species and season. In addition to the modifying the statewide survey, we also continued developing a process for intensively sampling specific geographic areas or populations using modified hunter surveys focused on sampling in select DAUs and/or GMUs. These surveys were constructed such that the probability of failure to detect at least 1 �case of brucellosis in the selected population was ~O.l even if herd prevalence is 1%. We will compare return rates and sample usability among seasons and collection methods, and use these comparisons to guide future survey efforts. Data from this year's survey will be analyzed in combination with those from previous years to compare sampling strategies. Results of survey modifications will be reported in future annual Job Progress Reports. We mailed about 9,825 blood sampling kits to elk hunters in selected GMUs statewide to gather samples for CDOW's annual brucellosis surveillance program conducted in cooperation with the Colorado Department of Agriculture's State/Federal Brucellosis Laboratory in Denver. Kits went to sportsmen with antlerless elk permits for second or late seasons in mountain GMUs statewide. Returned samples were identified by GMU of harvest. Usable samples were centrifuged, and sera were tested for antibodies to Brucella spp. using a standard card test. Unused sera were banked and stored at -20 C for future use. Chronic Wasting Disease Survey: To obtain reliable estimates for distribution and prevalence of CWO in wild cervids, we continued to survey for CWO in select deer and elk populations throughout Colorado. Brains from mule deer and elk harvested in various seasons during October 1994-January 1995 in GMUs 19 and 20 (DAUs D4 ,DIO/E4, E9) were collected for examination for CWO. Brains from hunter harvested mule deer and elk were collected, usually within 24-48 hrs of death, and fixed in 10% buffered formalin for at least 3 months. sections of medulla at the obex and frontal portion of the brain including basal ganglia, olfactory cortex and tract, and some frontal cortex were processed routinely for paraffin embedment. Histologic sections were cut at 56 ~m, stained with hematoxylin and eosin, and examined under a light microscope. In addition to formal surveys, we continued to encourage increased surveillance efforts by field personnel statewide and.submission of carcasses from deer or elk showing clinical signs resembling CWO, and held a workshop to inform field personnel about CWO and train them to in recognizing and properly submitting field cases. Bovine Tuberculosis Survey: Bovine tuberculosis was diagnosed in captive elk held on a second game ranch near Hesperus, CO in March 1994. We began investigating the possibility that tuberculosis might have spread to freeranging wildlife outside the infected premises. Retropharyngeal and other cranial lymph nodes and tonsils from mule deer and elk harvested in 74/741 were examined for gross lesions of bovine tuberculosis and collected for histologic evaluation and culture. Subsamples of parotid, mandibular, and retropharyngeal lymph nodes and tonsils, as available, were preserved in 10% buffered formalin and frozen and submitted to the Wyoming state Veterinary Laboratory in Laramie for histologic examination (and culturing, when warranted). When possible, eviscerated carcasses were also examined for gross lesions suggestive of tuberculosis. Disease Investigations Assistance was provided in investigating and Granite during July 1994-June 1995. Experimental pneumonia epizootics near Loveland Approaches We began developing a generalized, stochastic, individual-based simulation model of infectious disease in wild ungulate populations (Fig. 1). We plan to use this model in predicting consequences of disease introductions, improving understanding of the epizootiology of select disease problems, and evaluating potential disease management strategies. In this model, populations display density-dependent sigmoid growth in the absence of disease or other limiting processes. We employed a novel mathematical approach for estimating pathogen �transmission within simulated populations, and assumed transmission probabilities are a function of prevalence. Initially, we incorporated parameters to simulate introduction of bovine tuberculosis into a wild elk population and examined probable consequences of such introductions. As a preliminary step, we examined results of replicated 50-year simulations (n 500/parameter set) where 2 infected elk were introduced into a population of 500 wild elk. Our model incorporated population parameters estimated from a lightly hunted elk population (Forbes Trinchera Ranch). We assumed a constant cow-calf transmission rate (0.95) and a 2-year incubation period before newly infected animals became infectious. We then made replicated simulations, varying transmission coefficient (tc = 0.3 or 0.5 new infections/infected individual/year) to assess the influence of transmission on potential outcome of tuberculosis introductions. RESULTS Disease Statewide AND DISCUSSION Surveillance surveillance At least 60 carcasses and/or tissue samples representing 55 wildlife cases were submitted for diagnostic examination during July 1994-June 1995. Bronchopneumonia and chronic wasting disease were the most common diagnoses in wild_cervids submitted (although case submissions were undoubtedly biased by intensive monitoring for both diseases); all bighorn sheep submitted showed gross and/or histologic lesions of bronchopneumonia. Aside from pneumonia epizootics in bighorn herds near Loveland and Granite and salmonellosis in songbirds, all cases completed to date appeared to represent isolated cases of trauma or disease. Among carnivore cases, trauma, bronchopneumonia, and malnutrition were were diagnosed. Salmonellosis cases in passerine birds continued through July 1994, and cases were submitted from 10 locations throughout Colorado during April 1994-June 1995. Other mammalian and avian cases appeared to represent isolated incidents of unusual maladies (e.g., hairball in a mountain lion, carcinoma in an elk). For 15 cases, cause of death could not be determined. We will continue adding new accessions throughout the coming fiscal year to our computerized database for diagnostic case information, as well as data from archived reports as they become available. Surveys Brucellosis Survey: Of 9,825 elk hunters surveyed, 1,337 (14%) returned blood samples for brucellosis screening from animals harvested throughout Colorado during October 1994-January 1995. Of samples returned, 662 (49%) were usable; marked hemolysis and/or contamination precluded evaluation of the remaining samples. All elk sera tested were negative for antibodies to Brucella spp. on the standard card test. Overall, about 7% of the survey kits distributed to deer or elk hunters in 1993-1994 provided usable samples, as compared to 7% in 1993-1994, 7% in 1992-1993 and 5% in 1991-1992. Chronic Wasting Disease Survey: Seventeen cases of chronic wasting (CWO), a spongiform encephalopathy, were confirmed in free-ranging elk in Larimer County during FY 1994-1995. Forty-nine free-ranging have been confirmed in Colorado since 1981 (Fig. 1); to date, all these cases have been from Larimer County (GMUs 9, 191, 19, or 20) disease deer and CWO cases but 2 of (Fig. 2). Brains from about 150 mule deer and 65 elk harvested in GMUs 19 and 20 (DAUs D4, D10/E4, E9), were collected for examination for CWO. Histologic evaluation of all samples has not been completed, but of 343 deer and 212 elk brains from hunter-killed animals collected during 1991-1994 and examined to date, 3 deer were spongiform encephalopathy suspects; ancillary tests for confirmation are in progress. Based on survey data collected to date (and • �147 assuming all 3 suspects are confirmed positive), estimated prevalence of CWO in mule deer in DAUs D4/D10 combined is about 0.009 (95% CI 0.002-0.025); 95% CI for prevalence in DAU E4/E9 elk is 0-0.01 based on survey data (0/212) collected to tate. We prepared a manuscript manuscript follows: summarizing these cases; the abstract SPONGIFORM ENCEPHALOPATHY IN FREE-RANGING MULE DEER, ROCKY MOUNTAIN ELK IN NORTHCENTRAL COLORADO. of that WHITE-TAILED DEER AND T. R. Spraker, M. W. Miller, E. s. Williams, D. M. Getzy, W. J. Adrian, Gene G. Schoonveld, R. A. Spowart, K. I. O'Rourke, J. M. Miller, and P. A. Merz. Abstract: diagnosed Between March 1981 and June 1995, spongiform encephalopathy was in 49 free-ranging cervids from northcentral Colorado. Mule deer (Odocoileus hemionus) appeared to be the primary species affected and accounted for 41 of the 49 (84%) cases, but 6 Rocky Mountain elk (Cervus elaphus nelsoni) and 2 white-tailed deer (0. virginianus) were also included among these cases. Forty-seven cases originated from Larimer County, and mule deer submissions were clustered near two population centers (Fort CollinsLoveland and Estes Park). Clinical signs, when observed, included emaciation, excessive salivation, behavioral changes, ataxia and weakness. Emaciation with total loss of subcutaneous and abdominal adipose tissue and serous atrophy of remaining fat depots were the only consistent gross findings. Spongiform encephalopathy characterized by microcavitation of gray matter, intraneuronal vacuolation and neuronal degeneration was found microscopically in all cases. Scrapie-associated prion protein or an antigenically indistinguishable protein was demonstrated in brains from 16 affected animals, 10 using an immunohistochemical staining procedure, 9 using electron microscopy, and 7 using Western blot. Clinical signs, gross and microscopic pathology, and ancillary test findings in affected deer and elk were indistinguishable from those reported in chronic wasting disease of captive cervids. Prevalence estimates, transmissibility, host range, distribution, origins and management implications of spongiform encephalopathy in freeranging wild deer and elk remain undetermined. To obtain reliable estimates for distribution and prevalence of CWO in wild cervids, we continued to survey for CWO in select deer and elk populations throughout Colorado. Brains from about 25 mule deer and 29 elk harvested in GMUs 19 and 20 (DAUs D4 ,D10/E4, E9), from about 67 mule deer and 32 elk harvested on the Forbes Trinchera Ranch near Ft. Garland (DAU D31/E33), and from about 80 mule deer and elk harvested in GMUs 66 and 67 (GMU D25/E25) were collected for examination for CWO. Histologic evaluation of samples has not been completed, but all brains from hunter-killed deer and elk examined to date have been negative for spongiform encephalopathy. Bovine Tuberculosis Survey: Retropharyngeal and other cranial lymph nodes and tonsils were collected from deer and elk heads through game processing establishments in Durango and Hesperus for gross and histologic examination and culture of cranial lymph nodes; examination of 36 elk and 49 mule deer heads revealed no gross evidence of bovine tuberculosis infection, but abscessed tonsils and/or lymph nodes from 2 elk and 1 deer have been submitted for further microscopic evaluation; no microscopic lesions compatible with bovine tuberculosis have been observed in similar samples from previous years examined to date. Although no evidence ot tuberculosis has been detected in this survey, present sample sizes are insufficient to rule out the possibility of infection; for the number of negative samples examined to date, upper 95% confidence limits for prevalence estimates are about 0.08 for elk, about 0.06 for deer, and about 0.03 for deer and elk combined. Plans for additional surveillance are uncertain. �148 Disease Investigations No significant Experimental disease outbreaks were investigated during July 1992-June 1993. Approaches In examining preliminary results of 500 SO-year simulations where 2 infected elk were introduced into a population of 500 wild elk, transmission coefficient (tc) assumptions markedly influenced outcomes. Under conservative assumptions (tc = 0.3 new infections/infected individual/year), the probability that tuberculosis became established (i.e., infection still present SO years after initial introduction) in simulated populations was about 0.2 (Fig. 2), and prevalence in infected populations averaged about 0.03 (Fig. 3). Using a slightly higher tc (0.5 new infections/infected individual/year), the probability that tuberculosis became established increased to about 0.6 (Fig. 2), and mean prevalence in infected populations reached about 0.7 (Fig. 3). Our preliminary results suggest introduction of bovine tuberculosis into wild elk populations could represent a significant obstacle to national eradication goals. We plan to further refine parameter estimates for elk-tuberculosis simulations, and to explore application of this modeling approach to other real and potential disease problems affecting wild ungulate populations. Acknowledgments The statewide wildlife health monitoring and surveillance program described above relies heavily on efforts of dedicated field personnel throughout the Colorado Division of Wildlife, and truly represents a division-wide effort to improve our understanding and management of wildlife disease problems. In addition to those specifically listed, we collectively thank all of those regional and area biologists, district and area wildlife managers, and others who assisted by submitting diagnostic cases throughout the year. In particular, we thank personnel from areas 2, 4, 10, and 16, and from the Forbes Trinchera Ranch for assistance and logistical support in tuberculosis and CWO surveys and surveillance activities, and personnel from the StateFederal Cooperative Brucellosis Laboratory for their continued cooperation, assistance and logistical support in conducting annual brucellosis surveys. Prepared by __~~ __~~~~~ Michael W. Miller wildlife Research _ Veterinarian Table 1. At least 60 carcasses and/or tissue samples representing 55 wildlife cases were submitted for diagnostic examination during July 1994-June 1995. 1994 - 1995 Diagnostic DATE AGE SPECIES 8-9-94 3 Avian gulls 8-10-94 8-94 9-15-94 9-24-94 9-29-94 10-4-94 10-5-94 Cervine Elk Avian GH owl Cervine Ant. Cervine MD Cervine MD Bear Coyote REGION Imo/A Yng A 8-10 wks F F M M yng F NW NE SW NE NE SE NW Report CAUSE OF DEATH ACCESSION Pneumonia 1 pos. for botulism C Cerebral abscessation Enteritis and hepatitis Septicemia 945-03349 945-03975 945-06412 Wasting disease Undetermined Gastric foreign 945-7474 945-8007 945W1l39 body/gastritis 945-03306 # �149 Table 1. continued DATE SPECIES AGE REGION 10-10-94 Squirrel NA SE 10-3-94 10-3-94 10-10-94 10-10-94 10-10-94 10-10-94 10-22-94 11-10-94 11-4-94 11-4-94 11-4-94 11-21-94 11-17-94 11-11-94 11-17-94 11-21-94 MD Squirrel MD Pronghorn Squirrel 4 yrs F adult M NA NA M NA NE NE SE SE NA ? ? ? 3.5 yrs M Adult M NA NE NW SE NE NE SE NE 12-1-94 12-13-94 MD Elk Avian Avian Swan Avian Squirrel Avian Owl MD Elk Avian Starling Avian duck Mt. lion 12-15-94 12-15-94 12-15-94 12-21-94 12-27-94 12-19-94 12-23-94 12-26-94 12-27-94 12-28-94 1-09-95 1-09-95 1-10-95 1-9-95 1-27-95 2-2-95 2-11-95 Elk Mt. lion Golden eagle Mt. lion G. eagle Elk MD Elk Eagle Mt. lion CER CER MD Elk Moose Fox squirrel MD 2-17-95 3-20-95 3-20-95 4-95 4-14-95 4-24-95 Elk MD MD Don geese Beaver Avian Wh throated Pine siskins Elk Mule deer Prairie dog Raccoon 5-5-95 5-95 5-95 5-31-95 6-15-95 A Adult M Adult NA M 5 mos Adult F Yearling NA 2 yrs 10+ F NA F SE SW SW A M NA 10+ F NA Head Brain NA NA F A HEAD 7 mo M A F Immature 4-5 M NE NE NE NE NE NA SW NW NE NE A NW SE NE NE NE NA C A M/F 3 yrs M 5 yrs F A NA NA NW NE NE Calf 10 mo F M A 3/M/F C CAUSE OF DEATH Dermatitis, 1ymphocytic/mastocytic Wasting disease Ruptured mediastinal artery Unknown Unknown Unknown Neg. bluetongue Wasting disease Unknown 1 gout 6 unknown Eaten alive Unknown ? at time Malnutrition Unknown Gunshot Tubular epithelial degeneration and necrosis necrosis Staphylococci with no lesions Bronchopneumonia with abscessation Carcinoma Shot Poisoning Hairball Pneumonia Blunt trauma Unknown No wasting disease Pneumonia Emaciation starvatin Neg. Chlamydial Neg. Chlamydial Neg. Wasting d. Unknown Unknown Unknown Fractured liver due to fighting Starvation Most likely scours Unknown Salmonella Unknown Neg. Salmonella more tests pend. Salmonella Elaeophora Spongiform Encephalopathy Unknown Unknown ACCESSION 945-08323 945-7731 945-07741 945-08324 945-08322 945-08323 945-08321 945-09296 945-10818 945-10455 945-10460 945-10460 945-11640 945-11301 945-10905 945-11371 945-11639 945-12360 945-13253 945-13371 945-13370 945-13372 945-W1246 945-14155 945-13707 945-13987 945-14063 945-14155 945-14258 945-92744 945-92745 945-15270 945-15173 945-16622 945-17286 945-17988 94W1329 945-21274 945-21292 945-23109 945-23929 945-24647 945-25985 945-27169 945-25902 945-28483 945-29765 # �150 15 ..,' rtTh' . ....~ ..M.I),'~..d.~~J CU 10 o o L.. Q) E . ···:I~~I.Eik:::::····· en Q) en .c . 5 ::J Z . ..W ..................................................................................... White-tailed deer···· . ................................................................................................... .................................................................................... ................ ... . ........................................................................ .................................................................................... ............................................................................ ............................................................................ o 1981 1983 1985 1987 1989 1991 1993 1995 Year Fig. 1. Forty-nine cases of spongiform encephalopathy were diagnosed in freeranging deer and elk from northcentral Colorado between March 1981 and June 1995. Forty-six of these 49 cases were submitted since 1990; this pattern may he A nrnduct of intensified detection efforts, increasing prevalence, or both. e Larimer Count ® Mule deer (n-41) ft) Colorado Elk (n - 6) ® White-tailed deer (n - 2) Fig 2. All but 2 of the 49 documented CWO cases in wild deer and elk have originated in Larimer County (inset); most mule deer cases were clustered around Estes Park (n = 16) or in the foothills between Fort Collins and Loveland (n = 17). Black diamonds indicate locations of 2 wildlife research facilities where chronic wasting disease was described previously. Bar = 10 Jan. �151 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS Colorado State of Project Work No. Plan No. Job No. Period REPORT Covered: Mammals W-153-R-8 Research 2A Mountain 4 Strategies for Managing Infectious Disease in Mountain Sheep Populations July Sheep Investigations 1, 1994 - June 30, 1995 Authors: M. W. Miller, J. M. Bulgin B. J. Kraabel, Personnel: P. E. Bleicher, C. R. Kolus, J. A. Conlon, A. C. S. Ward, H. J. McNeil, and and M. A. Wild ABSTRACT We used ribosomal RNA fingerprinting and in vitro measures of cytotoxin production to compare Pasteurella haemolytica isolates from eight indigenous Rocky Mountain bighorn sheep herds in Colorado. Using ribosomal RNA gene restriction patterns, at least 26 distinct strains of P. haemolytica were identified among isolates (n = 59) from these herds; we identified one to seven distinguishable ribotypes within individual herds. Of the 26 ribotypes identified, 21 appeared unique to individual herds, four others (E, N, T, BB) were shared by only two herds, and one (A) was common to 3 herds. In vitro evaluation of-cytotoxin production by genotypically-distinct P. haemolytica isolates revealed further differences among strains: 4 ribotypes (AA, B, E, 0) showed marked cytotoxin production -- bighorn neutrophil death rate @ 150 ~g culture supernatant was 4-9 times that of an Enterobacter sp. control (7.1±0.4% neutrophil death @ 150 ~g supernatant); cytotoxicity of the other 22 strains examined approximated control levels. These findings support hypotheses that strains of P. haemolytica carried by healthy bighorn sheep may vary within and among wild populations. Our preliminary results also suggest the combination of genomic fingerprinting and cytotoxicity determination may offer a useful approach for studying th,e epizootiology of pasteurellosis within and among bighorn herds and may provide insights into strategies for effectively preventing or managing pneumonia epizootics. We examined effects of an experimental Pasteurella haemolytica toxoid-bacterin (AI, A2, TIO) on humoral immune responses and P. haemolytica carriage/shedding rates in bighorn sheep (Ovis canadensis) in a randomized, complete block experiment with a repeated measures structure. Thirty captive bighorns were divided into trios on the basis of age, sex, and previous history of pneumonic pasteurellosis; one bighorn from each trio was randomly assigned to receive 0, 1, or 2 doses of toxoid-bacterin. Because our experiment is ongoing, data and analyses reported here are preliminary. Mild, transient lameness in most vaccinated bighorns I day after initial vaccination was the only adverse effect observed. We identified 32 distinguishable biogroup variants among 266 P. haemolytica isolates from bighorns, but carriage and shedding rates did not differ among treatment groups (P ~ 0.53). In contrast, bighorns receiving I or 2 vaccine doses showed marked elevations in P. haemolytica cytotoxin neutralizing antibody titers 1 wk after vaccination (P 0.0001); mean = �152 responses peaked at 2 wks and titers remained elevated at least 6 wks after vaccination (P = 0.0001). Titers of agglutinating antibody to P. haemolytica serotype A1 capsular antigen were also elevated in vaccinated bighorns beginning 1 wk after vaccination (P ~ 0.0014) and showed similar response patterns. Preliminary data suggest this experimental P. haemolytica toxoidbacterin is safe and may stimulate protective immunity in bighorn sheep. Based on our findings, further evaluation of this, vaccine as a tool in preventing and managing pasteurellosis in bighorn sheep appears warranted. In a separate experiment, peripheral blood neutrophi1s from 10 bighorn sheep were exposed to P. haemolytica culture supernatants derived from one domestic (WSU-1) and three bighorn (CDOW-100, -521, -725) isolates. In vitro cytotoxicity of isolates was measured in both the presence and absence of prior administration of long-acting adrenocoticotrophic hormone (ACTH) gel to each bighorn. Mean serum cortisol concentrations in treated bighorns remained elevated about 10 hrs after ACTH administration (P ~ 0.05). For 3 of 4 isolates, neutrophil dp.ath rates were higher (P ~ 0.05) after bighorns received ACTH. White blood cell counts from ACTH-treated bighorn sheep demonstrated increased number of neutrophils, and decreased number of lymphoctyes and eosinophils when compared to white cell counts of controls (P = 0.02). Our data suggest cytotoxin-dependent killing of bighorn sheep neutrophils was enhanced by prior administration of ACTH. It follows that if similar processes occur in vivo, they could contribute to increased susceptibility of stressed bighorn sheep to pneumonic pasteurellosis. �153 EXPERIMENTS M. W. Miller, TO IDENTIFY AND MANAGE STRESS IN MOUNTAIN B. J. Kraabel, J. A. Conlon, SHEEP POPULATIONS B. J. McNeil, and J. M. Bulgin P. N. OBJECTIVE To develop strategies for managing population performance. infectious diseases affecting bighorn sheep SEGMENT OBJECTIVES 1. Analyze and report on data Comparing rates for tonsillar carriage and nasal shedding of Pasteurella spp., serum antibody titers to Pasteurella spp., and phenotypic and genotypic characteristics of Pasteurella spp. isolates among different indigenous bighorn populations. 2. Design and conduct an experiment evaluating select humoral and cellular immune responses of captive bighorn sheep to a multivalent Pasteurella haemolytica vaccine. 3. Use computer simulation managing pasteurellosis MANAGEMENT OF BACTERIAL modeling to examine alternative in bighorn sheep populations. AND VIRAL DISEASES IN MOUNTAIN strategies for SHEEP POPULATIONS Inability to control infectious disease outbreaks and subsequent mortality in mountain sheep populations represents a significant obstacle to long-term success in their management. Although the "bighorn pneumonia complex" has been studied intensively for over 3 decades, little is known about many aspects of its etiology and epizootiology. Moreover, management interventions recommended for preventing or controlling this problem remain untested. Most previous efforts to improve understanding and management of the epizootiology of pneumonia in bighorns involved post hoc investigations of dieoffs occurring in free-ranging sheep herds. These studies identified various etiological agents associated with known mortalities and attempted to determine predisposing causes and population consequences of individual outbreaks. From these investigations, comparisons of real or perceived patterns became the basis for hypotheses on the epizootiology of pneumonia in bighorns. Recognition of similar patterns in other outbreaks served as evidence supporting these as unifying hypotheses. Unfortunately, several of these hypotheses have failed to withstand rigorous experimental testing. And, despite our best management efforts, bighorns continue to die. Our strategy for developing a better understanding of the epizootiology and management of bacterial and viral diseases in bighorn populations differs -generally, we propose to take an adaptive environmental assessment approach for studying the bighorn pneumonia complex. As a foundation for our research strategy, we have assimilated existing knowledge on bighorn population dynamics (including the epizootiology and consequences of infectious disease) into a computer simulation model (Hobbs and Miller 1991). Because pasteurellosis appears to underlie virtually all respiratory disease problems reported for bighorns, our modeling efforts have focused on the epizootiology of pasteurellosis in sheep populations. We have constructed a model that reflects dynamics of bighorn populations seen in nature using the simplest assumptions necessary to reproduce those behaviors. We plan to conduct simulation experiments to identify variables that might be particularly sensitive to management perturbations in altering the dynamics of disease in bighorn populations. Those results will serve as the basis for designing management level experiments in the future. • �154 In parallel with our modeling efforts, we are conducting a series of experiments to develop, improve and standardize methods for collecting and interpreting diagnostic data to provide better estimates of key parameters driving our models. In particular, we have been developing tools for identifying strains of Pasteurella haemolytica and quantifying immunological responses of bighorns to infection by these pathogens. These tools will be key components of laboratory and field experiments designed to evaluate potential tactics (including vaccination and/or treatment) for managing pasteurellosis in wild sheep, and appear prerequisite to initiating management level experiments. To this end, our recent efforts have focused on both simulation modeling and on improving tools available for use in future management experiments that will be designed to study etiology, epizootiology, and prevention or control of disease outbreaks in bighorn populations: METHODS AND MATERIALS Management of Bacterial and Viral Diseases in Mountain Sheep Populations In conjunction with numerous cooperators, we continued developing and improving tools available for use in studying etiology, epizootiology, prevention or control of disease outbreaks in bighorn populations: and Epizootiology of pasteurellosis in indigenous biahorn populations (Miller, Spraker, Mills, Snipes, and Kraabel): We used ribosomal RNA fingerprinting and in vitro measures of cytotoxin production to compare Pasteurella haemolytica isolates from eight indigenous Rocky Mountain bighorn sheep herds (Almont/Taylor River, Avalanche Creek, Chalk Creek, Cottonwood Creek, Grant, Tarryall Mountains, Texas Creek, waterton Canyon) in Colorado. Genomic fingerprinting (Snipes et al. 1992) of remaining untyped isolates (n ~ 50) was completed. We also continued evaluating potency of cytotoxins derived from genotypically-distinct P. haemolytica isolates in vitro using methods described by Silflow et al. (1993). Experimental evaluation of a multivalent Pasteurella haemolytica toxoid-bacterin (AI, A2, T10) in captive bighorn sheep (Miller, Conlon, McNeil, Bulgin, and Ward): We used captive Rocky Mountain bighorn sheep (0. canadensis canadensis) (n = 30) (Table 1) in this experiment. All bighorns were housed at the Colorado Division of Wildlife's Foothills Wildlife Research Facility (Fort Collins, Colorado 80526, USA; 40035'N, 105010'W) throughout the study. We subdivided bighorns into groups by age and sex «1 yr, >1 yr rams, >1 yr open ewes, >1 yr pregnant ewes) (Table 1), and individuals within these subgroups resided together in 3-7 ha pastures throughout the study. In addition to natural forage, grass/alfalfa hay mix and a pelleted high-energy supplement were provided as prescribed under established feeding protocols for bighorn sheep in respective age/sex classes throughout the study (Miller 1990); fresh water and mineralized salt blocks were provided ad libitum. The general health of all bighorns was evaluated immediately after vaccination, as well as daily thereafter, and observations recorded throughout our experiment; particular attention was given to detecting respiratory signs (depression, segregation, anorexia, nasal discharge, coughing, labored breathing) in long-term observations. Injection sites were also examined weekly for 4 weeks after vaccine administration to assess local reactions to vaccine. All study animals were weighed at least monthly in conjunction with sampling. Health problems were evaluated and treated by attending veterinarians as necessary. Bighorns that died during our experiment were submitted to the Colorado State University Diagnostic Laboratory (Fort Collins, Colorado 80523, USA), where they were necropsied and ancillary diagnostic tests performed to determine cause of death. The experimental P. haemolytica toxoid-bacterin (Langford Laboratories, Inc., 131 Malcolm Road, Guelph, Ontario N1K 1A8, Canada) used here was an inactivated bacterial cell-free biologic extracted from culture supernatants • �155 of three serotypes (Al, A2, Tl0) of P. haemolytica; it contained leukotoxoid and serotype-specific surface antigens, and also incorporated the MUNOKYNIN~ adjuvant system. Methods for preparation and serotype Al components were the same as those used in a commercially-available bovine vaccine (PRESPONSE~, Langford Laboratories, Inc.). We examined the effects of this experimental P. haemolytica toxoid-bacterin administration on humoral immune responses and P. haemolytica carriage and shedding rates in captive bighorn sheep. Resistance to experimental challenge with pathogenic P. haemolytica was not tested in this study, but resistance to naturally-occurring pneumonic pasteurellosis in study bighorns and neonatal lambs was observed, recorded, and compared among groups. Our study was designed as a randomized complete block experiment with a repeated measures structure. In order to distribute treatments equally across the study population, our captive herd (n = 30 bighorns) was stratified by age «1 yr, >1 yr), sex (>1 yr rams, >1 yr open ewes, >1 yr pregnant ewes), and previous history of pneumonic pasteurellosis (health history) (Table 1). Within strata, individual sheep were assigned to blocks (n = 3 animals/block) (Table 1). One bighorn within each block was then randomly assigned to each of 3 treatment groups: 0 (control, no vaccination), 1 (1 vaccine dose), or 2 (2 vaccine doses 14 days apart) (Table 1). On day 0, we aseptically injected 2 ml of experimental toxoid-bacterin intramuscularly (1M) into bighorns in treatment groups 1 and 2; controls (treatment group 0) received 2 ml 0.9% saline, aseptically injected 1M. On day 14, bighorns in treatment group 2 received a second 2 ml vaccine dose (booster) injected 1M; bighorns in treatment groups 0 and 1 received 2 ml 0.9% saline injected 1M. All bighorns received vaccine or saline in the right hind leg on day 0 and in the left hind leg on day 14. Blood (about 10-12 mL) for serology was collected from each bighorn on wks 0 (prior to vaccination), 1, 2, 3, 4, 6, 8, 12, and 16. All blood samples were held for 1-4 hr at about 22 C, centrifuged, and serum collected. Serum was stored at -20 C until analyzed at Ayerst Veterinary Laboratories (131 Malcolm Road, Guelph, Ontario N1K lA8, Canada). We collected oropharyngeal and nasal swabs from each bighorn in conjunction with sample collections for serology on wks 0, 2, 4, 6, 8, and 12. Swabs were placed in transport tubes containing modified Cary and Blair medium (Port-A-Cul~, Becton Dickinson Microbiology Systems, Becton Dickinson and Company) and shipped overnight on ice packs to the Caine Veterinary Teaching and Research Center (CVTRC; 1020 East Homedale Road, Caldwell, Idaho 83605-8098, USA) for culture and analysis. Levels of cytotoxin neutralizing antibodies in sera were measured using an in vitro leukotoxin neutralization assay (Shewen and Wilke 1988). We expressed neutralization titers as the highest reciprocal 1092 dilution that yielded ~50% neutralization of toxicity. Levels of serum antibody against serotypespecific capsular antigens were measured using an indirect microagglutination assay (Shewen and Wilke 1988) that incorporated washed formalinized P. haemolytica serotype A1 as antigen. We expressed agglutination titers as the reciprocal 1092 of endpoint dilutions. Nasal and oropharyngeal swabs from sheep were cultured for detection of Pasteurella spp. (CIT). Isolates identified as P. haemolytica were further characterized by biochemical profile and serotype using CVTRC protocols (A.C.S. Ward, CVTRC, unpublished). Phenotypic differentiations of P. haemolytica isolates were based on biogrouping (Bisgaard and Mutters 1986, A.C.S. Ward, CVTRC, unpublished data) and serotyping by rapid plate agglutination (Frank and Wessman 1978). We calculated rates for isolating P. haemolytica, both in general and for biochemically distinguishable strains, from both nasal and oropharyngeal sites. Although resistance to experimental challenge with pathogenic P. haemolytica was not tested in this study, we evaluated resistance to naturally-occurring �1$ pneumonic pasteurellosis in study bighorns. Bighorns were observed daily and signs of respiratory disease (nasal discharge, depression, segregation, anorexia, coughing, dyspnea) recorded; we defined clinical pneumonia as a combination of signs including mucopurulent nasal discharge, depression (with/without segregation), anorexia and/or failure to gain weight, and coughing and/or dyspnea, accompanied by estimated resting body temperature ~ 39.5 C. The probable etiology of all pneumonia cases was determined by ancillary diagnostic tests. We compared 1) serum levels of neutralizing antibody titers to P. haemolytica leukotoxin, 2) serum levels of antibody to P. haemolytica capsular antigens, 3) rates of oropharyngeal carriage and nasal shedding of Pasteurella spp. and phenotypic traits of P. haemolytica isolates, and 4) rates of naturallyoccurring pneumonic pasteurellosis among treatment groups. We analyzed serology data using least squares ANOVA for General Linear Models (Freund et ale 1986) and the SAS Interactive Matrix Language. Responses to treatments were analyzed with analysis of variance for a randomized complete block design with a repeated measures structure. We used vaccine doses (0, 1, 2) as treatments and bighorn trios grouped by age/sex and health history as blocks; factors in the analysis were treatment, time, age/sex, and health history. Time was treated as a within subject effect using a multivariat~ approach to repeated measures (Morrison 1976). We calculated prevalence rates for tonsillar carriage and nasal shedding of P. haemolytica and compared these among treatments using categorical modeling; we also used Fisher's exact test to compare distributions of phenotypically distinct strains of P. haemolytica among treatments. We compared rates of naturally-occurring pneumonic pasteurellosis among study bighorns in respective treatment groups using Fisher's exact test. Effect of simulated stress on Pasteurella haemolytica cytotoxin-dependent killing of bighorn sheep neutrophils (Kraabel and Miller): Ten captive Rocky Mountain bighorn sheep were used as sources of neutrophils in this study: 4 adult ewes, 4 lamb ewes, and 2 lamb rams. All bighorns sampled were from a healthy captive herd held at the Colorado Division of Wildlife's Foothills Wildlife Research Facility (Fort Collins, Colorado, USA; 4003ss'N,10so10'. Animals were paired by sex and weight and placed in isolation pens (approximately so m2), and sampled in 2 groups (n = 6 and 4). One bighorn from each pair was randomly selected and administered adrenocorticotrophic hormone (ACTH) gel (Vedco, 40 Units/ml; 0.5 units/kg)subcutaneously; the same quantity of saline was administered subcutaneously to the other of the pair. Twenty-two ml of blood was collected from the jugular vein ten hr later. Total and individual sampling times were recorded to be correlated to cortisol levels in each animal (Miller et al., 1991a). Twenty ml of blood was placed in 5 ml of citrate phosphate dextrose solution (Sigma Chemical Company, St. Louis, Missouri, USA) and the remaining 2 ml was placed in ethylendiaminetetracetic acid (EDTA) vacutainer® tubes (Becton Dickinson and Company, Cockeysville, Maryland, USA). Pharyngeal swabs were collected at the same time, placed in Port a cul® transport tubes, containing modified Cary and Blair (BCB) media (BBL microbiology systems, Becton Dickinson and Company, Cockeysville, Maryland, USA), and cultured for Pasteurella spp. Total and differential white blood cell count were performed on the 2 ml of blood in EDTA tubes. The 20 ml of blood in citrate phosphate dextrose was centrifuged ate 1500 x g for 15 min. Plasma was saved for cortisol assay, and the buffy coat discarded. Hypotonic lysis of red cells was accomplished with the addition of 25 ml 0.001 M sodium phosphate monobasic for SO sec followed by the addition of 12 ml of O.OOlM sodium phosphate and 2.7% sodium chloride to stop the reaction. Following centrifugation at 600 x g for 10 min, the lysis and centrifugation steps were repeated, and the final pellets were resuspended in Hanks Balanced Salt Solution (HBSS) (Gibco Laboratories, Grand Island, New York, USA) containing 1% fetal bovine serum (FBS) (Hyclone Laboratories, Logan, Utah, USA). Cells were quantitated using a hemocytometer (American • �157 Optical Corporation, Buffalo, New York) and cell viability was determined by trypan blue exclusion (Boyse et al., 1964). Typical yields were >96% neutrophils, and these cells exhibited >92% viability. Cells were adjusted to a final concentration of 5 X 108 cellsjml in HBBS and 1% FBS. Four distinct strains of P. haemolytica (Table 1) were used as sources of cytotoxin. Cytotoxins were isolated from culture supernatant using methods described by Shewen and Wilkie (1982). Individual P. haemolytica isolates were streaked onto 5% sheep blood agar plates (Beckton Dickinson Microbiological Systems, Cockeysville, Maryland, USA) and incubated for 18 hr at 37° C. Several morphologically similar colonies were used to inoculate 100 ml of brain-heart infusion broth (Difco Laboratories, Detroit, Michigan, USA) and incubated at 37° C. Logarithmic phase growth was determined by an optical density reading of 1 at 600 nanometers using a model U-1000 spectrophotometer (Hitachi, Ltd., Tokyo, Japan). Bacteria were centrifuged at 6,000 x g for 10 min, and the culture supernatant were removed and filter sterilized in a 0.22 ~m filter (Sigma Chemical Company, st. Louis, Missouri, USA). Culture supernatant were dialyzed to exhaustion against distilled water using dialysis tubing with pore size of 6 - 8,000 molecular weight (spectrum Medical Industries, Inc., Los Angeles, California, USA). The remaining supernatant was then lyophilized. We compared the neutrophil susceptibility from treated and untreated bighorn sheep by incubating neutrophils with the cytotoxins from our P. haemolytica isolates. Lyophilized bacterial supernatant was resuspended in HBBS ad 1% FBS at concentrations of 150, 100, 50, 25, 5, 0.5 ~gj50 ~l, respectively. Fifty ~l of each supernatant preparation containing cytotoxin was added to the wells of 96-well plates (Corning Glass Works, Corning, New York, USA) followed by the addition of 2.5 x 105 neutrophils in 50 ~l of HBBS and 1% FBS to each well. Following 1 hr incubation at 37° C, 100~1 of lactate dehydrogenase substrate was added to quantitate the cytotoxicity of the bacterial supernatant (Korzeniewski and Callewaert, 1983). Quantification of the reduced LDH substrate was based on a model 450 96-well plate reader (Biorad Labs, Hercules, California, USA). All samples were compared to neutrophils treated with 0.05% Triton® detergent (Sigma Chemical Company, st. Louis, Missouri, USA) (maximal release) and to untreated cells (background release). The results were recorded as a percentage of LDH released from treated cells. Plasma cortisol was measured using an extracted double-antibody radioimmunoassay (RIA) (Hasler et al., 1976; Miller et al. 1991). All cortisol assays were conducted by the Endocrine Laboratory, Department of Physiology, Colorado State University. RESULTS Management of Bacterial and Viral AND DISCUSSION Diseases in Mountain Sheep Populations Epizootiology of pasteurellosis in indigenous bighorn populations: Using ribosomal RNA gene restriction patterns, at least 26 distinct strains of P. haemolytica were identified among isolates (n = 59) from these herds; we identified one to seven distinguishable ribotypes within individual herds. Of the 26 ribotypes identified, 21 appeared unique to individual herds, four others (E, N, T, BB) were shared by only two herds, and one (A) was common to 3 herds. In vitro evaluation of cytotoxin production by genotypically-distinct P. haemolytica isolates revealed further differences among strains: Four ribotypes (AA, B, E, 0) showed marked cytotoxin production -- bighorn neutrophil death rate @ 150 ~g culture supernatant was 4-9 times that of an Enterobacter sp. control (7.1±0.4% neutrophil death @ 150 ~g supernatant) (Fig. 1); cytotoxicity of the other 22 strains examined approximated control levels. All three indigenous bighorn herds that yielded markedly cytotoxic P. �158 haemolytica strains have recent histories of pneumonia epizootics: pasteurellosis outbreaks occurred in the Taylor River herd in 1979 and again in 1991, in the waterton Canyon herd in 1980, and in the Chalk Creek herd in 1981. One of these strains (E) was also recovered from dead bighorns during recent epizootics in the Alamosa Canyon (1989) and Rock Creek (1990) herds these latter herds can be linked to Taylor River by bighorn translocation activities during the last decade. Our findings support hypotheses that strains of P. haemolytica carried by healthy bighorn sheep may vary within and among wild populations. Our preliminary results also suggest the combination of genomic fingerprinting and cytotoxicity determination may offer a useful approach for studying the epizootiology of pasteurellosis within and among bighorn herds and may provide insights into strategies for effectively preventing or managing pneumonia epizootics. We are currently preparing a manuscript summarizing the findings reported here. Experimental evaluation of a multivalent Pasteurella haemolytica toxoid-bacterin (AI, A2, TI0) in captive bighorn sheep: Mild, transient lameness in most vaccinated bighorns 1 day after initial vaccination was the only adverse effect observed. We identified 32 distinguishable biogroup variants among 266 P. haemolytica isolates from bighorns, but carriage and shedding rates did not differ among treatment groups (P ~ 0.53). Bighorns receiving 1 or 2 vaccine doses showed marked elevations in P. haemolytica cytotoxin neutralizing antibody titers 1 wk after vaccination (P = 0.0001); mean responses peaked at 2 wks and titers remained elevated at least 6 wks after vaccination (P = 0.0001). Titers of agglutinating antibody to P. haemolytica serotype Al capsular antigen were also elevated in vaccinated bighorns beginning 1 wk after vaccination (P ~ 0.0014) and showed similar response patterns. Preliminary data suggest this experimental P. haemolytica toxoid-bacterin is safe and may stimulate protective immunity in bighorn sheep. Based on our findings, further evaluation of this vaccine as a tool in preventing and managing pasteurellosis in bighorn sheep appears warranted. Effect of simulated stress on Pasteurella haemolytica cytotoxin-dependent killing of bighorn sheep neutrophils: Mean serum cortisol concentrations in treated bighorns remained elevated about 10 hrs after ACTH administration (P ~ 0.05). For 3 of 4 isolates, neutrophil death rates were higher (P ~ 0.05) after bighorns received ACTH. White blood cell counts from ACTH-treated bighorn sheep demonstrated increased number of neutrophils, and decreased number of lymphoctyes and eosinophils when compared to white cell counts of controls (P = 0.02). Our data suggest cytotoxin-dependent killing of bighorn sheep neutrophils was enhanced by prior administration of ACTH. It follows that if similar processes occur in vivo, they could contribute to increased susceptibility of stressed bighorn sheep to pneumonic pasteurellosis. LITERATURE CITED Bisgaard, M., and R. Mutters. 1986. Re-investigations of selected bovine and ovine strains previously classified as Pasteurella haemolytica and description of some new taxa within the Pasteurella haemolytica-complex. Acta Path. Microbiol. Immunol. Scand. Sect. B 94:185-193. Boyse, E. A., L. J. Old, and I. Chouroulinkov. 1964. Cytotoxic test for demonstration of mouse antibody. Methods in Medical Research. 10:39-47. Carter, G. R. mycology. 1984. Diagnostic procedures in veterinary C. C. Thomas, Springfield, Ill. 515 pp. bacteriology and �159 Frank, G. H., and G. E. Wessman. 1978. Rapid plate agglutination procedure for serotyping Pas~eurella haemoly~ica. J. Clin. Microbiol. 7:142-145. Freund, R. J., R. C. Littell, and P. C. Spector. models. SAS Institute, Cary, NC, 187-201. 1986. SAS system for linear Hobbs, N. T., and M. W. Miller. 1992. Interactions between pathogens and hosts: simulation of pasteurellosis epizootics in bighorn sheep populations. Pp. 997-1007 in Wildlife 2001: populations. D. R. McCullough and R. H. Barrett, eds., Elsevier Science Publishers, Ltd., London, England, 1163 pp. Korzeniewski, C., and D. M. Callewaert. 1983. An enzyme-release assay for natural cytotoxicity. Journal of Immunological Methods. 64:313-320. Miller, M. W. 1990. Animal and pen support facilities for mammals research. Pages 45-63 in Pittman-Robertson Job Progress Report, Project W-153-R-4, WP2a, J4, Wildlife Research Report, Part 2. Colorado Division of Wildlife, Fort Collins, Colorado, USA. Miller, M. W., N. T. Hobbs, and E. S. Williams. 1991. Spontaneous pasteurellosis in captive Rocky Mountain bighorn sheep (Ovis canadensis canadensis): clinical, laboratory, and epizootiological observations. J. Wildl. Dis. 27:534-542. Morrison, D. F. 1976. Multivariate Co, New York. pp. 145-194. Shewen, P. E., and B. N. Wilkie. acting on bovine leukocytes. statistical methods. McGraw-Hill Book 1982. Cytotoxin of Pas~eurella haemoly~ica Infection and Immunity 35:91-94. Shewen, P. E., and B. N. Wilkie. 1988. Vaccination of calves with leukotoxic culture supernatant from Pas~eurella haemoly~ica. Canadian Journal of Veterinary Research 52:30-36. Snipes, K. P., R. W. Kasten, M. A. Wild, M. W. Miller, D. A. Jessup, R. L. Silf1ow, W. J. Foreyt, and T. E. Carpenter. 1992. Using ribosomal RNA gene restriction patterns in distinguishing isolates of Pas~eurella haemoly~ica from bighorn sheep (Ovis canadensis). J. wildl. Dis. 28: 347354. Silflow, R. M., W. J. Foreyt, and R. W. Leid. cytotoxin dependant killing of neutrophils sheep. J. Wildl. Dis. 29:30-35. 1993. Pas~eurella haemoly~ica from bighorn and domestic Wild, M. A., and M. W. Miller. 1991. Detecting nonhemolytic Pas~eurella haemoly~ica infections in healthy Rocky Mountain bighorn sheep (Ovis canadensis canadensis): Influences of sample site and handling. J. Wildl. Dis. 27:53-60. Wild, M. A., and M. W. Miller. 1994. Effects of modified Cary and Blair medium on recovery of nonhemolytic Pas~eurella haemoly~ica from Rocky Mountain bighorn sheep (Ovis canadensis canadensis) pharyngeal swabs. Wildl. Dis. 30:16-19. Prepared by Michael W. Miller Wildlife Research Veterinarian J. �160 Table 1. Block and random treatment assignments for captive bighorn sheep used in evaluation of an experimental Pasteurella haemolytica toxoid-bacterin (A1, A2, T10). Strata Treatment Age Sex Health historyb 0 1 2 <1 yr either H L894c C994 A94 P C294 Q94 L794 H M87 A93 M92 P M86 E992 L92 H L289 E83 T88 P M91 A82 E88 H M88 C89 E89 M93 L93 C92 Q92 A8S E392 L87 L88 Q86 >1 yr male female female - open - pregnant P 8 b c Grou128 o (control, no vaccination); 1 = (1 vaccine dose); 2 (2 vaccine doses 14 days apart). H = (no history of pneumonic pasteurellosis); P = (history of pneumonic pasteurellosis). Individual bighorns were identified by alphanumeric ear tags. �161 A.. Cytotoxin Neutralization 8 '" C) 0 6 .......... ~J~I·········································· 'CD c 0 -0- 0 doses -A- 1 dose ~~::2'd~~~~" T~···································· ............ -:-~*~~ ~ 4 <IS . I~ ·······················),·······t~~T··················· N - ........................................... ~I~~. <IS '- 2 :J CD .................................. Z T .,.. .,.. 0--2--2 0 o 2 T 4 l~± :r T__ 2 6 !. :To •••••••••••••••• O -0 l. 8 10 12 14 16 Week B. Pasteurella haemolytica A 1 8 N C) 0 6 ...... + T T T'--;'~ ~..................... . c 4 0 ..Y~.... ·i£lr·········{~~~r:~s~·~·· 'I~l ·1 <IS .! C :J C) C) 0 doses -A- 1 dose ~~:.' 2'd~~~~' . T--y 'CD - -0- T ··.:~·~~·······6~~·················Il.· - - - - - 2 6 . 2 < o o 4 8 10 12 14 16 Week Figure 1. (A) Bighorns receiving 1 or 2 vaccine doses showed marked elevations in P. haemoly~ica cytotoxin neutralizing antibody titers 1 wk after vaccination (P = 0.0001); mean responses peaked at 2 wks and titers remained elevated at least 6 wks after vaccination (P = 0.0001). (B) Titers of agglutinating antibody to P. haemoly~ica serotype A1 capsular antigen were also elevated in vaccinated bighorns beginning 1 wk after vaccination (P ~ 0.0014) and showed similar response patterns. ��163 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS REPORT State of Colorado Project No. W-153-R-8 Mammals Research Work Plan No. 2A Mountain Job No. 7 Experimental Evaluation of Mountain Sheep Transplanting and Disease Treatment Period Covered: Authors: Personnel: Sheep Investigations July 1, 1994 - June 30, 1995 M. W. Miller, J. Vayhinger, and S. Roush F. Barnes, R. Dobson, J. Duran, B. Elkins, W. Fey, J. George, Getzy, V. Jurgens, R. Hancock, M. Lamb, R. Myers, S. Ogilvie, Roberts, B. Thornton, A. Torres, T. Verry, R. zaccagnini. D. G. ABSTRACT We continued monitoring lamb survival among radiocollared ewes from 4 freeranging bighorn herds as part of a 4-year management experiment to examine effects of alternative lungworm treatment strategies on lamb survival and population performance. Since December 1991, 2 bighorn herds in the Tarryall Mountains [Sugarloaf Mountain (SL) and Twin Eagles (TE)] and 2 herds in the Collegiate Peaks [Chalk Creek (CH) and Cottonwood Creek (CW)] have been managed under 1 of 4 alternative lungworm treatment regimes: baiting with alfalfa hay and apple pulp treated with fenbendazole, baiting with alfalfa hay and apple pulp without fenbendazole, placing fenbendazole-treated salt blocks on bait stations, and withholding bait and fenbendazole. Treatments have been rotated annually under a predetermined, randomly-selected schedule. We monitored lamb production and survival among radiocollared ewes in each herd from May through October 1994 to complete the third field season. Year 4 started in mid-December 1994 with baiting and treated salt block distribution at scheduled sites. We began monitoring lamb production and survival among radiocollared ewes in each herc;lto assess· year 4..treatment responses. in May 1995.. Because this experiment will not be completed until October. 1995, data presented here relative to treatment effects·are preliminary and we have made no attempt to analyze or interpret them. Both production and mortality affected recruitment (lambs/marked ewes) through October among lamb cohorts monitored during May-October 1994. Lamb production, as estimated by observations of marked ewes with new (~ 2 wk old) lambs at heel, ranged from 0.65 at SL to 1.0 at CH, CW, and TE. In addition to reproductive/perinatal losses, some lambs disappeared in each of the experimental herds during the summer. Lamb survival (lambs in October/lambs born to marked ewes) through October ranged from 0.20 at CH to 0.73 at SL. Although sick and coughing lambs have been observed at CH every summer since 1991, respiratory signs appeared to be more severe and widespread in 1994, and lamb survival was about 1/3 that observed in previous years; overall recruitment of lambs through October appeared to be relatively high across the other 3 study herds in 1994, ranging from 0.59-0.88 lambs/marked ewes. �As many as 72 sheep fed at the TE bait site and as many as 76 sheep fed at the CW bait site during December 1994-February 1995. Marked ewes at TE (n = 16) averaged 44 days on bait (sd = 1); in contrast, marked ewes at CW (n = 14) averaged only 23 days on bait (sd = 6.3), and 2 additional marked ewes were never observed on bait. Radiocollared ewes visited the TE site almost daily; in contrast, most marked ewes at CW visited the bait site infrequently during December-February. In addition to bait, 3 of 16 marked ewes at CW also received 1 fenbendazole treatment, and 5 others received 2 treatments. Four 15 kg fenbendazole-treated salt blocks were available to sheep at 2 sites within the CH winter range between January and May 1995. All 4 blocks disappeared and were apparently consumed by late May; by comparison, a control block lost about 4 kg via environmental effects during January-May. Adjusting for estimated environmental losses, about 48 kg of treated blocks (about 79 g fenbendazole) were apparently consumed at CH. Block consumption equated to about 13 ewe treatments at CH in 1995, compared to 6.5 at TE in 1994, 26.5 at CW in 1993, and 6 at SL in 1992; under a more moderate dosing regime, block consumption equated to about 35 ewe treatments at CH in 1995. We observed marked and unmarked CH sheep using blocks in late January-April, but mule deer may also have used treated blocks at CH. Field data for the fourth lambing season span only May-June 1995, and consequently are quite preliminary. Observed lamb production through June 1995 ranged from 0.82 at SL to 0.93 at CW; lamb survival through June ranged from 0.71 at CW to 1.0 at TE. Perinatal lamb mortality associated with an extended period of cold, wet weather may have contributed to somewhat lower lamb production and survival through June 1995 -- 2 ewes seen in lambing areas in May-June showed behavior and physical traits typical of periparturient dams but were never observed with live lambs, and 4 other ewes were seen with lambs only once. Relatively consistent and predictable range use and movement patterns for each of the 4 study herds have emerged since monitoring began in 1991. Plots of May 1991-March 1994 location data (UTM coordinates) for radiocollared ewes revealed apparent differences in distribution and movement patterns among herds; although location data gathered since March 1994 remain unplotted, range use and movement patterns have not deviated appreciably from data plotted earlier. Ewes from the CW herd showed widest distribution and greatest movements; CH ewes were the most limited in their range use and movement. Although ranges of the SL and TE herds appeared to overlap considerably, to date we have observed no exchange of radiocollared ewes between these 2 herds. Disturbances by hikers (CW) and hunters (CH, SL) appeared to influence movements of ewe/lamb groups on occasion. Overall, noncapture mortality rates of adult ewes in the 4 study herds have averaged about 0.08 (se = 0.01) annually over the past 52 months, but causes and annual rates (ranging from 0 to 0.28 at CW in 1994) of ewe mortality appeared to vary among herds. Six of 62 marked ewes (3 at CW, 2 at CH, 1 at SL) died or disappeared during July 1994-June 1995. No consistent health problems have been detected- among marked ewes. In total, 20 radiocollared ewes (2 at CH, 5 at TE, 5 at SL, and 8 at CW) have died of noncapture causes since 1991. Of these, lion predation appeared to have caused 6 losses in the Tarryalls (3 at TE and 3 at SL), injuries from falls may have killed 2 ewes (at CW), pneumonic pasteurellosis killed 1 ewe (at CW), and lightning claimed 1 ewe (at CW); causes of death or disappearance for 10 other ewes (2 at CH, 2 at TE, 2 at SL, and 4 at CW) have not been determined, although we speculate lightning strikes also may have been involved in 2 deaths and 2 disappearances at CW and 1 death at CH during late June-early July. Despite observed variation in recruitment and adult mortality rates, winter range counts during 1991-1995 suggest all 4 bighorn herds under study have remained stable or grown since our experiment began in 1991. �165 EXPERIMENTAL EVALUATION OF MOUNTAIN M. W. Miller, SHEEP TRANSPLANTING J. Vayhinger, AND DISEASE TREATMENT and S. Roush P. N. OBJECTIVE Design, conduct, and report on management experiments to evaluate efficacy transplanting and disease treatment practices for managing mountain sheep populations. AGREEMENT continue parasite a management level experiment control program. of OBJECTIVE evaluating Colorado's mountain sheep We continued monitoring lamb survival among radiocollared ewes from 4 freeranging bighorn herds as part of a management experiment to examine effects of alternative lungworm treatment strategies on bighorn lamb survival and population performance. Year 3 of this 4-year study ended with completion of the summer field season in October 1994; year 4 began in mid-December 1994 with baiting and treated salt block distribution at scheduled sites and will continue through October 1995. Our study will be completed in October 1995. MATERIALS AND METHODS Beginning in December 1991, we began managing each of 4 study herds [Tarryall Mountains: Twin Eagles (TE) and Sugarloaf (SL); Collegiate Mountains: Chalk Creek (CH) and Cottonwood Creek (CW)] under 1 of 4 alternative lungworm treatment regimes: Control Treat OnlyBait Only Bait/Treat- no treatment -- bait and fenbendazole withheld; fenbendazole-treated salt blocks placed on bait stations; baited with alfalfa hay and apple pulp but not treated with fenbendazole; baited with alfalfa hay and apple pulp and treated with fenbendazole. Treatments were assigned to study herds as prescribed rotating schedule (Table 1.; Year 1 = 1992). by a randomly selected, Experimental treatments during the winters of years 3 (1994) and 4 (1995) were applied at scheduled sites (Table 1). In year 3, we baited for 55 days (ending 7 February) at SL and 76 days (ending 28 February) at CW; sheep at the SL site were also treated with fenbendazole (about 3 g/adult ewe) added to apple pulp on 31 January and 7 February. Treated salt blocks (1.65 g fenbendazole/kg, 15 kg/block; 4 blocks total) were available to sheep at TE during January-May 1994; 1 block held in a wire cage during that same period was used as an environmental control. During year 4, we baited for 56 days (ending 10 February) at TE and 82 days (ending 7 March) at CW; sheep at the CW site were also treated with fenbendazole (about 3 g/adult ewe) added to apple pulp on 16 February and 7 March. Treated salt blocks (1.65 g fenbendazole/kg, 15 kg/block; 4 blocks total) were available to sheep at CH during January-May 1995; 1 block held in a wire cage during that same period was used as an environmental control. We assessed effects of winter treatments on lamb production and survival by observing radiocollared ewes from all 4 herds about once every 2 weeks from May through October to determine whether they produced lambs, and whether their lambs were still alive. In addition to lamb survival data, we recorded approximate UTM coordinates, habitat type, and group size and composition for �100 each radiocollared ewe observed. All field data were transcribed into a computerized database to aid in mapping seasonal range movements and determining annual lamb production and survival rates. Radioco11ared ewes were also monitored every 2-4 weeks to detect mortality and movements during November through April in conjunction with a USFS/CDOW cooperative project to identify critical winter and transitional ranges of these 4 herds. Sixty-six radioco11ared ewes (15 at CH, 17 at CW, 17 at SL, and 17 at TE) were available for biweekly observation when assessment of year 3 treatment effects began in May 1994. Eight marked ewes died during May 1994-April 1995. We radioco11ared 3 additional ewes at CW during January 1995 to replace losses; all were immobilized with a combination of carfentanil HCl (1.5 mg), ketamine HCl (100 mg), and xy1azine HC1 (20 mg) delivered via syringe darts over bait and reversed with naltrexone HCl (50 mg IV + 100 mg SC). Consequently, 61 radiocollared ewes (13 at CH, 15 at CW, 17 at SL, and 16 at TE) were available for biweekly observation when assessment of year 4 treatment effects began in May 1995. In addition to ground observations, radiocol1ared ewes in the 2 Tarryal1 herds (SU, TE) were used as marked animals in an aerial mark-resight inventory exercise for estimating bighorn numbers in the Tarryall and Kenosha Mountains (J. George et a1., unpublished data). Sheep were counted during 3 helicpoter flights conducted in March-April 1995, and populaton sizes estimated using a joint hypergeometric maximum likelihood estimator for mark-resight data. Details of this inventory excercise will be reported separately. RESULTS Because this experiment presented here relative no attempt to interpret AND DISCUSSION will not be completed until October 1995, data to treatment effects are preliminary and we have made them. Treatment Rates Year 3: As many as 73 sheep fed at the SL bait site and as many as 90 sheep fed at the CW bait site during December 1993-February 1994. All marked ewes fed on at least 7 days. Marked ewes at SL (n = 17) averaged 45 days on bait (sd = 1.1) and marked ewes at CW (n = 15) averaged 26 days on bait (sd = 15.1). Radiocol1ared ewes visited the SL site almost daily. In contrast, most marked ewes at CW visited the bait site infrequently during DecemberJanuary. Many ewes in this herd stayed on alpine winter ranges until heavy snows apparently forced them to lower elevations in February, when CW bait site attendance increased markedly. In addition to bait, all 17 marked ewes at SL also received 2 fenbendazole treatments (31 January and 7 February). Four 15 kg fenbendazo1e-treated salt blocks were available to sheep at 3 sites within the TE winter range between January and May 1994. Block consumption ranged from 3.2-15 kg; by comparison, a control block lost about 2.3 kg via environmental effects during January-May. Adjusting for estimated environmental losses, about 23.7 kg of treated blocks (about 39.1 g fenbendazo1e) were consumed at TE. Using Schmidt et a1.'s (1979) dose recommendation (3 g fenbendazo1e/ewe/day, twice), block consumption equated to about 6.5 ewe treatments at TE in 1994, compared to 26.5 at CW in 1993 and 6 at SL in 1992; under a more moderate dosing regime (about 0.75 g fenbendazo1e/ewe/day for 3 consecutive days; Foreyt and Coggins 1990), block consumption equated to about 17 ewe treatments at TE in 1994. We observed 33 marked and unmarked TE sheep using a single block in mid-February; mule deer may also have used treated blocks at TE occasionally. Year 4: As many as 72 sheep fed at the TE bait site and as many as 76 sheep fed at the CW bait site during December 1994-February 1995. Marked ewes at TE (n = 16) averaged 44 days on bait (sd = 1); in contrast, marked ewes at CW (n = 14) averaged only 23 days on bait (sd = 6.3), and 2 additional marked ewes �167 were never observed on bait. Radiocollared ewes visited the TE site almost daily. In contrast, most marked ewes at CW visited the bait site infrequently during December-February. As in 1994, many ewes in this herd stayed on alpine winter ranges throughout the baiting period -- of 14 ewes seen at the CW bait site, 5 were observed ~9 times during the 82-day baiting period. In addition to bait, 3 of 16 marked ewes at CW also received 1 fenbendazole treatment, and 5 others received 2 treatments. Four 15 kg fenbendazole-treated salt blocks were available to sheep at 2 sites within the CH winter range between January and May 1995. All 4 blocks disappeared and were apparently consumed by late May; by comparison, a control block lost about 4 kg via environmental effects during January-May. Adjusting for estimated environmental losses, about 48 kg of treated blocks (about 79 g fenbendazole) were apparently consumed at CH. Block consumption equated to about 13 ewe treatments (2 X 3 g/ewe) at CH in 1995, compared to 6.5 at TE in 1994, 26.5 at CW in 1993, and 6 at SL in 1992; under the moderate dosing regime (3 X 0.75 g/ewe), block consumption equated to about 35 ewe treatments at CH in 1995. We observed marked and unmarked CH sheep using blocks on several occasions in late January-April, but mule deer also may have used treated blocks at CH. Apparent differences in salt block consumption between study herds in the Collegiate Peaks and Tarryall Mountains are supported by field observations suggesting marked differences in affinity for natural mineral licks between ewes in these 2 discontinuous mountain ranges. Whether these observed differences reflect natural mineral deficiencies in some ranges and/or greater abundance of salt associated with domestic stock grazing remains undetermined. However, our data suggest such differences could influence the potential use and efficacy of fenbendazole-treated salt blocks among diverse bighorn sheep ranges in Colorado and elsewhere. Lamb Production and Survival Year 3: The average proportion of ewes producing lambs in 1994 (91%) was high and essentially equivalent to average production in previous years (86-87%); 1994 lamb production ranged from 65% at SL to 100% at CH, CW, and TE (Fig. 1). Although most lambs observed were apparently born in May and June, new (~ 2 wk old) lambs were observed after 1 July in both Collegiate Peaks herds. Lambs disappeared from all 4 experimental herds during the course of the summer. Losses ranged from 1 of 11 known lambs in SL to 12 of 15 in CH (Fig. 1). Numerous sick and coughing lambs were observed with both marked and unmarked ewes in CH beginning in late June. One lamb collected in June 1994 was affected by pneumonic pasteurellosis that probably arose secondary to a congenital heart defect; no other lambs were collected and no lamb carcasses were recovered for postmortem examination later in the summer. Lamb losses apparently continued after our intensive observation period ended in October - by February 1995, only 2 lambs were known to be alive in the entire CH winter range. The cause of widespread respiratory disease and mortality in CH lambs in 1994 remains incompletely understood; pneumonic pasteurellosis likely played some role, but whether protostrongylosis was also a contributing factor could not be determined. Although sick and coughing lambs have been observed at CH every summer since 1991, respiratory signs appeared to be more severe and widespread in 1994, and lamb survival was about 1/3 that observed in previous years. Overall, lamb production and survival appeared remarkably high (86-91%) across the other 3 monitored herds. Year 4: Field data for the fourth lambing season span only May-June 1995, and consequently are quite preliminary. Observed lamb production through June 1995 ranged from 0.82 at SL to 0.93 at CW; lamb survival through June ranged from 0.71 at CW to 1.0 at TE (Fig. 1). Perinatal lamb mortality associated with an extended period of cold, wet weather may have contributed to somewhat lower lamb production and survival through June 1995 -- 2 ewes seen in lambing areas in May-June showed behavior and physical traits typical of periparturient dams but were never observed with live lambs, and 4 other ewes were seen with lambs only once. No sick or coughing lambs were observed at CH or elsewhere through June. �168 Range Use and Movement Patterns Relatively consistent and predictable range use and movement patterns for each of the 4 study herds have emerged since monitoring began in 1991. Plots of May 1991-March 1994 location data (UTM coordinates) for radiocollared ewes revealed apparent differences in distribution and movement patterns among herds (Fig. 2). Ewes from the CW herd have consistently shown widest distribution and greatest movements; CH ewes were the most limited in their range use and movement. Although ranges of the SL and TE herds appeared to overlap considerably, to date we have observed no exchange of radiocollared ewes between these 2 herds. Disturbances by hikers (CW) and hunters (CH, SL) appeared to influence movements of ewe/lamb groups on occasion. Location data gathered since March 1994 will be added to further define key ranges and migration corridors for these 4 herds. population Parameters and Performance Overall, noncapture mortality rates of adult ewes in the 4 study herds have averaged about 0.08 (se = 0.01) annually over the past 52 months, but causes and annual rates (ranging from 0 to 0.28 at CW in 1994) of ewe mortality appeared to vary among herds. Six of 62 marked ewes (3 at CW, 2 at CH, 1 at SL) died or disappeared during July 1994-June 1995. No consistent health problems have been detected among marked ewes. In total, 20 radiocollared ewes (2 at CH, 5 at TE, 5 at SL, and 8 at CW) have died of noncapture causes since 1991. Of these, lion predation appeared to have caused 6 losses in the Tarryalls (3 at TE and 3 at SL), injuries from falls may have killed 2 ewes (at CW), pneumonic pasteurellosis killed 1 ewe (at CW), and lightning claimed 1 ewe (at CW); causes of death or disappearance for 10 other ewes (2 at CH, 2 at TE, 2 at SL, and 4 at CW) have not been determined, although we speculate lightning strikes also may have been involved in 2 deaths and 2 disappearances at CW and 1 death at CH during late June-early July. Six of 10 ewe mortalities in the Collegiate Peaks herds since 1991 have occurred in apparently healthy ewes during mid June-mid August and may have been lightning-caused; whether telemetry collars are a predisposing factor in these losses is uncertain. Despite observed variation in recruitment and adult mortality rates, winter range counts during 1991-1995 suggest all 4 bighorn herds under study have remained stable or grown since our experiment began in 1991. Using the JHE generated from 3 counts flown during March-April 1995, the current estimate for bighorns in the Tarryall Mountains (SL and TE combined) is 173 animals (90' CI 158-194) (George et al., unpublished data); similar estimates are unavailable for either Collegiate Peaks herd. Intensive monitoring will continue through October 1995 with emphasis on documenting survival of known lambs, as well as range use and movement patterns. Experimental data will subsequently be analyzed and presented as a Job Completion Report in July 1996. Additional monitoring of marked sheep in the Tarryall Mountains is tentatively planned in conjunction with other upcoming field experiments. Table 1. Treatment assignments for 4 bighorn herds included in a 4-year management experiment to examine effects of alternative lungworm treatment strategies on bighorn lamb survival and population performance. HERD TARRY ALL MOUNTAINS COLLEGIATE MOUNTAINS YEAR 1992 1993 1994 1995 CHALK CREEK Bl B/T C T COTTONWOOD C T B B/T CREEK SUGARLOAF T B B/T C MTN. TWIN EAGLES B/T C T B 'Treatment assignments: BfT = bait with alfalfa hay and apple pulp treated with fenbendazole; B = bait with alfalfa hay and apple pulp without fenbendazole; T = fenbendazole-treated salt blocks on bait stations; and C = withhold all bait and fenbendazole (control). �169 ..•....•.. COTTONWOOD (Ceft" •• , 8UQARlOAF (T••• ,Oooy, CMAlKCAEEK (kI'OtIIy' 1W1. EAOlD (kI'/T ••• " ~ ....._, ees > > '::l en "'C c: canOHWOOD (Tr •• l~y) ees IUOAftLOA' (lai'Drlly, (1IaII/T••• " EAGLEI (Con,,", CIIAlKCllaK (Con••••, TWIN EAGLES (lr ••• Only) CHALK TWINEAOLU CHAUCC"~ TWIN c: 0 +-' o ::l "'C 0 'a. .c E COTTONWOOD (1IaII DrIly, (lailIT ••• " C01TONWOOD (kiIlTr •• t) "OARLO., tc ••••.••' IUOA"LOA' ees _J 'M •• •• •• •• C'U!DC (T••• 'Ooly, (kI'Oooy, Herd (treatment) Figure 1. Both production and mortality affected lamb recruitment during years 1 and 2 of our 4year study. Lamb production (open bars) was estimated by observations of marked ewes seen with lambs at heel; lamb survival (shaded bars) was estimated by following survivorship of lambs born to marked ewes through October. Values in parentheses are the number of marked ewes in each herd during the May-October observation period. Because data for year 4 (1995) cover only May-June. they probably underestimate lamb production and overestimate lamb survival. �170 BIGHORN SHEEP TREATHEm STUDY COTTONWOOD CREEK HERO (JIA. Y 1991 - MARCH 199.) BIGHORN SHEEP TREATMEm STUDY CHALK CREEK HERO (MAY 1991 - MARCH 199.) ........... '111 ••• + ~E + + ++++ " + + BIGHORN SHEEP TREATHEm STUDY TWIN EAGLE HERO (MAY 1991 - MARCH 1994) BIGHORN SHEEP TREATMEm STUOY SUGARLOAF Moum AIN HERO (MAY 1991 - MARCH 1994) cCUROY CREEK x X BISCH P£AK c: X ~X MCCUROY CREEt?< X X X " )«CUlDY ••••••••• 1')( ~"'P"""""" " X ICAl.E 6 X s: .0000 MC:UITAlt«P'EAKS ~LOCATt"" " PILOT PEA X Figure 2. Plots of May 1991-March 1994 location data (UTM coordinates) for radiocollared ewes revealed apparent differences in distribution and movement patterns among herds. Ewes from the CW herd showed greatest distribution and movements; CH ewes' range use and movements were the most limited. Although ranges of the Sl and TE herds overlap, no exchange of radiocollared ewes between these 2 herds has been observed. Jl(.u ••• t •••.• �171 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS state of project Work Colorado No. W-153-R-4 Plan No. Job No. Period Author: REPORT COvered: July Mammals Research 3A Pronghorn Investigations 2 Habitat Selection and Population Performance of a Pioneering Pronghorn Population 1, 1994 - June 30, 1995 T.M. Pojar ABSTRACT The rate of increase (ROI) for the Middle Park pronghorn population continues the downward trend as the population size increases. The projected late summer 1995 population size is 524 (Table 3) which is the 9th consecutive year of positive but diminishing growth. Using the linear regression technique (ROI on population size) to estimate carrying capacity, the projected K-value for this population is 547 animals (Figure 1). There are distinct differences in winter distribution of the population for the past 2 years, which may be due to density approaching carrying capacity. The second cohort of fawns (10 males and 10 females) were equipped with radio collars in December 1994 as part of the objective for estimating differential natural mortality between males and females. Forty radios were put on fawns of the year (20, Dec. 1993 and 20, Dec. 1994). Of the 20 2-year-olds (1993 fawns) 17 are alive with functioning radios. Two radios have been recovered; 1 collar slipped off the animal (female) and 1 collar broke (male). The third radioed animal (female) "disappeared" and is presumed to be dead. Three animals (2 females and 1 male) of this cohort emigrated to North Park in 1994. Eighteen of the 20 yearlings (1994 fawns) are alive and have functioning radios. One male and 1 female died within 10 days of capture; the male died of capture injuries and the female was hit by a vehicle. All of the yearlings have remained in Middle Park.' . �172 �173 HABITAT SELECTION AND POPULATION PERFORMANCE PRONGHORN POPULATION Thomas OF A PIONEERING M. ~ojar P.N. OBJECTIVE Describe population dynamics pronghorn population. and habitat SEGMENT and annual use of a pioneering, expanding OBJECTIVES 1. Describe seasonal population. distribution 2. Monitor natural mortality males and females. 3. Map areas of habitation 4. Monitor population dynamics of Middle Park pronghorn with: a. Ground counts to describe changes in population size. b. Ground counts to quantify population sex and age composition. and movement using of the Middle patterns Park pronghorn of radioed yearling the GIS format. STUDY AREA The study area (1993) is described in Pojar METHODS SEASONAL AND ANNUAL (1988) and a map of the area is in Pojar AND MATERIALS DISTRIBUTION Tracking was done mostly from the ground; fixedwing aircraft was used if an animal could not be located after a reasonable effort from the ground. Legal descriptions of animal locations were recorded to the nearest quarter mile then converted to UTM (U.S. Army 1973) coordinates for computer processing. All radioed animals have been located biweekly (with very few exceptions) since January 1, 1987. POPULATION SIZE AND STRUCTURE Herd structure estimates were obtained by classifying all animals that accompanied the animals that are radioed. The herd structure estimate used in population projections is the one with the largest sample size obtained in August or September. Total counts are made during winter by counting all animals associated with radioed animals. With the increased population size, it is not always possible to get an accurate count of total mature bucks (1.5 yrs and older) in the population. However, it is still possible to get very accurate counts of bucks in 60-80% of the population. The proportion of bucks in this portion of the population is then extrapolated to the total population to estimate total mature bucks. Total population count during winter, estimated number of mature bucks from the winter count, and recruitment based on fawn to doe ratios from late summer are used for the population projection. Population 1. projections Winter number are based on the following assumptions: counts represent the total population of mat~re bucks in Middle Park. and the estimated �174 2. Late summer age ratio estimates represent "recruitment" into the population. Annual survival of mature bucks and does and female fawns is 92.5%. Annual survival of males in their first year (after weaning) is 50%. (This severe mortality on male fawns is arbitrary, however, it allows the number of mature males in subsequent years to match fairly well with winter counts.) 3. 4. NATURAL MORTALITY OF MALES AND FEMALES The methods for estimating differential natural mortality between female pronghorn are outlined in Appendix I of pojar (1994). male and RESULTS SEASONAL AND ANNUAL DISTRIBUTION The winter distribution during 1994-95 was similar to the previous winter distribution, which exhibited some distinct differences from past years. A group of 115 ± spent the entire winter in the Sulphur Gulch area (T1N,R79W,S6). This group included all radioed animals that migrate east to Corral Creek and Granby in summer. As in the winter of 1993-94, another somewhat distinct group of 150 ± spent part of the winter north of Antelope Pass (T2N,R80W,S6) but returned to the Kremmling area when snow depth increased (ca 30 cm) in late winter. The remainder of the population wintered in the customary area north and east of the town of Kremmling. POPULATION SIZE AND STRUCTURE Total population size estimates are obtained during winter and herd structure estimates are obtained in late summer (Table 1). The annual changes in population size are used to calculate the rate of increase which is regressed on population size to project the K-value for the population. The ROI is calculated as where PI is the population size at time 1 and P2 is the population at time 2 (Table 2). The rate of increase for 1994-95 is 0.10 (Table 2), which is one of the lowest observed for this population. Based on the relationship of population size and ROI, the projected K-value is 547 (Figure 1). The population projection for late summer 1995 is presented in Table 3. In this projection it is assumed that the 1995 fawn production is the same as for 1994 at 46 fawns:100 does. The fawn to doe ratio is a critical assumption in the population projection and a subsequent ratio estimate based on a late summer 1995 sample may have a significant impact on this projection. It is possible the age ratio for 1995 will be higher than recent years because of very good growing conditions for forbs during spring and early summer. The herd structure estimates obtained in this study should not be subject to the shortcomings of herd structure estimates discussed by McCullough (1994) because of accurate population size data. The winter of 1994-95 was relatively mild with low snow fall and accumulation, and no extended periods of sub-zero (~) temperatures but spring and summer were above normal for precipitation resulting in excellent forb production. These conditions may have contributed to the expanded distribution of the wintering population and the potential for higher than expected 1995 recruitment. Ellis (1970) hypothesized that forb production during the last trimester of gestation influences recruitment. �175 Table 1. Herd structure of Middle Park pronghorn based on a sample obtained by locating radioed animals in late summer. The population size is from the subsequent winter counts with harvest added back into the population to get the pre-hunt population size, e.g. 1994 pre-hunt population was 466, 453 winter count plus 13 harvest. YEAR POP. SIZE NO. RADIO % RADIO B: 100D RATIO F: 100D RATIO SAMPLE % OF POP. 19861 1987 1988 1989 1990 1991 1992 1993 1994 80 122 160 223 261 308 347 425 466 7 24 22 17 13 39 31 58 5.7 15.0 10.2 6.5 4.2 11.2 7.3 12.4 36 54 40 56 22 23 26 10 29 77 77 32 50 47 65 48 66 46 47 63 108 161 148 148 286 266 332 59 52 68 72 66 48 82 63 71 1 This year's data based on the sample of the population trapped 16 December 1986. Table 2. Population size of the Middle Park pronghorn herd during winter and the calculated rate of increase. Population size reflects the removal of 1315 animals per year by harvest beginning in 1990, i.e. the 1994-95 winter population was 466 before harvest and 453 after harvest. YEAR POP. SIZE 1986-87 1987-88 1988-89 1989-90 1990-91 1991-92 1992-93 1993-94 1994-95 1995-96 (Projected) 80 122 160 223 246 292 332 410 453 524 NATURAL MORTALITY MALES OF YEARLING RATE OF INCREASE .52 .31 .39 .10 .19 .14 .23 .10 .16 AND FEMALES A capture operation, using the net-gun technique (Helicopter Wildlife Management, Salt Lake City, Utah) was used to radio 10 male and 10 female fawns in December, 1994. Since this is a relatively new technique for capturing pronghorn, my protocol for the operation is included in Appendix I. The radios were on expandable collars developed by Dick Bartmann (CDOW). The collars put on female fawns had an original circumference of 16.0 inches (40.6 cm) and were designed to expand to 17.5 inches (44.5 cm). Male fawn collars were 16.00 inches (40.6 cm) originally and will expand to 20.5 inches (52.1 cm) • �176 Table 3. Population projection text for the assumptions. for the Middle Park pronghorn population. POPULATION' BUCKS DOES FAWN'S TOTAL WINTER '94-95 96 245 112 453 WINTER MORTALITY 96 X .075 = 7 MORT 245 X.075 = 18 MORT 56X.5=28B 56X.075=4D 57 PREFAWNING 1995 96 - 7 = 89 MATURES + 28 YRLS TOTAL =117 245 - 18= 227 MATURES + 52 YRLS TOTAL = 279 LATE SUMMER 1995 MATURE 89 YRLS 28 TOTAL 117 MATURE 227 YRLS 52 TOTAL 279 See 396 @ 46F:100D 279 X .46 = 128 FAWNS 524 The radio packages used on the fawns in this study weighed 250 g. The lighter weight was accomplished by using lighter belt material for the collar, smaller batteries (2 LTC30 batteries rather than 2 C or 1 D battery), and circuitry that turns the radio off during nighttime to preserve battery life. This set of radios are set to turn on at 0700 hours and off at 1900 hours (MST). As of this writing, 19 of the 20 1993 radios can be accounted for (Table 4). The animal (female) with radio (149.510) was last seen in North Park (T9N,R79W,S32,NW) on June 21, 1994. Two other radios are functional but have fallen off the animals, 1 buck (149.450) and 1 doe (149.490). The buck's collar apparently caught on a fence and tore the belt material and the doe collar slipped off. It is presumed that both of these animals are still alive, however this has not been verified by positive ear tag identification. Movement of this cohort (1993 fawns) reflects that of the population with summer distribution following that of older radioed animals. The exceptions are the 3 (2 females; 149.510 and 149.730, and 1 male; 149.650) that emigrated to North Park. Two animals (1 male and 1 female) of the 1994 cohort died within 2 weeks of capture. One mortality was attributed to capture stress or injury and the other to being struck by a motor vehicle. Therefore, 18 (9 males and 9 females) of the 1994 cohort are still alive and being radio tracked (Table 5). None of this cohort has left Middle Park. �1n Table 4. Record of sex, ear tag numbers, radio frequency, and status as of June 30, 1995 of fawns radio collared on December 14, 1993 in Middle Park, Colorado (T2N,R80W,S36). Ear tag designation Y=yellow and B=blue. Sex Ear Tag F Y3 Y5 Y8 Y9 Y10 Yll Yl2 B53 B54 B55 Y1 Y2 Y4 Y6 Y7 B56 B57 B58 B59 B60 F F F F F F F F F M M M M M M M M M M Radio Frequency 148.500 149.230 149.150 149.272 149.502 149.512 149.490 148.760 148.730 149.430 149.650 149.172 149.450 149.410 149.470 149.390 149.190 149.550 149.530 149.132 Status Alive Alive Alive Alive Alive N. Park - Dead? Radio slipped off, recovered Alive N. Park, Last 4/10/95 Alive N. Park, Last 3/15/95 Alive Collar broke and recovered Alive Alive Alive Alive Alive Alive Alive Table 5. Record of sex, ear tag numbers, radio frequency, and status as of June 30, 1995 of fawns radio collared on December 12-13, 1994 in Middle Park, Colorado. Ear tag designation Y=yellow and B=blue. Sex Ear Tag F F F F F F F F F F M M M M M M M M M M B61 Y20 Y13 B62 Y15 Y22 Y23 Y24 B64 Y19 B65 Y21 B67 B66 Y18 Y25 Y16 B63 Y14 Y17 Radio Frequency 148.030 148.050 148.060 148.100 148.150 148.160 148.190 148.280 148.300 148.320 148.010 148.020 148.040 148.070 148.080 148.090 148.120 148.140 148.170 148.180 Status Alive Alive Alive Alive Auto kill, 12/23/94 Alive Alive Alive Alive Alive Alive Alive Alive Alive Alive Alive Alive Alive Alive Trapping mortality, 12/17/94 �178 REFERENCES Ellis, CITED J. E. 1970. A computer analysis of fawn survival in the pronghorn antelope. Ph.D. Thesis, Univ. of Calif., Davis. 70 pp. McCullough, D. R. 1994. What do herd composition Soc. Bull. 22:295-300. Pojar, Wildl. 1993. Habitat selection and population pronghorn population. Colo. Div. Wildl. performance of a pioneering Res. Rep. July, pp 199-207. 1994. Habitat selection and population pronghorn population. Colo. Div. Wildl. performance of a pioneering Res. Rep. July, pp 125-136. ,:'-,:{ I{:: ,:: :: {::: ':':,:'"':':''''' 0.9 [8 )t :: ": ») .\ L? :: .....,:') co :: . >/ .. :' ):: , .:•.•• :? • " ..,.:,}""'.:' ~ 0.7 :: .. :< :" (J :: C } }\ } 0.5 :~:.:. , > > 0.4 +oJ .< , }' .> :: :::::':' '.f." ::' i> '\ .. ::' :'::::::" 0.2 / ..\ < o o •••••••• )} ::} )) -: :1 ~1~; :::': r",<: '1 / < .:V.';=; ~7'iI': t F I )' 100 ::: tf: ..•.. :. " /':: , II :: i ": ( :. :,::,: ::':'::' ) ': ..... '.•••.••..•.•.•... ( g::'t: }' \:? /:'.:"'.'.' ,: .,. \ . / ) •.•. '.' ::: )\( :: : it '::? {.:. :?'" <t ::'{:}i'. }i:: 7':'::': )~, ?: i/ i,) ,?)' ~~ '.. ({ ... :: Ii> r 300 Population · ( '} .," :: 400 [,i Ii Hi : { .~~r~ ..:...'. ? it / ':{:: 500 :>/ :: ::::l:'.~r~ .:',~ I >.~' ,i>}{: ) Wt::~f:: ? }:}: :: :::, ~ii·~~~<.ii .'~.QJ . ":::":':'.: :: :: ':: ::::: ..... ':.::: ~l\\i!.~~i ( iE: J em r \1ji0~~iI 200 ...•. :.' ;!>~j ,}}:"~ -"E?' i :•••••••••• ,.,.:.:.: .•. :.:..••••.•• :. I •.• : •t";•:)r 1~~"'l~:<'<:h~ ::.•••••••••••••• ::•:•:.. •• /i. :::: >: # t.;; 0: 0.3 0.1 ::::' :':::: .... \/ :: r .:" ..: : •••••••• ,: :.:::.:: )::". •••••••• ••.••.• :ii' :::::: » , ':, :?': : : )< ::) :;: .. ". :::':::: co , ...,.,:" > ) ~; •••• g // \\::'} :':::'/::::: ::: :: t }: :::?, 11r~1?d ". ,::::'::':'. ',:< \.:: ",_",>:< ,}? ':::-',. > :,.,•• _,. :,,:,:'. :::::) ~ 0.6 '., '. :: :? ){ ..••'...•.•.... , )U ., : "".{:::.,,:,:, :':}}"'} :::.:::' :"':':' },,"':"":' :.:}:: .",/':': ::'_'." pp Universal Transverse Mercator Grid. Washington D.C. ™ No. 5-241-8, 64 pp. .:",;;/ ,·.·,·x .•':. ? 0.8 Q) tell us? T.M. 1988. Habitat selection and population performance of a pioneering pronghorn population. Colo. Div. Wildl. Res. Rep. July, 181-192. u.S. Army. 1973. Technical Manual: Headquarters, Dep. of the Army, '+o counts ;: ? it .... :1:.•••••••••••••.•• ::) , •.••••. : +.;. 600 ·t it I' 700 Size Figure 1. Rate of increase related to population size of the Middle Park, Colorado pronghorn population. �179 APPENDIX PRONGHORN Prepared I CAPTURE FOR NATURAL MORTALITY PROTOCOL FOR NET-GUN CAPTURE by: Thomas M. Pojar Date: INVESTIGATION METHOD December 20, 1994 Background: The Colorado Division of Wildlife is investigating the sex difference in natural mortality in free ranging pronghorn (Antilocapra americana). The Program Narrative prescribes a total sample of 60 radioed animals - 30 males and 30 females - captured over a 3 year period at a rate of 20 per year. In December 1993, the first year of the study, 20 animals were captured using the conventional drive trap method. The target animals are fawns of the year (about 5 months old). Using the drive trap method, it was necessary to process about 130 animals through the trap to obtain the 20 target animals. This put over 100 animals through the stress of trapping and handling to obtain the sample animals. During the trapping operation, 1 animal was killed and 1 suffered a broken leg; there were no post-capture mortalities of radioed animals. December 12 and 13, 1994 20 target animals were captured using the helicopter net-gun method. There were no direct capture injuries or mortalities but 1 radioed animal died 5 days after capture from capture-related injuries. Net-gunning will be used to capture 1995 from the Middle Park pronghorn 10 male and 10 female population. fawns in December capture protocol: Thomas M. Pojar is the project's principal investigator and will coordinate the capture operation. All persons involved in the capture operation, including the helicopter net-gunning crew, will be instructed on the care and handling of captured animals to minimize stress and injury to both the animals and crew members. The following protocol will be followed. 1. CAPTURE TIMING AND CONDITIONS. Early to mid December may be the best time to capture and handle pronghorn in Middle Park. It is usually before very deep or crusted snow conditions prevail and there are usually some relatively mild days during mid to late December to allow the animals to recover from the stress of capture. For net-gun capture, snow depths of 15-30 cm is surmised to be desirable to help slow the speed of the animals and to break their fall when netted. Capture operations will not be attempted if crusted snow conditions exist because of skin abrasion and injury to pronghorn's legs. 2. NO-FLY radius 3. NOTIFICATION OF AFFECTED PARTIES. Local rural residents and public agencies, including federal, state, and local agencies, will be notified of the time and general area of the capture operation. 4. EMERGENCY SERVICES. Personnel will be instructed as to the contact location of the nearest medical and emergency services. 5. RADIO COLLARS. Expandable collars will be used because target animals are young of the year and growth needs to be accommodated. The initial circumference of collars put on female fawns will be 40.6 cm (16.0 inches) expandable to 45.7 cm (18.0 inches); male collars will be 40.6 cm (16.0 inches) initially and will expand to 52.1 cm (20.5 inches). 6. COMMAND POST. The principal investigator and handling crew will be at a predetermined command post. The principal investigator will be in charge of all decisions for the care and welfare of the animals. ZONES. Pursuit and capture of a residence or developed will not take place area. within a 1 mile and �100 7. CHASE TIME. A capture will not be attempted without running the animal 1-2 minutes to "warm" animals to prevent capture myopathy. Upon location of a large group (50-100), a smaller group with 1 or more target animals will be separated from the large group within 1-2 minutes. The maximum pursuit time to maneuver the target group to an acceptable capture site will be 4-5 minutes. An individual pronghorn will not be actively pursued (active pursuit intense pursuit to capture) by the helicopter for more than 1 minute. Total pursuit of the target animal (and accompanying animals) will not exceed 10± minutes. Every effort will be made to take no more than 4-6 captures out of any single large group to avoid excessive chase time of non-target animals. Care will be taken by the helicopter crew to avoid chasing animals into fences, roads, or rivers. = 8. ANIMAL CARE AND HANDLING. Upon capture, the animal will be blindfolded, restrained (hobbled), and loaded into the back compartment of the helicopter. It will be ferried to the command post, unloaded and carried to the processing point (safe distance from helicopter) by 2-3 handlers per animal. The animal will be examined for injuries before the radio collar is installed; after processing it will be released on site (See item #10). Command posts will be located within 5 km (3 miles) of capture sites. Total time for handling will not exceed 2 minutes. 9. INJURED ANIMALS. Any debilitating injury or mortality of a captured animal will be reason to suspend the capture operation to assess the cause of the injury. The capture crew will be interviewed to determine if preventative measures are possible. Topics to consider include: pursuit techniques, weather conditions, visibility, fences, animal locations, groupings, etc •• The principal investigator will determine if the capture operation will resume. The severity of any animal injury will be assessed by the principal investigator and he will decide whether to release the animal or euthanize it. An animal with head, spinal, or upper extremity injuries will be euthanized immediately following guidelines of the CDOW Animal Care and Use Committee (1992, page 2 and Figure 1). The carcass will be processed for human consumption. Any other injury will be assessed on an individual basis with the primary criterion for euthanasia being the survivability of the animal. (Pending debate of ACUC euthanasia policy.) 10. RELEASE OF ANIMALS. While still hobbled and restrained, the animal will be again examined for injuries - skin abrasions (treat with disinfectant), neck, back, head, and extremities. First the hobbles will be removed then the blindfold. Handlers will continue to "hold" it (not restrain) until its eyes adjust to light and it is oriented enough to attempt to escape. The command post will be located such that released animals can be observed for about a half mile. The escape path will be clear of potentially dangerous obstacles such as vehicles, fences, and roads. upon release, the posture and gait will be observed, paying particular attention to how the head is carried (possible neck or back injury) and the use of all four legs as the animal runs. If an injury appears to meet euthanasia criteria, the animal will be immediately re-captured and euthanized according to guidelines in #9 above. Use of a video camera to document the behavior and condition of released animals would be desirable and could provide clues if postcapture mortality occurs. 11. POST-CAPTURE MONITORING. All radioed animals will be located 4-6 times within 10 days after capture to monitor for post-capture mortality. If a mortality is found, a necropsy will be performed and cause of death determined if possible. All injuries and mortalities will be recorded for future reference to the capture technique. �181 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS state of Project REPORT Colorado No. Mammals W-153-R-7 Research Work Plan No. 3A Pronghorn Job No. 5 Detecting Density Dependence Natural Populations Period Author: Covered: Investigations. in July 1, 1994 - June 30, 1995 T. M. Shenk ABSTRACT Logistic regression was evaluated as a feasible method to detect density dependence from temporal trends in demographic parameters. Data were generated from a 3-age class population growth model designed to mimic mule deer (Odocoileus hemionus) population growth. To estimate Type I error rates all demographic parameters were held constant, resulting in densityindependent growth. To estimate Type II error rates, overwinter fawn survival was modeled as density-dependent. Monte Carlo simulation was used to evaluate the effect of sampling variance in both the response variable, fawn survival rate, and the explanatory variable, population density. Fawn survival rate estimates were determined from a binomial distribution for a sample of fawns from the population. Density estimates were generated by adding a lognormally distributed sampling variance to the population densities. Increasing process variation, creating extra-binomial variation, was also added to fawn survival. Data were analyzed using logistic regression aSBuming (1) only binomial variation in the fawn survival rates and (2) assuming survival rates were overdispersed. Process variation (overdispersion) increased Type I error rates beyond the expected 5% when extra-binomial variation was not accounted for in the analysis, artificially inflating power (1 - Type II error) if overdispersion is not accounted for. Type I error rates remained at 5% for increasing process variation if an overdispersion parameter was estimated and used in the analysis. When overdispersion of the fawn survival rate was considered in the analysis, power increased as (1) estimates of fawn survival and density became more precise, .(2) time series length increased, and (3) time series included wider ranges of densities. Manipulations were suggested to best achieve (3). In summary, the stability of the Type I error rate when sampling variance was added to the response and explanatory variables as well as overdispersing the response variable, suggests logistic regression, adjusted for an overdispersed response variable is a valid method for detecting density dependence. Given the validity of the test, guidelines were suggested to maximize power. ��183 DETECTING DENSITY DEPENDENCE Tanya IN NATURAL POPULATIONS M. Shenk P. N. OBJECTIVE Evaluate the feasibility of detecting density populations through Monte Carlo simulation. SEGMENT dependence OBJECTIVES 1. Present an oral paper (INTECOL) held August England. 2. Evaluate logistic regression as a feasible dependence from temporal trends in density dependent life history parameter. 3. Complete annual report in natural at the Sixth International Congress of Ecology 24-30, 1995 at the University of Manchester, to the CDOW, technique to detect density and a potentially density- due August 1, 1995. STUDY AREA The study will take place in the Department of Fishery Colorado State University, Fort Collins, Colorado. and Wildlife Biology at METHODS PRESENT AN ORAL PAPER AT THE SIXTH INTERNATIONAL CONGRESS OF ECOLOGY HELD AUGUST 24-30, 1995 AT THE UNIVERSITY OF MANCHESTER, ENGLAND. (INTECOL) Effects of sampling variance on detecting density dependence in trends, by T. M. Shenk, G. C. White, and K. P. Burnham was presented The paper, temporal by myself (INTECOL) on August 24th at the Sixth International Congress held at the University of Manchester, England. EVALUATE LOGISTIC REGRESSION AS A FEASIBLE DEPENDENCE FROM TEMPORAL TRENDS IN DENSITY LIFE HISTORY PARAMETER. of Ecology TECHNIQUE TO DETECT DENSITY AND A POTENTIALLY DENSITY-DEPENDENT Simulations were conducted to evaluate logistic regression as a feasible method for detecting density dependence from temporal trends in density and a potentially density-dependent life history parameter. Results are presented as a draft manuscript, Detecting density dependence from temporal trends in demographic parameters using logistic regression. The draft manuscript is presented in the Results section of this Annual Report. COMPLETE An annual ANNUAL report REPORT TO THE CDOW, DUE AUGUST to the CDOW was completed 1, 1995. and submitted on August 1, 1995. • �184 RESULTS EVALUATE LOGISTIC REGRESSION AS A FEASIBLE DEPENDENCE FROM TEMPORAL TRENDS IN DENSITY LIFE HISTORY PARAMETER. TECHNIQUE TO DETECT DENSITY AND A POTENTIALLY DENSITY-DEPENDENT The following is a draft of the manuscript evaluating logistic regression as a method for detecting density dependence from temporal trends in density and a potentially density-dependent life history parameter. �185 DETEC'rING DENSITY DEPENDENCE USING LOGISTIC REGRESSION FROM TEMPORAL TRENDS TANYA M. SHENK, Department of Fishery and Wildlife University, Fort Collins, CO 80523 GARY C. WHITE, Department University, Fort Collins, of Fishery and Wildlife CO 80523 IN DEMOGRAPHIC PARAMETERS Biology, Biology, Colorado Colorado State State INTRODUCTION The concept of density-dependent population growth is central to our current understanding of population dynamics, especially processes such as compensatory mortality and management of harvested populations. The general acceptance of density-dependent regulation of population abundance has not, however, been paralleled by detection of such regulation from temporal trends in natural populations although numerous tests have been developed attempting to do just this (Varley and Gradwell 1960, Eberhardt 1970, Bulmer 1975, Royama 1977, Slade 1977, Berryman 1978, Vickery and Nudds 1984, Pollard et al. 1987, Reddinguis and den Boer 1989, Turchin 1990, Holyoak and Crowley 1993, Dennis and Taper 1994). Validity and power of these tests to detect density dependence in natural populations as well as interpretation of test results have proven controversial (Strong 1986, Gaston and Lawton 1987, Hassell et al. 1987, Lomnicki 1987, Mountford 1988, solow 1990, Bartmann et al. 1992, Holyoak 1993). Part of this controversy may stem from the lack of robustness these tests exhibit when test assumptions are violated (see Shenk et al. in prep). A statistical test is robust if test validity is not severely affected when test assumptions are violated. Test assumptions are often violated when data used to perform the tests come from field-collected samples. Two inherent problem with field-collected data are: (1) the associated sampling variance of any parameter estimate and (2) temporal and spatial (process) variation in demographic rates (i.e. birth, death, immigration and emigration rates). Although sampling variance is readily recognized and often accounted for as a source of variation in field data, process variation is often ignored. However, the assumption of only binomial variation in life history parameter estimates can rarely be met with data estimated from natural populations (Eberhardt 1978). Extra-binomial variation can result not only from environmental variation, but from lack of independence in the data (Anderson et al. 1994) or heterogeneity of demographic rates among individuals (Lebreton et al. 1992). Through Monte Carlo simulations 3 of the most commonly used tests for detecting density dependence from time series of population abundances (Bulmer 1975, Pollard et al. 1987, Dennis and Taper 1994) have been shown to be invalid when only the assumption of no sampling variance is violated (Shenk et al. in prep). Type I error rates were inflated beyond the nominal value for even small sampling variances, resulting in detection of density dependence more often than it occurred. Considering the poor performance of these tests when only 1 of the inherent sources of variation was added to the data, the approach of evaluating time series of population abundances to detect density dependence appears futile. • Fluctuations in population abundances are however, a result of changes in birth, death, immigration, and/or emigration rates. These changes in birth, death, or migration rates may be density-dependent responses. Therefore, logistic regression (see Cox 1970) could be used to support the hypothesis of a density-dependent response if density, as an explanatory variable, �186 significantly improves model fit for estimation of any of the life history parameters. For example, Clutton-Brock et al. (1987) used logistic models to investigate how changes in population density interacted with age, reproductive status, dominance rank, and matriline size to affect fecundity or calf survival in a resource limited population of red deer (Cervus elaphus). Selection of a model inclusive of density over a simpler model without density was support for a density-dependent response variable. The objective of this paper is to investigate logistic regression as a feasible (valid and powerful) technique to detect density dependence from temporal trends in density and a potentially density-dependent life history parameter when both sampling and process variation occur in the data. Monte Carlo simulations were first used to validate the methodology through establishing robust Type I errors consistent with the set nominal value. Because of the sensitivity of earlier tests to the addition of sampling variance (see Shenk et al. in prep), robustness of this technique when sampling variance occurs in the explanatory variable (density) as well as the response variable was investigated. Also investigated was the effect of process variation in the response variable on Type I error rates. Process variation results in overdispersion of the response variable, violating a critical assumption of the standard logistic regression technique. A valid test however, must be complemented by high power if test results are to be informative. Therefore, once validity of the test was established, power was estimated for varying conditions of sample size, precision of the estimated parameters, process variation, and initial population densities. Methods are suggested to increase power and strength of inference based on a series of simulated population manipulations. METHODS Logistic regression Regression methods describe the relationship between a response variable (Y) and one or more explanatory variables (x). In logistic regression the response variable is binary, thus 2 assumptions of logistic regression vary from those of linear regression. The first assumption of logistic regression defines the distribution of the conditional mean of the response variable given a value of the independent variable as [E(yIX)]. Using n(x) to represent E(yIX), the specific form of the logistic regression model is n t x) (1) following Hosmer and Lemeshow (1989). This model, n(x), bounds the conditional mean of the regression equation between zero and 1. A logit transformation of n(x) results in logit(n(x)) = lOgl 1 - rr t n(x) x) The logit(n(x» is linear in its parameters, to +00, depending on the range of x. J 130 + I3Ix continuous, (2) and may range from The second assumption of logistic regression which varies from linear regression concerns the conditional distribution of the response variable (Hosmer and Lemeshow 1989). An observation of the response variable may be -00 �187 expressed as y = E(yIX) + e. Linear regression assumes that e follows a normal distribution with mean zero and a constant variance across all values of x. Logistic regression assumes the response variable follows a binomial distribution with probability given by the conditional mean, n(x). An observation of the response variable is expressed as y = n(x) + e where e is distributed with mean a and variance equal to n(x) [1 - n(x)]. In a logistic regression analysis, maximum likelihood estimates are computed for Po and P10f Eq. 2. A test of the null hypothesis, Pl = 0, determines significance of the explanatory variable x. Significance of x is determined through likelihood ratio tests, G = - 210 l 9 (likelihood (likelihood without the variable) wi th the variable) J (3) where the distribution of G follows a chi-square distribution with 1 degree of freedom under the hypothesis that Pl equals zero (Hosmer and Lemeshow 1989). If the probability associated with the resulting G value is less than a the explanatory variable, x, is considered significant in predicting the response variable, y. If the conditional distribution of the response variable is not binomial and the data contain extra-binomial variation the data are overdispersed. Estimators of model parameters often remain unbiased in the presence of overdispersion, but the model-based, theoretical variances are underestimated (McCullagh and NeIder 1989). Thus, instead of var(n) = n(l - n), var (n) = $n(l - n) to account for the overdispersion of the data where $ is referred to as the overdispersion parameter. Deviance is defined as D = - l 210g (likelihood (likelihood of the current model) of the saturated model) J (4) where the saturated model contains as many parameters as there are data points (Hosmer and Lemeshow 1989). Deviance (D) provides a comparison of observed to predicted values and serves as a goodness-of-fit chi-square statistic (X2). An estimate of the overdispersion parameter $ is then defined as (5) where the difference in number of parameters between the saturated and current model being tested determine the degrees of freedom, rdf (Anderson et ale 1994) • The likelihood ratio test (LRT) of Eq. 3 will tend to be inflated by extrabinomial variation because of the underestimated variance of ~l' To compensate, instead of comparing the LRT statistic (on degrees of freedom) to the distribution of X2~f' one would use as a test statistic the LRT/df statistic divided by ~ treated as an F statistic with df and rdf degrees of freedom: F df,rdf = LRT/df ~ (6) (Lebreton et ale 1992). If the probability associated with the resulting F is less than a, the explanatory variable is considered significant in predicting the response variable. �188 Population growth models To evaluate the effect of both sampling and process variation on Type I error rates, densities were generated from a density-independent, constant finite growth population model. To evaluate the effect of sampling and process variation on power of the tests, densities were generated from a densitydependent population growth model. Density-dependent population growth was modeled from Bartmann et al. (1992) to mimic mule deer (Odocoileus hemionus hemionus) population growth. Density is hereafter defined as number of mule deer per square kilometer. Density-independent growth ~.--Population growth was generated for an agestructured population with 3 age classes (juvenile, yearling, and adult) as based on demographic information for mule deer from Bartmann et al. (1992). To generate density-independent growth, values for all demographic parameters were held constant over all years and all densities. The following equations were used to generate deterministic population densities for n = 5, 10, 25, 50, and 100 years. Density of juveniles (fawns) in time t+l: time series of (7) where Dy t+l is the density of yearlings at time t+1, DA t+l is the density of adults at time t+1, by is the birth rate for yearling females, and bA is the birth rate for adult females. A sex ratio of 50:50 is assumed. Density of yearlings in time t+l: Dy t+l = DJ t * SJ t * (1 - e} + iJ (8) ' where eJ is the emigration rate of juveniles, iJ is immigration of juveniles, and SJt, is juvenile survival rate at time t. Density of adults (i.e., individuals greater than 1 yr old) in time t+l: (9) where Sy and SA are the survival rates of yearlings and adults respectively, ey and eA are the emigration rates of yearlings and adults respectively, and i and iA are immigration of yearlings and adults respectively. y The following parameter values were used to generate the age-specific densities from the above equations: by = 0.75, bA = 0.75, sJ = 0.357, Sy = 0.862, SA = 0.862 from Bartmann et al. (1992). As no information was available, emigration rates and immigration were held at 0.00 for all age classes. Finally, total population density at time t+l: (10) An initial population density of 2 adult individuals, given the demographic parameters defined above, results in density-independent growth. Density-dependent growth ~.--Density-dependent growth was generated for the same age-structured population as that for the density-independent growth. However, survival of juveniles was modeled as density-dependent. The function for the density-dependent juvenile survival was taken from Bartmann et al. (1992). Bartmann et al. (1992) report a 3-year average overwinter survival of fawns as a logit-link function: 10ge[S / (1 - s)] = 1.1906 - 0.0195 * (Dec. density) (11) • �189 where s is overwinter fawn survival and Dec. density is total population density in December. The logit-link function is used to relate the parameters in a linear formula and to keep the survival estimates within the interval (0,1). The back transformation to the survival rate is given as s = (1 + exp [-(1.1906:" 0.0195 * Dec. density) ] )-1. (12) All other demographic parameters remain constant (i.e., density-independent) over all years, sex and population densities because data were not available to model these parameters as density-dependent functions. A density-dependent population growth model with only a single density-dependent response may over-simplify the effect of density on population growth. To compensate, the density-independent parameter values are conservative estimates based on values reported by Bartmann et al.(1992). For example, the model estimate for adult survival is 0.862, based on the mean annual (1982-88) adult female survival ranging from 0.760 to 1.000 (from Bartmann et ale 1992). If adult survival is in reality density-dependent, the model under-estimates survival at low densities and over-estimates survival at high densities. Using a constant mean value over all densities should dampen the incorrect response to density on population growth. These conservative density-independent parameter values coupled with the fact that overwinter fawn survival is the most density-sensitive population growth parameter in mule deer should together provide a reasonable projection of population growth. An initial population density of 2 adult individuals, given the demographic parameters defined above, results in sigmoid growth. Population density approaches K carrying capacity in approximately 50 years (Fig. 1a). To evaluate effect of initial population density on power of the test a series of simulations were conducted varying initial population density (Do = 2, K/4, K/2, 3K/4, and K). Stable age distributions were assumed for initial populations regardless of population density. Stable age distributions are defined as those that would occur if the population were allowed to grow from a density of 2 mature individuals per square kilometer (Table 1). Population growth models following density manipulations.--In an attempt to increase power, data were generated to mimic manipulations decreasing population densities away from K. Initial population densities (Do) were set at K followed by Dl = K/4 or K/2. Following this single-year decrease, population growth continued as defined by the density-dependent population growth equations above. Number of years of pre-treatment data (x) also varied from x = 1, 2, 3, 4, to 5. For example, if x = 2 then both Do and Dl were set at K followed by DJ = K/4 or K/2. After year 3, population growth continued as defined by the density-dependent population growth equations above. Parameter estimates The population growth models described above generate deterministic fawn survival rates and population densities. Data on survival and population density for natural populations are not usually from censuses. Both parameters are more typically estimated from sampling the total population and thus have associated sampling errors. To evaluate the robustness of logistic regression to the effects of sampling variance in both the response (fawn survival rate) and explanatory variable (density) estimates of both parameters were generated by adding appropriate sampling variances to the parameters. Process variation, defined as temporal and spatial variation in a life history parameter in response to stochastic environmental conditions, was also added to fawn survival rates to mimic natural population growth. Density estimates.--The skewed, non-negative nature of the lognormal distribution makes it an appropriate distribution for mimicking sampling variance when estimating population density. Therefore, sampling errors were �190 generated as lognormal errors with mean 1 and variance c2D; where c is the coefficient of variation for sampling and 0 !S: C s 1. Sampling errors were added to the total annual population densities generated from either the density-independent growth models or the density-dependent growth model (see Fig. 1b for effects of sampling variance on density estimate). Magnitude of sampling variance was increased from c = 0.00 to 1.00. Juyenile survival ~ estimates.--Annual juvenile (fawn) survival rate was estimated by using k Bernoulli trials to mimic the fates of k radio-collared fawns. Each Bernoulli trial represented a radio-collared fawn with probability p of survival as defined in the population growth model. For density-independent growth models p was held constant at 0.357. For densitydependent growth models p was determined from the density-dependent function of Eq. 12. The number of Bernoulli trials was set at either k = 20 or 50 to mimic the fates of either 20 or 50 radio-collared fawns in the population. The number of radio-collared fawns "surviving" the Bernoulli trials was then divided by the total number of radio-collared fawns (trials) to calculate annual fawn survival rate. Thus, sampling error on the fawn survival rate was binomially distributed (see Fig. 2b for effects of sampling variance on fawn survival rate estimates). To mimic natural conditions however, process variation was added to annual fawn survival rates. Annual fawn survival rates were determined as described above for either density-independent growth or density-dependent growth. Process variation was included in these annual survival rates by adding a random error to the survival rate. White and Bartmann (1995) report a process variation of 0.040 for 30 estimates of mule deer fawn survival in the Piceance Basin in northwest Colorado. Therefore, the normally distributed random error term was set with mean zero and variance of either 0.02 or 0.04. The addition of process variation to the annual fawn survival rate resulted in extrabinomial variation (overdispersion) of the estimated fawn survival rates (Fig. 2c). To avoid unrealistically low or impossibly high fawn survival rates when process variation was added, survival rates were truncated at 0.02 and 0.99. Annual fawn survival rate estimates were determined as above by dividing the number of radio-collared fawns 'surviving' the Bernoulli trials by the total number of Bernoulli trials (k = 20 or 50). Future reference to radio-collared fawns represent outcomes of these Bernoulli trials. MONTE CARLO STUDY RESULTS The Monte Carlo study was designed to first validate the methodology through establishing robust Type I errors consistent with the set nominal value when: (1) sampling variance was added to both the response (overwinter fawn survival) and explanatory (density) variable and (2) process variation was added to the response variable. Secondly, once validity of the test was established when both sampling and process variation were added to the data power (1 - Type II error) was estimated under varying conditions of (1) sampling variance in both the response and explanatory variable, (2) process variation in the response variable, (3) time series lengths, and (4) initial population density. Power was estimated when populations were simulated for both unmanipulated and manipulated population growth. Simulations were standardized at 1000 repet.itions for each set of parameters to estimate either Type I or Type II error rates, depending on the underlying population model (density-independent or density-dependent growth). Type I error rates were estimated as the percentage of rejections of the null hypothesis ~ O. PROC GENMOD (SAS Inst. Inc. 1993) was used to perform the logistic regressions. All tests were conducted at a nominal significance level of a = 0.05. = �191 Type I Error Rates Data were generated under the null hypothesis of density-independent growth as described above. A logistic regression model was fit where the response variable was overwinter fawn survival and the explanatory variable was density. Type I error rates were determined for both standard logistic regression and logistic regression adjusted for extra-binomial variation in the response variable. To evaluate the robustness of Type I error rates, parameter'estimates varied by: (1) increasing sampling variance in the explanatory variable (density, c = 0.0 to 1.0) and the response variable (fawn survival rate, k = 20, 50), (2) increasing process variation in the response variable (PV = 0.0, 0.02, and 0.04), and (3) increasing time series lengths (n = 5, 10, 25, 50, and 100 years). Initial population densities for all simulations estimating Type I error rates were 2 mature individuals per square kilometer followed by density-independent growth (i.e., all population growth parameters were held constant over all years and population densities). standard logistic regression.--Type I error rates were not shown different than the expected 5% when process variation was not added to the overwinter fawn survival rates (PV = 0.0, Fig. 3) for all number of years of data available and for increasing sampling variance in the explanatory variable, population density. Precision of the response variable estimate, fawn survival rate, also did not affect the Type I error rate when PV = 0.0. A process variation of 0.02 in fawn survival rates increased the Type I error rates above the expected 5% to 24%-45% when precision of the response variable was determined by 20 radio-collared fawns (Fig. 3). As process variation was increased to 0.04, Type I error rates increased to as high as 63% when c = 0.00. The trend of increasing Type I error rates with increasing process variation held when number of radio-collared fawns used to estimate the annual fawn survival rate was increased from 20 to 50. The effect of increasing the number of radio-collared fawns to estimate annual fawn survival rate was to decrease sampling variance. However, the Type I error rates were even greater when fawn survival rate was estimated more precisely. Thus, logistic regression was incorrectly detecting a relationship between fawn survival and population density more often than the nominal value of 5%. Increased number of years of data available (n = 5, 10, 25, 50, and 100) further inflated the Type I error rates when sampling variance on the explanatory variable was low. The inflated Type I error rates in the presence of extra-binomial variation in the response variable, invalidates standard logistic regression as a method for detecting a relationship between a response and explanatory variable when overdispersion occurs in the response variable. If no process variation exists, Type I error rates are equal to a. Logistic regression adjusted ~ oyerdispersjon.--Type I error rates were not found different than the set nominal value of 5% for all years of data available, over all values of process variation and sampling variation in the response variable, and for all values of sampling variation in population densities. Simulations conducted in this study support logistic regression, adjusted for overdispersion of the response variable, as a valid method for detecting a relationship between the response and explanatory variables in either the presence or absence of process variation. Power Power of a statistical test is the probability of obtaining a significant result when the null hypothesis being tested is not true. The null hypothesis of both the standard and overdispersion-corrected logistic regression is no significant relationship between the response and explanatory variables. �192 Thus, performing both methods on series of population variable) generated from density-dependent overwinter (response variable) allows the estimation of power. densities (explanatory fawn survival rates Power estimates were determined: (1) to compare power between standard logistic regression and logistic regression adjusted for overdispersed response variables, (2) to estimate power with unmanipulated population growth as time series length, process variation, and sampling variance in both the explanatory and response variable increased, as well as varying initial population density in relation to carrying capacity, and (3) to estimate power for manipulated population densities as treatment (population reduction) effect increased and number of years of pre- and post-treatment data varied. Comparison Qf ~ between methods.--Estimating power of a test is only useful if the test has been shown to be valid. Standard logistic regression has been shown to be valid only when no process variation exists in the response variable. However, Type 1 error rates for logistic regression adjusted for overdispersion of the response variable remains stable at the nominal 5% for process variation as high as 0.04. To avoid potentially violating the assumption of only binomial variation in the response variable, necessary for valid standard logistic regression results, it would seem more appropriate to always use the adjusted method. However, to evaluate the loss of power associated with the overdispersion adjustment power was estimated for both methods where process variation was set at 0.00 (to assure valid power estimates), Do = K/4, and precision of the response variable was increased from k = 20 to 50. Although lower than that for standard logistic regression, power remained high (> 80%) for n > 10 and c < 0.4 for the adjusted method (Fig. 4). However, power decreased more rapidly for adjusted logistic regression as sampling variance on the density estimates increased. In general, power increased for increasing time series lengths, and increasing precision of the response variable. Low power for n = 5 and 10 years results from the slow growth rate of the population (see Fig. 1a) and thus too few data are available to detect a relationship between fawn survival and population density. Power of 5% when n = 5 actually reflects a 5% chance of detecting a relationship when none exists, i.e., Type I error rate. Power is lost by including the overdispersion adjustment when no process variation occurs. However, the alternative of an inflated Type I error if process variation is assumed to be negligible and in reality is not is far more problematic. Therefore, the remaining power simulations were conducted to more fully evaluate logistic regression adjusted for an overdispersed response variable (hereafter referred to as logistic regression) as a feasible method for detecting density dependence in natural populations. Unmanipulated population growth.--In general, while varying a single parameter and holding all others constant, the following trends (see Figs. 5-9) were consistent throughout the simulations using data generated to mimic unmanipulated, density-dependent population growth: (1) power increased as number of years of data available increased, (2) power decreased as sampling variance increased on the explanatory variable, and (3) power increased as sampling variance decreased for the response variable. All 3 of these trends are as expected. The strengths and weaknesses of using logistic regression to detect density dependence in unmanipulated populations were however, more readily highlighted by several inconsistent power trends. These inconsistent power trends resulted from: (1) the interactions between initial population density and time series length and (2) the interaction between initial population density and increasing process variation. Density-dependent population growth as determined by Eqs. 6-11 results in sigmoid growth (Fig. 1a). Fawn survival remains high when population density • �193 is very low and remains low when population density is near K making it more difficult to detect a relationship between the two at either of these extremes (Fig. 2a). Consequently, lowest power occurred when the time series captured only the narrowest range of survival responses either because the series was too short to allow ample growth over a wide range of densities or because initially high densities prevented observation of low density responses (Figs. 5-9). For example, given the growth parameters used to generate the densitydependent data when Do = 2 it takes approximately 50 years to approach K. Time series of n = 5 or 10 years only capture the low densities associated with very high fawn survival. Twenty-five years of data only begins to approach the low fawn survival rates resulting from higher densities and thus has high power only when PV = 0.00 and fawn survival rate is estimated with higher precision (k = 50). When n > 25, the time series includes the entire range of densities and maximum power results (-100%, decreasing only in response to high values of c). The interaction between time series length and initial population density is made clear when comparing power for n = 25 years and Do = 2 to Do = K/4 or K/2 and n = 25. Power is significantly improved for time series of 25 years when initial densities are high enough to capture a wider range of fawn survival responses. Extremely low power for Do = 3K/4 and Do = K for any time series length reflects the narrow range of densities achieved by an unmanipulated population near K. The effect of time series length and initial population density illustrate inherent problems in detecting density dependence given the strength and function of the relationship between density and overwinter fawn survival. Capturing the widest range of densities improves power when these 2 parameters are considered. Increasing sampling variance in either the response or explanatory variable or both served to mask any relationship and as such decreased the power of the test. Power was less affected by sampling variation with longer time series or when the time series captured a wider range of population densities (e.g., Do= K/4). Decreasing sampling variance in the explanatory variable substantially increases power when a relationship exists. Time series length improves power when process variation increases and initial densities are suboptimal (e.g., Do = 3K/4, K) if sampling variance is low « 0.20). In the extreme case of Do = K and PV = 0.00, both population density and overwinter fawn survival remain constant over all time series lengths. Therefore, it appears no relationship exists between density and fawn survival and the 5% power of the test (Fig. 9) represents a Type I error rate. The effect of increasing process (environmental) variation on test power was influenced by initial population density. For a given time series length and number of radio-collared fawns, power decreased as process variation increased if initial population density was < 3K/4. If Do ~ 3K/4 power increased as process variation increased and all other parameters were held constant. When population densities are near or fluctuate around K fawn survival rate remains nearly constant prohibiting any relationship being detected by logistic regression. When this is the case, process variation in fawn survival rates serves to provide a wider range of both overwinter fawn survival and as a consequence densities than would exist without the process variation. Because the range of values for overwinter fawn survival and population density is narrow near K, even small increases in sampling variance in either the survival or density estimates results in significantly lowering power. Manipulated population growth.--Power increased as the range of population densities and their density-dependent overwinter fawn survival rates increased. This can be accomplished either by chancing to monitor population growth throughout this area of the growth curve or by manipulating population densities to ensure coverage over a large range. For populations with a growth rate similar to mule deer the first approach requires time series > 25 �194 years to achieve high power for even the best case scenario (Do= K/4) as demonstrated above. Manipulating populations over a wide range of densities offers an alternative to achieving high power over a shorter time period. Power was estimated for a series of feasible density manipulation approaches. Assuming the response variable could be measured to the precision achieved by radio-marking 50 fawns, power was estimated for studies of either 5 or 10 years. These simulations were also conducted for increasing sampling variance in the explanatory variable (0 = 0.0 to 1.0) and increasing process variation in the response variable (PV = 0.0, 0.02, 0.04). The range of densities resulting in the greatest change in fawn survival rate for a period of 10 years was determined by plotting annual overwinter fawn survival as a function of population density (Fig. lOa). Assuming initial population density near K, population densities resulting from single year density reductions to D = 2, K/4, K/2, and 3K/4 were generated followed by unmanipulated growth for the remaining 8 years (Fig. lOb). Superimposing the desired density range over the resulting density ranges highest power should be achieved if densities were reduced to K/4. Higher power estimates from a single second year reduction to K/4 (Fig. 11) to a single second year reduction to K/2 (Fig 12) support this hypothesis. Power for both reduction simulations when n = 5 or 10 years was significantly improved over power achieved in the unmanipulated time series simulations with initial densities equivalent to the reduction values of either K/4 or K/2. Both treatments resulted in a wider range of densities than if the population were simply observed from initial densities of K/2 or K/4 because of the initial K density. To most effectively use either a 5 or 10 year time frame in which to conduct the manipulation the effect number of pre- and post-treatment years (x = 1, 2, 3, 4, 5) had on power was also investigated. Number of pre- and posttreatment years had a greater affect on power when population densities were reduced to K/4 than K/2 (Figs. 11 and 12 respectively). The effect of the pre-treatment data is to ensure at least one data point near K. In the case of the reduction to K/2 the population returned to densities near K, decreasing the range of survival rates observed within the 10 year time frame (Fig. lOb). Thus, data collected at densities near K after the treatment were redundant. During a 5-year study this is not the case and power is affected by number of pre-treatment years (3 years pre-treatment data yielding the highest power when PV = 0.0). Pre-treatment data provide no redundancy when population density is reduced to K/4. In general, when densities were reduced to K/4, power was greatest when number of pre- and post-treatment years was equal. For n = 5 years, power was highest for 2 years pre-treatment; for n = 10 years power was highest for 5 years pre-treatment. Such a design for this treatment (K- K/4) ensures the widest range of densities and thus fawn survival rates. For all density-manipulation simulations power decreased increased and as time series length decreased. as process variation DISCUSSION Logistic regression is not a new technique to detect density dependence (see Clutton-Brock et al. 1987). However, given the inflated Type I error rates demonstrated in this paper when overdispersion is not accounted for in the analyses, caution is suggested in interpreting the results if overdispersion of the response variable was not considered in the analysis. Alternatively, logistic regression, adjusted for overdispersion of the response variable, is a valid and potentially powerful technique to detect density dependence from trends in density and an associated demographic parameter. The stability of a nominally set Type I error rate with increasing process variation in the response variable and increasing sampling variation in the explanatory • �195 variable (density) supports the validity of the test. Type II error rates suggest high test power if there are sufficient data available to track the density-dependent response. Tracking of the density-dependent response requires time series that capture a wide range of densities which may require long (> 10 years for the mule deer population modeled here) time series of unmanipulated populations where densities begin well below K or manipulating populations over a wide range of densities. If parameter estimates are obtained from unmanipulated populations initially well below K, the required time series length needed to detect an existing density-dependent relationship will vary with rate of population growth. As population growth rate increases, shorter time series are required to detect a significant relationship because the population approaches K in fewer years (Shenk, unpubl. data). Conversely, as population rate decreases the required time series length to detect a significant relationship would increase because the population would take longer to approach K. Simulation results from this study clearly demonstrate the feasibility of detecting a significant relationship between density and a density-dependent demographic response from temporal trends in observational data under conditions conducive to tracking the density response. Interpretation of test results using observational data however, can never establish a cause and effect relationship and thus limit inference of test results. A significant result may only indicate a covariation by chance or both variables may be functions of another variable (Sokal and Rohlf 1995). Logistic regression applied to data from experimental manipulation of the independent variable (density) can however, be used to establish cause and effect. Thus, not only will density manipulations ensure increased test power as shown from the Monte Carlo simulations but stronger inference can be made in interpreting test results. The density manipulations simulated here demonstrate only one alternative to maximizing power in logistic regression to detect a density-dependent relationship with a given demographic parameter. White and Bartmann (1995) also demonstrated increasing test power when the range of response variable values widened as a function of more severe density manipulations. The fundamental concept is to manipulate densities over the widest range thus eliciting a wide range of response values. One alternative to the experiments suggested, would be annually increasing density reductions each year over the desired range of densities (i.e., reduce to 3K/4 in year 1, K/2 in year 2, etc.). Such a design would be advantageous for detecting density dependence in game species because hunting could continue throughout the study. Temporal density manipulations could also be replaced with spatial manipulations. Bartmann et ale (1992) used 3 spatial replicates, each with different densities (low, medium, and high) to estimate response in overwinter fawn survival rates. Replication of density manipulations will further strengthen inference from test results by increasing external validity. In summary, logistic regression adjusted for overdispersion of the response variable is a valid and powerful tool to detect density dependence in natural populations if care is taken to obtain precise estimates of both density and the response variable over a wide range of densities. Inference to cause and effect of density on a given demographic parameter can be further strengthened by experiments with replicated density manipulations. LITERATURE CITED Anderson, D. R., K. P. Burnham, and G. C. White. 1994. AIC model selection in overdispersed capture-recapture data. Ecology 75(6) 1780-1793. �196 Bartmann, R. M., G. C. 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Oecologia 85:419-423. White, The effects effects �198 Table 1. Stable age distributions for varying initial population densities (Do). Stable age distributions are defined as those that would occur if population growth was initiated with 2 mature individuals and allowed to grow from the density-dependent population growth equations defined in the text. Do DA Dy DJ 2 2 0 0 K/4 = 22 6 4 12 12 7 26 18 9 40 24 9 56 45 K/2 3K/4 = K = 89 67 �199 TRUE SAMPLING VARIATION SAMPLING AND PROCESS VARIATION (a) 200 K t 150 100 50 0 W le::( 0 10 20 40 30 en W >Ien 90 100 K 200 t 150 , 100 I 50 , 1\ / .' '" I, 1" I , ',' I " '" , •... , " \ 1/ , ~ " -" , ," ' "I, .I , •.•• \ II •.•• \ \ , " / I' , " I " I I , 1-, 'I I \,I " " , i ; I' v , , ," ,, , \ ,' "~, ',, ,<: I,' "\ " " . v , '" , '.1 > \ " " Z W 80 70 (b) :2E I- 60 50 0 C 0 10 20 30 40 50 60 70 80 60 70 80 100 (c) 200 K t 150 100 50 ft-. . o _ . o 10 ....•... / ............ 20 30 40 50 90 100 YEARS Fig. 1. Population densities generated by the density-dependent growth model with (a) no sampling variance (c = 0.00) in the density estimates, (b) sampling variance in the density estimates (c = 0.20), and both process variation in the fawn survival rates (PV = 0.04) and sampling variance (c 0.20) in the density estimates. • �200 TRUE SURVIVAL SAMPLING VARIANCE SAMPLING AND PROCESS VARIATION (a) K 0.8 t 0.6 0.4 0.2 (b) K 0.8 " I ••••--', \/ 0.6 I /' 'I t ~ \ I' ....'I' - ,\ .... 1 V , '/ I I \ I \ ,~ \ I I \' 0.4 " , / I \ .I,', \ I "I " " /, \ I \ •.. " ,- \ - " I \, '/ I ' ,. ,', '-'\. r-;» I ,_ - ..••, I \ " I \.1 0.2 o 10 20 30 40 (c) 50 60 70 80 90 100 60 70 80 90 100 K t 0.8 0.6 0.4 0.2 o 10 20 30 40 50 YEARS Fig. 2. Fawn survival rate estimates over time as population density approaches K carrying capacity with (a) survival rates following a deterministic density-dependent population growth model, (b) sampling variance in fawn survival estimates (k = 50), and both process variation (PV = 0.04) and sampling variance (k = 50) in fawn survival rate estimates. �201 20 FAWNS 50 FAWNS pv=o.O PV = 0.0 0.9 0.9 0.8 0.8 0.7 0.7 0.6 0.6 O.S O.S 0.4 0.4 0.3 0.3 0.2 0.2 0.1 0.1 ~ 0 0.0 0.2 0.4 0.6 0.8 0.2 1.0 0.4 0 ~ ~ W W a. >I- 0.9 0.9 0.8 0.8 0.7 0.7 0.6 0.6 0.5 O.S 0.4 0.4 0.3 0.3 0.2 0.2 0.1 0.1 0 0 0.0 0.2 0.4 0.6 0.8 1.0 0.0 0.2 PV = 0.04 1.0 0.4 0.6 0.8 1.0 PV = 0.04 0.9 0.9 0.8 0.8 0.7 0.7 0.6 0.6 0.5 0.5 0.4 0.4 0.3 0.3 0.2 0.2 0.1 0.1 0 0.8 PV = 0.02 PV= 0.02 ~ 0.6 0.0 0.2 0.4 0.6 0.8 0 1.0 0.0 0.2 0.4 0.6 0.8 1.0 CV (DENSITY ESTIMATE) ···············0··········.----- 5 years 10 years ----_-- .. - .. _--.- 25 years ----0---- 50 years _··_·····_-CJ'·_··_··_··_ 100 years Fig. 3. Type I error rates of standard logistic regression analyses as sampling variance increases in the explanatory variable, density. Annual fawn survival rate is the response variable. The left column of graphs result from a less precise estimate of fawn survival rate (k = 20), the right column of graphs depicts results from the more precise estimates (k = 50). Process variation in the fawn survival rate estimates increase from top (PV = 0.0) to bottom (PV = 0.04). Each graph contains 5 curves depicting increasing time series lengths (n = 5, 10, 25, 50, and 100 years). Initial population density is 2 mature individuals per square kilometer followed by density-independent growth (i.e., all population growth parameters held constant over all densities) . • �202 ADJUSTED STANDARD FAWNS=20 FAWNS=20 1. 0.9 0.9 0.8 0.8 0.7 0.7 0.6 0.6 0.5 0.5 0.4 0.4 0.3 0.3 0.2 0::: w 0.1 o 3: o a. 0.2 ·····0 ....." .. __ 0.0 •. ····:· ..··:·· ...~ .....:.h..;.h..: ..... ~...h; 0.2 0.4 0.6 0.8 o 1.0 0.0 0.2 FAWNS=50 -,,--0--0--0 0.9 0.4 0.6 0.8 1.0 FAWNS=50 "'il:::i~:~;.~: '0- -0 0.9 0.8 0.8 . ~I!l. 0.7 -: g 0.7 'a 0.6 0.6 0.5 0.5 0.4 0.4 '0 .•.... 0.3 III 0.2 .... ta ••.• 0.3 0.2 ·s.... 0.1 0.1 Ow_~--~~~~~--~~~~ 0.0 0.2 0.4 0.6 0.8 " -,0. .•.. ·0 ......••.....••.... -ra......•,•......."......., __. • • =···..g.....'IlI...h-:...~ .: ····:· .... 1.0 CV (DENSITY ESTIMATE) ----0---- •...•.•••...••.. -0- •.......•••..• 5 years 10 years 25 years 50 years ·"·"·---'-·0·"·""_· 100 years Fig. 4. Power of standard logistic regression and logistic regression analyses, adjusted for overdispersion of the response variable, as sampling variance increases in the explanatory variable, density. Annual fawn survival rate is the response variable. The top graphs result from a less precise estimate of fawn survival rate (k = 20), the bottom graphs are results of the more precise estimates (k = 50). No process variation occurs in the fawn survival rate estimates (PV = 0.0). Each graph contains 5 curves depicting increased time series lengths (n = 5, 10, 25, 50, and 100 years). Initial population density is K/4. �203 2 20 FAWNS 50 FAWNS pv= 0.0 0.9 0.9 0.8 0.8 0.7 0.7 0.6 0.5 0.4 .- ...• .. .. - .•..... .. 'w' w· • .. ..•....• 0.2 0.4 PV 0.6 , 0.8 0.2 ,..0.. 0.2 1.0 = 0.02 0.4 0.6 .-.I>.-.G·- .-.g.'-'D'-'G.-.e·:.:·a:':·~:·::·..a, ....a. 0.8 ..,.- .••. -.0"·:a:.:·:Iil:::·~..a... 0.9 "'1:3, 0.8 . 'u,. '." 'B •••• 0 0.8 '0... '0 0.7 0.7 0.6 0.6 0.5 0.5 0.4 0.4 0.3 0.3 ••.•. 0.2 0.2 e-, c. .•-."~''''''-G.. "0'" 0.1 0 1.0 = 0.02 PV 0.9 0.0 0.2 0.4 0.6 0.8 0.8 1.0 PV= 0.04 -Q--ow-Q:, ""9':.;'8':..:' PV g:-'O ...."13... "0 0.9 •.••.•• g 1.0 = 0.04 -e , "0 0.7 0.9 'm, "tl, 0.8 • 0.3 0.1 0.0 0 '., - &'-0 • OUL~_L~~~~L_~L_~~ a.. •.. 0 --D' -'0' 0.4 0.1 ~ -. 0.5 - ...• 0.3 W .. ' 0.6 0.2 0:::: = 0.0 PV .-tJ-. -0-. -CJ_. -(3_.-a --o._.I!t'-'C)--'C'--""-'(lI-_.e-"':Q:':'::A:t.:,"'m ,, 0- 0.6 a, '0. .., 0.8 , 0.7 0.6 0 0.5 0.5 0.4 0.4 0.3 0.3 0.2 0.1 0 0.0 0.2 0.4 0.6 0.8 o 1.0 0.0 0.2 0.4 0.6 0.8 1.0 CV (DENSITY ESTIMATE) ...........-0 5 years . 10 years _- ... __ .•.. _-_-_ 25 years ----0---50 years ""-""""1::1""""" 100 years Fig. 5. Power of logistic regression, adjusted for overdispersion of the response variable, as sampling variance increases in the explanatory variable, density. Annual fawn survival rate is the response variable. The left column of graphs result from a less precise estimate of fawn survival rate (k = 20), the right column of graphs are results of the more precise estimates (k = 50). Process variation in the fawn survival rate estimates increases from top to bottom (PV = 0.0, 0.02, 0.04). Each graph contains 5 curves depicting increased time series lengths (n = 5, 10, 25, 50, and 100 years). Initial population density is 2 mature individuals per square kilometer. �204 Kl4 20 FAWNS pv= 50 FAWNS pV= 0.0 0.0 0.9 0.9 0.8 0.8 0.7 0.7 0.6 0.6 0.5 0.5 0.4 0.4 0.3 0.3 0.2 0.2 • ··:·····:·····: .....:·····;·· ..·'1·..··Ii····:jl 0.1 •....• o o 0.0 0.2 0.4 0.6 0.8 0.0 1.0 0.2 0.4 ·~a·:::." .. rr. 0.8 0:: W s: 0 a. \ \ II! 0.6 0.6 II , 0.4 ~, 0.5 "13., c 0.4 0.3 0.3 D. 0.2 0.1 0.2 0.1 "'····.0 ......••..... o .. ~ 0.7 \ ill 0.5 . 1.0 0.8 ),. • 0.7 0.9 0.8 = 0.02 PV PV = 0.02 0.9 0.6 0 ...•.'0 6 . OUL~-L--~~~L-~~~-L 0.0 0.2 0.4 PV 1 0.9 0.6 0.8 0.0 1.0 0.2 0.4 = 0.04 PV 0.6 0.8 1.0 0.8 1.0 = 0.04 O:::':~" , 0.8 Q. 0.9 b\ ,-, \ 0.7 ... 0.8 ,b 0.7 Q'. 0.6 " 0.6 v, q.,. '" 0.5 ". 0.4 0.3 0.2 0.1 0.2 0.4 0.6 0.8 1.0 0.2 .04 0.6 CV (DENSITY ESTIMATE) •••••••••••••• 5 years 1)- ••••.••••••••• 10 years •.•... _.a_._._,. 25 years ----0---50 years ············'0·········· 100 years Fig. 6. Power of logistic regression, adjusted for overdispersion of the response variable, as sampling variance increases in the explanatory variable, density. Annual fawn survival rate is the response variable. The left column of graphs result from a less precise estimate of fawn survival rate (k = 20), the right column of graphs are results of the more precise estimates (k = 50). Process variation in the fawn survival rate estimates increases from top to bottom (PV = 0.0, 0.02, 0.04). Each graph contains 5 curves depicting increased time series lengths (n = 5, 10, 25, 50, and 100 years). Initial population density is K/4. • �205 Kl2 50 FAWNS 20 FAWNS pV= 0.0 pv=o.o .. 0.9 \ 0.8 0.7 , 0.9 0.8 -. 1 0.7 0.6 0.6 0.5 '.j:] 0.5 0.4 0.4 0.3 0.3 0.2 0.1 0.2 0.4 0.9 -, W :: 0 a. ta \ \ o. , '1 \ \ 0.7 c::: q 0.8 Q 0.8 0.6 a 0.5 0.3 , 0.7 b\ ",\ ",'. • 0.4 = 0.02 PV PV= 0.02 0.9 0.6 , 0.6 , C " 0.5 P. ", , 0.4 ,': '. 'I.... \ ' Q.', -', 0.3 0.2 0.2 0.1 0.1 0.2 0.4 0.6 0.8 0.2 1.0 0.4 PV= 0.04 0.9 c. 0.9 0.7 '\ ". 0.7 \. 0.6 \ '1 , Q, ',_ \ ,~ , ...' 'II, , m 0.4 0"" , . '. \ e, 0.5 \ 0.4 ". D , ta '. '. 0,3 0.2 0.1 'q 0.8 b 0.6 0.5 = 0.04 1 e 0.8 0.3 PV 0.6 0.2 c:::~::±:±±~~~ 0.2 0.4 0.6 0.8 0.1 O~~~~~~~~ 1.0 0.0 0.2 0.4 0.6 0.8 1.0 CV (DENSITY ESTIMATE) •.............•.. Q 5 years . 10 years ._._._ •. 15._._ •.• 25 years ----0---50 years "-""""'-0-"""'" 100 years Fig. 7. Power of logistic regression, adjusted for overdispersion of the response variable, as sampling variance increases in the explanatory variable, density. Annual fawn survival rate is the response variable. The left column of graphs result from a less precise estimate of fawn survival rate (k = 20), the right column of graphs are results of the more precise estimates (k = 50). Process variation in the fawn survival rate estimates increases from top to bottom (PV = 0.0, 0.02, 0.04). Each graph contains 5 curves depicting increased time series lengths (n = 5, 10, 25, SO, and 100 years). Initial population density is K/2. �206 DO 3K14 20 FAWNS 50 FAWNS PV = 0.0 pV= 0.0 0.9 0.9 0.8 0.8 "Q 0.7 0.7 ( 0.6 0.6 I .,\ 0.5 0.5 0.4 q.'; 0.3 \". ..." 0.2 0.1 a • ·····IiI·:~·:!il""·e·-a 0 .,\ 0.3 '..,~ 0.1 .'\ 0.4 • t~ 0.2 I 0.0 0.2 :e:····g.,-I!f~ 0.4 0.6 0 0.8 0.2 0.0 0::: W s:0 a.. 0.8 1.0 = 0.02 PV PV = 0.02 0.9 0.9 0.8 0.8 " 0.7 0.6 0.4 1.0 '" 0.7 0.6 0.6 0.5 0.5 III 0.4 0.4 i.. \ \ 0.3 '. 0.3 0.2 0.2 0.1 0.1 0.2 0.4 PV 0.6 0.8 0.2 1.0 = 0.04 PV 0.9 0.8 0.9 0.6 0.8 1.0 0.8 1.0 = 0.04 III 0.8 0. 0.7 0.7 0.6 0.5 0.4 ~ 0.6 0.5 q 0.4 0.4 0.3 0.3 0.2 0.2 0.1 0.1 0.2 0.4 0.6 0.8 1.0 ·0 q \ \ \ 0.2 0.4 0.6 CV (DENSITY ESTIMATE) ----0---- ················0·············· 5 years 10 years 25 years 50 years ···_··_··_···m···''''··' 100 years Fig. 8. Power of logistic regression, adjusted for overdispersion of the response variable, as sampling variance increases in the explanatory variable, density. Annual fawn survival rate is the response variable. The left column of graphs result from a less precise estimate of fawn survival rate (k = 20), the right column of graphs are results of the more precise estimates (k = 50). Process variation in the fawn survival rate estimates increases from top to bottom (PV = 0.0, 0.02, 0.04). Each graph contains 5 curves depicting increased time series lengths (n = 5, 10, 25, 50, and 100 years). Initial population density is 3K/4. �DO 207 K 20 FAWNS 50 FAWNS pv= 0.0 0.9 0.9 0.8 0.8 0.7 0.7 0.6 0.6 0.5 0.5 0.4 0.4 0.3 0.3 0.2 0.2 0.1 0.1 "-0 0 0.6 PV 0::: W ~ 0 a.. 0.8 0.0 1.0 0.9 0.8 0.7 0.7 0.6 0.6 0.5 0.5 0.4 0.4 0.3 0.3 0.2 0.2 0.1 0.1 0.4 PV 0.6 0.8 0.6 0.8 1.0 '! 0. -, b, 0...•. 0.2 1.0 = 0.04 0.4 PV 0.9 0.6 0.8 1.0 0.8 1.0 = 0.04 0.9 0.8 0.8 0.7 I)] 0.7 0.6 0.6 0.5 0.5 0.4 0.2 0.4 PV = 0.02 0.9 0.2 0.2 = 0.02 0.8 0.3 = 0.0 PV 0. " 0.4 , Cl. 0. 0.3 0. , 0.2 0.1 0.1 0.2 0.4 0.6 0.8 0 1.0 0.0 0.2 0.4 0.6 CV (DENSITY ESTIMATE) ----0---- ......•........•. 0- .....••••..•.• 5 years 10 years 25 years 50 years ·············0·········· 100 years Fig. 9. Power of logistic regression, adjusted for overdispersion of the response variable, as sampling variance increases in the explanatory variable, density. Annual fawn survival rate is the response variable. The left column of graphs result from a less precise estimate of fawn survival rate (k = 20), the right column of graphs are results of the more precise estimates (k = 50). Process variation in the fawn survival rate estimates increases from top to bottom (PV = 0.0, 0.02, 0.04). Each graph contains 5 curves depicting increased time series lengths (n = 5, 10, 25, 50, and 100 years). Initial population density is K. �208 (a) 0.8 10 years 0.7 •• •• ns > •~ 0.6 =s C/) a. II. s::::: ~ 0.5 ns -. II u, 0.4 0.3 ~--~ _L o W W- L___~ _L__~~ ~ t L-__~ ~ 00 Population density ~ 100 K (b) 100 >. ..•... 80 ......... -0 ...--.-0 -······0·-- - _O---·O ....-··O ...---~--·--~ t/) c (1) "'C 60 c \'. 0 \ :;:; C'CS \'. 40 0 ', '. \ ...... * ' \ \ ::J a. a, '; \ \ ', '. $ >$<. • • .$ $ $ 20 ----0---- ----0----0 0 2 4 6 8 10 Years 0------ -0 2 .--_. ._.-. __ Kl4 )]( Kl2 0-----·-0 3K14 Fig. 10. (a) Annual overwinter fawn survival and population densities generated from the density-dependent population growth model described in the text. Vertical dashed lines enclose the desired range of population densities to capture in a la-year time series for high power. (b) Ten year population densities resulting from single year population reductions in year 2 (all populations in year 1 are near K). Each line on the graph depicts decreased treatment effect (K - 2, K - K/4, K - K/2, and K - 3K/4). Horizontal dashed lines enclose the desired range of population densities to capture in a 10year time series for high power as determined from (a). �209 K ••• Kl4 5 YEARS 10 YEARS pV= 0.0 pv= 0.0 -0-:.::-;.:-.:-:- ~. 0.9 0.9 0.8 0.8 0.7 0.7 'b II 0.5 \ 0 0.4 , 0.3 • -, 0.1 0.0 0.2 b··>tl· , II- 0.4 "U'\~";":'i";:.'1iI 0.3 'D ••• 'fJ_ 0 .... _. .... 0.6 0.8 0.2 0.1 0 1.0 0.0 0.2 W s: 0 a. 0.9 0.9 0.8 0.8 0.7 0.7 '''"8.. 0.6 0.6 -'_11.:"8.., 0.5 0.5 0.4 0.4 0.3 0.3 0.2 0.2 0.1 0.1 0 0 0.0 0.2 0.4 0.6 0.4 0.6 0.8 1.0 PV = 0.02 PV = 0.02 ~ ';lJ -,-,n.,.: 0.4 , n "->~, -Do. 0.5 .. 0 0.2 h. '.:0 0.6 -, \ 0.6 \-,'> "q~ 0.8 1.0 . o ::!;t1::-::'a- __ . ~ 0.0 0.2 PV = 0.04 0.4 0.6 0.8 1.0 PV = 0.04 0.9 0.9 0.8 0.8 0.7 0.7 0.6 0.6 0.5 0.5 0.4 0.4 0.3 0.3 0.2 0.2 0.1 0.1 0.2 0.4 0.6 0.8 1.0 0 0.0 0.2 0.4 0.6 0.8 1.0 CV (DENSITY ESTIMATE) DO = K14 1 YEAR 2 YEAR 3 YEAR 4 YEAR 5 YEAR ---0--- ·······m······- Fig. 11. Power of logistic regression, adjusted for overdispersion of the response variable, following a single year manipulation from K - K /4 as sampling variance increases in the density estimates (c = 0.0 to 1.0). The left column of graphs have time series length of 5 years; each graph has 4 lines depicting increasing number of years of pre-treatment data (x 1,2,3,4) compared to Do = K/4 and n = 5. The right column has 10 years of data; each graph with 6 lines depicting number of years of pre-treatment data (x = 1,2,3,4,5) compared to Do = K/4 and n = 10. Process variation in the fawn survival rate estimates increases from top to bottom (PV = 0.0, 0.02, 0.04). = �210 K • Kl2 10 YEARS 5 YEARS pV= 0.0 pv=o.O 0.9 0.9 0.8 0.8 0.7 0.7 0.6 0.6 0.5 0.5 0.4 0.4 0.3 0.3 0.2 0.2 0.1 0.1 0 0.2 0.4 0.6 0.8 0.0 0.2 0.7 0.7 0.6 0.6 0.5 0.5 0.4 0.4 0.3 0.3 0.2 0.2 a. 1.0 0.8 0.8 s: 0 0.8 0.9 0.9 W 0.6 PV = 0.02 PV = 0.02 ~ 0.4 1.0 0.1 0.1 0 0.2 0.4 0.6 0.8 1.0 0.2 0.0 PV = 0.04 0.6 0.4 PV 0.8 1.0 0.8 1.0 = 0.04 1 0.9 0.9 0.8 0.8 0.7 0.7 0.6 0.6 0.5 0.5 0.4 0.4 0.3 0.3 0.2 0.2 0.1 0 m ·s 0.1 0.0 0.2 0.4 0.6 0.8 0 1.0 0.0 0.2 0.4 0.6 CV (DENSITY ESTIMATE) oo = Kl2 1 YEAR 2 YEAR ......... {) . 3 YEAR _._- - _.- ... 4 YEAR --~--- 5 YEAR •..•..• 6- ..•..• Fig. 12. Power of logistic regression, adjusted for overdispersion of the response variable, following a single year manipulation from K - K /2 as sampling variance increases in the density estimates (c 0.0 to 1.0). The left column of graphs have time series length of 5 years; each graph has 4 lines depicting increasing number of years of pre-treatment data (x = 1,2,3,4) compared to Do = K/2 and n = 5. The right column has 10 years of data; each graph with 6 lines depicting number of years of pre-treatment data (x = 1,2,3,4,5) compared to Do = K/2 and n = 10. Process variation in the fawn survival rate estimates increases from top to bottom (PV = 0.0, 0.02, 0.04). = �211 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS state of project REPORT Colorado No. W-1S3-R Mammals Research Work Plan No. 3A Pronghorn Research Job No. 6 Pronghorn Winter Period Covered: Authors: July Wheat Damage study 1, 1994 - June 30, 1995 D. C. Strohmeyer, G. C. White, and R. B. Gill ABSTRACT Wildlife managers have responded to winter wheat damage complaints by reducing pronghorn (Antilocapra americana) numbers via hunting and trapping removals. Recent research (Torbit et al. 1993) has suggested pronghorn may not damage winter wheat, implying reducing pronghorn populations may not be necessary. For this suggestion to be true, free-ranging pronghorn must stop foraging on wheat as wheat enters the jointing stage. Our research focuses on evaluating this suggestion. For two years, marked, free-ranging pronghorn have stopped using wheat prior to when winter wheat became vulnerable to grazing damage. Preliminary results of our feeding trials suggest that nutrition is not sole criteria tame pronghorn use to select different developmental stages of wheat. ��213 PRONGHORN WINTER D. C. Strohmeyer, WHEAT DAMAGE G. C. White, STUDY and R. B. Gill P.N. OBJECTIVES Elucidate pronghorn movement 1. Analyze spring the telemetry 1994. 2. Conduct preliminary patterns concerning SEGMENT OBJECTIVES data and vegetation feeding winter wheat data collected use. during the trials. METHODS AND MATERIALS 1995 TELEMETRY The study area encompassed the Pawnee National Grassland in Weld County, Colorado. It was at the northern edge of the shortgrass steppe region. Fourteen females were radio-collared in February 1993. vegetation use of marked animals was monitored via aerial and ground telemetry. Logistic regression modalled pronghorn behavior. pronghorn use of vegetation types (native range or winter wheat) was the dependent variable. This discrete, nominally-scaled variable was recorded as use versus non-use and was binomially distributed. Time (t), a continuous variable, and collar (C), a discrete, nominally-scaled variable, were the predictor variables: logit (1tN) = Po In( ::) = + P1 t + P2c , 1 1tw = 1- 7tN ' where ~w = = ~N probability probability that pronghorn that pronghorn use wheat, and use native range. The null hypothesis was that there were no changes in the vegetation type used by pronghorn over time, Uo: PI = O. The alternative hypothesis was that the logit of the ~w increased linearly over time, Ha: PI > o. 1994 NUTRITION DATA Vegetation data were collected in wheat-prairie pairs in Weld County, Colorado. One set of data from each pair was collected at a marked-animal relocation. The second half of the pair was selected by closest proximity. Sampling periods were 1-week intervals. Vegetation was clipped, then ovendried. Standard chemical analyses were done by the Colorado Division of Wildlife nutrition lab. Nutritional dynamics were quantified via 2 diet compositions. The first was a diet of 100% winter wheat. The second was a 100% shortgrass prairie diet taken from Schwartz (1977:29). We related pronghorn logistic regression. habitat use patterns to pronghorn diet quality via This analysis was repeated separately for dietary �214 neutral detergent fiber logit(1tR) (NDF) and dietary = l~::) = crude protein. The model was: [io + [i1P + [i2W + [i3P•W, where ~v = probability that pronghorn use wheat (~v = 1 - ~R)' ~R = probability that pronghorn use shortgrass prairie, W = winter wheat diet quality, and P = shortgrass prairie diet quality. We used this model to quantitatively relate the period when there were no differences in pronghorn use of vegetation types, ~R = ~v 0.5, to diet quality values for each marked pronghorn. = FEEDING TRIALS We compared preferences of female, hand-reared pronghorn for two phenological stages of wheat via two-way feeding trials. Twelve replications (36 experimental trials = 1 treatment * 12 animals * 3 replications) were done. The animals were in outside pens and were on a standard, nutritionallybalanced diet with ad libitum water. The wheat was grown in a greenhouse after vernalization. We examined whether or not pronghorn had a preference or behaved randomly (no preference). Pronghorn preference data were binomially distributed, use or non-use of tillering wheat. RESULTS AND DISCUSSION 1995 TELEMETRY Five pronghorn were relocateable during the 1995 field season. Five deaths were confirmed during the study. Two transmitter failures were visually verified. The fates of two marked pronghorn were not determined. Pronghorn shifted from winter wheat fields to shortgrass prairie as we expected (~ < 0.0001). The last marked-animal relocation on wheat was March 23rd, which was about one month earlier than previous years (Table 1). More importantly, this behavioral shift occurred prior to when winter wheat became vulnerable to grazing damage. All wheat had entered the jointing phenological stage and became vulnerable to grazing damage on April 15th. Table 1. Date of last relocation on winter wheat of marked pronghorn, county, Colorado, 1993-1995. Collar Year Date of Most Recent Wheat Relocation 292 April 10 1993 April 10 570 598 March 21 660 March 20 108 March 27 1994 171 April 12 292 March 27 340 March 13 550 April 16 570 April 14 April 598 8 660 March 3 1995 108 March 18 292 March 18 550 March 21 598 March 23 660 February 9 Weld �215 1994 NUTRITION DATA Neutral detergent fiber data showed slight negative trends for both prairie and wheat diets (Figure 1). Crude protein data showed a negative trend for wheat diets while increasing then decreasing for prairie diets (Figure 2). The period of no difference in pronghorn vegetation use corresponded to mean crude protein contents of 17% and 15% for wheat and prairie diets, respectively. FEEDING TRIALS The results of the preliminary trials were the opposite of what we expected (Table 2). The decreasing selection of tillering wheat during the trials suggested that pronghorn choices were not motivated solely by nutrition. For example, the pronghorn may have been more interested in plant height. Table 2. Wheat feeding trials. Animal Bolero DD Nicki Sena Yoyo Phlox Typha Athena 236 Joanie Casey Ellie stage selected by initial muzzle contact Trial 1 joint joint joint tiller tiller joint joint Trial 2 tiller joint joint tiller joint joint joint joint tiller tiller tiller joint tiller joint joint during preliminary Trial 3 joint joint joint joint joint tiller joint joint joint joint joint joint NEXT QUARTER'S OBJECTIVES There are 2 objectives for next quarter: (1) finish the laboratory for the 1995 telemetry data, and (2) conduct more feeding trials. LITERATURE CITED Schwartz, C. C. 1977. Pronghorn grazing strategies prairie, Colorado. Ph.D. Thesis. Colo. State 113pp. Torbit, S. C., R. B. Gill, of pronghorn grazing 57: 173-181. analyzes on the shortgrass Univ., Fort Collins. A. W. Alldredge., and J. F. Liewer. 1993. Impacts on winter wheat in Colorado. J. Wildl. Manage. �216 100 Postjointing Prejointing 80 - • • • • ~ ~ l- Q) u: .•... 60 - • I ~ 0 I 40 - 0 • I.I!I·~~ ~ I CJ ~ Q) ro t \I 0 Q) +-' Q) I El c: 0 • • 0 ..0 0 § ~ 0 0 bl 0 •• 0 B I- ::; Q) z 20 - O~--~--~I---.L_-'-I-'~-'I---'--~ o 5 o Figure 1. Neutral detergent pronghorn shortgrass prairie Grassland, March-June 1994. 10 Time (weeks) 20 15 • Prairie Diet Wheat Diet fiber contents for a 100% winter wheat diet and a diet. Data were collected on the Pawnee National 40 Postjointing Prejointing 0 I 30- --- ~ ~ c: ·iii +-' 0 I- a, 0 0 I 0 ~ I • I • • • 20- Q) "0 ::) 0 I I I 0 § 0 0 IBi •• ~I I I- o • 10 - I 0 I • • !~•• 5 0 0 I ~ ~ Iii bl 0 • • ~ • I I 10 Time (weeks) [J Wheat Diet Figure 2. shortgrass Grassland, El • • I 0 0 0 15 20 • Prairie Diet Crude protein contents for a 100% winter wheat diet and a pronghorn prairie diet. Data were collected on the Pawnee National March-June 1994. �217 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS State of Project REPORT ~----~C~o~l~o~r~a~d~o~---------No. W-153-R-4 Mammals Research Work Plan No. 3A Pronghorn Job No. 7 Experimental Pronghorn Surveys Fixedwing Line Transects and Helicopter Quadrats Period Author: Covered: July Investigations Using 1, 1994 - June, 30, 1995 T.M. Pojar ABSTRACT The second year of data collection was completed to compare pronghorn population density estimates obtained by fixedwing line transect and helicopter quadrat surveys. The line transect estimate for 1995 is not available for this report because the data was not received from Western Air Research Incorporated in time for analysis. There were modifications in both the line transect and quadrat surveys in 1995 designed to reduce the variance of both methods. Line transects were run on 1.609 km (1 mile) intervals increasing the sample from 4 to 16 transects for the surveyed area. The' UTM coordinates of all groups seen on the line survey were recorded and used to re-stratify the quadrat survey. There was little correlation between number of subjects seen on quadrats during the line transect survey and the number seen during the quadrat survey (R2=0.0905). The regression was different from Q (~=0.0592) but has little predictive power. The estimated population was 7,708 ± 32.5% from the helicopter quadrat survey. The cost of the fixedwing line transect survey was approximately $1200. and the helicopter survey cost was $3,750.00 (excluding ferry time). A Global position System (GPS) was used for navigation. The flight route was recorded (via GPS) during the search of 32 of the 40 quadrats. The actual area searched was 14.8% larger (~ < 0.0001, paired t-testJ n=3~) than the designateq area. ,The mean designated area was 2.564 Jtrti2 (0.9899 mi2) was not different 'from 2 ~59 km2 (1 mi2) f~ = 0.105) and the actual area searched was '2.944 krit2' (1'.1367 mil)., ��219 EXPERIMENTAL PRONGHORN SURVEYS USING FIXEDWING HELICOPTER QUADRATS Thomas LINE TRANSECTS AND M. Pojar P.N. OBJECTIVE Compare fixedwing line transect pronghorn density. and helicopter SEGMENT quadrat in estimating OBJECTIVES 1. Compare pronghorn density estimate and precision transect survey with helicopter quadrat survey. 2. Evaluate 3. Test accuracy of a Global geographic points. consistency surveys of line transect Positioning of fixed-wing line data analysis. System (GPS) in locating known STUDY AREA The study area is 1,171 km2 (452 mi2) of sagebrush steppe pronghorn north and west of Craig. It is described in Pojar et ale (1995). METHODS The methods are outlined habitat AND MATERIALS in the Program Narrative, see Pojar (1994), Appendix I. There were modifications in both the line transect survey and helicopter quadrat survey in 1995. The intent was to improve the precision of both survey results. Line transects were run north and south at 1.609 km (1 mile) intervals, which increased the sample of transects from 4 to 16. The increased sample size will most likely improve the precision of the line estimate (17% coefficient of variation of li reduced to < 5%). The UTM coordinates of all groups seen on the line survey were recorded and the relative density across the area was used as a stratifying factor for the subsequent helicopter quadrat survey. Flying 16 lines "sampled" every quadrat in the area because the area is 25.7 km (16 miles) wide. From this relative density information, the 40 sample quadrats for the quadrat survey were stratified. Only 1 of the original 40 quadrats was changed with the new stratification. Strata boundaries, however, were changed in accordance with the relative densities observed from the line transect survey. A Global Position System (GPS) was used for locating quadrat corners and to "track" the actual route of the search. The total number of quadrats remained unchanged from the previous survey in 1994 (40 quadrats). The quadrats were searched using a Bell-Soloy helicopter. The GPS external antenna was mounted on the rear boom behind the engine and a 12 volt connection to the helicopter electrical system was used to power the GPS. The GPS was used exclusively to locate quadrat corners and navigate the quadrat perimeter; if old quadrat markers were observed they were ignored. The navigation was done via GPS by the second observer/navigator communicating directions to the pilot while the primary observer counted and recorded pronghorn groups on a tape recorder. RESULTS The line transect survey was done on May 20 and 21, 1995 by Western Air Research Incorporated (Driggs, Idaho) with Jeff Madison and Mike Bauman (CDOW) as observers. Methodology followed that described by Johnson et ale 1991. Analysis procedures for line transect data will follow those described by �220 Buckland et a1. (1993) and Laake et a1. (1993). In addition, the data will be subjected to analysis by the program, TRANSAN (Routledge and Fyfe 1992), which implements the shape-restricted estimator of Johnson and Routledge (1985). The helicopter survey was done on May 25 and 26, 1995 and took 7.5 hours of flight time to complete (excluding ferry time). The new stratification alignment (Table 1) was used in the analysis. The population estimate was 7,708 with a 90% confidence interval of ± 32.5%. The lower population estimate in 1995 compared to 1994 (8,465) was expected because of increased harvest in 1994 (Jeff Madison, Pers. corom.). Tracking of the actual route around the perimeter of a quadrat by GPS indicated that, on average, the actual area surveyed was 14.8% larger that of the designated quadrat. The designated quadrat area was not different (~=0.105) from 2.59 km2 (1 mi2). If the population estimate were adjusted for the actual area surveyed, it would be 6,712. stratification based on relative densities during the line transect survey was not helpful in reducing the variance of the quadrat estimate. The regression coefficient of quadrat counts on line counts was different from Q (~=0.0592) but with a very faint correlation (R2=0.0905) resulting in no practical predictive power (Figure 1). REFERENCES CITED Buckland, S. T., D. R. Anderson, K. P. Burnham, and J. L. Laake. 1993. Distance sampling: estimating abundance of biological populations. Chapman and Hall, New York, N.Y. 471pp. Johnson, B. K., F. G. Lindzey, and R. J. Guenzel. 1991. Use of aerial line transect surveys to estimate pronghorn populations in Wyoming. Wildl. Soc. Bull. 19:315-321. Johnson, E. G., and R. D. Routledge. 1985. The line transect method: a nonparametric estimator based on shape restrictions. Biometrics 41:669679. Laake, J. L., S. T. Buckland, D. R. Anderson, and K. P. Burnham. 1993. DISTANCE user's guide. Version 2.0. Colo. Coop. Fish and Wildl. Res. Unit, Colorado State Univ., Fort Collins. 72pp. Pojar, T. M. 1994. Experimental pronghorn surveys using fixedwing line transects and helicopter quadrats. Colo. Div. Wi1d1. Res. Rep. July, pp163-172. _______ , D. C. Bowden, and R. B. Gill. 1995. Aerial counting experiments to estimate pronghorn density and herd structure. J. Wildl. Manage. 59:117-128. Routledge, R. D., and D. A. Fyfe. 1992. TRANSAN: Line transect estimates based on shape restrictions. Wildl. Soc. Bull. 20:455-456. �221 Table 1. Strata used in the 1995 helicopter quadrat survey for pronghorn in the Craig, Colorado study area 1,171 km2 (452 mi2). These strata were based on relative density observed during a line transect survey conducted 5 days earlier. Total count represents the total number of animals seen on each quadrat and was used in the estimation of population size. Stratum Sample Quads Total count 124.3 (48) 1 7 8 9 6 49 3 0 II 93.2 (36) 2 4 5 6 24 2 39 42 III 132.1 3 11 12 2 6 9 IV 98.4 10 16 17 4 16 7 V 251.1 (97) 13 14 15 22 23 24 25 26 27 36 37 22 32 9 5 12 19 6 28 3 26 10 VI 106.2 (41) 18 19 20 21 29 0 4 2 35 62 VII 194.3 (75) 28 30 31 32 33 30 53 8 14 9 VIII 170.9 (66) 34 35 38 39 40 57 0 13 12 20 I (51) (38) �222 70r-------------------------------------------~ o 60 o 0.0905 o ...., 50 o C ::J o 40 o ...., ""C co •... co 30 ::J o 20 0 10 8 8 8 0 0 0 I 0 5 0 I I I I 10 15 20 25 Line count Figure 1. Regression of quadrat sample unit counts on corresponding line counts, Craig, Colorado, 1995. The regression is different from 0 (~ = 0.0592) but has little predictive power. 30 �223 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS State of Project Work Colorado No. Job No. Author: Covered: Mammals W-153-R-8 Plan No. Period REPOR~ July Research 4A Mountain 3 Mountain goat numbers, distribution, and dispersal in the northern Collegiate range. Goat Investigations 1, 1994 - June 30, 1995 D. F. Reed ABS~RACT This study involving mark-recapture and the pioneering of mountain goats in the northern Collegiate range required an additional $13,500 if it were to go forward. These moneys were made available from the bighorn sheep and mountain goat auction and raffle funds and were allocated near the beginning of the segment. Approval was obtained from the animal welfare committee and 50 mountain goats were radio-collared 10-15 August 1995. A helicopter count was conducted 25 August 1995 during which a low number of the marks (radiocollars) were observed and identification of distinguishing marks .(numbered or color coded collars) was poor. Costs of the helicopter net-gunning (capturing and attaching radio-collars) exceeded allocated budget resources - hence field work was curtailed during winter and spring. ��225 MOUNTAIN GOA~ NUMBERS, DIS~IBUTION, AND DISPERSAL IN THE NORTHERN COLLEGIAm RANGE Dale F. Reed P •N. OBJE~IVE To improve estimates of mountain goat populations by mark-resight methodology and to estimate dispersal rates in an increasing mountain goat population. SEG~ OBJE~IVES 1. Mark (radio-collar) Plata) • 2. Conduct a preliminary helicopter count to establish an aerial route, obtain a resighting estimate, and examine the sightability of distinguishing marks. 3. Conduct fixed-wing 50 mountain flights goats in the North to estimate locations Collegiates of telemetered (Quail-La animals. STUDY AREA The study area is described in the Program Narrative (Appendix A). ME~BODS AND MA~ERIALS The methods are outlined in the Program Narrative (Appendix A). For net-gun capturing a Hughes 500C (Helicopter Wildlife Management) was used. Once the pilot and gunner located and netted an animal, the gunner was quickly landed to handle the animal while the pilot often left to bring in another person waiting nearby on the mountain to assist and finish collaring, taking samples, and releasing the animal. This allowed the gunner to be off netting another animal. Count techniques included For counting a Sou loy (High country) was used. observers, the middle person counting the left side and forward, and the person on the right side, counting right. The middle observer also did orienteering and recording of observations on layed-out quad maps. two For estimating locations by telemetry a Cessna 185 was used. The technique involved flying relatively high (15,000-16,000 ft) listening to the strength of signals from two external antennas. Forty-five and five of the transmitters had frequencies of 165.20-165.72 MHz and 173.0372-173.2127 MHz, respectively. A LOTEK (Suretrack STR1000) receiver was used for the 165 frequencies and a Telonics (TR-1) was used for the 173 frequencies (Channels 7-8, 16-18). Collars supporting the transmitters were either color coded or color coded and numbered for individual identification. RESUL~S Consistent with the plans in the Program Narrative, 50 mountain goats were captured and radio-collared from 10-15 August 1995. Most of the animals collared (n = 30) were adult females (Table 1). The efficiency of capturing and radio-collaring mountain goats via helicopter net-gunning may be calculated as 2.4 animals/hour but variables likely influence such efforts (e.g. pilot and net-gunner performance, weather, animal distribution and group size, etc.). �226 The helicopter count conducted 25 August 1995 yielded 107 mountain goats, 74 north of Clear Creek and 33 south of Clear Creek. Of the 50 collared animals only 13 were observed. Gusty winds prevented classification of 21 animals and searching the head of Willis Gulch and the west side of Hope Mountain. Furthermore, we were unable to identify collar numbers from the helicopter. Hopefully, weather conditions in the future will permit better sightability and closer viewing for collar number identification. Table 1. Number of mountain goats captured and radio-collared 1995 by sex/age group and approximate hours of flight. 10-15 August Date AQQroximate hours of flight Male 10 11 14 15 Total Adult Female 2-~r old Male Female 4 6 2 0 0 12 3 1 (waited two days for animals 0 8 0 0 6 4 0 0 10 30 5 1 Yearling Male Female 1 0 1 0 to adjust) 0 0 0 2 0 4 Total 13 17 6 7 8 12 4 4 50 21 Measured from beginning in morning to ending for day -- includes capturing, animal handling, and replenishing. searching, Four fixed-wing flights were conducted. Most of the telemetered animals were estimated to be near where they had been collared in August, but several had moved up to 7 km (Table 2). Locating efforts will begin in ernest during the next segment. �227 Table 2. Collar frequency or channel (color/number designation), location when collared (10-15 Aug 94), and estimated locations during fixed-wing flights 29 Nov 94 - 27 Jun 95. Frequency/Channel (color/number) 10-15 Aug 94 165.200 (Blu 20) Galena (Grn 21) (Grn 22) (Grn 23) (Grn 24) (Grn 25) (Grn 26) (Y/B 27) (Grn 28) (Yel 29) (Grn 30) (Blu 32) (Or 33) (Yel 34) (Blu 35) (Yel 36) (Yel 37) (Yel 38) (Yel 39) (Blu 41) (Blu 42) (Blu 43) (Or 45) (Or 46) (Blu 47) (Blu 48) (Or 49) (Blu 50) (Blu 51) (Blu 52) (Blu 53) (Blu 54) (Or 55) (Blu 59) (Blu 60) (Blu 61) (Blu 62) (Blu 63) (Blu 64) (B/Y 65) (Orange) (Yellow) W 210 220 230 240 250 260 270 280 290 300 320 330 340 350 360 371 380 390 410 420 430 450 460 470 480 490 500 510 520 530 540 550 590 600 610 620 630 640 650 660 670 680 710 720 7 8 16 17 18 (B & W) (Black) (Blu/Or) (Grn 7) (Grn 8) (Blue) (Grn 17) (Grn 18) .. .. .. .. 29 Nov 94 Locations 9 Dec 94 23 Dec 94 27 Jun 95 (- same) " Cirque .. E Crystal L SE Hope Middle Mtn (A) Galena (?) Willis Cirque (- same) .. (none - harvested 09/25/94 7 km SE of A) (- same) (- 2-3 km W) (- same as 300) (? ) NE Ellingwood " W Galena " (?) (- same as 340) NW Twin Pks (- same) SW Willis Cirque (- same) Galena (?) (W Twin Pks) (?) (none - harvested 09/ /94 W Galena (- same) (S Clear Ck, NE Waverly Mtn) .. ( (? ) Middle Galena Mtn .. (S Clear (- same) (? ) (? ) .. •• .. ) (S & low on Quail, (?) ( .. ( .. 5-6 km NE) Ck, NE Waverly Mtn) •• .. ) .. .. ) W Galena (? (? (? (? •• ) ) ) ) •• Upper Galena Galena Middle Mtn (?) (?) (? ) SE La Plata •• (none - harvested 09/06/94 E Galena) (S Clear Ck, NE Waverly Mtn) (Low & SW Quail) (S Independence Pass) ) (" " .. (" ( Pk .. (? ) Middle Mtn (A) SE La Plata Pk S Willis Lake W Galena Cirque Sayres Gulch SE Twin Pks SW " •• N Middle Mtn SE La Plata Pk W Willis Lake NE Ellingwood Middle Mtn (?) ) .. ) 7km /94 (none - harvested 09/ (S Independence Pass) (- same) ( .. ) ( .. South A) ) (mortality) (none - not found) (S Silver King Lake) (Missouri Mtn) (.. (S Silver .. ) King Lake �228 �229 Appendix A PROGRAM NARRATIVE state project No. Colorado W-153-R Mammals Research Work Plan No. 4A Mountain Job No. 3 Mountain goat numbers, distribution, and dispersal in the northern Collegiate range. A. Goat Investigations NEED Problem Mountain goats (Oreamnos americanus) generally have not been considered indigenous to Colorado. However, the Wildlife Commission passed a resolution on 11 March 1993 proclaiming the mountain goat "native." Whichever the case, they were either introduced or re-introduced into the collegiate Range in 1948 and subsequently into other areas including Mt. Evans, San Juan Mountains, Gore Range, and Marcellina Mountain (Denney 1977). These areas were often within the historical, if not present, range of mountain sheep (Ovis canadensis canadensis). The problem is that as mountain goat populations increase, they may expand their range into areas managed specifically for mountain sheep. The Division's policy in regard to areas managed for mountain sheep and areas where mountain goat populations will be permitted or considered for trans locations has been debated. The concern has been and continues to be that at some threshold density of one or both of these species, "competition" may occur resulting in a competitive advantage for mountain goats (Bassow 1985). Furthermore, a concern has developed that mountain goats in the northern portion of G-3N (Quail Mountain-La Plata Peak area) are increasing and dispersing at unknown rates. Consequently, it is important that methods for obtaining better population estimates, distribution, and dispersal rates of mountain goats be developed. Literature There is a substantial volume of literature concerning the study of wild animal populations. Mark-recapture theory, models, and various statistical treatments are involved in methods frequently used (see reviews by Caughley 1977; otis et aL 1978; Overton and Davis 1969; Seber 1973, 1982; White et al. 1982). The amount of literature is greatly reduced when considering only the "size of the population" versus 8 other properties that can be investigated by some form of mark-recapture (Caughley 1977:133) and when limiting the application to wild ungulates (Strandgaard 1967, Woolf 1973, Rice and Harder 1977, Bartmann et al. 1987). Aspects of mark-recapture techniques most applicable to a study of free-ranging wild ungulates include the importance of experimental design, of equal catchability of all individuals, marked and unmarked, where truth of the assumption can be tested in a pilot experiment or have such a test built into the experimental design (Caughley 1977:134), and of adequate sample sizes including the proportion of marked animals in the population. concerning the latter aspect, large proportions (>67% Strandgaard 1967, >45% - Bartmann et al. 1987) of small populations need to be marked to generate reliable population estimates (White et al. 1982). .. �230 Approaches to the study of population size in free-ranging wild ungulates might include 1) the Petersen estimate (Petersen 1896, Lincoln 1930) or the simplest mark-recapture procedure (Minta and Mangel 1989) which calls for marking on one occasion and recording proportion of marked animals captured (or "sighted" since actual capture is not necessary if marks can be recognized [Eberhardt 19691) on a second occasion (Caughley 1977:141), 2) Schumacher's method which calls for marking on several occasions and is estimated from the rate at which the proportion of marked individuals rise as progressively more are marked (Caughley 1977:145), 3) methods involving the assumption of "closure" - the simplest appropriate model of 8 possibilities (models Mo, Mt, Mb, Mh, Mtb, Mth, Mbh, and Mtbh) (Otis et al. 1978, White et al. 1982), 4) methods involving the assumption of "open" populations where the process of birth, death, and migration are allowed to operate--essential elements are incorporated in the classic Jolly-Seber model (Seber 1978:196-232, Seber 1982:196-232), and 5) procedures involving maximum likelihood estimates (MLE) used in combining repeated Lincoln-Petersen estimates (Bartmann et al. 1987). Related to increases in the number and changes in the distribution of mountain goats is the question of dispersal. Dispersal-rate studies by Caughley (1970), Clarke (1971), and Davidson (1973) of populations of thar, red deer, and sika deer introduced into New Zealand indicate that dispersal rates for these three ungulate species varied by species, with sika deer dispersing slowest (1 mi/yr), followed by thar (3 mi/yr), and red deer (7 mi/yr). Caughley (1970) proposed two models to account for observed dispersal rates of thar: one was density-dependent, while the other was a model of random dispersal. His data suggested that dispersal rates were most closely approximated by the random dispersal model. It would be valuable to conduct "goodness of fit" tests between observed mountain goat dispersal rates and predicted dispersal rates from Caughley's (1970) two models, because if the predicted pattern closely fit either of the two predictive models it would allow managers to predict the rate of spread of mountain goats from new introduction sites and adjust management strategies accordingly. B. OBJECTIVES The overall objectives of this study are: To improve estimates of mountain goat populations by independently testing the "one helicopter count method" against mark-resight methodology and to estimate dispersal rates in an increasing mountain goat population. Specific objectives may be stated as null hypotheses: 1. The population of mountain goats on Quail-La Plata (G-3N) as estimated mark-recapture methods is not significantly different than the minimum number as determined from established helicopter counts. 2. Mountain goat dispersal rates are not significantly different than Caughley's (1970) "random" dispersal model or his "density-dependent" dispersal model. C. EXPECTED RESULTS by OR BENEFITS Obtaining better population estimates should improve the Division's ability to match harvest regulations with the number of mountain goats. Information on dispersal rates of mountain goats should provide a better basis from which harvest and translocation management strategies can be formulated. �231 D. APPROACH 1. Methods: The methods will generally follow the capture-resight technique of Minta and Mangel (1989). About 50 percent of the estimated mountain goat population of 100 will be marked with radiotelemetry collars to assure that marked animals are alive and in the population (population closure assumption) during counts. Marking a high percent of the population will provide relatively narrow confidence intervals at the 95 percent confidence level. Capture-resight will require independence among both captures (marking) and resights of the animals. Marking - Mountain goats (n = 50) will be captured by net-gun from a helicopter and marked with individually identifiable (unique) telemetry collars equipped with mortality sensors. The capture and marking will be done during 5-10 days during July or August in 1994 (first year), after the time adult females return from kidding areas with their neonates and the alpine areas are relatively free of snow. No new marks will be added to the population until the following year. During the second and third years, an estimated additional 5-10 goats maybe radio-collared each year to replace any losses due to natural or hunting mortalities. Counts - Either during the second year ('95) or during the second and third years ('95 and '96), 5 helicopter counts will be conducted in the Quail Mountain-La Plata Peak area during July and August in order to sight marks in the population of mountain goats. Which option will depend on the rate of increase and the rate of dispersal of mountain goats as determined in 1995. The objective here is to obtain capture-resight estimates of the mountain goat population in order to independently verify single ground count methods and to monitor movements and dispersal. The counts will cover the previously established helicopter flight routes and areas where mountain goats may have pioneered. This means that the counts will cover the entire area and, hence, will not be predicated on sampling or small-sample biases (White et al. 1982). An approximate schedule of the counts for the 2 years are as follows: 1995 6 13 20 27 3 Jul Jul Jul Jul Aug 1996 5 12 19 26 2 Jul Jul Jul Jul Aug It is estimated that a minimum of 2 hours per count will be required to count the area. In addition, immediately after each count about 1 hour of flight time will be needed to verify with telemetry that the marks (radio-collared animals) are located in or out the population. An additional hour may be needed to check the location of dispersing animals. 2. Analysis: Ho 1: Mark-recapture analyses for purposes of estimating population size will generally follow the technique of the Petersen estimator for mark and resight data considered by Minta and Mangel (1989). Models assuming demographic closure (closed populations) are based on the relatively short period between marking and recapturing (sighting) and the verification of telemetry collars is designed for this. A simple Monte Carlo simulation will be run to provide a full probability distribution for the population of mountain goats. From these probability distributions, maximum likelihood estimates and likelihood �232 intervals for the populations Mangel [1989] for description Basic program). will be computed (see Appendix in Minta and of Monte Carlo simulation and availability of PC Ho 2: Tests of "goodness of fit" between observed and predictive models of dispersal rates will follow the work of Caughley (1970). Both of his models, "random dispersal" and "density-dependent dispersal," will be tested against observed rates of dispersal in mountain goats. A graphical framework similar to that of Waser et ale (1994) will be used to describe dispersal rates, rates of survival of dispersers, and rates of survival of philopatric animals. Schedule Activity Fiscal Year Capture and mark, monitor dispersal Conduct counts and monitor dispersal Same as 1995-96 Data analysis and publication 1994-95 1995-96 1996-97 1997-98 Personnel Principal Investigator Co-investigator Co-investigator D. F. Reed J. Vayhinger S. R. Ogilvie Estimated Cost Months (01) Personal Services D. F. Reed D. Hall 12 1 Supplies and Services Helicopter capture Helicopter flights Vehicle (4x4) Misc (n (n = = 50 @ 450 ea) 1 @ 4 hr & 455/hr) 930 (28) Travel Expenses GEOGRAPHIC (Equipment) 83,500 TOTAL E. 22,500 1,820 878 400 25,598 Total (31) Capital Expenditures 54,372 2,600 56,972 Total (02) Operating Amount LOCATION The principal area of study will involve the range between Lake Creek on the north and Clear Creek on the south, and east of Quail Mountain to Grizzly Peak (on the continental divide) in the west, excluding areas designated as wilderness. other prominent features include La Plata Peak (14,336 ft in elevation and located about 23 miles northwest of Buena Vista and 4 miles �northwest of Winfield), Twin Peaks, and Mount Hope. Adjoining areas including the south slopes of Mount Elbert (14,433 ft) will be included depending on the presence or dispersal of mountain goats. Geology of the area includes gneiss and schist, a tertiary intrusion, and typical talus slopes and glaciated valleys. F. RELATED Colorado FEDERAL Federal LITERATURE PROJECT Aid Project FW26P. CITED Bartmann, R. M., G. C. White, L. N. Carpenter, and R. A. Garrott. 1987. Aerial mark-recapture estimates of confined mule deer in pinyon-juniper woodland. J. Wildl. Manage. 51:41-46. Bassow, S. 1985. Potential Rocky Mountain bighorn (Oreamnos americanus). competition between the Mt. Evans populations of sheep (Ovis canadensis) and mountain goats B.A. Thesis. Princeton Univ •• Princeton. NJ. 77pp. Caughley, G. 1970. Liberation, dispersal, and distribution of HIMALAYAN (Hemitragus jemlahicus) in New Zealand. N. Z. J. Sci. 13:220-239. Caughley, G. 1977. Analysis 234pp. of vertebrate populations. J. Wiley Clarke, C. M. H. 1971. Liberations and dispersal of red deer Island districts. N. Z. J. For. Sci. 1:194-207. thar and Sons, in northern NY. South Davidson, M. M. 1973. Characteristics, liberation, and dispersal of sika (Cervus nippon) in New Zealand. N. Z. J. For. Sci. 3:153-180. Denney, R. N. 1977. Status and management of mountain goats in Colorado. Pages 29-36 in W. Samuel and W. G. MacGregor, eds. Proc. First Int'l. Mtn. Goat Symp. 243pp. Eberhardt, Wildl. L. L. 1969. Population Manage. 33:28-39. estimates from recapture Lincoln, F. C. 1930. Calculating waterfowl abundances returns. U.S. Dep. Agric. Circ. 118. 4pp. frequencies. on the basis J. of banding Minta, S. and M. Mangel. 1989. A simple population estimate based on simulation for capture-recapture and capture-resight data. Ecol. 70:17381751. otis, D. L., K. P. Burnham, G. C. White, and D. R. Anderson. 1978. Statistical inference from capture data on closed animal populations. Wildl. Monogr. 62:1-135. OVerton, W. S., and D. E. Davis. 1969. Estimating the numbers of animals in wildlife populations. Pages 403-455 in R. H. Giles, ed., Wildlife Management Techniques, 3rd Ed. The Wildl. Soc., Washington, DC. 623pp. Petersen, C. G. J. 1896. The yearly immigration of plaice into the Limfjord from the German Sea. Rep. Dan. BioI. Stn. 1895 6:1-77. Rice, W. R., and J. D. Harder. to a mark-recapture census 41:197-206. 1977. Application of multiple aerial sampling of white-tailed deer. J. Wildl. Manage. �Seber, G. A. F. 1973. The estimation of animal parameters. Hafner Press, NY. 506pp • • 1982. The estimation of animal abundance -----ed. MacMillan Publishing Co., NY. 654pp. Strandgaard, H. 1967. Reliability population. J. Wildl. Manage. abundance and related and related of the Petersen 31:643-651. method parameters. tested 2nd on a roe deer Waser, PM, S R Creel, and J R Lucas 1994 Death and disappearance estimating mortality risks associated with philopatry and dispersal. Behav Ecol5(2):135-141. white, G. C., D. R. Anderson, K. P. Burnham, and D. L. otis. 1982. Capture-recapture and removal methods for sampling closed populations. Los Alamos National Laboratory. LA-8787-NERP. Los Alamos, NM. 235pp. , and R. A. Garrott. 1987. Analysis of biotelemetry -----Colo. State Univ., Unpubl. Draft. 225pp. data - a primer. Woolf, A. 1973. Population dynamics and remote censusing white-tailed deer herd. Ph.D. Thesis, Cornell Univ., of a large, captive Ithaca, NY. 168pp. �Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS State of Project REPORT Colorado No. W-153-R-8 Mammals Research Work Plan No. SA Black Job No. 2 Development of black Inventory Technigyes Period Author: Covered: Thomas July Bear Research bear 1, 1994 - June 30, 1995 D. I. Beck Personnel: T. Beck, R. Hays, S. Lechman, M. McLain, J. Olterman, L. Willmarth, CDOW; R. Stevens, Colo. State Patrol; T. Holland, USFS; D. Bowden, G. White, Colo. State Univ. ABSTRACT· A black bear (Ursus americanus) resighting system utilizing cameras activated by active infra-red sensors was developed for a 465 km2 study area. Fortyfive cameras were distributed in 45 10.4 km2 quadrants. Six resighting sessions of approximately 14 days were conducted. Five-hundred forty-four photographs of black bears were obtained. Loss of unique ear-tags was a major impediment to population estimation. Of 119 photos of collared black bears, only 30 had ear-tags; representing 14 different black bears. An estimate of 168 black bears, yearling and older, was obtained using a Lincoln-Peterson estimator. The 95% C.I. was 134-204 (mean ± 20%). Black bear density was estimated to be 36 bears/100 km2• Problems with the camera system and modifications used are discussed. Limited efforts to call black bears in with predator calls during september were mostly unsuccessful. ��237 DEVELOPMENT OF BLACK BEAR INVENTORY Thomas TECHNIQUES D. I. Beck P.N. OBJECTIVE 1. Evaluate a capture-sight program utilizing for estimating black bear density. 2. Document age and gender hunting seasons. 3. Obtain density estimates of black bears in 3 heavily markedly different vegetation communities. bias cameras in vulnerability SEGMENT of black Evaluate the use of infra-red triggered cameras bears on a 465 km2 study area on the Uncompahgre 2. Evaluate 3. Evaluate predator Colorado. techniques calling to estimate as a method METHODS bears hunted stations during areas autumn of OBJECTIVES 1. 2 mark-resight set on bait to resight Plateau. black marked black bear density. for observing/hunting black bears in AND MATERIALS Study Area Description The Uncompahgre Plateau study area was located on the northern end of the plateau, in parts of GMU' 61 and 62. The 465 km2 area was divided into 45 quadrants, each 10.4 km2• The area is roughly 22 km north-south and 25 km east-west. East and west boundaries coincide with vegetative changes, the north boundary is an escarpment, while the south boundary is nontopographical. The area was not subjectively chosen as the best black bear habitat on the Plateau, but rather as a representative block of varied habitats. The vegetation composition was determined with the use of USFS timber-type maps. Using a dot grid, 50 points were recorded for each 10.4 km2 quadrant. Vegetation types recorded were: oak brush (Quercus gambelii), aspen (Populus tremuloides), ponderosa pine (Pinus ponderosa), Douglas fir (Pseudotsuga menziesii), pinyon-juniper (Pinus edulis - Juniperus spp.), non-forest (grasses-Artemisia spp.-Chrysothamnus spp.). Resighting Black Bears Resighting of black bears was accomplished by setting out 45 active-infrared triggered recorders and cameras. The recorder-camera system used was manufactured by Trailmaster (10614 Widmer, Lenexa, KS 66215) and consisted of the 1500 Recorder coupled with the TM 35-1 Camera Kit. The camera was an Olympus Infinity 35-mm point-and-shoot. ™ The recorder and camera were housed in a metal box (50mm ammo box; 28X18X15 cm). The box had a 6X9 cm hole cut out of the bottom for the camera field. Two 3-cm diameter holes were drilled for the infra-red beam and the beamalignment light. The recorder was mounted with 2 machine screws while the camera was secured with a single large-thread tripod screw. Approximately 50 3-mm pilot holes were drilled in the box in areas where a bear would likely rest a paw on the box. Sharp-pointed drywall screws, 5 cm long, were drilled into these holes from the inside; thus creating a "porcupine" effect. The box • �was affixed to a tree by cam-lock to reduce slipping. straps with 2 saw-edged brackets on the box The infra-red transmitter was housed in a protective box made from 19-cm diameter plastic irrigation pipe. The pipe was split to make half-circle sections, each cut 20 cm long. Redwood lumber was used to construct a back plate and top. Again, drywall screws were mounted facing out to deter bear activity. Two U-bolts were mounted in the back plate for affixing the unit to 2 75-cm long steel stakes, which were driven into the ground. A 40-mm diameter hole was drilled in the pipe for the infra-red beam. Initially, burlap was glued to the plastic pipe so that a habanero pepper sauce could be sprayed over the unit. It was hoped this hot sauce would deter black bear inquisitiveness. The recorder unit was programmed so that the infra-red beam had to be broken for 0.25 sec to be recorded (value of -P = 5). All beam breaks were recorded by date and time of day. The data back on the camera was programmed to imprint day of month and time of day on each picture. Konica 400 print film with 36 exposures was used for all surveys. The camera was set in CONTINUOUS mode. Each time the recorder triggered the camera, the electrical circuits were complete for 2.5 seconds, resulting in 3 or 4 pictures being taken consecutively. The recorder was programmed to have a 10-minute delay between series of pictures. Once a series of pictures was taken, the recorder continued to count the beam breaks but would not activate the camera until 10 minutes had elapsed. Each series of 3 or 4 pictures resulting from a single beam break was considered to be ONE picture. Only 1 roll of film was used during each session, thus at some sites not all visits by black bears were recorded on film. A single frame was exposed after set-up to insure the system was properly configured and to provide a beginning date and time on the film roll. Fresh batteries ("C"-size) were installed in the recorders and infra-red transmitters prior to sessions 1, 3, and 5. Six resighting sessions were conducted during 1994 at approximately biweekly periods beginning 12 June and ending 31 August. Resighting was not attempted in September because a preponderance of collared bears left the study area during the fall mast period in 1993. Three different sites were used in each quadrant, with 2 consecutive resighting sessions occurring at each site. Attractant baits were varied between sessions, in the following order: rotting fish, rotting beaver, honeycomb and anise oil, rotting fish, cherries and peaches, and a synthetic lure called essence of shellfish. The baits were placed in burlap bags (30X51 cm), and most baits weighed less than 5 kg. The synthetic lure was mixed with lard and the bags soaked in liquid lard. The altering of baits was intended to reduce the impacts of attenuation. Curiosity, not hunger, was the key element in attracting black bears to a camera site. Within each quadrant, specific sites were selected based on availability of black bear foods, terrain, proximity to water, availability of trees for securing bait and cameras, and probability of livestock and/or human interference. It was believed that the resighting procedures would be most effective if we could locate the sites where black bears would be naturally traveling rather than relying on the baits to attract black bears from long distances. The camera set-up for sessions 1 and 2 consisted of the camera-recorder mounted on a tree (DBH > 15 cm) at a height of 2.5-3.5 m with the infra-red transmitter mounted on the ground at a distance of 5-9 m. The bait was suspended from a 30-mm diameter cable strung between trees with the bait height at least 3.5 m. The bait was aligned directly over the infra-red beam, at a distance of 3-6 m from the camera. The ideal situation was to have the bait 4 m from the camera; based on the assumption that the black bears would most likely be photographed directly under the bait. For sessions 3-6, camera height was reduced to 2 m and distance from camera to infra-red transmitter was reduced to an ideal of 4 m. During sessions 1-4 the Infinity camera was �239 set at the normal telephoto setting to prevent direct camera box as such (35mm) focal length but during sessions 5 and 6 the (70mm) was used. All cameras were mounted facing WNW-to-ENE sunlight from hitting the infra-red receiver unit in the direct sunlight hits will trigger the camera. At the end of each session all film was removed from cameras and developed as negatives. Negatives were examined on a light table with the aid of a 12X magnifier. A record of all exposures was kept for each quadrant. All bear exposures, unknowns, and a variety of other exposures were subsequently printed. Specific identification of marked black bears and the unknowns was made from the prints. In addition, a day and time record of all beam breaks was obtained from the Trailmaster recorder and matched with the photographs. Photographed bears were recorded as: unknown classification, tag number, collared but no ear tag, untagged, tagged but unable to read tag, cub. Subjective evaluations of untagged black bears were made in an attempt to categorize yearling bears, since these bears were not available to be tagged in 1993. Series of pictures separated by 10 minutes were considered as separate bears, even if it appeared to be the same bear. Other wildlife was recorded by species. Often no animal was observed in the photographs and these pictures were labeled as SITE. In most cases where photographs were taken during dark hours, the first of the 3 or 4 exposures·was dark (a result of the camera electronic configuration and consequent battery drain). If no animal was visible in the subsequent pictures the series was labeled as DARK. Each series of 3-4 pictures was considered one picture in the summaries. A master file was kept for each session which includes of operation, day and time of all beam breaks, summary negatives, and all prints. site description, date of photographs, all A total of 65 black bears were collared at the end of the 1993 field season. Preliminary radio-tracking indicated that a significant portion may temporarily reside outside the defined study area. Aerial radio-tracking was conducted every Tuesday (with 1 exception) during the months of June, July, August, and early September. Since work schedules began on Tuesdays, this meant tracking data was available for the 1st, 8th, and 15th days of a camera session. Black bears were recorded as either IN or OUT of the study area. If a black bear was located in the area during any session, it was considered to have been available for resighting during that session. Not all bears were located each flight and subjective decisions as to IN or OUT were made based on locations during previous and subsequent flights, prior knowledge of movements, and age and sex class. These decisions were recorded uniquely to distinguish them from definite locations. Density Estimation The population estimation techniques initially chosen for evaluation require unique identification of each marked animal. Unfortunately, the ear tag/streamer combinations used during this study were pulled out by the bears at a relatively high rate. Thus a version of the Lincoln-Peterson estimator for sampling with replacement was used (Seber 1982:61). The estimator is N = n1 (n2+1) m2+1 where n1 is the number of marked bears in the area during the resighting session, nz is the total number of bears seen on the resighting occasion, and mz is the number of marked bears seen on the resighting occasion. Estimates were derived for each resighting session and the final estimate is the average of the 6 sessions. �242 ups were done with the camera-to-bait distance as near to 4 m as possible (4.3 m avg). However, nearly all animals were photographed at the infra-red transmitter housing staked in the ground (avg. distance = 6.1 m). This resulted in pictures where tag status of the bear was difficult to determine. Therefore, we changed protocol so that the camera-to-housing distance was approximately 4 m (avg. = 4.0 m for sessions 3-6). Photographic scale was much improved with the shortening of this distance. Another change was in the lowering of the camera unit. When the camera unit was high enough that a bear had to climb to investigate it we found that nearly all such investigations led to the camera unit being disturbed and knocked out of alignment. By lowering the camera unit so that such investigations could be done by a bear standing up, fewer cameras were disabled. We lowered the cameras beginning with session 2. Thus our idealized protocol became a set-up with camera height of 2 m, camera-totransmitter housing distance of 4 m, camera-to-bait distance of 3-4 m, and bait height of 3.5 m. The use of habanero sauce sprayed on the ground unit was discontinued. Rather than inhibit animal interference, it seemed to encourage both black bears and red squirrels (Tamiasciuris hudsonicus) to prolong their investigations of the unit. A number of film rolls were exposed because of these animals pulling the burlap loose and breaking the infra-red beam or by bears pulling the entire unit from the ground. Nearly all such interference with the ground unit ceased upon cessation of the hot sauce use. Our initial protocol was set-up with approximately 14-day camera sessions. This was based mostly on work schedules and a reasoned guess as to efficacy of the small baits. Actual days of operation varied from 10 to 18; with median values of 12, 15, 12, 15, 12, and 13 respectively for sessions 1-6. New sites were established for sessions 1, 3, and 5 whereas only film was changed for sessions 2, 4, and 6; thus explaining the shorter periods of operation for the 3 odd sessions. Of 526 pictures of black bears for which we have the date of photo, 51.3 % were taken in the first 5 days of a session, 30.4% in the second five days, 16.0% in the third 5 days, and 2.3% after day 15. The most noticeable declines in activity occurred after Days 6 and 12. Therefore, a 14-day session appears to be optimal. It is of interest to note that 88.5% of 522 black bear photos for which we have the time of day were taken during daylight hours. Bear activity appeared to be fairly uniform throughout the day with a definite peak period of the 3 hours at dusk. However, this pattern may be biased because of the high number of DARK category pictures. If a high proportion of these photographs was caused by black bears then the proportion of nocturnal activity was underestimated. If all of the DARK pictures had been black bear caused then the distribution of bear activity would have been uniform throughout the dark and light hours except for the peak at dusk. The time interval between sequential photographs of black bears was calculated to evaluate the camera time-delay setting. Of 515 sequential photo pairs, 465 (90%) were separated by >60 minutes. Only 36 (7%) were within 30 minutes of each other, while 14 (3%) were between 30 and 60 minutes. Of the 36 pairs between 10-30 minutes, 30 appeared to be the same bear in both photos, 3 definitely involved different bears, and 3 were unclear. Thus a 10-minute delay appears acceptable and possibly a 30-minute delay would be better. The 10-minute delay will be maintained as the protocol during the Middle Park study area in order to evaluate changes in a different habitat type. The rate of unknown bear classifications was greater during sessions 5 and 6. This was primarily because of the switch to the telephoto setting on the camera. By the end of the 4th session, it was apparent that the high proportion of missing ear tag/streamers would prohibit the use of the preferred statistical estimators. It was decided to try the telephoto setting to see if close-up photos of the head of bears could be consistently obtained. �243 If so, then possibly ear tags without streamers could be successfully used to identify black bears. The major failing of the telephoto setting was that a high proportion (31-35%) of photos were not usable because only portions of the bear were visible, and usually not the head. Blurring and field of view problems also occurred. The most significant problem encountered was the loss of ear tag/streamers. Only a quarter of the collared bears photographed still had ear tags. No eartagged bears without collars were photographed even though 24 black bears had been tagged but not collared in 1993. The ear tag/streamer combinations were similar to what has been used in other states (Montana, Wyoming, New Mexico; R. Mace, R. Grogan, C. Hunt, pers. comm.) but tag loss does not seem to be a problem elsewhere. The collars were readily observed in all the photographs of collared bears. Therefore, a marking system based on the collar rather than an ear tag will be developed prior to the Middle Park portion of the study. Density Estimation Use of a Peterson-Lincoln estimator provided a population estimate of 168 bears, with the 95% Confidence Interval being 134-204 (Table 2). The density of black bears for the 465 km2 study area was estimated to be 36/100 km2 (93/100 mi2). Table 2. Lincoln-Peterson estimates Plateau, 465 km2 study area, 1994. Session 1 2 3 4 5 6 Mean SD of black bear population, UNMARK MKSEEN MARKS 65 80 26 75 42 31 17 38 15 29 14 16 52 51 52 56 43 42 Uncompahgre POPEST 239.8 155.6 136.5 196.0 163.4 118.6 168.3 43.7 With the number of resightings made, had the bears been identifiable uniquely, the preferred estimators (Minta-Mangel and Bowden revision) would likely have produced a much more precise estimate (G. White, pers. comm.). Initial simulations using the Minta-Mangel estimator suggested that an initial capture rate of 0.5 and a resighting rate of 0.3 would be optimal for a precise estimate. Had the tagging system worked, would we have attained these rates in an actual field situation? We captured 89 black bears, age yearling or older, in 1993. We estimate the study area population to be 168 black bears. This estimate includes a number of yearlings, which were cubs in 1993 and thus not marked. Our subjective ratings suggest that at least 26% of the unmarked photographed bears were yearlings. This is likely conservative based on the difficulties encountered in sessions 5 and 6 (32% based only on sessions 1-4). A reasonable estimate for the 1994 study area population would be 89 tagged bears, 59 untagged 2+yr bears, and 20 untagged yearlings. Thus our capture rate in 1993 would have been 0.6. Similar calculations which allowed for 9% of the captured bears not inhabiting the area in 1994 still resulted in a capture rate of 0.6. Calculating a resighting rate is much more speculative because of the ear tag loss. The 30 photos of tagged bears were of 14 different bears. If that �244 proportion was true for the 89 photos of collared, untagged bears then another 41 bears may have been photographed. Thus we could have resighted 55 of the 65 collared bears, of which only 59 were located in the study area during 1994. It appears that developing a reliable marking system is the primary impediment to this technique providing precise estimates of black bear density. A system of affixing colored dowels (2X10 cm) to the top of the collars will be attempted in Middle Park. A 2-dowel system using 8 colors will allow for unique marking of sufficient numbers of bears. Predator Calling to Attract Bears Over 30 attempts were made to call in black bears, with only 1 bear observed to respond favorably. The evaluation was curtailed in favor of trying to observe bears without calling in an effort to determine which bears had lost ear tags. The one favorable response was by a subadult female which came within 12 m of the caller. Once she observed the caller she moved off without vocalizing. Oddly, she was not the target bear of the calling set. The female that was targeted did not respond to the call in any noticeable way. The .'. most common response recorded was no response ·apparent but in some- cases the target bear left the area rapidly. The principal investigator did call in an adult female with cubs while hunting in a different area in September. This female walked out on a canyon rim to investigate the sound. Her interest was maintained throughout 5 minutes by sporadic squalling. She kept looking down into a canyon rather than across to the source of the call. Approximately 5 minutes after first observing the adult female, 2 cubs appeared. They did not respond at all to the call but wandered aimlessly even while their mother searched for the source of the call. Calling will be attempted again in a much different habitat during september 1995. It will also be evaluated during May-July 1996 as an aid to observing bears in a non-hunting situation. LITERATURE CITED 1982. The estimation of animal abundance Seber, G.A.F. parameters. MacMillan, New York. 654 pp. and related �245 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS state of Project REPORT Colorado No. W-153-R-4 Mammals Research Work Plan No. 9A Elk Investigations Job No. 3 Spatial Period Author: Covered: July K. R. Wilson, Analysis of Elk Survival 1, 1994 - June 30, 1995 'D. A. Werle, and N. T. Hobbs ABSTRACT We developed a landscape simulator to evaluate model and logistic regression to detect annual in relation to habitat use. a proportional hazard differences in animal rate survival ��247 SPATIAL ANALYSIS P. N. OF ELK SURVIVAL OBJECTIVES The objective of this project is to evaluate methods for the statistical analysis of effects of habitat use on survival rates of mammals. We will determine the feasibility of detecting differences in population processes attributable to variations in landscape patterns. This will be accomplished by evaluating various survival analysis models using spatial simulation modeling. SEGMENT OBJECTIVE Evaluate a proportional hazard rate model and logistic regression to detect annual differences in animal survival in relation to habitat use. INTRODUCTION Biotelemetry techniques, such as triangulation, are common in wildlife research and management. Radio telemetry studies attempt to determine the habitat components important to wildlife. Common aims are to determine whether a species uses habitats available randomly, to rank habitats in order of relative use, to compare use by different animal groups, etc. (Kenward 1993). These type of studies attempt to show a preference or an avoidance for particular habitat types by comparing a habitat's availability to its use (Alldredge and Ratti 1986; Kenward 1987; Pietz and Tester 1983). Springer (1979), Hupp and Ratti (1983), and Lee et ale (1985) all discuss biases and sampling error problems with this technique. Telemetry locations are not perfect (Ables 1969). Tracking using radio telemetry often involves triangulation which only provides an estimate of an animal's location. A telemetry location, therefore, is not an exact point, but lies somewhere within an area delimited by triangulation error (Heezen and Tester 1967; Springer 1979). This error can result in locations which encompass more than one habitat, and the ratio of telemetry error to habitat sizes affects the efficiency of testing for habitat selection (Nams 1989). Telemetry error can be estimated, but Pietz and Tester (1983) and Kenward (1987) suggest that telemetry location should be disregarded when error areas, or error polygons, cover more than one habitat type. At an extreme, when error areas are large, observed habitat use can be biased toward random use of habitats. For example, when towers have been used in telemetry, White and Garrott (1986) have shown that the power ofax2 goodness of fit test to determine random versus nonrandom use is related to tower locations, the location of the animals relative to the towers and the precision to the bearings. Power of preference tests also decreases with increasing habitat complexity, decreasing sample sizes, and decreasing precision of bearings. Porter and Church (1987) recognize an additional limitation with the observed vs expected approaches. Almost all methods compare habitat use with a measure of habitat availability. The entire study area, chosen by the researcher, and not the animal, may not be available due to the presence of other animals, or the animal's use of an area may be the outcome of choices at different levels (Senft et ale 1987). Johnson (1980) found that analytical procedures were sensitive to a priori decision concerning the distribution of habitat types. Researcher decision on what is available, therefore, can severely affect results in habitat use studies. Second, these approaches do not lend themselves to examine habitat characteristics that may be most important to habitat preference analysis, such as juxtaposition and interspersion. (Porter and Church 1987). . �248 So, there are many criticisms of the use of use/availability data to infer biological relationships between animals and their habitats (Van Horne 1983; Hobbs and Hanley 1991). The goal of science is to learn from our past in order to find better ways to study the present. What can be done to better study the animal-habitat relationships that will not be sacrificed by the limitations mentioned above? At the least, as concluded by White and Garrott (1986), ••a habitat experiment using triangulation should be simulated on a computer ••••• Fortunately, newer technologies and more powerful computers allow for quicker, more in-depth, and more accurate studies of animal-habitat relationships. Geographic informational systems (GIS) are computer systems capable of assembling, storing, manipulating, and displaying geographically reference information, ie. data identified according to their locations. GIS can be used for scientific investigations, resource management, and planning. Global positioning systems (GPS) are space-based radio positioning systems that can provide 24 hour three-dimensional positions anywhere on Earth. The costs of these systems are relatively low, which may allow animals to be continuously tracked if needed. By combining GPS and GIS, management and scientific investigations of natural resources can utilize very accurate and precise tools. Selective availability of GPS signals by the U.S. military now limits locations of animals to within a few meters (still far better than telemetry), but the technology, for example using GPS base stations at known locations) is there to acquire GPS locations within millimeters. In fact, a Canadian telemetry company has recently marketed and used a GPS collar on large ungulates (Rodgers and Anson 1994). Both GPS and GIS can create large volumes of data. The current computer technology of personal and UNIX based computer systems allow these data sets to be manipulated and returned much faster than earlier computers. Landscape ecology has developed in response to our desire to understand the structure, function, and change inherent in nature (Forman and Godron 1986). Habitat patch characteristics and their effect on population size and survival have been studied in white-footed mice, Peromycus leucopus, (Fahrig and Merriam 1985), birds of the California oak woodlands (Block et al. 1994), Bachman's sparrow, Aimophila aestivalis, (Pulliam et al. 1992), and Kirtland's warbler's, Dendroica kirtlandii (Probst and Weinrich 1993). Structures of habitat fragments were found to interact with autecologies of these species, and influenced patterns of species richness. Several theoretical studies have demonstrated the importance of spatial heterogeneity for overall population persistence (Lomnicki 1980; Hastings 1982) and abundance (Taylor and Taylor 1977; Hanski 1982, 1985). Spatial population structures and the dynamics of species are greatly affected by the physical structure of the environment (Hanski 1994). These studies have shown that spatial heterogeneity is important to understanding the dynamics of individuals. Our objective, therefore, is to delve into the relationship of how survival may vary as a function of landscape heterogeneity. We will attempt to detect variations in animal survival as variations in landscape characters (habitat, slope, elevation, distance to water, etc.) occur. Our assumption follows that variations in the use of this space will affect population processes (e.g. survival) • GPS can continuously track animals through a landscape. Getting this information is relatively inexpensive, but too much information can be counter-productive, due to the time and budget constraints. We will also determine what type of sampling schemes are appropriate. We will weigh the advantages of the amount of information available through GPS and GIS with what is needed to discover the relationships between animals and their habitat. This will be accomplished with computer simulation. Computer simulations offer complete knowledge of systems. They eliminate uncertainty created by the fact that "natural" systems are never truly known. They also allow complex systems to be modeled and tested under a variety of scenarios with relative speed and ease. Our simulations will consist of two �249 parts. The first is a landscape simulator. This program will allow user input to create landscapes. By creating landscapes on a computer, all information pertaining to the landscape is known perfectly. Secondly, animal movements will be simulated through the known landscapes. A complete record can be kept to determine how the simulation animal's population processes change as its environment (changes in landscape characteristics such as slope, aspect, habitat type, distance to water, etc.) changes. RESULTS Landscape Simulator The program was created on an IBM-compatible personal computer using the c++ programming language. It is a DOS based program which dynamically allocates and deal locates memory, and creates output files for future references. The landscape simulator is an interactive tool used to create landscapes. The user is prompted for all input regarding the desired landscapes. Input includes: 1) the dimensions of the grid, 2) the number of habitat types (up to five), 3) the number of habitat centers for each habitat type, 4) the spread associated with each center, 5) the annual survival associated with each habitat type, 6) the overall percentage of cells for each habitat type, and 7) critical elevation points. The user is also prompted for the type of habitat clumping desired. The choices include random allocation, a semiclumped and a clumped landscape. These data will be implemented in creating a landscape grid. The landscape will have different features associated with each grid cell. Each habitat type mayor may not provide vital resources to the animal. The assumption being that different habitats provide different resources at different levels. Therefore, each habitat type has a survival associated with it. A vertebrate, such as an elk, Cervus elaphus, would find better forage in an aspen grove than it would in desert-like habitat, and survival would thus be higher in cells of aspen rather than in cells of desert. Characters such as slope and aspect may also be important to foraging vertebrates. Areas may be too steep to support vegetation, or inaccessible. Aspect is important in areas of snowpack. South-facing slopes tend to melt earlier and offer forage to vertebrates before north facing slopes. The habitat type, distance to water, the slope, and the aspect could all affect animal survival. The landscape simulator will include the capability for inclusion of characters according to user input. These landscape will then be used as a grid to track animal movements. Clumped Landscape: The data for each grid cell are determined one by one, beginning in the upper left hand corner of the entire area. The function: Gi.f21t is used, where the dependent variable, Yixr' is the probability of grid cell (x,y) being included with habitat center I-l ••n, where n is the number of habitat centers. Here, distj is the straight-line distance of the grid cell from the habitat center I, and OJ is the spread associated with the center. This results in an array of probabilities (l ••n). These are then normalized. The largest probability determines with which habitat center, and hence which habitat type, the cell is included. This type of habitat determination leads to the clumped landscape. Random Habitat: Random allocation of grid cells to habitat types uses a psuedo-random number generator with a uniform (0,1) distribution. This routine returns a random number which is used to randomly choose the habitat �250 type of the grid cell. The routine also checks to ensure that the number cells desired by the user for a particular habitat type is not exceeded. of Output: The landscape simulator creates two output files. GRID.OUT stores all information for each grid cellon a separate line. The x and y coordinate, the habitat type, the elevation, and the distance from water are all recorded. Other data that may be important, such as slope and aspect, will be included later. GRID.ASC is an output file that is created for input into program DISPLAY (C. Flather, U.S. Forest Service, Personal Communication). This program is a tool used to evaluate landscape scenes. spatial statistics, such as perimeter-area and a grid analysis are completed. The landscape is also visually displayed using VGA graphics. By utilizing both programs, the user can create landscape scenes to desired specifications, and know all information pertaining to those landscapes. Movement Models: The movement model begins by placing the animal at a random location on the grid created with the landscape simulator. The animal is given the choice of moving in one of eight directions, or remaining in its present location. The location of the animal ( x and y coordinate), the habitat type of the location, and the elevation of the cell, are recorded for each movement. An animal moves through its landscape, for example with one movement per day, and at each location, a (0,1) random number is generated to determine the survival of the animal. If the random number is greater than the current daily survival rate of the animal, based on the current cells survival attributes (i.e. based on survival probability of the current habitat type, elevation, etc.), the animal is assumed to have died, and the movement simulation is completed. This type of movement model is not biologically realistic, but serves as a null model to which future movement models can be compared. Future movement models will include the correlated walk, and the memory model. The correlated walk assumes that step I will be somehow related to step I-i. The direction of a step is chosen from a normal distribution of directions centered on the bearing of the previous step. Using this type of movement model will eliminate unrealistic back-and-forth movements. It will also force the animal to use more of the landscape grid. The memory model allows the animal to "search" the immediate area. A sight radius will be selected such that within that radius, habitats can be detected and determined. The circle of detectable habitat cells will then be divided into equal arcs. The arc containing the larger number of preferable habitats will be favored and be directed toward. This model could also allow animals to recall favorable locations. When an "appropriate" decision for the next direction of movement cannot be reached, the animal could recall the nearest favorable location that was previously visited, and direct itself toward it. By adding the memory model, more realism can be obtained. Preliminary Model Verification: This year's work has focused on creation of the simulation model. The landscape model is currently functional and includes the random movement model. Verification of the components is currently underway, as well as the programming of the additional movement routines. One of the goals of verification is to show that all parts of the model work, and uses the correct data at the correct time (Pegdon et al. 1990). Verification of the movement simulator consisted of trials using different habitat types. Fifty individuals were tracked through separate homogeneous landscapes. The habitats types had either 90%, 75% or 50% annual survival rates (i.e. each cell had the same survival 90%, 75%, or 50% for each trial). The random movement model would predict a 90% survival rate in a homogeneous landscape made of cells with 90% survival. A 75% survival rate, and a 50% survival rate would likewise be expected for the remaining trials. Each trial consisted of 50 simulation runs as follows. A homogeneous landscape with each cell having a 90% survival rate was created and an animal was randomly placed in the landscape. The animal moved through the landscape in a random walk �251 until death (see above) or 365 time steps (days) had occurred. This was completed 50 times (i.e. for 50 individuals), and at the end of the trial the number of animals surviving was tabulated. The process for the trials of the other survival rates was identical. The estimated survival rates for the movement simulations in the three landscapes were 90%, 78% and 48%, respectively (See Table 1). These results are reasonable considering the stochastic nature of the Monte Carlo simulations, and the small differences may be due to the low sample size (50). In addition, the GRID.ASC output files from several of the simulations were used as input to the DISPLAY program. Verification that these results are correct is currently underway. Future Work: Future work will include inclusion of the rema~n~ng movement models (the correlated walk and the memory model) and the completion of model verification. Finally, detailed simulations will be run to determine the feasibility of detecting changes in animal survival as a function of movement through heterogeneous landscapes under various sampling schemes. Table 1. Estimated average survival rates for animals using a random walk movement through three habitats types of a homogeneous landscape. Expected survival rates for habitat types 1, 2, and 3 were 0.90, 0,75, and 0.50, respectively. Observed survival results are based on 50 simUlation runs (repetitions). Habitat Type Observed Survival 1 0.90 Expected Survival 0.90 2 0.75 0.78 3 0.50 0.48 LITERATURE Ables, CITED E.D. 1969. Home-range Studies of Red Foxes (Vulpes vulpes). J. Mammal. 59:108-119. Alldredge, J. R. and J. T. Ratti. 1986. comparison of Some statistical Techniques for Analysis of Resource Selection. J. Wildl. Manage. 50(1):157-165. Block, W. M., M. L. Morrison, J. Verner, and P. N. Manley. 1994. Assessing Wildlife-Habitat-Relationships Models: A Case Study with California Oak Woodlands. Wildl. Soc. Bull. 22:549-561. Fahrig, L. and G. Merriam. 1985. Habitat Patch connectivity and Population Survival. Ecology 66(6):1762-1768. Forman, R.T.T., and M. Godron. 1986. Landscape ecology. John Wiley and Sons, New York. Hanski, I. 1982. Dynamics of Regional Distribution: the Core and Satellite Species Hypothesis. Oikos 210-221. Hanski, I. 1985. Single-species Spatial Dynamics May Contribute to Long-term Rarity and Commoness. Ecology 66:335-343. Hanski, I. 1994. Spatial Scale, Patchiness, and Population Dynamics on Land. Phil. Trans. R. Soc. Land. 343:19-25. Hastings, A. 1982. Dynamics of a Single Species in a Spatially Varying Environment The Stabilizing Role of High Dispersal Rates. J. of Mathematical Biology 16:49-55. �252 Heezen, K. L. and J. R. Tester. 1967. Evaluation of Radio-tracking by Triangulation with Special Reference to Deer Movement. J. Wildl. Manage. 31(1):124-141. Hobbs, N. T. and T. A. Hanley. 1990. Habitat Evaluation: Do Use/ Availability Data Reflect Carrying Capacity? J. Wildl. Manage. 54(4):515-522. Hupp, J. W. and J. T. Ratti. 1983. Evaluation of Radio-Tracking Triangulation Accuracy in Heterogeneous Environments. Proc. Int. wildl. Biotelem. Conf. 4:31-46. Johnson, D. H. 1980. The Comparison of Usage and Availability Measurements for Evaluating Resource Preference. Ecology 61(1):65-71. Kenward, R. E. 1987. Wildlife radio tagging: equipment, field techniques and data analysiAcademic Press, Orlando, Florida. Kenward, R. E. 1993. Compositional Analysis of Habitat Use from Animal RadioTracking Data. Ecology. 74(5):1313-1325. Lee, J. E., G. C. White, R. A. Garrott, R. M. Bartmann, and A. W. Alldredge. 1985. Accessing accuracy of a Radiotelemetry System for Estimating Animal Locations. J. Wildl. Manage. 49:658-663. Lomnicki, A. 1980. Regulation of Population Density due to Individual Differences and Patchy Environment. Oikos 35:185-193. Nams, V. o. 1989. Effects of Radiotelemetry Error of Sample Size and Bias when Testing for Habitat Selection. Can. J. Zool. 67:1631-1636. Pegden, C. D., R. E. Shannon and R. P. Sadowski. 1990. Introduction to Simulation Using SIMAN. Mc-Graw-Hill Inc. New York. Pietz, P. J. and J. R. Tester. 1983. Habitat Selection by Snowshoe Hares in North Central Minnesota. J. Wildl. Manage. 47(3):686-696. Porter W. F. and K. E. Church. 1987. Effects of Environmental Pattern on Habitat Preference Analysis. J. Wildl. Manage. 51(3):681-685. Probst, J. R. and J. Weinrich. 1993. Relating Kirtland's warbler Population to Changing Landscape Composition and Structure. Landscape Ecol. 8(4):257271. Pulliam, H. R., J. B. Dunning, and J. Liu. 1992. population Dynamics in Complex Landscapes: A Case Study. Ecol. Appl. 2(2):165-177. Rodgers, A. R. and P. Anson. 1994. News and applications of the global positioning system. GPS World (July): 20-32. Senft, R. L., M. B. Coughenour, D. W. Bailey, L. R. Rittenhouse, o. E. Sals, and D. M. Swift. 1987. Large Herbivore Foraging and Ecological Hierarchies. Bioscience 37:789-799. Springer, J. T. 1979. Some Sources of Bias and Sampling Error in Radio Triangulation. J. Wildl. Manage. 43(4):926-935. Taylor, L. R. and R. A. J. Taylor. 1977. Aggregation, Migration, and Population Mechanics. Nature 265:415-421. Van Horne, B. 1983. Density as a misleading indicator of habitat quality. Journal of Wildlife Management 47:893-901. White, G. C. and R. A. Garrott. 1986. Effects of Biotelemetry Triangulation Error of Detecting Habitat Selection. J. Wi.ldl. Manage. 50(3):509-513. �253 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS State of Project Colorado No. Work Plan No. Author: W-153-R-7 Mammals lOA Covered: July Research Kit Fox Studies 1 Job No. Period REPORT Kit Fox (Vulpes macrotis) in Colorado 1, 1994 to June Status 30, 1995 J. P. Fitzgerald Personnel: J. P. Fitzgerald, L. Dent, J. Eussen, M. Link, J. Prather, Reddy, D. Watson, R. Basagoitia, S. Boyle, R. Hays, T. Beck, B. Gill., Olterman M. J. ABSTRACT In FY1994-95, 2326 trap nights of effort resulted in 46 captures of 31 individual kit foxes. Eighteen were animals not previously captured. Since work began in March 1992, 6805 trap nights of effort have resulted in capture of 38 kit foxes: 10 adult males, 7 juvenile males, 14 adult females, and 6 juvenile females. Hundreds of square kilometers of what appears to be suitable habitat is unoccupied or occupied at low levels. Trapping was conducted in parts of Mesa, Garfield, Delta, Montrose, and Moffat counties. Captures were made in Mesa and Garfield Counties (10 kit fox), and in Montrose and Delta Counties (21 foxes). Most captures were in the Peach Valley and Montrose East areas. No captures were made in Brown's Park, Moffat County in 171 trap nights of effort. Attempts to locate foxes during the winter using ATV's were not successful due to lack of snow. Of 37 collared or ear tagged foxes caught since 1992, the fate of 23 is unknown, 7 are dead, 7 are alive. Movement of 3 foxes between Montrose East and Peach Valley was documented. Home ranges of Peach Valley and Montrose East foxes were estimated to be from 1.5 to 7.7 km2• Foxes used an average of 3.7 dens per individual during the tracking period. Reproductive succ~ss is low •.Of 14 adult females captured since 1992 only 6 litters of pups.have been produced. The Wildlife Commission exp~nded the area with trap restrictions to portions of Mesa, Garfield, and western Delta counties. Plans for the next fiscal year are to complete the search effort, finalize reports, and make recommendations for any continued research on the species. ��255 KIT FOX (VULPES MACROTIS) James STATUS IN COLORADO P. Fitzgerald P. N. Objective Document the geographic western Colorado. distribution and relative Segment 1. 2. 3. 4. continue Garfield continue Complete Complete abundance of kit fox in Objectives to monitor radio-collared foxes in Montrose, Delta, Mesa counties. search for new kit fox populations. write-up of earlier field study effort (M. Link Thesis). plans for 1995-96 field efforts. and Methods Field methods for live-trapping were similar to those reported previously (Fitzgerald and Link 1993, Fitzgerald and Verbeck 1993). Live-trapping effort was concentrated in the Gunnison and Colorado River drainages. Boyle radiotracked foxes in fall and winter in the Montrose East and Peach Valley areas to estimate home ranges and nightly movements. Boyle, Basagoitia, and Hays searched for fox tracks in fall and winter months using ATV's. Dent, Eussen and Hatch have continued the search effort and monitored radioed animals during this spring and early summer. Results LIVE TRAPPING and Discussion EFFORTS: A total of 2326 trap nights of effort resulted in 46 captures of 31 different individuals (Table 1). sixteen were animals captured for the first time. Table 1. Trapping effort and numbers of individual kit foxes captured county and area, 1 July 94 to 30 June 95, westcentral Colorado. County and Area Mesa/Garfield West and North of Grand Junction Mesa/Delta SE of Grand Junction to Delta Montrose/Delta Delta to SE of Montrose Moffat Browns Park-Vermillion Creek Totals Trap Nights by Captures 1125 8 528 2 502 21 171 0 2,326 31 Since May 1992 we have captured, marked, and released 38 individual kit foxes. Baits have usually been road-killed prairie dogs, cottontails, and ground squirrels supplemented with turkey chicks. A commercial attractor scent (chicken and fish, Rob Erickson's on Target A.D.C.) has been used in conjunction with the baits since mid-May ~995 in all traps. Since March 1992 trapping efficiency and capture rates have increased (Table 2) although much of the increase is probably the result of more concentrated effort in areas where foxes are known to be present. Seventy percent of • �256 Table 2. Trapping effectiveness and capture 1995, Kit Fox survey western Colorado. Trap Period # Individual Trap Nights Fox Captured rates, March Total Fox captures* 1992 to July TNE/Indiv Fox 1, TNE/AII Capt*. Mar 92-June 93 Peach Valley only 2725 836 9 9 9 9 303 93 303 93 July 93-June 94 1754 18 21 97 88 July 94-June 95 2326 31 46 75 50 6805 38** 75 179 91 Total *TNE = Trap nights of effort, all captures includes recaptures. ** Total of all individuals captured during the entire project. captures have been in the months of May, July, August and September, when recaptures that occur within 2 weeks of the last capture are excluded (trap happy animals were often caught on consecutive days) (Table 3). Table 3. Trapping success by age, sex and month of capture, 1992 - July 1, 1995, for foxes captured or recaptured when recaptures occurred at least 2 weeks post capture. No captures were made in october or November. Month Jan Feb Mar Apr 3 1 0 3 Male Adult Juv Female Adult Juv Unknown Tot May Jun Jul Aug Sep Dec 1 1 3 3 2 4 0 1 4 2 1 12 5 0 1 1 1 4 2 1 1 3 6 4 3 2 4 3 8 3 11 9 Total 17 6 1 23 10 1 57 Of 37 foxes (10 adult males, 7 juvenile males, 13 adult females, 6 juvenile females) captured and collared or ear tagged we have recaptured 14 of them (4 adult males, 2 juvenile males, 6 adult females, 2 juvenile females) at least once (range 1-6). Of 23 animals captured once,S were adult males, 6 adult females, 6 juvenile males, 5 juvenile females, and 1 sex unknown. Forty-three percent of our adult animals have been recaptured compared to 31% of the juveniles. In late spring-early summer 1995 we have not captured any foxes in almost 400 trap nights of effort including areas known to have foxes. This may reflect increasing trap wariness as well as a possible decline in populations. Different trapping methods, including selective use of padded jaw traps will be tried in some of the areas where we are finding sign of foxes but not making any catches. Mesa/Garfield (Wand N of Grand Junction): Eight kit foxes were captured during 732 trap nights of effort in July and August (Fig 1, Table 4). An additional 393 trap nights of effort in May and June 1995 have not resulted in any captures. These 8 fox are the first captured in 3 field seasons and 2059 total trap nights of effort since 1992. Seven of the 8 foxes were radio-collared, none have been located since April 1995. �257 Fig 1. Length of radio-contact and status of collared Garfield Counties, July 1 1994 - June 30 1995. Month 1994 SON and Year J A D J F kit foxes, 1995 M A M Mesa and J Location Rabbit Valley M (194/195) F (196/197) F (198/199) x-----------------------x--- unknown 151.244* 151.221 151.034 Prairie Canyon/Baxter U (153/154) 151.082 x- unknown Pass x- Corcoran Point F(J126/127) 150.834 M(130/131) 150.468 F(132/133) 150.638 unknown x---------------------- unknown x--------------------------x--------------------------- unknown unknown Cheney Res. M(J101/102) * Ear tags unknown xin ( ) followed unknown by radio-collar frequency Three of the 8 animals (1M,2F) were captured in Rabbit Valley close to the utah border. Neither female has been located since September 1994. The male was last located on 10 April 95. An animal of unknown sex was captured and radio-collared on 21 Aug in Prairie Canyon, Garfield county. It was accompanied by another fox which was not captured. The collared animal has not been located since its date of capture. Four foxes (2M, 2F), 2 of them juveniles (1M, 1F), were captured north of Grand Junction near Corcoran Point. Three of them were radio-collared. Two (M150.468, F150.638) were last located 9 April 95. The third has not been located since 12 Dec. Two other pups were observed but not captured at Corcoran Point, Basagoitia photographed kit foxes visiting guzzlers in this same area in the summer of 1994. Field crews this spring found fresh kit fox sign at Prairie Canyon and Corcoran Point but did not capture any foxes. The radio-collars on the 7 animals in this portion of the study region are close to their expected battery life and may no longer be working. Hundreds of square kilometers of what appears to be suitable range is unoccupied or inhabited by scattered individuals difficult to detect and capture. Mesa/Delta Counties (Grand Junction to Delta): Two animals, 1 juvenile male and 1 adult female were captured near Cheney Reservoir (near the Mesa-Delta County line) in 528 trap nights of effort. The female broke her jaw in the trap and died during rehabilitation efforts. The male has not been located since it was collared. A juvenile male (M150.980, ear tag 17) captured 22 Nov 93 by Verbeck in T14S/R95W/S33, northeast of Delta north of the Gunnison River has not been located since its capture. These 3 foxes are the only captures made in this section of the study area in 1597 trap nights of effort since 1992. Basagoitia reported on 5 Sept 94 seeing a live fox along Highway 50 south of Cheney Reservoir. On 3 Sept he observed 2 dead kit foxes on the highway in Delta County, 1, 2 km NW of Adobe Flats reservoir the other by the Alkali Flats road. Basagoitia photographed a kit fox at a guzzler in Wells Gulch during the summer of 1994. Basagoitia and Hays spent considerable time live-trapping and searching for fox tracks and sign using ATV's from Jan-April. They placed 4 road killed deer as baits but found no evidence of fox visiting the carcasses. Effort suggests few foxes �258 Table 4. Locations of radioed foxes in the lower Gunnison and Colorado Valleys, Mesa and Garfield Counties July 1, 1994 to July 1, 1995. Capture Site Date, and Date of Last Radio-Contact Sex Rabbit Valley 8/2/94 12/19/94 4/10/95 8/2/94 8/10/94 9/9/94 Cheney Res. 7/11/94 7/13/94 Radio Collar Frequency M 194/195 151.244 F F 196/197 198/199 151.221 151.034 Prairie Canyon/Baxter U 8/21/94 Corcoran Point 7/15/94 12/19/94 3/20/95 7/15/94 7/17/94 12/19/94 3/20/95 4/9/95 7/19/94 12/19/94 3/20/95 4/9/95 Ear Tag Number Location T10S T10S T10S T10S T10S T10S R104W R104W R104W R104W R104W R104W S20 S20 S17 12 13 13 S12 Pass 153/154 151.082 T7S R105W F(J) 126/127 150.834 M(J) M 128/129 130-131 F 132-133 T10S T10S T10S T10S T10S T10S T10S T10S T10S T10S T10S T10S M(J) F(A) 101-102 Jaw Broken not radio-collared 150.468 150.638 - Died 150.851 in Rehab R100W R100W R100W R100W R100W R100W R100W R100W R100W R100W R100W R100W T13S R98W T13S R98W 35 35 35 35 35 35 35 35 35 35 35 35 30 30? live in this expanse of the Gunnison River basin with many kilometers suitable habitat unoccupied or occupied at very low densities. Peach Valley and Montrose River of East Populations: We have spent 2240 trap nights of effort in the region from Delta south of the Gunnison River to south and east of Montrose since the study began in 1992. This is the center of our only population of foxes and the site of most of our radio-tracking effort. Twenty-seven of the 38 individual kit foxes have been taken from this area. During the 94-95 reporting period, in 452 trap nights of effort, a total of 8 new foxes were captured. Four were captured in Peach Valley: adult female (F151.083), adult male (M151.257) and 2 female pups too small to radio-collar (they were collared with pink nylon collars). At Montrose East, 3 adult (MI51.160, M151.107, M151.042) and 1 juvenile (M151.131) male were captured. In Peach Valley 11 adult kit foxes (SF, 6M) have been captured and radioed since 31 May 1992 (Fig 2, Table 5). Five of those animals, 3 females and 2 males are dead. Two were killed by coyotes, the cause of death of the others is unknown. The status of 4 animals is unknown. Two animals, male 151.095 and female 151.083) are still alive. Two radio-collared males (M151.209, M151.056) have moved from Montrose East to Peach Valley. These 4 animals are the only radio-collared animals known to be alive in Peach Valley. �259 Fig 2. Length of radio-contact with collared kit foxes, Peach Valley, 1995. * = Animals that have moved to Peach Valley from Montrose East. 1992 M Animal FO.238 MO.309 FO.873 FO.889 F1.009 M1.095 MO.037 M1.177 MO.499 F1.083 M1.257 M1.209* M1.056* J S N 1993 1994 J F M A M J J A SON J M M J S N D 1992- 1995 JFMAMJ X-------Dead X-Unknown x-----------------------------------------------------Alive X----------------------------------------------Dead X-------------------------------------------------Dead X--------------------------------------------------A1ive X----------------------Dead X--------------------------------Unknown X--Unknown X------------Unknown X--------Dead X--------------------Alive X--------------------Alive Table 5. Kit fox radio-collared in Peach Valley, 1992 - July 1, 1995, and dates of last sightings/radio-signals. New radio-collar frequencies are shown as they are replaced. ETR = ear tag replaced. capture or Date Located Sex 5/31/92 11/23/92 F 9/28/92 M 2 2.7 9/28/92 4/6/94 4/14/95 6/28/95 F 5 2.5 9/28/92 3/2/93 9/22/93 1/6/94 3/15/95 F 2/23/93 12/6/93 9/5/94 2/28/95 4/14/95 Ear Tag Number Weight Kg 1 2.0 6 2.4 Radio Collar Frequency 150.238 7 ETR18 2/23/93 12/6/93 9/9/94 1/25/95 6/28/95 M 4/21/93 9/24/93 3/26/94 M 8 2.5 2.6 2.0 3.0 3.1 ETR147/148 12 ETR15 Dead 150.309 T50N/R9W/S9 Unknown 150.338 150.956 150.873 T50N/R9W/S9 T50N/R9W/S9 T51N,R9W/S32 T51N,R9W/S32 Alive T50N/R9W/S9 T50N/R9W/S9 T50N/R9W/S9 T50N/R9W/S17 T49N/R8W/S? Dead T51N/R9W/S29 T50N/R9W/S5 T50N/R9W/S10 T50N/R9W/S5 T51N,R9W,S29 Dead T51N/R9W/S29 T50N/R9W/S5 T50N/R9W/S10 T51N/R9W/S29 T51N/R9W/S29 Alive T50N/R9W/S22 T50N/R9W/S8 T50N/R9W/S8 Dead 150.379 150.850 150.889 150.638 150.594 151.009 150.468 150.189 151.095 2.5 2.8 Fate T51N/R9W/S29 T15S/R94W/S27 (moved from PV to SE of Montrose) F Location 150.708 150.037 (C) (C) �260 Table 5 continued. 9/20/93 6/19/94 1/27/95 M 9/30/93 11/22/93 M 7/14/94 9/7/94 12/19/94 1/27/95 F 7/18/94 9/9/94 11/18/94 M(J) 1/27/95 13 2.7 ETR46/47 16 151.177 2.3 1.9 192/193 1.6 M 30/151 (First captured on 8/26/94 M 26/27 (First captured on 8/24/94 6/20/95 * Dead 150.499 151.083 190/191 4/18/95 6/29/95 4/13/95 150.813 (C) = killed 151. 257 T50N/R9W/S16 T51N/R9W/S29 T51N/R9W/S29 Unknown T50N/R9W/SI7 T50N/R9W/SI7 Unknown T51N/R9W/S31 T51N/R9W/S31 T51N/R9W/S29,30 T51N/R9W/S29 Unknown T50N/R9W/S10 T50N/R9W/SI0 T50N/R9W/S21 Dead 151.209 T51N/R9W/S29 in Montrose East as MI51.107) T51N/R9W/S32 T51N/R9W/S32 Alive 151.056 T50N/R9W/S9 in Montrose East as MI51.131) T50N/R9W/S9 Alive by coyote Adult female (F150.873) was captured by Dent in mid-April. He reported she was lactating but that an injury to her right rear leg was causing her to limp. She is still alive but not believed to have any pups. Dent recovered the remains of female 151.009 in Peach Valley on 14 April and judged she had been killed by a coyote. Peach Valley female (F150.889) first captured 28 Sept 1992 was recovered dead on 15 Mar 1995, 19 km south of her capture area and south of the Montrose East population. The area her carcass was recovered in had been trapped by field crews in the summer of 1994 with no fresh fox sign reported. Trapping by Hays in the area the carcass was recovered in did not yield any other foxes. The male foxes alive in Peach Valley include 2 animals (M151.209, M151.056) that moved from the Montrose East site to Peach Valley after early January 1995. Basagoitia and Hays on 27 Jan captured a male with only 1 ear tag (30) and no radio-collar they recollared the animal (M151.209). This animal was radio-collared (M151.107) and ear tagged 30/29 on 26 Aug in Montrose East by Boyle. Between 10 Jan and 27 Jan he moved from Montrose East to Peach Valley and lost his collar. Dent recovered collar 151.107 on 18 April close to the Water Tank in Peach Valley not far from the area Hays and Basagoitia recaptured and collared the fox as MI51.209. Male 151.056 moved to Peach Valley from Montrose East between 10 Jan and 13 April when Dent recaptured him in Peach Valley. He replaced radio-collar 151.131 with a new unit (M151.056). Boyle first captured this fox in Montrose East on 25 Aug. The third male in Peach Valley, M151.095 is an animal that has stayed in the Valley since first captured on 23 Feb 1993. In 1994, the Montrose East population included at least 13 animals (4 adult females, 3 adult males, 3 female pups, 3 male pups) clustered in a 5km2 area (Fig 3, Table 6). As of 28 June 1995, 5 of 11 animals collared in 1994 are alive. Three, F151.288, F151.037, M151.237, are at the Montrose East site. Two males (M151.056, M151.209) have moved to Peach Valley. Two are dead, the fate of 4 is unknown. Olterman has made several flights over the study area �261 trying to locate radio-collared foxes. He last flew on 27 June and will making other flights in the next few weeks to attempt to locate missing animals. Brown's Park, Moffat be County: In the project year we spent 171 trap nights of effort in Brown's Park with no captures. Clait Braun of the Colorado Division of Wildlife reported seeing a kit or swift fox near Vermillion Creek in July 1994. Dent in late August spotlighted a small fox in that area but could not make an identification. These observations along with reports in 1993 indicate kit or swift foxes live in Moffat County. Three summers of trap effort with a total of 616 trap nights have not resulted in any captures. Figure 3. Length of radio contact and fate of radio-collared Montrose East population, May 1994 to July 1, 1995. Month and Year and Area M Animal ID F151.288 F151.146 F151.037 F150.037 F150.940 F151.019 (.020) M151.237 F151.056 M151.160 M151.209 M151. 042 X---------------------------------------Alive X------------------------------Unknown X-------------------------Unknown X------------------------------Unknown X------Dead X---------------Unknown (Collar recovered) X-----------------------------------Alive X----------------------------Alive X--------------Unknown X----------------------------Alive X-----Dead J J 1994 A SON D J F 1995 M A kit foxes, M Table 6. Kit foxes trapped in the Montrose East population NL = non-lactating females; L = lactating. on July 1 1995. Capture or Date Located Sex 5/29/94 8/26/94 12/22/94 1/10/95 4/17/95 6/27/95 F(NL) 5/29/94 8/25/94 1/10/95 F(NL) 5/29/94 8/28/94 9/15/94 F(NL) 5/29/94 M pup Ear Tag Number Weight Kg Radio Collar Frequency 2.5 2.4 150.947 2.8 151.288 150.403 ETR22/38 2.5 2.4 2.3 23 2.3 150.338 21/32 F(NL) 22/28 (recaptured 24 on 7/7/95 1. 75 151.146 - now F151.037) not-collared J in PV in PV in 1994 and status Location Fate T49N/R8W/S7 T49N/R8W/S7 T49N/R8W/S6,7 T49N/R8W/S6 T49N/R8W/S7 Alive T49N/R8W/S7 T49N/R8W/S7 T49N/R8W/S7 T49N/R8W/S7 Unknown T49N/R8W/S7 T49N/R8W/S7 T49N/R9W/SI2 Alive T49N/R9W/S7 Unknown �262 Table 6 continued. 5/30/94 8/25/94 1/10/95 F pup 176/177 5/30/94 F pup 178/179 1.4 not-collared T49N/R8W/S7 Unknown 5/31/94 6/7/94 7/28/94 F (L) 2.3 150.940 T49N/R8W/S7 T49N/R8W/S7 T49N/R8W/S9 Dead T49N/R8W/S7 T49N/R8W/S7 T49N/R8W/S8 T49N/R8W/S8 Unknown T49N/R8W/S7 T49N/R8W/S7 T49N/R8W/S7 T49N/R8W/S7 T49N/R8W/S6 Alive T49N/R8W/S7 T49N/R8W/S7 T49N/R8W/S7 T50N/R9W/S9 T50N/R9W/S9 Alive T49N/R8W/S7 T49N/R8W/S6,7 T49N/R8W/12 Unknown 5/3/94 8/28/94 9/16/94 11/11/94 180/181 F pup 182/183 Collar recovered 6/3/94 8/25/94 1/10/95 4/18/95 6/27/95 M Ad 8/25/94 12/22/94 1/10/95 4/13/95 6/27/95 M Pup 26/27 not-collared 150.037 not-collared 151.019(.020) no trace of carcass 3.0 2.5 not-collared 150.708 151. 237 Moved 8/26/94 12/22/94 1/10/95 1/27/95 4/18/95 M Ad Moved 2.0 to Peach Valley M Pup Behaviors 1.8 2.2 184/185 8/27/94 12/29/94 1/4/95 8/27/94 11/9/94 11/22/94 1.5 2.3 33/34 29/30 1.5 2.4 to Peach Valley M Ad of foxes 48/49 in Montrose 2.3 151.131 151.056 151.160 151.107 151.209 151.042 T49N/R8W/S7 T49N/R8W/S6 T49N/R9W/S7 Unknown T49N/R9W/S7 T49N/R8,9W/S7,12 T49N/R9W/S12,7 T51N/R9W/S29 T51N/R9W/S32 Alive T49N/R9W/S7 T49N/R8W/S8 T49N/R8W/S18 Dead(C) East and Peach Valley: A report on kit fox activity patterns is appended to this report (Boyle 1995). Boyle estimated home range size for 7 kit foxes in the Montrose East population and 2 animals in Peach Valley (Table 7). Animals at Montrose East averaged 3.6km2 (range 1.5-5.9), those in Peach Valley 7.5-7.7km2• Home range sizes may reflect habitat and prey abundance differences between the 2 sites. Three animals have moved from Montrose East to Peach Valley (M151.209, M151.056) or out of Peach Valley (F150.889). This is the first evidence of foxes moving between the 2 areas. Boyle (1995) reported fox tracks north of Flattop Mesa between Peach Valley and Montrose East and did not believe they were made by collared animals. scattered individuals may inhabit the area between these 2 sites but to date we have not captured any and several landowners have refused permission to cross their properties. Most radio-collared animals have not moved far from their original capture locations (Table 5, 6). The eight foxes which we have had radio-collared for the greatest length of time (Table 8) show little movement from their original �2~ Table 7. Estimated home range size for nine kit foxes, Peach Valley and Montrose East populations, October-January 1994-1995. From Boyle (1995). Animal No. Montrose 150.037 150.403 150.947 150.708 151.131 151.160 151.107 Age/Sex Home Range ~ # of Locations East Peach Valley 151.009 151.083 sites of capture. predation. Juv Ad Ad Ad Juv Juv Juv F F F M M M M 5.9 4.7 1.5 4.7 2.3 2.5 3.9 56 52 53 50 36 38 36 Ad Ad F F 7.7 7.5 22 18 Fidelity towards home ranges may reduce the probability for Home ranges of animals in the Montrose East study area overlapped and the individuals using different dens varied over the field season (Boyle 1995). Boyle reported 9 individual foxes used an average of 3.7 dens (range 2-6) from late September to mid-January. A den complex adjacent to the main road through the study area was continuously used by 2-4 foxes from late November to January 10. Previous work by Verbeck (Fitzgerald and Verbeck 1993) and Link (1995) agree with Boyle's observation of frequent den changes with a tendency to favor some dens more than others. Boyle (1995) estimated total minimum distance travelled and straight-line maximum distances traveled during single night movements of 5 individual foxes in the Montrose East group. He estimated total minimum distance to average 6.2 km and maximum straight line distance to average 2.0 km. Movements of the observer in tracking foxes may have caused them to increase their movements more than might occur if undisturbed. Link, Beck and Dent (personal communications) reported kit foxes being radio-tracked on foot were aware of their presence and moved rapidly away from them. In the narrow Montrose East Valley it would be hard to avoid harassing the foxes when tracking their movements. Boyle's effort confirms our working assumption that foxes do not travel more than a few km's in their nightly forays and traps have to be set close together and left out for several days. Reproductive Success: In December Boyle reported transitory paLrLng of F150.947 and M 151.131, followed by F150.947 then denning with F150.403. Dent in April could not locate any paired animals but did trap a lactating female. Field crews working in late spring and summer have not located any dens with pups. Since May 1992, we have captured 14 adult females at times of the year when reproductive status could be determined. Four have shown evidence of lactation or contained uterine scars (F150.238). In Montrose East in spring and summer 1994, 4 of 5 adult females were non-lactating. This field season the only female (150.873) in Peach Valley does not appear to have pups. In Montrose East female 151.288 did not show signs of pregnancy or lactation in mid April. Female 150.338 (captured on 7 July 1995) shows no sign of bearing pups. Since 1992 we know of only 6 litters of pups, 3 in Peach Valley each with 2 pups, 2 in Montrose East with 7-8 pups using a single den, and 1 at Corcoran Point with 4 pups. The low numbers of adult females that are reproducing may be due to a number of different causes including: lack of �Table 8. Estimated size of range of 8 kit foxes radio-collared months, Peach Valley and Montrose East populations. Fox # Months Peach Valley F150.873 F151.009 F150.889 M151.095 M151.177 Montrose F151.288 F151.037 Ml51.237 Radioed Estimated 33 25+ 29+ 28 16 Range for at least (km2) 7 8 8 8 7 East 13 7+ 12+ 3 5 4 males, lack of adequate food, disturbance from field crews radio-tracking them, or, in the case of the Montrose East animals high population density a limited spacial area. other 7 in Activity: Link has completed her M.A. Thesis detailing the first 2 years of the project. Copies are enclosed with this report. The Wildlife Commission acted in September 1994 to expand the area with trapping restrictions to protect kit fox. Field plans have been completed for the 95-96 year: 1. We will continue to radio-collar animals captured in Peach Valley, Montrose East or at Corcoran Point but will not radio individuals taken elsewhere. All captured animals will be ear-tagged. 2. We will conclude our extensive trapping efforts with work to concentrate on the 64 km area along the base of the Book Cliffs from Corcoran Point to the Utah border and the 38 km long area at the base of Grand Mesa from Cheney Reservoir to Wells Gulch. 3. We will spend additional time in Brown's Park trying to resolve if kit or swift foxes live there. 4. From September 1995 through June 1996 we will use a monthly aerial search as the primary method for monitoring radioed animals in the Gunnison and Colorado river drainages. 5. Depending on weather we will conduct a limited amount of snow-tracking during the winter of 95-96. 6. By April, 1996 we will complete reports and recommendations for continued research on kit fox in western Colorado. By: James P. Fitzgerald Contractor, University of Northern Colorado. �265 Colorado Division Wildlife Research July 1995 of Wildlife Report JOB PROGRESS state of Project Colorado No. 1 COvered: July N. T. Hobbs, Research Multi-species llA Job No. Author: Mammals W-153-R-4 Work Plan No. Period REPORT Investigations Predicting the Impacts of Environmental Change: Simulations of Genetic and Species Diversity at Landscape and Regional Scales 1, 1994 - June 30, 1995 J. Miller, and J. A. Wiens ABSTRACT Loss of intact, natural habitat is the foremost threat to wildlife diversity in Colorado and the West. Historically, the prevailing source of habitat loss in the Rocky Mountain region has been harvest of natural resources (e.g., logging, mining, and agriculture). However, changing demographic and economic trends will drive a fundamental shift in the source of environmental change affecting wildlife habitat. As a result of these trends, residential development is likely to become the predominant human influence on the diversity of Colorado's wildlife during the coming decade and beyond. It follows that protecting wildlife habitat will depend in a pivotal way on developing wise policy on land use in the places where people live. To foster such policy, we propose to develop a System for Conservation Planning (hereafter, SCoP). The initiating idea of SCoP is that the success of habitat protection will depend on offering wise alternatives for land use, alternatives that meet human needs for economic vitality as well as the needs of wildlife for natural landscapes. Providing information needed to develop these alternatives is the primary goal of the SCoP project: The goal of SCoP is ~o obtain, assemble, and dis~ribu~e sta~e of ~e ~ informa~ion on effec~s of land use on wildlife diversi~y, p~icularly land use associa~ed wi.i;h.resid_e.n~~ale~pifi!.Ilsion in· Colorado and ~e Jies~. Meeting this goal requires' that'we enhance access to cur;rent 'kno~ledge needed to support decisions on land use while we simul.taneously strive to improve that knowledge. To that end, the SCoP has intitated pilot efforts in three Colorado counties, Larimer, Summit, and Boulder. In Larimer and Summit Counties, we are working with local citizens and planners to design . information systems that support land use decisions with regard to preserving and protecting wilidfe habitat. In Boulder County, we have initiated studies of effects of residential development on avian communities in riparian z.ones. ��267 PREDICTING THE IMPACTS OF ENVIRONMENTAL CHANGE: SIMULATIONS OF GENETIC AND SPECIES DIVERSITY AT LANDSCAPE AND REGIONAL SCALES P. N. Objective 1. Develop analytical tools to support decisions on management of wildlife diversity in Colorado. These tools will include simulation models and research results that predict consequences of changes in land cover types and land use for maintaining wildlife diversity. Segment Objectives 1. Develop a vertebrate population simulator land use change on population persistance. for predicting the effects of 2. Prepare grant proposals to the National Science Foundation and the Nature Conservance to support research on the effects of residential development on vertebrate species richness and community composition. 3. Choose Planing. areas for pilot projects of initiating a System for Conservation Results One of the overarching goals of the Colorado Division of Wildlife Long Range Plan is to "Lead statewide efforts involving federal, state, county and local governments, private landowners and other organizations to define and identify high-priority habitats in the state." A System for Conservation Planning (hereafter, SCoP, pronounced "scope") is an experimental project that is helping set priorities for habitat protection. . SCoP was motivated by the realization that rapid growth of the human population will be the most important source of change in wildlife habitat Colorado during the coming decade and beyond. In the face of such change, Colorado counties and municipalities need ways to identify particularly significant wildlife habitats. They also need tools to protect those habitats. in The mission of the SCoP project is to help local communities set biologically sound priorities for habitat conservation and to inform those communities of the economic and regulatory mechanisms available to achieve habitat conservation. To accomplish this nlission, the goal of the SCoP project is to obtain, assemble, and distribute state of the art information on effects of land use on wildlife diversity, particularly land use associated with rapid population growth. SCoP is working toward this goal in three parallel efforts: • Development of a collaborative process to help decision makers, planners, and citizens work together to set conservation priorities. • Production of accessible information systems that will help citizens and decision makers foresee large scale, cumulative effects of changes in land use on wildlife diversity. • Publication of a Wildlife Protection Handbook summarizing the legal and free-market mechanisms for restoring, preserving, and enhancing wildlife habitat in Colorado counties. Support for these efforts is provided by the Great Outdoors Colorado Trust Fund with matching support from the Colorado Division of Wildlife, Clarion �268 Associates, the American Planning Association, and the National Biological Service. Grant proposals for additional matching funds are currently under review by the National Science Foundation. The project is housed and staffed at the Natural Resource Ecology Laboratory of Colorado State University. Work in Summit County A pilot project was initiated in Summit County in January of this year. The first step in the project was to form a "Collaborative Design Team." Collaborative design is an approach borrowed from industry. It is based on the simple idea that the most successful products are designed in teams composed of clients with a wide range of needs and producers with a wide range of expertise. In Summit County the producers include wildlife ecologists (Tom Hobbs and Tanya Shenk, CDOW Research), a geographer (Bill Riebsame, CU), land use attorneys (Chris Duerksen, and Erin Johnson, Clarion Associates), and GIS analysts (Pam Schnurr, CDOW NW Region and Dave Theobald, CU). These experts work together to meet the needs of a group of representative clients in Summit County including Joe Sands and Marsha Osborn (County Commissioners), Brian Peters (Planner), Alex Chappell (DWM), and a diverse group of citizens including developers, landowners, and wildlife advocates. The collaborative approach used in Summit County has been exceeding successful. We began with a series of 1/2 day sessions to share knowledge of important issues in habitat conservation. These sessions reviewed material on land use planning and land use law, as well as basic principles of conservation biology, habitat modeling, and human geography. Subsequently, the Design Team worked together to set goals and objectives for development of an information system designed to support decisions on land use in Summit County relative to wildlife habitat. The Team decided that a particularly important goal was to develop an objective procedure that would categorize the landscape in Summit County in terms of its ability to support all species of wildlife. These categories could be used in master planning efforts county wide to guide zoning and/or habitat acquisition. In addition, it was decided that the information system produced by SCoP should allow a "coarse screening" of wildlife concerns that would likely occur during the development proposal and review process. Such a screening would be useful for developers who could use it to learn "up front" what wildlife issues would likely arise in getting a proposal approved. This screening would also be useful to citizen advocates who could use the information to assure that the proper concerns were considered during development review. Planners and Commissioners emphasized the importance of predicting and interpreting impacts to wildlife habitat that accumulate over time. The Team also asked that the system educate citizens about wildlife species, their habitat requirements and distribution, status, life history, and so on. Above all, the Team asked that the system be usable and accessible to citizens. The producer subgroup presented a design of the information system to the Design Team on May 9. Completion of an initial prototype of the information system is projected for late August. Work in Larimer County An additional pilot project will be initiated in Larimer County this summer. Tom Hobbs, SCoP project leader, has had several meetings with the Larimer county commissioners and planning staff. An outcome of these meetings is that SCoP will work in Larimer County to support their Partner in Land Use System, which is a master planning effort focussing on the Front Range corridor. The SCoP project in Larimer County will proceed as follows. In late May, we will form a Collaborative Design Team similar in composition to the one we used in Summit County. During the summer we will have 2 full day information sessions covering the Larimer County planning process, relevant aspects of �2~ land use law, and some basic concepts of human geography and conservation biology. In September, we will hold a goal setting session with the Design Team to give us input on how to usefully link an information system on wildlife habitat to the land-use decision process in Larimer County. In October, we will present a sketch of the system to the design team and to Partnership Land Use System symposium #3. Work in Boulder county Riparian areas have been widely recognized as key landscape features and centers of biological diversity (Naiman et ale 1993). In particular, riparian zones in the western United States have been described as critical sources of diversity (Thomas et ale 1979) with unusually high value for vertebrate faunas (Harris 1984, Johnson 1989, Terborgh 1989, Finch and Ruggiero 1993). Although western riparian zones at lower elevations comprise less than 1% of the total land area (Bottorff 1974, Knopf et ale 1988), up to 80% of terrestrial vertebrate species depend on them for at least part of their life cycles (Chaney et ale 1990). These zones provide habitat for more species of breeding birds than any other western plant community, and contain some of the most diverse avifaunas in North America (Johnson et ale 1977, Stevens et ale 1977, Stamp 1978, Knopf and Samson 1994, Ohmart 1994). In northeastern Colorado, 82% of breeding bird species occur in riparian vegetation (Knopf 1985). Intensive human use of riparian areas, however, may compromise their conservation value. Lowland riparian systems receive the heaviest human use of any of the vegetation communities in western North America (Thomas et ale 1979). Human activities that impact riparian areas, include water development, agriculture, domestic livestock grazing, timber harvest, and recreation. Because western riparian areas at lower elevations exist primarily as narrow, linear strips of vegetation along water courses (Knopf et ale 1988), they are also likely to be affected substantially by activities that occur in the surrounding landscape matrix. From prehistory to the present, successful human settlements in the arid North American west have been located along riparian systems (Johnson 1989, Ohmart 1994). As western states are currently experiencing unprecedented population growth (U.S. Bureau of the Census 1993), impacts on riparian areas related to residential development are likely to increase. Although increases in the human population are virtually certain, it remains unclear how these increases will affect bird communities in riparian zones. Traditionally, ecologists have avoided areas of human habitation when selecting research sites (Pickett et ale 1992) and, as a consequence, little is known about the ecology of populated areas (McDonnell and Pickett 1993). Based on the relatively few studies describing urban avifaunas, the general trend is a decrease in species richness concurrent with an overall increase in density (Batten 1972, Emlen 1974, Geis 1974, Aldrich and Coffin 1980, Beissinger and Osborne 1982, Bezzel 1985, DeGraaf and Wentworth 1986). Increases in density are often accounted for by a few species, primarily alien and pest species such as the European starling (Sturnus vulgaris), rock dove (Columbia livia), and house sparrow (Passer domesticus) (Geis 1974, Aldrich and Coffin 1980, Beissinger and Osborne 1982, Bezzel 1985, DeGraaf and Wentworth 1986). Increases in western populations of both native pests and exotic species associated with urbanization (Johnston and Garrett 1994, Marzluff et ale 1994) may also influence avian assemblages in riparian areas near residential development. Another potential impact of urbanization involves nest predation. Predation is commonly the primary cause of nest mortality (Ricklefs 1969, Best and Stauffer 1980, Martin 1988) and some evidence suggests that nest predation rates are higher near suburban development (Wilcove 1985). A common explanation is that because human-associated predators (e.g., domestic cats �270 and dogs, raccoons, skunks, and blue jays) attain higher densities in residential environments (Hoffman and Gottschang 1977, Churcher and Lawton 1987, Haspel and Calhoon 1989, Rosatte et al. 1991, Coleman and Temple 1993), elevated predation rates will result (Soule et al. 1988, Harris and SilvaLopez 1992, Noss 1993). The mere presence of certain predators, even in the absence of actual predation, may be sufficient to exclude certain sensitive species from otherwise suitable habitat (Engels and Sexton 1994). Although riparian systems are often the centerpiece of urban planning and residential development (Adams and Dove 1989, Binford and Buchenau 1993), few studies have addressed the indirect human influences on these areas. The first objective of this study is to examine changes in species composition and relative species densities for lowland riparian avifaunas across a gradient of human development from low to high residential densities (McDonnell and Pickett 1990, 1993). The second objective is to describe predator assemblages for lowland riparian areas across the same gradient and to assess relative predation rates among members of these assemblages. Methods Avian species richness and densities were quantified using point transects (Buckland et al. 1993) on each of 16 study sites. Study sites are located on South Boulder Creek, Coal Creek, Boulder Creek, and the Cache la Poudre river and have been chosen to represent a gradient from high-density to low-derisity residential areas. Each site was visited three times during the breeding season in 1995. Point counts were conducted using standard protocol (see Ralph et al. 1993) and individual observers rotated visits to a given site in order to minimize observer bias (Verner 1985). The same methods will be employed in 1996 and 1997. Avian species richness and composition are strongly correlated with vegetation structure in riparian areas (e.g., Stauffer and Best 1980). Thus, a variety of vegetation measurements were collected at each avian census point. At each avian census point, we assessed average tree height, diameter, number of layers, crown cover, shrub density and composition, ground cover, and snag numbers using a slightly modified version of the point-centered quarter method (Cottam and curtis 1956, also see Ralph et al. 1993). Features of the surrounding landscape mosaic will be quantified using aerial photographs and a geographic information system. Here, we will measure housing density and tree/shrub density in three concentric 1 km bands centered on each study site. Cumulative totals of the variables of interest will be derived. We will also measure distance to the nearest development and its residential density. Finally, average width of the woody riparian vegetation has been shown to strongly influence avian species (Stauffer and Best 1980) and will be assessed at each site. The ideal approach to studying nest predation is to locate active nests and determine the fates of nestlings during the breeding season. This is logistically difficult, however, when the goal is to compare a number of sites for many species. As an alternative, artificial nests have been used to describe patterns of nest predation across landscape gradients (e.g., Andren et al. 1985, Wilcove 1985, Ratti and Reese 1988). Two important criticisms have been made of this approach: 1) identification of predators is important to management, yet such identification if often made on the basis of signs at the nest - an often crude and imprecise method (Major 1991), and 2) the method is biased in that it attracts certain species of nest predators but not others (Angelstam 1986, Martin 1987, Major 1991). In 1995, we placed 16 transects at 12 of the study sites, using 20 artificial nests per transect. Nests are left in the field for 15 days, approximately the length of the egg-laying and incubation period for small passerine birds, �271 or until both eggs are gone, whichever comes first. During this time, they are checked every third day. Determination of predation rates based on analysis of this data should provide an indication of where to focus subsequent efforts to identify predators. Here, we will employ the use of automatic cameras in conjunction with artificial nests (Picman 1987, Major 1991), and trackbeds in conjunction with odor attractants (Roughton and Sweeney 1982, Shepard and Greaves 1984). We are unaware of any studies on predator assemblages using automatic cameras or artificial nests that have been conducted in riparian zones. Finally, we will compare predation rates based on artificial nests with rates of nest success based on real bird nests in 1996 and 1996. preliminary Results As of this writing, our avian censuses are complete for 1995, but the data have not been analyzed. Measurement of vegetation structure and composition has just begun. Some artificial nest transects are still being run and predation rates for the completed transects have not been determined. Initial inspections of the data, however, show some unexpected trends. Predation rates appear to be highest for sites that have the lowest surrounding residential density. Also, there is a tendency for predation rates on nests near recreational trails to be lower than rates in areas with no trails. While these results are intriguing, we urge caution in their interpretation. There are certain biases associated with the use of artificial nests (see "Methods") and rates obtained with artificial nests mayor may not correlate with predation on real nests. The use of artificial nests, however, provide an indication of where to concentrate future efforts regarding the monitoring of real nests, the use of cameras, scent posts, etc. When data based on artificial nests are combined with these other types of data, we expect to obtain reliable information on changes in predator assemblages and predation rates across a gradient of urbanization. Literature Adams, Cited L., and L. Dove. 1989. Wildlife reserves and corridors in the urban environment. National Institute for Urban Wildlife, Columbia, Maryland. Aldrich, J. W., and R. W. Coffin. 1980. Breeding bird populations from forest to suburbia after thirty-seven years. American Birds 34:3-7. Andren, H., Angelstam, P., Lindstrom, E., and P. Widen. 1985. Differences in predationpressure in relation to habitat fragmentation: an experiment. Oikos 45:273-277. Angelstam, P. 1986. Predation on ground-nesting birds' nests in relation to predator densities and habitat edge. Oikos 47:365-373. Batten, L. R. 1972. Breeding bird species diversity in relation to increasing urbanisation. Bird Study 19:157-166. Beissinger, S. R., and D. R. Osborne. 1982. Effects of urbanization on avian community organization. Condor 84:75-83. Best, L. B., and D. F. Stauffer. 1980. Factors affecting nesting success in riparian bird communities. Condor 82:149-158. Bezzel, E. 1985. Birdlife in intensively used rural and urban environments. ornis Fennica 62:90-95. Binford, M. W., and M. J. Buchenau. 1993. Riparian greenways and water resources. Pages 69-104 in D. S. Smith and P. C.Hellmund, editors. Ecology of Greenways. University of Minnesota Press, Minneapolis. Bottorff, R. L. 1974. Cottonwood habitat for birds in Colorado. American Birds 28:975-979. Buckland, S. T., Anderson, D. R., Burnham, K.P., and J. L. Laake. 1993. Distance Sampling -- Estimating Abundance of Biological Populations. Chapman & Hall, London. Chaney, E., Elmore, W., and W. S. Platts. 1990. Livestock Grazing on Western Riparian Areas. U.S. Environmental Protection Agency, Washington, D.C. �272 Churcher, P. B., and J. H. Lawton. 1987. Predation by domestic cats in an English village. Journal Zoological London 212:439-455. coleman, J. S., and S. A. Temple. 1993. Rural residents' free-ranging domestic cats: a survey. Wildlife Society Bulletin 21:381-390. cottam, G., and J. T. Curtis. 1956. the use of distance measures in phytosociological sampling. Ecology 37:451-460. DeGraaf, R. M., and J. M. Wentworth. 1986. Avian guild structure and habitat associations in suburban bird communities. Urban Ecology 9:399-412. Emlen, J. T. 1974. An urban bird community in Tucson, Arizona: derivation, structure, regulation. Condor 76:184-197. Engels, T. M., and C. W. Sexton. 1994. Negative correlation of blue jays and golden-cheeked warblers near an urbanizing area. Conservation Biology 8:286-290. Finch, D. M., and L. F. Ruggiero. 1993. Wildlife habitats and biological diversity in the Rocky Mountains and northern Great Plains. Natural Areas Journal 13:191-203. Geis, A. D. 1974. Effects of urbanization and type of urban development on bird populations. Pages 97-105 in J. H. Noyes and D. R. Progulski, editors. Wildlife in an Urbanizing Environment. November 27-29, 1973, Springfield, Massachusetts. University of Massachusetts, Amherst. Harris, L. D. 1984. The Fragmented Forest. University of Chicago Press. Harris, L., and G. Silva-Lopez. 1992. Forest fragmentation and the conservation of biological diversity. Pages 197-237 in P. K. Fielder and K. J. Subodh, editors. Conservation Biology. Chapman and Hall, New York. Haspel, C., and R. E. Calhoon. 1989. Home ranges of free-ranging cats (Felis catus) in Brooklyn, New York. Canadian Journal of Zoology 67:178-181. Hoffman, C. D., and J. L. Gottschang. 1977. Numbers, distribution, and movements of a raccoon population in a suburban residential community. Journal of Mammalogy 58:623-636. Johnson, A. S. 1989. The thin green line: riparian corridors and endangered species in Arizona and New Mexico. Pages 35-46 in G. Mackintosh, G.W. Press, Washington, editor. Preserving Communities and Corridors. D.C. Johnson, R. A., and D. W. Wichern. 1992. Applied Multivariate Analysis, 3rd Edition. Prentice Hall, New York • .Johnson, R. R., Haight, L. T., and J. M. Simpson. 1977. Endangered species vs. endangered habitats: a concept. Pages 68-79 in R. R. Johnson and D. A. Jones, Jr., editors. Importance, Preservation and Management of Riparian Habitat: a Symposium. General Technical Report RM-43. U.S. Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colorado. Johnston, R. F., and K. L. Garrett. 1994. Population trends of introduced birds in western North America. Studies in Avian Biology 15:221-231. Knopf, F. L. 1985. Significance of riparian vegetation to breeding birds across an altitudinal cline. Pages 105-111 in R. R. Johnson, C. D. Ziebell, D. R. Patton, P. F. Ffolliott, and R. H. Hamre (technical coordinators). Riparian Ecosystems and Their Management: Reconciling Conflicting Uses. First North American Riparian Conference, April 16-18, 1985, Tucson, Arizona. General Technical Report RM-120. U.S. Department of Agriculture, USFS, Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colorado. Knopf, F. L., Johnson, R. R., Rich, T., Samson, F. B., and R. Szaro. 1988. Conservation of riparian ecosystems in the United States. Wilson Bulletin 100:272-284. Knopf, F. L., and F. B. Samson. 1994. Biological diversity - science and action. Conservation Biology 8:909-911. Major, R. 1991. Identification of nest predators by photography, dummy eggs, and adhesive tape. Auk 108:190-195. Martin, T. 1987. Artificial nest experiments: effects of nests appearance and type of predator. Condor 89:925-928. Martin, T. E. 1988. Habitat and area effects on forest bird assemblages: is nest predation and influence? Ecology 69:74-84. �273 Marzluff, J. M., Boone, R. B., and G. W. Cox. 1994. Historical changes in populations and perceptions of native pest bird species in the west. Studies in Avian Biology 15:202-220. McDonnell, M. J., and S. T. A. Pickett. 1990. Ecosystem structure and function along urban-rural gradients: an unexploited opportunity for ecology. Ecology 71:1232-1237. McDonnell, M. J., and S. T. A. Pickett. 1993. Introduction: scope and need for an ecology of subtle human effects and populated areas. Pages 1-5 in M. J. McDonnell and S. T. A. Pickett, editors. Humans as Components of Ecosystems. Springer-Verlag, New York. Naiman, R. J., Decamps, H., and M. Pollock. 1993. The role of riparian corridors in maintaining regional biodiversity. Ecological Applications 3:209-212. Noss, R. F. 1993. Wildlife corridors. Pages 43-68 in D. S. Smith and P. C. Hellemund, editors. Ecology of Greenways. University of Minnesota Press, Minneapolis. Ohmart, R. D. 1994. The effects of human-induced changes on the avifauna of western riparian habitats. Studies in Avian Biology 15:273-285. Pickett, S. T. A., Parker, V. T., and P. L. Fiedler. 1992. The new paradigm in ecology: implications for conservation, biology above the species ·level. Pages 65-88 in P. L. Fiedler and J. K. Jain, editors. Conservation Biology. Chapman & Hall, New York. Picman, J. 1987. An inexpensive camera setup for the study of egg predation at artificial nests. Journal of Field Ornithology 58:372-382. Ralph, C. J., Geupel, G. R., Pyle, P., Martin, T. E, and D. F. DeSante. 1993. Handbook of Field Methods for Monitoring Landbirds. General Technical Report PSW-GTR-144. U.S. Department of Agriculture, USFS, Southwest Research Station, Albany, California. Ratti, J., and K. Reese. 1988. Preliminary test of the ecological trap hypothesis. Journal of Wildlife Management 52:484-491. Ricklefs, R. E. 1969. An analysis of nesting mortality in birds. Smithsonian Contributions to Zoology 9:1-48. Rosatte, R. C., Power, M. J., and C. D. MacInnes. 1991. Ecology of urban skunks, raccoons, and foxes in metropolitan Toronto. Pages 31-38 in L. W. Adams and D. L. Leedy, editors. Wildlife Conservation in Metropolitan Environments. Proceedings of a National Symposium on Urban Wildlife, 11-14 November, 1990, Cedar Rapids, Iowa. National Institute for Urban Wildlife, Columbia, Maryland. Roughton, R. D., and M. W. Sweeny. 1982. Refinements in scent-station methodology for assessing trends in carnivore populations. Journal of Wildlife Management 46:217-229. Shepard, D. S., and J. H. Greaves. 1984. A weather-resistant tracking board. Pages 112-113 in D. o. Clark, editor. Proceedings of the 11th Vertebrate Pest Conference, University of California, Davis. Soule, M. E., Bolger, D. T., Alberts, A. C., Wright, J., Sorice, M., and S. Hill. 1988. Reconstructed dynamics of rapid extinctions of chaparral-requiring birds in urban habitat islands. Conservation Biology 2:75-92. stamp, N. E. 1978. Breeding birds of riparian woodlands in southcentral Arizona. Condor 80:64-71. Stauffer, D. F., and L. B. Best. 1980. Habitat selection by birds of riparian communities: evaluating effects of habitat alterations. Journal of Wildlife Management 44:1-15. Stevens, L., Brown, B. T., Simpson, J. M., and R. R. Johnson. 1977. The importance of riparian habitat to migrating birds. Pages 156-164 in R. R. Johnson and D. A. Jones, Jr., editors. Importance, Preservation and Management of Riparian Habitat: a Symposium. General Technical Report RM-43. U.S. Forest Service, Rocky Mountain Forest and Range Experiment station, Fort Collins, Colorado. Terborgh, J. 1989. Where have all the birds gone? Princeton University Press, Princeton, New Jersey. ter Braak, C. J. F., and I. C. Prentice. 1988. A theory of gradient analysis. Advances in Ecological Research 18:271-317. �274 Thomas, J. W., Maser, C., and J. E. Rodiek. 1979. Riparian zones. Pages 40-47 in J. W. Thomas, editor. wildlife Habitats in Managed Forests: the Blue Mountains of Oregon and Washington. u.s. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station, Portland, Oregon. u.S. Bureau of the Census. 1993. Statistical Abstract of the United States. U. S. Government Printing Office, Washington, D.C. Verner, J. 1985. Assessment of counting techniques. Current Ornithology 2:247-302. Wilcove, D. S. 1985. Nest predation in forest tracts and the decline of migratory songbirds. Ecology 66:1211-1214. � https://cpw.cvlcollections.org/files/original/15ca5d39d91e499fd7a8d45b9d25e306.pdf 411314a77d1a20d1ee90a4eda4e44ec6 PDF Text Text Colorado Division Wildlife Research April 1995 of Wildlife Report JOB FINAL REPORT state of Colorado Project Work W-166-R-4 1 Plan Job Title: Period Covered: Author: Personnel: Wildlife. James Game Birds Investigations 20 : Job Development Colorado Migratory and Evaluation 01 April 1994 through of Moist-Soil 31 March Management Techniques in 1995 K. Ringelman James K. Ringelman and Michael R. Szymczak, Colorado Division of ABSTRACT High soil permeability, timing of water availability, amount of water available, and cost of water are potential limitations to moist-soil management in Colorado. Most soils near eastern plains riparian areas are sand or sandy-loam, which make it difficult to maintain water and moist-soil conditions. Ditch water, the most cost-effective source of water for impoundments, is generally unavailable until mid-May. Conventional moist-soil techniques would require that water remain impounded over winter or be applied in March or April in preparation for May drawdown. However, these and other limitations are common.throughout eastern Colorado, therefore any suitable moist-soil management techniques must work within these constraints if such management is to be widely adopted. . In 1989, technical and popular literature was reviewed for information on moist-soil management techniques applicable to Colorado. Wildlife researchers in Missouri and Texas responded to letters of inquiry and provided details of ongoing moist-soil studies in those states. Personnel in Kansas and adjacent states were consulted during a waterfowl habitat management workshop hosted by the State of Kansas. Progress on the "expert system" computer software for moist-soil management was reviewed with personnel of the u.S. Fish and Wildlife Service. Based on this information, eight criteria were developed for screening potenti'al sites ·to conetizuct; experimental moist-soil impoundments. Four candidate sites were considered, and field sq.rveys.were conducted at each site. During 1991, Wellington Wildlife Management Area, a 6,669 ha property owned and operated by the Colorado Division of Wildlife, was selected as the site for moist-soil experiments. A levee plow and water control structures were purchased in late winter, and the proposed site for impoundment construction was surveyed for grade by a professional engineer on 26 February 1991. Anticipated water depths in each impoundment were expected to range from 40 cm at the distal end of the impoundment to near 0 cm at the proximal end. Based on this engineering survey, 5 impoundments, each ~0.5 ha in area, were constructed in March and April using 0.5 m high levees. A filling and drawdown schedule was established on 28 May, but ditch water was unavailable until 1 June because of wet spring weather and resultant insufficient call for irrigation water. Ditch water was again unavailable in mid-June as early .._., i.<,': rlilll~ il~I~lii~~rl~~lrl~mlilml BDOW011083 �2 summer rains continued. After initially filling all impoundments on 1 June, it became apparent that the initial engineering of impoundment grades was in error, because water only accumulated for 3-6 m behind the distal levee at maximum capacity. consequently, impoundments were effectively unusable for experimental purposes and no experiments were performed. The impoundment site was re-examined for grade in fall and winter 1991, and a decision was made to abandon the study site because of the high cost of impoundment rehabilitation. The Moist Soil Advisor computer package was obtained from D.B. Hamilton of the National Ecology Research Center, u.s. Fish and Wildlife Service. Preliminary evaluations of the software indicate that this expert system is applicable to moist-soil management in Colorado. Meanwhile, the search for suitable field sites to verify model expectations and plant responses continued. A wetland on private property just south-east of Fossil Creek Reservoir was identified as a potential moist-soil unit. The landowner indicated a willingness to cooperate with water manipulations and follow-up evaluations at this site. The wetland was flooded in late April into May 1992, and a drawdown performed about 1 June 1992. However, uncontrollable high water during the 1992 growing season in the 4 ha pond resulted in a vigorous stand of threesquare bulrush (Scirpus americana), with minimal germination of smartweeds (Polygonum spp.), curlydock (Rumex crispus), wild millet (Echinochloa crusgalli), and other moist-soil plants. In late summer, filamentous green algae became common, due in part to water stagnation. In 1993, an attempt to control bulrush by maintaining water at a comparatively high level resulted in limited success. Some smartweed germinated during the high water period. No other suitable experimental impoundment sites have been found. Late availability of ditch water, an arid climate, highly permeable soils in many eastern plains sites, and short growing seasons in intermountain valleys all present obstacles to moist-soil management in Colorado. Although generally not applicable throughout Colorado, moist-soil management may have potential applications on selected sites. Further development of this technique will require close coordination with managers in an adaptive management context. Accordingly, this Federal Aid project will be concluded, but further progress on moist-soil management will be pursued and reported under the auspices of "Cooperative Management Programs" (Work Plan 10, Job 1). �3 Colorado Division Wildlife Research October 1995 of Wildlife Report State of Colorado Project W-166-R-4 Work Plan 1__: Job Title: Period Author: Job Harvest western Covered: Michael JOB PROGRESS REPORT Migratory Game Bird 22 distribution Colorado 01 April Investigations of mallards 1994 through and pintails 31 March banded preseason in 1995 R. Szymczak Personnel: J. Gamble and staff, Brown's Park National Wildlife Refuge; J. Grode, K. Holzer, U. S. Forest Service; J. Broderick, R. Caskey, J. Corey, D. Coven, P. Creeden, R. Del Piccolo, J. Ellenberger, V. Graham, J. Gray, J. Gumber, W. Hegberg, T. Mathieson, J. Miller, J. Olterman, R. Olterman, N. Smith, M. Szymczak, J. Todd and K. Wagner, S. Wait, and S. Yamashita, Colorado Division of Wildlife ABSTRACT Ducks were trapped in modified Salt Plains bait traps and banded on 16 different wetlands located in 5 areas across western Colorado in August and September, 1994. About 2,000 mallards (Anas platyrhnchos) were banded, with the number captured generally well distributed between trapping areas. Only 31 northern pintail (Anas acuta) were banded in 1994. ��5 HARVEST DISTRIBUTION OF MALLARDS AND PINTAILS BANDED PRESEASON IN WESTERN COLORADO P. N. OBJECTIVE 1. Document captured 2. Determine if the geographic location of recovery of mallards pintails is dependent on the area of banding in Colorado. 3. Determine the relationship between the recovery distribution of western Colorado banded mallards and pintails and the distribution of recovery of those species banded in other areas of the Pacific Flyway. 4. cooperate in analysis of Pacific preparation of reports. SEGMENT 1. 2. 3. the distribution of band recoveries preseason in western Colorado. Flyway-wide of mallards band and pintails recovery and data and OBJECTIVES Trap and band mallards and pintail in 4-5 areas of western Colorado in late August-early September using salt plains bait traps (Szymczak and Corey 1976). Recommended areas are: (1) Browns' Park National Wildlife Refuge, (2) Yampa River Valley below Craig, (3) Colorado River Valley below Glenwood springs, (5) Uncompahgre-lower Gunnison River Valley, and (6) the Cortez - Mancos area. Submit banding schedules and recapture reports to the U. S. Fish and Wildlife Services' Bird Banding Laboratory. Summarize and file band .return reports. Contribute Alberta. manpower and equipment to cooperative duck banding crews in INTRODUCTION In 1990, the Pacific Flyway Study Committee formulated a S-year cooperative mallard and pintail preseason banding program that was endorsed by the Pacific Flyway Council. This program was designed to address banding needs throughout the western U. S., including Alaska, and in the provinces of British Columbia and Alberta. Through the first 3 years, over 305,000 d~cks were banded under this program. This report covers the fourth year of banding during the preseason period in western Colorado. Background information can be found in Szymczak (1992). METHODS Trap Area Selection Most breeding mallards in western Colorado are associated with small wetlands that are widely distributed throughout high elevation, mountainous areas. Only in Browns' Park, located in the extreme NW corner of Colorado, are there extensive wetland complexes capable of supporting breeding ducks. Trapping on widely distributed wetlands with low densities of breeding ducks was assumed to be an inefficient method for banding western Colorado mallards. Therefore, strategically located areas, to which post-breeding and fledged Colorado mallards would move, were selected as primary trapping areas. In �6 1994, trapping occurred on the Browns' Park National Wildlife Refuge (BPNWR), along the Colorado River Valley from Rifle to near Fruita (CRV), in the Uncomphagre River Valley from Montrose to Delta including some locations east of Delta in the Gunnison River Valley (URV), in the Cortez-Mancos area (CM), and at Gardner Park and Allen Basin reservoirs, about 5 miles west and 7 miles Northwest, of Yampa, respectively. Trapping Period Ducks were trapped and banded in each area for a consecutive period of about 10 days beginning on 18 August and ending on 18 September. The actual banding periods in 1994 were: BPNWR - 18 thru 31 August; Yampa Area - 6 thru 20 September; CRV - 1 thru 9 September; URV - 27 August thru 5 September; CM 3 thru 12 September. Trapping and Recording Technique All birds were trapped in modified Salt Plains bait traps (Szymczak and Corey 1976) using whole shelled corn for bait. Traps were visited daily. Mallards and pintails were the target species, but green-winged teal (Anas carolinensis) were also banded in all areas, ring-necked ducks (Aythya collaris) were banded near Yampa, blue-winged teal (Anas discors) and/or cinnamon teal (Anas cyanoptera) were banded near Yampa and in Browns' Park, and gadwall (A. strepera) and American wigeon (Anas americana) were banded near Yampa. Banded birds were recorded by wetland site. Band numbers of all birds captured that were banded in previous years or outside the specific area of trapping were recorded. Records were also maintained in some areas on the number of traps operated by wetland in order to evaluate capture/unit of effort .. at each trap site. Alberta Banding One volunteer was recruited from the Colorado Division of Wildlife's permanent staff to work on a cooperative duck banding crew in southern Alberta. The crew was active in Alberta throughout the month of August. All banding records were submitted to the U. S. Fish and Wildlife Service's Bird Banding Laboratory by the Canadian Wildlife Service. Therefore the results of the Alberta trapping effort will not be reported here. RESULTS Trap Locations Trapping was distributed over a total of 16 different wetlands in the 5 areas (Table 1). Fravert Pond (CRV) and Sanders' Pond (URV) were added as trap sites in 1994 while Hog Lake (BPNWR), Skipper's Island (CRV), Hall Pond (URV), Mancos Wetland (eM), and Hooten's Pond (CM), were dropped. Nearly all trap· sites were.located in Palustrine Emergent Persistent Wetlands, but some sites in the Colorado River Valley were in Riverine Upper Perennial Rock Bottom wetlands (Cowardin et ale 1979). Allen Basin is a high mountain water storage reservoir that contained mostly dead storage water at the time of trapping. Banding and trapping efficiency Nearly 2,000 mallards were banded during trapping in western Colorado in 1994 (Table 2) bringing the 3-year total to slightly over 7,000 mallards. The number of birds banded increased in the Yampa area, decreased in Browns' Park and remained similar in the other 3 areas. Nearly all mallards banded on the BPNWR were adults. However, immatures mallards comprised 75 %, 76%, 68% and 59% of the CRV, URV, CM, and Yampa samples, respectively. Throughout the trapping area 64.8% of the mallards banded were immatures compared to 61.9% in 1993, 62.6% in 1992, and 68.3% in 1991. �7 Only 31 northern pintail, the secondary target species, were banded (Table 3). Addition species banded were green-winged teal, American wigeon, blue-winged teal and/or cinnamon teal , gadwall and ring-necked ducks (Table 3). Band Reporting and Record Keeping All new banded birds and recaptures were submitted to the U. S. Fish and Wildlife Service's Bird Banding Laboratory on standard forms. computer files contairiing the number of birds banded by area, site, day, age and sex were constructed at the Colorado Division of Wildlife's Research Center. LITERATURE CITED Szymczak, M. R. 1992. Harvest distribution of mallards and pintails banded preseason in western Colorado. Job Prog. Rep., Colo. Div. Wildl., Fed. Aid Wildl. Rest. Oct. Pp. 49-53. Szymczak, M.R., and J. F. Corey. 1976. Construction and use of the Salt Plains. duck trap in Colorado. Colo. Div. Wildl., Div. Rep. 6. 13pp. Prepared by: Michael R. Szymczak Researcher/Scientist IV �8 Table 1. Trapping August- September, locations during preseason banding in western Colorado, 1994', wetland Area Name Location Flynn Marsh spitzie Slough T10N, R103W, Sec 16,SEl< T10N, R103W, Sec 15, sl,z Gardner Park Res. Gardner Park Res. Allen Basin Res. T1N, R86W, Sec 22, NElo T2N, R86W, Sec 9, Nl,z Colorado R. Valley ,Latham's Slough Morse's Pond Fravert Pond T8S, R97W, Sec 27, swlo T1S, R1E, Sec 34, NElo ute Meridian T1N, R2W, Sec 36, swlo ute Meridian T6S, R93W, Sec 8, NWlo Uncompahgre R. Valley Markley's Pond SWeitzer Lake Sanders' Pond T50N, R9W, Sec 30, NWlo T15S, R95W, Sec 28, swlo T14S, R94W, Sec 34, NW~ Cortez/Mancos Totten Res. Weber Res. T36N, R15W, Sec 20, NWlo T36N, R13W, Sec ,12, NE~ summit Lake williamson's Pond Nolan's Pond T37N, R14W, Sec 33, swlo T36N, R15W, Sec 3,1SE~ T37N, R16W, Sec 8, N).; Browns' Park Natl Wildl. Ref. Walker Wildl Area N. Table 2. Number of Mallards banded by area, site, age and sex and trapping efficiency during pre-season trapping in western Colorado, 1994. Area Brown's Park Yampa Area Colorado River Uncomp. River Cortez-Mancos site AM Age/sex AF IM IF Flynn Marsh 81 spitzie Slough ~ SUb-total 106 64 o 2 26 1. 90 1 1. 3 Gardner Park Allen Basin Sub-total 11 18 29 12 18 30 24 29 53 N. Walker Fravert Pond .Morse's Pond, 'Latham Slough Sub-total 69 23 o o 26 __1_ 99 -2. 30 95 4 52 __2_! 205 Markley's Sweitzer Lake Sanders' Pond Sub-total 35 40 18 93 22 25 20 67 27 Nolan's Pond 47 Totten Res. 10 Weber Res. 10 summit Lake Williamson's Pd. __2. 99 SUb-total 16 28 GRAND TOTAL ff!/ Exclude Yampa area sltes. 426 4 6 23 Total 147 __2.J_ 200 No. trap days 41 16 57 3.6 3.4 4.8 15.0 5.5 14.0 9.4 12.7 ~ 32 70 74 144 83 270 18 7 11 112 113 2 3:0 ~ No. banded per trap/day 8 172 506 12 40 135 75 ~ .304 107 35 __2.Q 299 175 60 20 182 192 656 _dQ 110 21 121 21 11 16 88 80 284 50 42 20 10 10 5.0 4.2 -2. 3.3 13 19 2 _Q. __1 52 175 140 _.lQ 466 269 738 539 1972 __1_ 10 53 260M 5.0 8.8 6.1 6.0 8.0 14.2 8.8 7.6M �9 Table 3. Number of northern pintail and other species banded by area, site, age, and sex in western Colorado, preseason 1993. Number of additional birds banded as locals in parentheses. Species Northern pintail Area site Colo. R. Uncomp. River CortezMancos AgelSex IM IF AM AF Latham Slough 0 1 0 0 Markley's Pd. sweitzer L. Sanders' Pd. 1 0 8 0 0 2 1 2 3 0 0 5 0 1 Q 7 1 4 .1 _! 7 31 Totten Res. Nolan's Pd. Total 2 _2 .1 13 •• Total 1· 3 1 18 Green-winged teal Brown's Pk. Spitzie Sl. 0 2 0 0 2 Yampa Gardner Park 0 1 2 6 9 Colo. R. Walker Wildl. Latham Slough Morse'S Pd 2 0 1 0 0 0 1 1 1 1 1 2 4 2 4 Sanders' Pd. Sweitzer L Markley's Pd. 34 5 10 3 1 2 4 4 20 4 3 6 45 13 38 Totten Res. Nolan's Pd. 1 _Q 53 1 _Q 10 Uncomp. R. CortezMancos Total Blue-winged/cinn. Teal 4 _2. __ 9 6 37 32 132 Brown's Pk. Flynn Marsh 1 1 0 0 2 Yampa Gardner Pk. Q Q 1 .1 Q 0 .1 1 0 Q 0 0 2 (1) Q 0 .1 Total Ring-necked Duck 0 _! Yampa Gardner Park Allen Basin Total 1 3 3 (1) 6 (1) Q 6 (1) 10 1 11 Gadwall Yampa Gardner Pk. 0 0 0 0(1) 1 Am. Wigeon Yampa Allen Basin 0 0 2 0 2 ��11 Colorado Division Wildlife Research April 1995 of Wildlife Report JOB FINAL REPORT state Colorado of project Work 1 Plan Job. Title: Period Author: Migratory W-166-R-4 Job Ecology Covered: Robert Investigations 23 of Waterfowl 01 April Game Birds in Montane 1994 through Wetland 31 March Communities 1995 L. Sanders Personnel: James K. Ringelman, Andrew Selle, Michael R. Szymczak, and Joseph S. Todd, Colorado Division of Wildlife; Leigh H. Fredrickson and Robert L. Sanders, University of Missouri-Columbia. ABSTRACT Wetland habitat characteristics and waterfowl use of 24 montane wetlands were recorded on the Big Creek Lakes study area in 1993-94. Wetland physical and chemical characteristics, aquatic invertebrate biomass and densities, and waterfowl population characteristics and habitat use were recorded during the period May - September in both years. Physical and chemical features of the 24 intensively sampled wetlands were recorded monthly during both field seasons. Physical parameters included basin size, shoreline length, and vegetative habitat composition of each wetland as determined by aerial photographs. Chemical parameters measured in the field included pH, conductivity, dissolved oxygen, hardness, nitrates, orthophosphates, sulfates and apparent color. Additionally, water samples collected on 1-2 August 1994 were submitted to the University of Missouri limnology laboratory for analysis of total and dissolved nitrogen, total and dissolved phosphorous, algal chlorophyll and true color. A total of 115,125 aquatic invertebrates (222.865 g dry wt.) were collected in 1,896 sample sweeps during the two field seasons. Invertebrates were sorted into taxonomic order, counted, dried and weighed. Values were entered into a database file and are currently being analyzed to determine if relationships exist among wetland origins and habitat types, water chemistry parameters and/or waterfowl use of wetlands. Waterfowl observations were conducted twice weekly throughout both field seasons~ Waterfowl species, number of birds, breeding status, habitat use and .activity data were. recorded for each wetland. Data collected are currently being analyzed to determine if relationships exist between waterfowl use of wetlands and habitat parameters measured under objectives 1 and 2. A thesis is being prepared, which will serve as the basis for several scientific publications. Therefore, final reporting of research results will be detailed in subsequent annual reports under the "Migratory Game Bird Publications" Job, .(Work Plan 22, Job 2). ��13 Colorado Division Wildlife Research August 1995 of Wildlife Report JOB PROGRESS State of Colorado Project W-166-R 1 Work Plan Job Title: Period Author: : Job Integrated Covered: James Avian Research REPORT - Migratory Game Bird Investigations 24 Waterbird 1 April Management 1994 through Studies 31 March 1995 H. Gainmonley Personnel: J. Gammonley, J. Ringelman, M. Szymczak, Colorado Division of Wildlife; D. Cooper, R. Durfee, A. Polonsky, Colorado State University; M. Laubhan, National Biological Service. ABSTRACT A pilot study was conducted 1 May to 12 August 1994 to examine waterbird ecology at Russell Lakes State Wildlife Area (RLSWA). Aerial photographs were taken in May and used t·o produce a map of 6 dominant vegetation cover types. We collected food habits and body condition data for mallards, gadwalls, cinnamon teal, blue-winged teal, American avocets, killdeer, and Wilson's phalaropes, and time-activity data for these species and white-faced ibis, snowy egrets, black-crowned night herons, and spotted sandpipers. We also collected aquatic invertebrates in 4 cover types, to determine seasonal changes in the abundance and diversity of invertebrate foods used by waterbirds at RLSWA. Data analyses will continue during 1995-96. Results from 1994 were used to prepare a detailed study proposal for 1995-96. Collections for food habits and body condition data, time-activity data collections, line-transect counts, and invertebrate collections will continue. Nest searches and habitat measurements in each cover type will also be conducted. ��15 INTEGRATED WATERBIRD MANAGEMENT STUDIES ,I P. N. OBJECTIVES 1. Map the location of wetlands and wetland 2. Document the hydrologic regime and water, charact'eristics of each wetland type. 3. Identify the aquatic invertebrates associated with each community, and document seasonal trends in invertebrate abundance and biomass. 4. Quantify the abundance, spatial and temporal use patterns, behaviors, and food habits of waterbirds in different wetland types. Relate the dynamics of endogenous lipid and protein reserves to food habits and migration and breeding ecology. 5. Determine the seasonal wetland habitat requirements for all waterbirds, and consolidate these needs into a conceptual design for an optimum wetland commuriity. 6. Determine the water management protocol and wetland development guidelines needed to produce the optimum wetland community. Prepare wetland development and water management plan for the RLSWA. SEGMENT communities on the RLSWA. soil and vegetation wetland diversity, a OBJECTIVES 1. Map wetlands photographs, and wetland communities using a combination of aerial USGS Quadrangle maps, NWI maps, and ground-truthing. 2. stratify wetland communities according to type and randomly select sites within each community. Identify the species composition and measure temporal trends in the standing crop of invertebrates at each site. 3a. Determine the abundance and habitat use of all waterbird early spring through late summer by periodic censuses. 3b. Determine behaviors through time-activity sampling of Canada geese, mallards, gadwalls, cinnamon teal, blue-winged teal, black-crowned night herons, snowy egrets, white-faced ibis, American avocets, killdeer, Wilson's phalaropes, and spotted sandpipers. 3c. Document the food habits of representative shorebirds and waterfowl by collecting actively feeding birds and examining esophageal and proventricular contents. Species representative of six foraging guilds will be sampled: gadwall (aquatic herbivore), cinnamon/blue-winged teal (aquatic, straining carnivore), mallard (aquatic, straining omnivore), killdeer (terrestrial/aquatic gleaner), avocet (aquatic, gleaner/sweeper), and Wilson's phalarope (aquatic/pelagic gleaner). 4. Sample the diversity and spatial arrangement of wetland and upland habitats to relate landscape features to the diversity and abumdance waterbirds. species from 5. Research alternative optimization methods that might be applicable to wetlands management (e.g., stochastic dynamic programming, expert systems, or graphical interpretation). Using one or more techniques, determine the optimal wetland management scheme for RLSWA. 6. Prepare annual reports appropriate scientific and publish journals. the results of the study of in ." �16 INTRODUCTION The San Luis Valley (SLV) is one of the most important breeding areas for waterbirds in Colorado (Ryder et al. 1979, CDOW 1989, Nelson and Carter 1990, Gilbert et al. in press). A wetland ecosystem can be managed for habitats that maximize requirements for a narrow group of avian species, or for more diverse habitats that optimize resources for a variety of avian species. The latter approach, which embodies a philosophy of integrated waterbird management, better fits the philosophy of increased emphasis on managing landscapes for species diversity. One goal of the Colorado State,Waterfowl Management Plan is to provide habitat of sufficient quality to maintain duck and goose populations at desired levels for maximum recreational opportunities (CDOW 1989). In addition, the SLV draft management plan for waterbirds recommends maintenance of ,diverse wetland habitats with 25% of the actively managed habitat on public lands managed for nongame waterbirds (Olterman 1993). In 1994, CDOW initiated a pilot study to examine the resource use by both game and nongame waterbird species on Russell Lakes State Wildlife Area (RLSWA) • STUDY AREA We studied waterbird ecology at RLSWA, a 1,550 ha wetland complex located in the SLV in Saguache County. we categorized habitats (cover types) at RLSWA according to hydrology (flooding depth and duration) and vegetation structure as follows: short emergent (SE), tall emergent (TE), shallowly flooded open sites with no emergent vegetation (SO), semi-permanently flooded sites with no emergent vegetation (SP), saltgrass (SG), and upland shrub (US). We created a GIS map of RLSWA based on these cover types, using aerial photos (scale = 1:4,000) taken on 5 May 1994 (cover type designations pased on photos were ground-truthed during summer of 1994). METHODS Habitat Use Each week, we used 15 established line transects to census waterbirds on RLSWA. The species, sex (when possible), and flock size of all waterbirds flushed, as well as their perpendicular distance from the transect line and the cover type they flushed from, were recorded during each count. We used counts as a crude index of waterbird abundance each week. We also used linetransect analysis techniques (Buckland et al. 1993) to estimate mean densities of species with adequate sample sizes using program DISTANCE (Laake et al. 1994) • Collections We shot foraging waterbirds from 1 May to 3 August; most birds were collected during a 10-day period each month in May, June and July. We observed foraging birds for >20 min prior to collection, to ensure that _"individuals contained food items obtained at the collection site and to determine the pair status of collected birds. Immediately following collection, the esophagus of each bird was removed and stored in 95% ethanol. Birds were weighed and stored in a freezer for later processing. In the laboratory, we identified and sorted food items contained in the esophageal contents of each bird. Each food item was dried (GOoC) and measured to the nearest 0.1 mg. Collected waterbirds were thawed, dissected, and categorized according to sex, age (plumage characteristics and/or the presence of a bursa), and reproductive status (prelaying, laying, incubating, post-breeding). Proximate analysis was used to determine the percent nutrient composition (lipid, protein, and mineral) of each plucked carcass and the reproductive tissues of laying females. . Time-activity Budgets We used focal sampling (Altmann 1974, Tacha et al. 1985) to determine the diurnal activities of selected waterbird species in each cover type. During each 10-minute focal session, observers recorded each time the subject �17 changed its activity (resting, preening, aggression) or cover type. Invertebrate locomotion, foraging, alert, or Collections Aquatic invertebrates were collected from 25 sites in SE, DO, SO, and SG habitats at approximately 2 week intervals from May through August. At each site, 3 replicate samples were collected in the water column, vegetation, and benthos. Invertebrates in each sample were identified to the lowest taxonomic. status and counted. RESULTS waterbird Community Characteristics A total of 31 waterbird species was recorded during weekly line transect censuses. Nine additional species were observed during the field season, but not counted during line transect censuses. Crude estimates of relative abundance indicate that cinnamon teal, mallards, gadwall, redheads, whitefaced ibis, avocets, killdeer, and Wilson's phalaropes (hereafter phalaropes) were the most abundant breeding waterbirds at RLSA in 1994. Peak waterbird numbers occurred in spring, when the area was used by both locally breeding pairs and m~grants en route to other breeding areas (Fig. 1). Transient shorebirds and nonbreeding species were present in small numbers throughout the summer, and species richness varied little throughout the field season. Tall emergent habitats were used by ducks, but avoided by shorebirds (Fig. 2). Ducks also .used DO areas to a greater extent than shorebirds. Mallards, teal, gadwa.lls, and phalaropes used SE sites to a greater extent than other cover types at RLSWA. Avocets used SO habitats, and killdeer used US and SG sites, more than other species. Exploratory analyses of count data have been conducted using program DISTANCE. Sample sizes were adequate to obtain density estimates for mallards, cinnamon teal, redheads, avocets, and killdeer, but were too small to obtain separate density estimates for each count or for each cover type. Furthermore, the layout of the transect lines likely resulted in the violation of some assumptions required for use of program DISTANCE. In September, we established 14 new transect lines that will be used in future counts at RLSWA. Food Habits A total of 163 (95%) of the birds we collected contained food in their esophagus, including 147 adults and 16 juveniles. Seasonal diets of each species were extremely variable, even at higher taxonomic levels (Table 1). Aquatic invertebrates comprised a substantial proportion of the diets of all species, although plant foods comprised an average of 45-75% of gut contents of duck species. Chironomid larva were an important food item for adults of most species, occurring in 33% (gadwall and killdeer) to 62% (avocet) of each species, and comprising 3.5% (gadwall) to 46.6% (avocet) of the diets of each species. Shorebird species consumed high proportions (15-43%) of coleopterans. Carcass Analyses We collected a total of 172 birds in 1994. Body measurements are presented in Table 2. Carcass analyses will be completed in 1995. Carcass lipid, protein, and mineral content will be compared among birds in different reproductive categories for each species. External body measurements will be used to correct for variation in nutrient composition due to individual differences in structural size. Time-Activity Budgets We collected 544 focal sessions (4,982 minutes) of time-activity data on 12 waterbird species in 1994. Time budget data will be analyzed to determine the effects of cover type, month, time of day, and social status on activities of each -species. �18 Invertebrate Collections processing of invertebrate samples was completed in January of 1995. A reference collection of aquatic invertebrate taxa at RLSWA was produced. Invertebrate data will be analyzed to examine how invertebrate diversity and abundance varies over time at individual sites, and h PLANS FOR 1995-96 Analyses of data collected in 1994 will continue. Results from the 1994 pilot year were used to develop a detailed study plan for 1995-96. Linetransect counts, collections, time-activity budgets, and invertebrate collections will continue in 1995. In addition, invertebrate and plant food samples will be taken at collection sites; water depth and conductivity will be measured at collection sites, time-activity sites, invertebrate collection sites, and random sites in each cover type at RLSWA throughout the 1995 field season; and randomly located plots will be searched monthly for waterbird nests. Nest vegetation measurements will be taken, and the fate of each nest will be determined. Field work will resume in April 1995. This project is expected to continue for 2-3 years. LITERATURE Altmann, J. 1974. Observational Behaviour 49:227-267. CITED study of behaviour: sampling methods. Buckland, S. T., D. R. Anderson, K. P. Burnham, and J. L. Laake. 1993. Distance sampling: estimating abundance of biological populations. Chapman and Hall, London. Colorado Division of Wildlife. 1989. Colorado statewide waterfowl management plan 1989 - 2003. Colo. Div. Wildl., Terrestrial Wildl. sect., Migratory Game Bird Program Unit. 97pp. Gilbert, D. W., D. R. Anderson, J. K. Ringelman, and M. R. Szymczak. In press. Response of nesting ducks to habitat and management on the Monte Vista National Wildlife Refuge. Wildl. Monogr. Laake, J. L., S.T. Buckland, D. R. Anderson, and K. P. Burnham. 1994. DISTANCE user's guide, version 2.1. Colorado cooperative Fish and wildlife Research Unit, Colorado State Univ., Ft. Collins, CO. 84pp. Nelson, D. L., and M. F. Carter. 1990. Birds of selected Luis Valley. Colo. Div. Wildl. Unpubl. Rep. 40pp. Olterman, J., ed. wild1- Rep. 1993. The San Luis Valley waterbird R. A., W. D. Graul, and G. C. Miller. movements of ciconiforms in Colorado. Conf. 3:49-58. Tacha, T. C., P. A. Vohs, and G. C. Iverson. 1985. and continuous sampling methods for behavioral Ornithol. 56:258-264. by: James H. Gammonley Researcher/Scientist III plan. of the San Colo. Div. 1979. status, distribution, and Proc. Colonial Waterbird Group Ryder, Prepared wetland A comparison of interval observations. J. Field �19 1,200 I I I I I 1,000 , , I LL o ,.," 600 \ \ , ,," \ \, , .. , , .. " ," '_22 .. .. ,,, I \ "'1' , .... \ ill m :::> ," , \ ," , , a: ~ .".. \ I I ' " a: m , '" 'I " o 24 , ,, ' " "\ , I (J) " ,.. • I I I I .. •• 400 " z 5-2 5-17 5-24 5-31 6-7 6-14 6-21 6-28 7-6 7-12 7-19 7-26 8-2 DATE Rg.1. Waterbird use recorded on line transects at RLSWA, 1994. Mallard Blue-winged teal Redhead Cinnamon teal Gadwall Rg.2. Killdeer Avocet • Short emergent ~ .Tall emergent [J Saltgrass • Shallow open IZ] • Upland Deep open Phalarope Use of RLSWA habitats by waterbirds censused in 1994. �20 Table 1. Diet composition in adults of 7 species of breeding waterbirds collected at RLSWA in 1994, based on esophagus and proventriculus contents. Values are expressed as aggregate percent dry weight (frequency of occurrence). Trace amounts «0.05 g) tr. :2l2ecies (n} MALL (l5} foo!1 lt~m GADW CITE (U} (;ll} .AMPHIPODA BWTE (8} NEMATODA CLADOCERA COPEPODA OSTRACODA 0.1 (7) 1.6 (5) tr (5) 12.8 (38) tr (10) 0.8 (24) tr (5) 2.5 (38) tr (10) tr (33) (2Z} 0.5 (4) tr (7) 0.5 (4) HYDRACARINA tr (7) 0.7 (13) tr (13) tr (7) 0.6 (13) 0.8 (27) GASTROPODA Lymneaidae Physidae Planorbidae COLEOPTERA curculionidae larv. ad. 0.8 (20) tr (7) larv. ad. Haliplidae WIPH (JO} tr (13) EUBRANCHIPODA Dytiscidae KILL (~~} 0.1 (4) tr (4) 0.5 (10) OLIGOCHAETA AMAV 0.2 (5) 0.2 (5) 1.4 (33) 0.1 (5) tr (5) 0.3 (24) Hydraenidae Hydrophilidae 1arv. tr (7) ad. larv. tr (5) tr (5) 0.2 (5) 0.8 (14) 15.2 (40) 42.6 (73) 25.1 (41) 12.2 (23) 15.7 (57) 0.7 (10) 0.6 (3) 3.3 (6) 1.1 (7) 13.2 (33) 5.9 (11) 0.4 (4) 3.7 (28) 4.3 (12) 0.1 (10) 23.8 (53) DIPTERA 0.8 (4) 64.2 (68) tr (7) 1.3 (17) 3.3 (7) 5.5 (20) 41.3 . (57) tr (4) 46.6 (64) 3.8 (12) 4.0 (8) tr (3) tr (3) 17.3 (33) 4.0 (7) 0.8 (17) 0.5 4.0 6.1 (20) ad. Agromyzidae 0.2 (13) 5.1 (8) 5.1 (8) ad. ad. Unidentified tr (5) 0.8 (19) 20.8 (63) 14.0 (50) 4.0 (38) 2.8 (50) 1.0 (25) tr (3) 4.3 (13) 2.5 (17) 1.8 (3) 0.1 (5) larv. Heteroceridae tr (19) 5.0 (24) 4.1 (14) 0.1 (10) 0.8 (19) 1.0 (29) 20.8 (67) 40.7 (71) tr (5) 2.8· (33) 0.6 (14) 17.3 (62) 5.0 (14) tr (5) 21.4 (57) 1.3 (19) 0.9 (19) 0.8 (13 ) 13.1 (75) ad. 1.9 (4) 2.6 (11) 2.5 (7) 46.2 (59) tr (4) Ceratopogonidae larv. ad. Chironomidae larv. pup. ad. CUlicidae larv. Dolichopodidae ......... _ larv. tr (7) tr (7) 22.8 (53) 0.1 (7) 0.8 (20) tr (7) tr 12.9 (50) 0.2 (25) tr (13) .................................................................................................................................................................................................. tr (4) 3.7 0.5) 12.7 (41..) 5.6 (22) 16.0 (41) 4.4 (7) - ....................................... �21 .Table l. (Continued) [QQ9 it!ilm Dolichopodidae pup. MALL GADW (1:2) (n) CITE (All) ~ngci!ils(n) BWTE UI) (13) tr (7) AMAV KILL (~::i) (JQ) (,8 ) (10) 4.0 (4) 1.3 (7) 4.0 (13) 0.8 (4) 4.5 (12) 1.3 (7) 8.0 (13) 0.6 (7) ad. Ephydridae larv. tr (7) tr (5) 12.0 (24) ad. Muscidae larv. Stratiomyidae larv. tr (13) tr (7) Tabanidae larv. Unidentified larv. tr (13) 0.1 (5) 0.1 (5) 0.4 (14) 5.3 (14) tr (5) 0.7 (10) 4.6 (10) larv. C1:\llib1:\gt;Ls 0.4 (14) ad. ~ ad. HEMIPTERA Corixidae larv. ad. Notonectidae 1.0 (13) 0.6 (7) 0.4 (7) 0.1 (19) 0.1 (14) tr (5) 1.6 (14) 1.6 (14) HOMOPTERA HYMENOPTERA 0.4 (7) 0.4 (7) 1arv. tr (9) tr (5) tr (5) Limnephilidae larv. 4.8 (7) 4.8 (7) 7.4 (32) 2.1 (12) 5.3 (20) TOTAL ANIMAL SEEDS IUIilQcbsu::i§ 1.3 (7) 16.2 (33 ) 10.1 (26) 6·.1 (22) 4.6 tr (4) 1.3 (7) tr (3) 3.1 (4) 3.1 (4) 3.4 (10) 2.3 (10) tr (5) tr (10) larv. SPIDER ni~t1cbli§ 5.8 (25) 5.8 (13) 1.1 (5) tr (5) larv. TRICHOl>TERA Leptoceridae (7) 0.3 (4) 5.0 (13) Coenagrionidae 1.0 (7) 1.0 (7) LEPIDOPTERA Unidentified 0.3 (4) 0.5 (5) 0.6 (5) Lestidae larv. 1.9 (4) tr (13) larv. ODONATA 1.9 (4) tr (4) tr (4) tr (4) tr (4) ad. EPHEMEROPTERA WIPH (~Zl tr (7) tr (4) 31.5 (80) 25.4 (81) 55.4 (90) 45.8 (88) 95.0 (100) 97.7 (100) 90.2 (96) 68.4 (100) 10.6 (53) tr 19.2 (62) 3.9 (19) 44.6 (90) 9.1· (43) 5.1 (10) 53.8 (88) 26.4 (75) 4.2 (13) 2.5 (12) 1.1 (7) 7.9 (14) (7) ..................... :...................................................................................................................... -.............................................................................. -........................ .;' �22 Table 1. (Continued) I [QQ!;)1:t!ilm JUDCUS Eotilmom~toD Sdr:pus' MALL (l::l) GADW (ill) VEGETATION (II) tr (7) 0.6 (7) 39.4 (93) 17.8 (33) 0.1 (7) ~ EQti:lmog!:!toD AMAV {il::i} KILL {JQ} WIPH {ilZ} 0.1 (4) 13.3 (48) Zi:lnnlQbdlii:l Unidentified CITE (ill ) fll2~Qi!i!1Z (D) BWTE 2.0 (29) 55.4 (76) 6.8 (10) 4.6 17.0 (57) 1.1 (14) 12.3 (43) tr (5) 0.5 (13) 3.9 (50) 2.0 (4) 0.7 (3) 4.2 (7) tr 18.8 (50) 0.4 (13) 0.4 (3) 1.2 (10) (4) 3.7 (4) 1.9 (4) (4) 1.2 (10) 1..9 (4) 0.4 (4) 2.5 (20) (5) 2.4 (16) 0.1 0.1 (7) 40.3 (71) 3.7 (24) tr (5) .0.4 (13) TOTAL PLANT 68.5 (100) 74.6 (100) 44.6 (90) 54.2 (88) 5.0 (24) 2.3 (13) 9.8 (19) Mean g food (dry weight) 0.3'28 0.090 0.134 0.063 0.049 0.014 .0.005 Zi:lnniQb!illlli:1 Unidentified �23 Table 2• .Body measurements of adults of 7 species of breeding waterbirds collected at RLSWA in 1994. Values are.expressed as mean (standard error). Body mass (g) Males Females Length (mm) Males Females Wing length (mm) Males Females Culmen (mm) Males Females Tarsus (mm) Males Females Right breast (g) Males Females Heart (g) Males Females Liver (g) Males Females Gizzard (g) Males Females Intestine (g) Males. Females Ceca (mm) Males Females Right leg muscle (g) Males Females MALL GADW (Hll) (6a5) SR~Q1~1:I(n CITE (5ll6) = mSll§l:Iifem511~1:!) BWTE KILL AMAV ( 3i5) (lU13) (2000) WIPH (lV14) 1117 (37) 946 (26) 855 (42) 734 (18) 374 (11) 362 (8) 3.69 (15) 362 (12) 317 (7) 303 (7) 81 (2) 97 (3) 45 (1) 58 (2) 564 (6) 517 (5) 5~2 (7) 466 (2) 404 (6) 375 (6) 399 (1) 368 (5) 448 (4) 426 (3) 251 (2) 254 (3) 222 (1) 240 (3) 291 (4) 271 (3) 281 (3) 249 (7) 191 (1) 180 (6) 190 (1) 175 (5) 254 (12) 262 (13) 165 , (2) 169 (2) 135 (6) 136 (2) 57.3 (0.2) 51.9 (0.6) 45.4 (1.2) 40.8 (0.4) 44.1 (0.9) 42.4 (0.4) 41,.5 (1.5) 39.7 (0.9) 94.4 (1.5) 86.0 (1.3) 19.7 (0.4) 19.9 (0.3) 29.4 (0.5) 32.6 (0.5) 45.8 (0.4) 43.7 (0.6) 42.3 (0.8) 39.9 (0.5) 32.9 (0.2) 32.3 (0.6) 32.3 (0.8) 30.8 (0.7) 97.0 (1.2) 90.8 (1.2) 36.6 (0.4) 36.3 (0.3) 30.9 (0.4) 33.0 (0.3) 130.9 (2.9) 108.6 (3.2) 96.0 (2.9) 76.2 (1.8) 41.6 (1.8) 37.6 (1.0) 40.3 (2.4) 39;1 (2.3) 27.2 (0.7) 25.3 (0.5) 8.4 (0.3) 9.5 (0.3) 4.6 (0.2) 5.8 (0.3) 13.5 (0.5) 11.0 (0.5) 12.3 (0.9) 9.0 (0.4) 3.7 (0.3) 3.E! (0.1) 4.1 (0.4) 3.6 (0.2) 3.4 (0.1) 3.4 (0.1) 1.1 (0.1) 1.2 (0.1) 0.6 (0.'0) 0.7 (0.0) 21.0 (1.3) 22.0 (1.3) 15.3 (1.1) 17.3 (1.0) 8.2 (1.1) 10.6 (0.7) 9.4 (1.7) 11.5 (1.0) 8.9 (0.6) 8.3 (0.5) 2.0 (0.1) 3.0 ,. (0.3) 1.4 (0.1) 1.·8 (0.2) 34.4 (5.1) 25.9 (1.6) 32.3 (2••) 23.6 (1.2) 8.1 (1.2) 8.6 (0.4) 10.2 (0.5) 11.6 (2.3) .6.2 (0.4) 5.6 (0.'2') 1.5 (0.1) 1.7 (0.'1,. 0.9 (0.0) 1.1 (0;1, 28.1 (2.4) 26.8 (1.0) 24.7 (1.4) 26•.1 (1.4) 13.4 (0.8) 15.1 (0.4) 13.4 (1.0) 13 •. 7 (0.7) 11.2 (0.6) 12.2 (0•.6,). 2.2 (0.1) 2.6 (0.1) 1.6 (0.1) 2.2 (0.1) 307 (38) 239 (9) 414 (27) 385 (16) 139 (11) 158 (6) 154 (1) 148 (9) 117 (3) 117 (3) 69 (2) 70 (6) 55 (2) 68 (4) 40.8 (1.3) 32.1 (1.5) 24.5 (1.2) 19.4 (0.7) 12.1 (0.3) 11.2 (0.4) 11.1 (0.5) 10.1 (0.8) 12.0 (0.4) 10.4 (0.3) 2.8 (0.1) 3.2 (0.1) 1.2 (0.1) 1.5 (0.1) ,'" ��25 Colorado Division Wildlife Research October 1995 of Wildlife Report JOB PROGRESS State of Colorado Project W-166-R-4 Work Plan Job Title: Period Cooperative Covered: Auth~r: Personnel: Michael Migratory : Job 10 REPORT Game Birds Investigations __ 1_ Management 01 April Programs 1994 through 31 March 1995 R. Szymczak Michael R. Szymczak, Colorado Division of Wildlife ABSTRACT Recommendations for wetland habitat improvements and/or management were provided for public and private land managers across the state. Proposals for funding projects with Duck stamp monies were evaluated and rated. Presentations on various aspects of waterfowl ecology and wetland reclamation were given at training schools and workshops. Initial organizational meetings on establishing Focus Areas and Focus Area partners in Colorado for the Intermountain West Joint Venture of the North American Waterfowl Management Plan were held. Responsibilities as Colorado's representative on Pacific Flyway Study Committee, including chairman of subcommittees for 4-Corners Band-tailed pigeons and Rocky Mountain Population Canada geese were fulfilled. Waterfowl surveys were conducted in North Park and on selected private lands in the San Luis Valley. A 3-5 year cooperative post breeding goose banding program was continued in the Cortez-Mancos and Durango areas. BDOWOlll00 ��27 COOPERATIVE MIGRATORY BIRD MANAGEMENT PROGRAMS Michael R. Szymczak James K. Ringelman In 1988, the Colorado Division of Wildlife (CDOW) created the Migratory Game Bird Program Unit (MBPU) within the Terrestrial Wildlife Section. This administrative change combined all individuals having statewide responsibilities for research and management of migratory game birds. Members of the MBPU work in concert to improve migratory bird management in Colorado. This job was created to allow team members to participate in these management programs. In November 1993, project personnel assumed additional responsibility for leading and administering the Duck Stamp wetland development program. In July 1994, the author became Colorado State Coordinator for the Intermountain West Joint Venture of the North American Waterfowl Management Plan. P. N. OBJECTIVES 1. Participate in developing and implementing habitat-based waterfowl management plans on a statewide~ habitat region, and project basis. 2. Advise and/or and/or habitat 3. Present information on the principles of waterfowl management to workshop attendees, educational clas.ses, and conservation organizations. 4. Participate levels. s. Cooperate in developing surveys and techniques of migratory bird management programs. state and federal land managers on beneficial habitat acquisitions developments and provide expertise in preparation of development management plans. Advise private land man~gers in developing management plans and assessing impacts on waterbird populations. in migratory bird management meetings at the state and flyway that will assess the impact SEGMENT OBJECTIVES 1. In conjunction with the Statewide Waterfowl on habitat region plans, when requested. 2. Serve as chairman of the Waterfowl Habitat Project Review Committee (WHPRC). Provide biological expertise on waterfowl biology and wetland development programs on public and private areas when requested. 3. Prepare and present requested. 4. Compile appropriate migratory bird population status information and represent Colorado at Western Migratory Upland Game Bird meetings and Pacific Flyway Study Committee and Council meetings. Attend migratory game bird program and biologist meetings in Colorado when requested. s. Provide methodology for wetland habitat surveys when requested. lectures on migratory Management Plan, continue game bird management and migratory 6. Cooperate in Canada goose trapping and banding Cortez area and in Brown's Park. work when game bird population operations in the Durango- 7. Conduc,t experimental surveys of waterfowl breeding pairs on wetland development units under management by the U. S. Fish and Wildlife through Partners in Wildlife program. .. . �28 RESULTS Waterfowl Management Plans Figures and Tables for breeding and wintering ducks and geese were updated for the San Luis Valley (SLV) Waterbird Plan. Additional recommendations were made for changes in the initial plan. wetland Developments, and WHPRC and IWJV Activities Potential sites for wetland developments and existing wetlands were visited and recommendations for development and/or management were made for: the Lone Dome State Wildlife Area(SWA,CDOW) near Dolores, the Jackso~ Lake SWA (CDOW) near Ft. Morgan, the Jackson Lake State Land Board Tract near Ft. Morgan (CDOW), the Kemp/Breeze SWA near Hot Sulphur Springs, the Horsethief SWA (CDOW) near Grand Junction, the Colorado River Wildlife Area (Bureau of Reclamation) near Grand Junction, Hi-Line Reservoir Colorado State Park (CSP) near Grand Junction, Barr Lake (CDOW/CSP) near Brighton, the Showcase project site near Leadville (USFS), ECO-TURF farm (Private) near Ft Collins, Eagle Ridge Ranch (Private) near Gunnison, and the 4-Eagle Ranch (Private) near wolcott. A~ chairman of the WHPRC, I: chaired 2 committee meetings for ranking and funding proposals submitted for the 1994-95 funding year; informed proposal submittees of the out come of their funding request; periodically monitored progress of project planning, construction, and money of new and previous years funded projects; coordinated Site Specific Agreements and fund reimbursement with the Ducks Unlimited MARSH program; solicited, reviewed, and obtained additional information needed to complete project proposals submitted for 1995-96 funding; distributed proposal packets to committee members, and scheduled the 1995 meeting to review proposals. As State Coordinator for the IWJV, I solicited potential partners in wetland acquisition and development in the IMJV area in Colorado (all of Colorado west of the eastern foothills), chaired a meeting of the partners in which 7 geographic Focus Areas were selected, recruited chairman for 5 of 7 Focus Areas, attended 1 Focus Area meeting, and attended 3 meetings of State Coordinators of the IWJV. In addition, I prepared and forwarded background information on preparing Implementation Plans to Focus Area chairman. Informational Programs Formal presentations were given at the Colorado State Extension Service's Reclamation Workshop in Grand Junction on Wetland Development and Enhancement, and Reclaiming Degraded Wetlands. A program on waterfowl use of wetlands, designing wetlands for waterfowl, and applying for Colorado Duck Stamp and Ducks Unlimited MARSH monies was presented to the CDOW District Wildlife Management Trainees. Waterfowl Technical Committee and Council Meetings I attended the July 1994 Pacific Flyway Study Committee (PFSC) and Council meetings. Waterfowl population status was reviewed along with characteristics of the 1993-94 waterfowl hunting season harvest, and proposed 1994-95 hunting season recommendations were formulated and forwarded through the Council to the USFWS Regulation Committee. Populations of specific interest to Colorado whose status was reviewed in July were (1) breeding and wintering mallards inhabiting western Colorado and (2) the Rocky Mountain Canada goose population. I chaired the Rocky Mountain-Canada goose population SUbcommittee. The winter meeting of the PFSC featured training on and discussions of the models developed for Adaptive Harvest Management of ducks by the Federal and State Adaptive Harvest Management Team; the development of a proposal for an experimental season to evaluate the effect of simplified duck bag limits on hunter recruitment; and a review of band-tailed pigeon wing aging techniques and test of wing aging skills of Study Committee members. �29 The March 1995 meeting of the Pacific Flyway study Committee allowed committee members to exchange general information on migratory game bird populations and formulate regulatory recommendations for the Flyway Council, for species hunted before Oct. 1, including doves, band-tailed pigeons, snipe, rails, and cranes. I chaired the 4-Corners Band-tailed pigeon subcommittee. Early season special Canada goose seasons were also considered. Of special interest was the review of duck regulation packages proposed for the 1995-96 hunting seasons. Three packages (restrictive, moderate, liberal) were offered, and the one selected would be based on a combination of the (1) mallard breeding populations in prairie U. S. and Canada, and (2)·number of ponds in prarie Canada, both as measured during the May 1995 breeding pair survey. For all meetings reports containing topics pertinent to Colorado were written, compiled and. distributed to appropriate CDOW personnel. Population Survey Methodology Surveys of nesting and brood rearing Canada geese on Walden Reservoir in North Park during spring 1994 recorded the lowest number of nests since surveys began in 1990; primarily because of nest destruction by badgers on one island that usually supports a high nest density. Detailed tables are submitted to CDOW Northeast Regional and BLM personnel annually. cooperative Canada Goose Banding For the third consecutive year, Canada geese were banded in the CortezMancos area. In addition geese were banded for the first time in the DurangoBayfield area. The trapping operation was conducted with the cooperation and personnel of the CDOW Southwest Region. A total of 126 goslings and 35 adults were banded at 5 locations in the Cortez-Mancos area, and 100 goslings and 106 adults were banded in the Durango-Bayfield area at five locations in the late June 1994 (Appendix A). The total number banded in three years in the CortezMancos area is 550. Because of other commitments, geese were not banded in Brown's Park in 1994. Duck Breeding Pairs - Partners for Wildlife Areas In May 1994, duck breeding pair counts were conducted from the ground and the air on 13 Partners for Wildlife wetland development areas in the San Luis Valley. These counts were designed to evaluate methods of enumerating breeding pairs on these small, intensively managed private land areas that are leased by the U. S. Fish and Wildlife Service. It is assumed that special surveys of these areas will be needed to evaluate the effectiveness of the program and provide reliable duck breeding pair numbers from these areas for the valley-wide breeding pair survey. Since initial analysis of 1993 counts showed little variation between ground or air count results obtained on specific areas on consecutive days, each area was counted only once from the ground and the air in 1994. Analysis of these data are not complete at this time. DISCUSSION Project personnel provide useful information in planning and evaluating waterfowl management and habitat enhancement programs in Colorado and educating land management agency personnel about the habitat requirements of waterfowl. We expect that with increased emphasis on habitat enhancement in Colorado as outlined in the statewide Waterfowl Management Plan, our services will be more in demand. Additional activity on the WHPRC will help insure that the money raised through the Colorado Duck Stamp program is spent in accordance with the objectives of the program. Initiation of the IWJV in Colorado presents an opportunitie to increase funding for wetland acquisition and development in Colorado. �30 Conducting and/or formulating surveys and banding efforts and informing. management agency personnel about various aspects of waterfowl and wetland ecology provides a valuable service to management agencies, the waterfowl resource and in 130me cases the hunting public. Continued participation on Flyway committees ensures that Colorado will remain informed on migratory bird matters, have input in migratory bird hunting regulations, and have influence on habitat programs affecting migratory game birds. Prepared by: ?l0L,jVP~ Michael R. Szymczak Researcher/Scientist IV �31 APPENDIX A Table 1. Numbers of Canada geese trapped and banded, by location, in the Cortez-Mancos area, 1992-94. Loc. F Loc. M Location 92 93 Thomas's Pond. 12 8 Verde' Ponds 1 Dolores' Hatchery 6 Baikie Ponds 5 6 7 35 Duddleson Ponds 9 34 McPhee Reservoir 1 92 93 17 14 11 93 8 2 2 18 7 30 9 26 Cortez Golf Course 6 93 5 4 2 3 3 4 8 4 3 6 8 3 16 16 27 Ad. M Ad. F 23 39 23 82 71 28 23 29 64 2 19 19 168 Total 18 23 4 12 57 O'Neal 17 31 16 18 83* Vallecito 4 5 2 0 11 Tay-Col Cattle Co. 9 14 6 5 34 Dove Ridge Subdivision 6 2 0 0 8 44 56 48 58 206 * One bird banded of unknown age and unknown sex. 16 8 James Ranch Subtotal .;.. 17 Table 2. Numbers of Canada geese trapped and banded, by location, in the Durango-Bayfield area, 1994. Loc. F 28 10 i Loc. M 42 5 0 23 94 34 4 9 64 93 22 2 9 92 9 3 5 2 7 88 3 4 2 Total 94 1 1 27 68 92 3 10 62 94 2 5 20 88 Ad. F 0 3 Forest Pond 50 92 13 14 5 Total 94 6 7 11 16 14 Colbert's Pond Browning Pond 94 Ad. M 221 161 ��33 colorado Division of Wildlife Wildlife Research Report October 1995 JOB PROGRESS REPORT state of Colorado Project W-166-R-4 Work Plan 22 Job Title: Author: : Job Migratory Period Covered: Migratory Game Birds Investigations __ 2_" Game Bird Publications 01 April 1994 through 31 March 1995 Michael R. Szymczak Personnel: Wildlife James K. Ringelman and Michael R. Szymczak, Colorado Division of ABSTRACT The following list contains those articles that were prepared submitted for publication or published during this segment: and/or Ball, I. J., T. E. Martin, and J. K. Ringelman. 1994. Conservation of nongame birds and waterfowl: conflict or compliment? Trans. N. Am. Wildl. Nat. ,Resour. Conf. 59:337-347. Gilbert, D. W., D. R. Anderson, J. K. Ringelman, and M. R. Szymczak. Response of nesting ducks to habitat management on the Monte Vista National Wildlife Refuge. Wildlife Monograph 131 (In press). Jeske, c.'W., M. R. Szymczak, D. R. Anderson, J. K. Ringelman, and J. A. Armstrong. 1994. Relationship of body condition to survival of mallards in San Luis Valley, Colorado. J. Wildl. Manage. 58:787-793. Prepared by: A~i!..JP'~£ , Michael R. SZymcak Researcher/Scientist IV -_/'." iiillf,u BDOWDll1Dl ��35 'colorado Division Wildl.ife Research October 1995 of Wildlife Report JOB PROGRESS state of Project: Work Colorado W-164-R-4 Plan __ 2_,_ Job Title: Period Forensics Investigations Job_3_ DNA 'ANALYSIS FOR SPECIES AND SEX IDENTIFICATION AND TISSUE MATCHING OF BLOOD, BLOOD STAINS AND MEAT SAMPLES OF COLORADO'S FORENSIC CASES. Covered: Personnel: REPORT 1 July, W.J. Adrian, 1994 30 June, R.P. Ellis, 1995. and R.J. Magnuson. ABSTRACT • -. DNA primers were used to amplify specific portions of the X chromosome and/or the Y chromosome. The amplification product for the X chromosome is 442 base pair (bp) and for the Y chromosome 224 bp. This technology is now on line for the Colorado Division of Wildlife and is a proven, court acceptable technique for determining the gender of deer and elk. This technique has not been perfected on pronghorn. Work is in progress on this species. "OM:; I :~ , '" ��37 DNA ANALYSIS FOR SPECIES AND SEX IDENTIFICATION AND TISSUE MATCHING OF BLOOD, BLOOD STAINS AND MEAT SAMPLES OF COLORADO'S FORENSIC CASES. SEGMENT OBJECTIVES 1. Obtain equipment, supplies, DNA-based technologies. primers 2. Use the DNA-based probes to assay game animal individual identification. 3. To place on-line, at Colorado State University, Dept. of Microbiology a laboratory for routine testing of game animal tissue samples for gender determinations, population analysis and wildlife enforcement. METHODS and probes necessary tissue for use in for sex, species and AND MATERIALS This work is a cooperative endeavor between Robert Ellis of Colorado State University. the Division of Wildlife and Dr. Wildlife forensics utilizing DNA assays have been analyzed by several investigators. These include assays utilizing PCR primers (short DNA sequences which initiate DNA amplification) for sex determination of deer, antelope and other mammals; species specific RFLP (restriction fragment length polymorphism) probes for discrimination of antelope, elk and mule deer; mitochondrial DNA probes to distinguish mule and white-tailed deer (1,2,3). Other commercial probes by AGTC (Analytical Genetic Testing Center, Inc.) are available for possible use in big game DNA analysis (4). The non-commercial probes are already available for use in objective 2 of this proposal. In November 1992, two papers by J. LeMay, S. Fain and J. Ruth were presented at the North West Association of Forensic Scientists Fall Meeting in Portland, Oregon on the use of DNA assays for game animals (5,6). These papers demonstrated the successful development and use of the DNA probes to identify from meat and blood, the gender of most ruminant species as well as routine "individualization" of white-tailed deer. Development of individual-specific RFLP banding patterns from elk genomic DNA using a non-radioactive protocol has allowed for the elimination of radioisotope handling in DNA analysis (7). These patterns arise by a conjugated antibody-enzyme mediated reaction. The use of non-radioactively labelled probes will allow for reduction of safety concerns associated with radioactively labelled. probes, easier disposal of used probes and longer probe storage life. The non-.isotopically labelled probes have at least equal resolution of the radioactive banding patterns. Though previous work has demonstrated adequate polymorphism using·RFL~ analysis, many forensic samples contain insufficient DNA to utilize RFLP methodology. The third objective is to utilize RAPD-PCR or similar techniques in differentiating between individuals of the same species. This technique would be used as a "fingerprinting" scheme to determine without doubt the identity of tissues collected from the saine animal. Likewise, tissues collected from different animals of the same species would have a different DNA "fingerprint" (8,9,10,11). In total, we would be able, in a matter of a few days, to produce irrefutable evidence of gender, species, and individual origin of tissue samples. ." �38 RESULTS AND DISCUSSION DNA primers were used to amplify specific portions of the X chromosome and/or Y chromosome. The amplification product for the X chromosome is 442 base pair (bp) and 224 for the Y chromosome. This gives us much better separation between the X and Y chromosome, therefore making it much easier to read. This technology is now on line for the Colorado Division of Wildlife and is a proven, court acceptable technique for determining the gender of deer and elk meat, blood and blood stains. This technique has not yet been proven for the pronghorn. Genetic "fingerprinting" of meat, blood and blood probes are now being stains. LITERATURE tested for the individualization CITED 1) Cronin, M.A. 1986. Genetic relationships between white-tailed deer, mule deer and other large mammals inferred from mitochondrial DNA analysis. M.S. thesis. Montana state University. 2) Menke, S.D. 1992. Protein synthesis during development of normal and mutant flight muscle in Drosophila melanogaster and Isolation and characterization of sex specific DNA in deer and antelope. Ph.D. thesis. University of Wyoming. 3) Blackett, R.S., and P. Keim. 1992. Big Game Species deoxyribonucleic (DN~) probes. Journal of Forensic 37:590-596. Identification Sciences. by 4) Analytical Genetic Testing 80231.7808 Cherry Creek #201. CO. 5) LeMay, J.P., and S.R. Fain. 1992. Gender Determination Wildlife Species from PCR Amplified Sex-Linked Genes. Scientist Fall Meeting. Portland, OR. 6) Ruth, J.L. 1992. The Forensic Identification of Individual Deer Using DNA Probes. Forensic Scientist Fall Meeting. Portland, OR. 7) Stern, C.M. 1992. State University. 8) Welsh, J., and M. Mcclelland. PCR and a matrix of pairwise Res. 19:5275-5279. 9) Kambhampati, S., W.C. Black IV, and K.S. Rai. 1991. RAPD-PCR for identification and differentiation of mosquito species and populations techniques and statistical analysis. J. Med. Entom. 29:939-945. Center, Non-Isotopic Inc. South Drive, DNA fingerprinting Denver, of Mammalian Forensic of Elk. Colorado 1991. Genomic fingerprinting using primed combinations of primers. Nucleic Acids 10) Orita, M., H. Iwahana, H. Kanayawa, K. Hayaski, and T. Sekiya. 1989. Detection of polymorphisms of human DNA gel electrophoresis as singlestrand conformation polymorphisms. Proc. Natl. Acad. Sci. USA. 86:2766-2770. 11) Hayaski, K. 1991. PCR-SSCP: A simple and sensitive method for detection of mutations in the genomic DNA in PCR methods and applications, Cold Spring Harbor LaB. Press, pp.34-38. Prepared by:~~~ ~~~ William J. Adrian Wildlife Researcher _ .", �39 Colorado Division of Wildlife Wildlife Research Report October 1995 JOB PROGRESS REPORT State of_~Co<o~IQ~rawdltloo:.,__ Project: _ __..(W ..•.•.... -...•. l ••. 50,._...•. R,,_-...,7),.___: Peregrine Period Covered: Falcon Restoration Program 1 July, 1994 - 30 June, 1995 Personnel: G.R Craig, Colorado Division of Wildlife and J.H. Enderson, The Colorado College. ABSTRACT In the 1995 peregrine breeding season, 71 territories were occupied by 61 breeding pairs that fledged 94 young. Productivity averaged 1.40 young fledged for those pairs that were monitored. Contents of 5 nonviable eggs were collected and have been preserved for future analysis. Shell fragments were collected at 12 sites and are awaiting measurement. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the author. ��41 PEREGRINE FALCON RESTORATION PROGRAM Gerald R Craig SEGMENT OBJECTIVES 1. Annually monitor the number of breeding pairs of peregrines and their reproductive success in Colorado. 2. Annually monitor organochlorine pesticide levels in wild breeding peregrines in Colorado. 3. Monitor breeding population turnover through band recoveries, presence of color markers, and telephotographic identification of individual breeding adults. 4. Augment poor wild production by placementof captive hatched wild young and captive produced young into occupied wild nests. 5. Release captive hatched and captive produced young at potential and vacant wild territories. 6. Monitor recruitment of reintroduced peregrines into the wild breeding population of Colorado. METHODS AND MATERIALS 1. Visit all known peregrine breeding territories throughout Colorado and observe them from a distance to establish the presence of breeding adults. Breeding pairs will be kept under surveillance to determine initiation of egg laying. Depending upon the individualfemale's reproductive history and eggshell condition (obtained through measurement of previous year's eggshell thicknesses) and availability of captive hatched young for release, breeding pairs either will monitored or manipulated as outlined in approach 4. Those pairs not designated to be manipulated will be revisited periodically throughout the nesting season to document reproductive success. When a pair's behavior indicates that egg laying has occurred and incubation is underway, the eyrie will be visited to document the number of eggs produced. The eggs will be candled to ascertain viability and approximate age. All nonviable eggs will be collected for chemical analysis. A second visit will be made to determine productivity, band nestlings, and collect eggshell fragments and unhatched eggs for thickness measurement and analysis under 2a and 2b. 2a. Eggshell fragments encountered during eyrie visits described in approaches 1 and 4a will be measured for index to thickness following standardized procedures. 2b. Whole, nonviable eggs which are encounteredduring eyrie visits will be collected, preserved and submitted to the appropriate Fish and Wildlife Service approved laboratory for pesticide analysis. Eggs collected from the wild in the course of Approaches 4a, 4b and 4c that are artificially at the Peregrine Fund's Boise, Idaho facility also will be submitted for shell thickness measurement and chemical analysis. 3. Peregrines present at breeding territories will be examined to determine the presence of bands or color markers. Band confirmation will be accomplished through observation from a distance with telescopes and concealed remote controlled cameras. When banded falcons are encountered, every effort will be made to read band numbers without trapping or handling the birds. It is possible this can be accomplished in most situations with a Questar field model telescope (SO-l3Ox). When band numbers cannot be discerned, attempts will be made to trap and examine the falcon at a time when capture will have least impact upon breeding activities. ..,. �42 4a. In accordance with an annual release plan developed and approved by the State, U.S. Fish and Wildlife Service, Bureau of Land Management, National Park Service, and the.Forest Service, a predetermined number of wild breeding pairs will be manipulated to augment natural productivity. Pairs with a history of reduced clutch size, cracked eggs, or infertile or dead eggs will be candidates for fostering efforts. 4b. On occasion, it may be necessary to recycle several early breeding pairs in order to delay them until captive hatched young of the proper age are available for placement into wild sites. No later than 10 days after the last egg has been deposited, the eyrie will be visited and the entire clutch removed without replacement. Approximately 14 days after removal of the clutch, the pair will recycle, select another nest ledge, and deposit a second clutch of eggs. If the eggs are thin shelled, they may be replaced with plastic replicas and treated as outlined in approach 4a This technique also works well to augment captive production with wild produced eggs. 4c. At times, pairs will select inferior eyrie ledges that may compromise nest success such as ledges that are too narrow to support a brood of large nestlings, the site may be vulnerable to predators, or it may be exposed to the elements. If the ledge cannot be mechanically improved, pairs can be relocated to other ledges through the recycling method described in approach 4b since they invariably relocate and select a new ledge when recycled. 5. In accordance with an annual release plan developed and approved by the State, U.S. Fish and Wildlife Service, Bureau of Land Management, National Park Service, and the Forest Service, a predetermined number of captive produced falcons will be released at unoccupied or potential sites through the technique of hacking. This technique is employed at locations that do not have the benefit of protection or care from adults. Young falcons of about 35 days of age will be placed in a hack box on a suitable cliff ledge at the reintroductionsite. They will be fed and cared for by attendants until they are flying and capable of fending for themselves. This technique assures that the young become familiar with their surroundings and hopefully will return to the site as adults and take up residency. Hacking requires constant attendance and observation to protect the vulnerable young and assure they have sufficient food while they are dependent upon the hack site. While the hack sites will be operated by the State, actual costs to operate the sites will be borne by the appropriate land administering agency (Forest Service, Bureau of Land Management, and National Park Service). 6. . Confirmedbreeding territories and selected potential breeding sites will be surveyed annually to document the presence of released falcons and ultimately determine the success of recovery efforts. RESULTS AND DISCUSSION Survey Effort In 1995,4 teams comprised of2 observers each were assigned particular regions of the state to monitor breeding activities and survey potential cliffs as time permitted. Three teams were assigned regions west of the Continental Divide and 1 team was located east of the Divide. Eighty-three sites were monitored by the teams. Two previously recorded territories were not visited due to their remoteness and time constraints. An additional 30 potential nesting cliffs were examined for presence of nesting peregrines as time permitted. Three previously unknown breeding territories were added, all of which were reported by other sources and confirmed by the teams. �43 Territoty OccUPanCY Breeding territory occupancy increased from 70 sites in 1994 to 71 in 1995. One site (site 84) remained vacant in 1995 after the apparent deaths of the adults in 1994. Site 87 was vacant in 1995 and site 81 was reoccupied after a vacancy in 1994. Three previously undocumentedsites( 87, 88, 89 and 90) were confirmed. Over the past 5 years, the rate of occupancy has fluctuated around the 80% level (Fig. 1). Figure 1 RATE OF OCCUPANCY 90% - 0 UI 0:: :::> g 7011. (1) UI !:: 60% (1) IL 0 >z UI 0 ffi Q. 50% - .-__ n n ----------_._----------------- N R ~ n n n ~ ~ Q M M M -------------------_. ~ ~ M § ~ ~ ~ ~ M ~ YEARS Reproduction In 1995, the number of breeding pairs (those that produced eggs) leveled off at 60 (Fig. 2.), while the productivity averaged1.40 young fledged per total pair which represents a continuing downward trend in recent years (Fig. 3). The average fledged brood size (young fledged per successful pair) was 2.42 which was a slight increase over previous years. This indicates that the lower productivity did not occur throughout the population, but was the result of the failure of pairs to produce young. In fact, 17 breeding pairs failed to produce young in 1995. The downward trend in successful pairs since 1987 (Fig. 4) mirrors the declining productivity over that period. Although the trend has been downward since the mid 1980's,productivity still remains above the threshold of 1.25 considered necessary for population stability. Until the breeding success stabilizes, productivity should continue to be monitored. Eggshell Condition Five whole, nonviable eggs were encounteredin the course of visits to 5 sites. Shell thicknesses from these eggs are not available at this time. Eggshell fragments were also collected from 12 additional nesting sites in the course of visits to band young. Thickness measurements are not available for these samples at this time. Until current thickness measurements are forthcoming, no conclusions can made about the 1995 thicknesses. Figure 5 shows the eggshell thickness trends through 1994. These values are highly variable due to small sample sizes, mixing of fragments from different eggs, and variation of thickness of fragments from the poles of the egg versus the waist. However, there appears to be a slight trend toward thicker eggs over the past 8 years with thickness averages less than 10010 thin since 1992. Greater variability among eggs is also evident since 1988 with some eggs being thicker than pre-DDT era eggs and others 16% thinner .;., �44 Figure 2 OCCUPANCY BY BREEDING PAIRS 10 ~·i.···········.· ••••••• ······················· •....•••.•••................•..•..•.••••..•.............•....•.••.....••.•......•...•................•••............................• 1 ~ ~·1··················································· n n u ~ n ···,~···I n n n 10 ~ ~ ~ ~ ~ ~ ~ N 5 ~ ~ ~ ~ N ~ ~ 00 ~ ~ ~ M ~ YEARS Figure 3 PERCENT OF TOTAL BREEDING PAIRS TIIAT WERE SUCCESSFUL 100% •• 5 80% •.•.••.•••..•.••.••••• LL ~ ~ 60% •••.••.••..••••••••.•• :::l til ffi 40% •••.••••••.••.•••••••• ffi0. 0% •• _ n 0 73 u --- 20% •••••••.•.•.•••••.•••• 74 --- 75 76 rr 78 79 eo 81 ~ __ ~ ~ YEARS ~ ~ ~ M Pan Organochlorine Residue in Eggs The 5 nonviable eggs collected during the 1995 season have been preserved along with eggs encountered in 1994, 1993, 1992 and 1991. This collection of 42 eggs is awaiting scheduling for pesticide analysis by the Fish and Wildlife Service when funding is available. Release and Augmentation Efforts Remedial management efforts such as fostering or hacking have not been undertaken since 1989. ." �45 Figure 4 PRODUCTIVITY OF COLORADO PEREGRINES 25- ~ III ~ 1_5 - ----------- --- ------------------------------ ------------------------------------------------_:p--<-:----------,----::'ci5-::-:~:~--------- ----------:--- -------------------- --------- -------------------------------------------------------------------/:~:-------------------------------------------------------------------------------------------------------_. ~ ~ ~ -------- ------- 0_5 - --.-- ···-····--······-··:··:·-.~~:::-·--·::'-.:.:6~~~~::·-:.:~:.:.:~~.::::.:.: :)/ ~ :.- - - - - - - - - - O-~r--,---r--,---~-,~~~~~~~~~~~--~--r--,---r--,---~~r-~---r-n ro ~ n ~ ro D 00 ~ ~ ~ ~ ~ ~ ~ ~ ~ 00 ~ ~ ~ ~ ~ YEARS _ Total Pairs ... ....,.... Unmanipulated Pairs Figure 5 CUMULATIVE EGGSHELL TIllCKNESSES 0.39 E .s _,.c:!i 0 0.37 Pre-DDT &a ThidaIcss 0.35 0.33 :E I- ~ s: ••coco .~ -IID%TlIIN .- .~ t- T 0.31 1,/ I\. ./ i..o""" 0.29 w 0.27 " \ \ / / :::r ,/ ......•... .L f- .....••.• -~ / - ~ ./ -- ~ "- ~ 0.25 I Years MOJdnun and Mlrinun Thlcknosses _ A_"ge Thlcknosses Prepared By: Gerald R Craig Life Science Researcher N ..~ ��47 Colorado Division of Wildlife Wildlife Research Report October 1995 JOB PROGRESS REPORT State of_--"Co<¥!llo""r...,ad ••• o,,--_ Project: _ ....• CW •...•.•... -....•• l=5..•.. 1-.,A;R._-7).r...J-_ : Bald Eagle Nest Site Protection and Enhancement Program Period Covered: 1 July, 1994 - 30 June, 1995 Personnel: G.R Craig, Colorado Division of Wildlife ABSTRACT Bald eagles occupied 21 Colorado nesting territories in 1995. One new territory was discovered and lne that had been vacant since 1979 was reoccupied. Fourteen territories hatched young and 13 pairs fledged 23 young. Productivity averaged 1.01 young per occupied territory. This Job Progress Report represents a preliminary analysis and is subject to change. For this reasoJ, information ,presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the author . .,;. . ," " ��49 BALD EAGLE NEST SITE PROTECTION AND ENHANCEMENT PROGRAM Gerald R Craig SEGMENT OBJECTIVES 1. Monitor nest site occupancy and reproductive success. 2. Document survival rates and mortality factors. 3. .Determine migration and wintering areas. 4. Determine if philopatry occurs in breeding eagles. 5. 6. 7. Document pesticide contaminationthrough eggshell measurement and chemical analysis of nonviable eggs. 8. Where necessary, implement actions to stabilize nests and maintain occupancy. METHODS AND MATERIALS This work. will be a cooperative endeavor between the Division and Dr. Richard Knight of Colorado State University. 1. Annually visit all documented 'breeding sites to determine the presence of bald eagles. Pairs fit territories will be documented by DWMs and other field personnel. Previously unrecorded pairs willjprobably be revealed in the course of aerial eagle and waterfowl flights. DWMs will confirm actual incubation from ground visits. 2. Occupied territories will be visited by DWMs periodically throughout the breeding season to determine hatch of young, nesting failures, etc. 3. In May and June, a Utility Worker will observe breeding eagles from a distance and endea or to follow their movements to locate important foraging areas. Responses of eagles to various human activities and land uses will be recorded. 4. In June, when the young are determinedto be old enough to band, sites will be visited by Craig and Knight to place a federal band on one leg and a colored, alpha numeric marker on the other. The color markers will permit identificationif the young return in subsequent years. During the same nest visit the fdUowing will be recorded: Physical parameters such as tree species, height, DBH, condition, and dominance. Nest condition, size, and location. Vegetative community and land use practices. In addition, collect prey remains, nonviable eggs and eggshell fragments. ';' �50 5. ~pproximatelY 5cc's of blood will be collected from each nestling. The blood will be analyzed at the SjavannahRiver Ecology Lab in Aiken, South Carolina. Electrophoretic examination will permit genetic comparison with samples collected from other populations in Saskatchewan, the Lake States and Arizona, well as determine the heterogeneity of the Colorado birds. i 6. ~ necessary, remedial actions will be taken to stabilize nests that are threatened by wind throw. Should ilie tree be decadent and in danger offalling, an artificial nest base may be placed in a suitable, adjacent tt;ee. Action will be taken only after it has been deemed desirable to encourage the eagles to nest at the "" location. RESULTS AND DISCUSSION Territruy OCCYPanCY Bald eagle nesting activitiesfor Colorado are summarizedin Table 1. In 1995,21 territories were occupied (Adams, Archuleta, Fremont, Jefferson, Grand, Gunnison, La Plata #1 and #3, Mesa #2, Mineral, Moffat #1, #2, #3 and #4, MonteZUlllfl#3, Morgan, Rio Blanco #3, #4, and #5, Routt, and Weld #3). One new territory (Moffat #5) was added in 1995. 1jhenest size and location suggests that this was a first nesting attempt. Two previously vacant sites were reoccupied (Grand and Fremont). The Fremont site was vacant the previous year and the Grand site was last occupied iJ 1979. Although the Routt site was reported as inactive in 1994, the pair was relocated in a new nest upstream drthe 1994 nest. It is possible that the pair had relocated to it in 1994 and a goose occupied the vacated nest that y~. The pair at Rio Blanco #4 occupied the territory, but did not appear to construct a nest to replace the one that slipped out of the tree in 1994. Land Status The new territory at Moffat #5 is on private property and livestock grazing is the primary land use. The Grand site is administered by the Forest Service and water related recreational activities dominate the land use. I .. Reproduction Reproductive efforts are summarized in Table 2. Twenty-six young were hatched by 14 pairs and 23were fledged by 13 pairs (1.77 young per successful pair) which yielded an overall productivity of 1.01 young per territory occupying pair.' Seven pairs either failed to prq<Juceeggs, or failed during incubation. Eight young were banded and color marked at 3 locations (Adams, La Plata Co. #3, Moffat #2 &3, and Morgan) Fish and "fildlife Service bands were affixed to the nestlings' right legs and red alpha-numeric bands with yellow vinyl flags rere affixed to their left legs. Culmen length and foot pad length measurements were obtained from the eaglets that were banded. Banded Adults Both adulJ at the Routt site were color marked. The male's color marker tag was still affixed while the female's had detached. However, the female's anodized band with the alpha numeric code was visible. The male had been produced at the Moffat #2 site in 1990 and the female had fledged from the same site in 1989. The Routt site is approximately ., 30 miles east of their natal nest. I .;" �51 The banded adult female at the Craig #2 site was present in 1995. This individual was first documented at this site in 1991. Although the band prefix was obscured, the following digits were visible 3(?)4402. In 1 5, the 402 digits were again read. Nest Stabilization Efforts No nest stabilization efforts were undertaken in 1995. The nest at Rio Blanco Co. #5 nest was Constructed using an old heron nest as a base. The nest blew down killing the single young in it. Since this was a first nesting effort, the pair will be monitored in the future to establish their nest preference. If necessary, an artificial nest base will be installed The Rio Blanco #3 nest tree continued to decay sufficiently that it was deemed too hazardous to attempt to climb. This year, it was noted that the eagles were frequenting an artificial nest that had been constructed in an adjacent tree a number of years previously. Several branches were trimmed to allow better access by the eagles. The Gunnison site slipped out of the tree overwinter. In 1994, it had bee judged to be too unstable tolclimb. The. pair relocated to a large nest in an adjacent tree, but failed to produce young. Should they return in subsequent years, it appears that no stabilization will be necessary. I The Adams site nest tree has completely died. Although it is inundated for at least half of the year, there is a proposal to REA attempt to shore up the tree by driving treated power poles into the earth on either side and anchoring it in the effort to keep it upright. The nest that the Adams pair constructed in 1994 slipped out sometime during the winter which validated the decision not to climb the nest that year. Prepared by: _ Gerald R Craig Life Science Researcher IV �U1 I\) Table 1. Bald Eagle Nesting Efforts in Colorado 1974' 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 1986 1987 1988 1989 Site legg IA IA IA IA IA ? ? ? ? ? ? eggs 2yng lyng 2yng La Plata Co. #1 1yng 2yng 2yng 2yng 1yng -- 2yng 2yng -- 2yng Oyng 1yng 2yng 1yng 3yng Moffat Co. #1 2yng 2yng 2yng Oyng IA IA IA IA IA IA IA ,A IA IA La Plata Co. #2 Oyng Oyng IA IA IA IA IA IA IA IA IA IA Grand Co. A A A A IA IA IA IA IA IA Montezuma Co. #1 1yng lyng ? eggs Oyng 2yng 2yng 2yng 2yng Rio Blanco Co. #1 3yng 2yng 2yng 2yng Oyng 1yng A 2yng Rio Blanco Co. #3 2yng 2yng eggs IA IA IA Weld Co. #1 2yng 1yng 1yng 1yng 1yng Montezuma Co. #2 lyng Oyng 2yng 3yng Moffat Co. #2 legg IA ? eggs Moffat Co. #3 eggs legg eggs 2yng Adams Co. eggs 2yng IA IA Archuleta Co. 1yng 1yng Montezuma Co. #3 eggs Weld Co. #2 La Plata Co. #3 Rio Blanco Co. #4 Morgan Co. #1 Mesa Co. #1 Fremont Co. Routt Co. Gunnison Co. Mineral Co. Weld Co. #3 Montezuma Co. #4 Jefferson Co. Mesa Co. #2 Moffat Co. #4 o '1 4 4 4 1 0 3 6 2 6 6 5 10 8 16 Total Young 1 1 2 2 3 3 1 3 4 2 4 5 10 9 8 10 .Tcte l Pairs Young/Occ. Terr. 0.00 1.00 2.00 2.00 1.33 0.33 0.00 1.00 1.50 1.00 1.50 1.20 0.50 1.11 1.00'1.60 A = Active IA = Inactive ~' 1990 Oyng 2yng IA IA IA 2yng 1yng Oyng 1yng 2yng IA 2yng IA 1yng IA 2yng 2yng 1991 1yng 2yng IA IA IA 2yng 3yng IA Oyng 2yng Oyng 3yng Oyng 2yng IA 2yng 1yng 2yng Oyng 1992 1yng 2yng IA IA IA 1yng 3yng IA lyng 3yng Oyng 3yng Oyng lyng IA ? 2yng 3yng eggs 13 19 20 10 13 14. 1.30 1.46 1.43 1993 2yng 2yng IA IA IA 2yng 2yng IA IA Oyng 2yng 2yng eggs 2yng IA 1994 1995 3yng IA 2yng 2yng IA IA IA 0 yng IA 2 yng 2yng 0 yng lyng 2 yng IA IA IA IA Oyng 3 yng 2yng 1 yng 3yng 3 yng IA 1 yng lyng 2 yng IA IA ? IA 1 yng 3yng Oyng IA 2yng Oyng 3 yng eggs Oyng IA 2yng IA A 2yng IA: 2yng lyng 2yr'1g0 yng A ? 0 yng A OYr:lgIA o IA: IA o Oyng Oyng 2ypg 2yng 1Yl')g1 yng 18 16 22 18 16' 21 1.00 1.00 1.05 �53 Table 2. Colorado Bald Eagle Nesting Efforts - 1995 Site AglilofBirds Male Female Adams Co. #1 Archuleta Co. Fremont Co. Jefferson Co. Grand Co; Gunnison Co. La Plata Co. # I La Plata Co. #3 Mesa Co. #2 Mineral Co. # 1 Moffat Co. # 1 Moffat Co. #2 Moffat Co. #3 Moffat Co. #4 Montezuma Co. #3 Morgan Co. Rio Blanco Co. #3 Rio Blanco Co. #4 Rio Blanco Co. #5 Routt Co. Weld Co. #3 Adult Adutl Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Total 21 Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult 21 Young Produced Young Fledged 3 2 0 0 1 0 0 3 1 0 0 Comments Pair in brooding posture late in season. Pair incubated, failed. 1 0 0 1 1 2 0 2 3 1 2 2 3 2 0 1 2 0 2 0 2 3 1 1 2 3 2 0 0 2 0 26 23 Nest fell overwinter, switched to adjacent nest Pair either failed early or did not nest. Pair built nest, failed. Pair present, failed to nest. 1st nest effort, nest blew out killing young. Pair disturbed at egg laying. .•. � https://cpw.cvlcollections.org/files/original/64a1ccc2a1d420e695963156bac39acd.pdf 2b96dee8d48202dbf464c5c35578cf45 PDF Text Text 1 JOB PROGRESS REPORT State of _--,C"",o""lo"..ra...,d.."o,--_ Project:_~vv~-1~5~Q-~R~-8~_ Work PIan _2_ : Job Job Title: Pere~e ENDANGERED \VILDLlFE INVESTIGATIONS _j_ Falcon Restoration Program Period Covered: 1 July, 1995 - 30 June, 1996 Personnel: G.R. Craig and C. Wigblman. Colorado Division of Wildlife and J.H. Enderson, The Colorado College. ABSTRACT In, the 1996 peregrine breeding season, 83 territories were occupied by 78 breeding pairs that fledged 140 young. Productivity averaged 1.65 young fledged for those pairs that were monitored. Contents of 6 nonviable eggs were collected and have been preserved for future analysis. Shell fragments were collected at 18 sites and are awaiting measurement. This. Job Progress Report reptesents a preliminary analysis and is subject to change. For this reason, infoImation presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the author. ��3 PEREGRlNE FALCON RESTORATION PROGRAM Gerald R. Craig SEGMENT OBlECTNES 1. Annually monitor the munber of breeding pairs of peregrines and their reproductive success in Colorado. 2. Annually monitor organochlorine pesticide levels in wild breeding peregrines in Colorado. 3. Monitor breeding population turnover through band recoveries, presence of color markers, and telephotographic identification of individual breeding adults. 4. Augment poor wild production by placement of captive hatched wild young and captive produced young into occupied wild nests. 5. Release captive hatched and captive produced young at potential and vacant wild territories. 6. Monitor recruitment of reintroduced peregrines into the wild breeding population of .Colorado. METHODS 1. Visit all known peregrine breeding territories tbroughoot Colorado and observe them from a distance to establish the presence of breeding adults. Breeding pairs will be kept under surveillance to determine initiation of egg laying. Depending upon the individual female's reproductive history and eggshell condition (obtained through measurement of previous year's eggshell thicknesses) and availability of captive hatched young for release, breeding pairs either will monitored or manipulated as outlined in approach 4. Those pairs not designated to be manipulated will be revisited periodically throughout the nesting season to document reproductive success. When a pair's behavior indicates that egg laying has occurred and incubation is underway, the eyrie will be visited to document the munber of eggs produced. The eggs will be candled to ascertain viability and approximate age. All nonviable eggs will be collected for chemical analysis. A second visit will be made to determine productivity, band nestlings, and collect eggshell fragments and unhatched eggs for thickness measurement and analysis under 2a and 2b. 2a. Eggshell fragments encountered during eyrie visits described in approaches 1 and 4a will be measured for index to thickness following standardized procedures. 2b. Whole, nonviable eggs which are encountered during eyrie visits will be collected, preserved and submitted to the appropriate Fish and Wildlife Service approved laboratory for pesticide analysis. Eggs collected from the wikl in the course of Approaches 4a, 4b and 4c that are artificially at The Peregrine Fund's Boise, Idaho facility also will be submitted for shell thickness measurement and chemical analysis. 3. Peregrines present at breeding territories will be examined to determine the presence of bands or color markers. Band confirmation will be accomplished through observation from a distance with telescopes and concealed remote controlled cameras. When banded falcons are encountered, every effort will be made to read band mnnbers without trapping or handling the birds. It is possible this can be accomplished in most situations with a Questar field model telescope (SO-130x). When band munbers cannot be discerned, attempts will be made to trap and examine the falcon at a time when capture will have least impact upon breeding activities. 4a. In accordance with an ammal release plan developed and approved by the State, U.S. Fish and Wildlife Service, Bureau of Land Management, National Park Service, and the Forest Service, a predetermined munber of wild breeding pairs will be manipulated to augment natural productivity. Pairs with a history of reduced clutch size, cracked eggs, or infertile or dead eggs will be candidates for fostering efforts. �6 Fig. 3. PERCENT OF TOTAL BREEDING PAIRS mAT WERE SUCCESSFUL 1000/0- '.'i; 80% -- - - - -- - - - -: .... ::. ,"; 86 87 ------ ~ 60%---------- :: ~ <n E- 400/0 -- - - - -- --- ~ c... 74 75 76 77 78 79 80 81 82 83 84 85 88 89 90 91 92 93 94 95 96 YEARS I<-~,·.;:.<·.I Successf"ul Pairs Fig. 4. PRODUCTIVITY 2.5 - ~ U> 2 - ------------------------------------- i\ Q <£) ~ 1.5 u - -----1-- ••.------------: u o ~ <xl P-. \ ---- ( 1 - C> Z ::::> :2 0.5 o ---f----- ------------------------------------------------------------------------------------ --i--------~·'-8-/--\b~-p I;tl ---.~R.-------------~./-------- ---- -. ----- ----. ------ ------------ -------------- -L,---r--~_.--._--._~--.__9~4r~~_.--.__.--_.--~_.--.__.r__.--~_.--~--r_ -, 73 74 75 76 77 78 __ , ," 79 80 81 82 Total Pairs 83 84 85 YEARS 86 --E3-- 87 88 89 90 91 92 93 94 95 96 Unmanipulated Pairs Eggshell Condition Six whole, nonviable eggs were encountered in the course of visits to 5 sites. These eggs were preserved whole for . contaminant analysis and shell thicknesses are not available at this time. Eggshell fragments were also collected from 18 additional nesting sites in the course of visits to band young. Thickness measurements have not been compiled for these samples at this time, so no conclusions can be made about the 1996 thicknesses but eggshell values are available through 1995 (Fig. 5). These values are highly variable due to small sample sizes, mixing of fragments from different eggs within the same clutch, and variation of thickness of fragments from the poles of the egg versus the waist. However, there appears to be a slight trend toward thicker eggs over the past 8 years with thickness averages less than 10% thin �7 since 1992. Greater variability among eggs is also evident since 1988 with some eggs being thicker than pre-DDT era eggs and others 16% thinner. Fig. 5. CUMULATIVE EGGSHELL TIllCKNESSES 0.39 0.37 ]: 0.35 ~ 0.33 ~ = 0.31 j -.-------------------------------------------------------- -------------- _~~_~_..~~~:'~~~_:~~~.:'_"_s_t-_ -----. ------- --- - 0.29 0.27 0.25 73 74 75 I 76 77 78 79 80 81 82 83 &4 85 86 87 82 89 90 91 92 93 94 95 Years MaxttnU%n and Minizn.u= Thickne"""" ~ Average Thickne""",,,' Qr~anochlorine Residue in E~~s The 6 nonviable eggs collected during the 1996 season were preserved along with eggs encountered in 1991-94. This collection of 47 eggs is awaiting pesticide analysis by the Fish and Wildlife Service when funding is available. Release and Au~ntation Efforts Remedial management efforts such as fostering or hacking have not been undertaken since 1989. Prepared By:___".C",--,,__._f{_f'..;:::;___;;_ },/II-tf [: __ Gera1dR.~ Life Science Researcher IV ��9 JOB PROGRESS REPORT State of Project: Work Plan ~C~o~l~o~r~a~d~o~ _ W-164-R-2 1 Laboratory Investigations Job l Job Title: DNA ANALYSIS FOR SPECIES AND SEX IDENTIFICATION. AND TISSUE MATCHING OF BLOOD. BLOOD STAINS AND MEAT SAMPLES FOR COLORADO'S FORENSIC CASES. Period Covered: 1 July 1995 30 June 1996. Personnel: W.J. Adrian, R.P. Ellis, and R.J. Magnuson. ABSTRACT Research was initiated to develop the ability in the .CDOW Wildlife Forensics Laboratory to "DNA fingerprint" biological speCimens-and to determine whether such specimens are from the same animal (identical DNA fingerprint) or different animals (nonidentical DNA fingerprint). In many instances each year, it is necessary to establish such identity or nonidentity in suspected illegal hunting activities. In some cases, physical examination and analysis of the evidence is sufficient to establish identity or nonidentity. In instances where physical techniques are not successful, DNA fingerprinting is the method of choice. Specimens from elk and grizzly bear were collected and stored at -20C until DNA extractions were performed. Each specimen included in the development of the DNA fingerprint technique was accurately identified as to species, age, gender, and location of animal when the specimen was collected. Small portions of tissue were extracted and the resulting genomic DNA was purified and quantified spectrophotometrically. Purified genomic DNA, within the recommended DNA concentration range, was selected for further analysis. We now have the capability to conduct DNA fingerprinting of game animals. ��11 SEGMENT OBJECTIVES in DNA-based 1. Obtain equipment, supplies, primers, and probes necessary technologies. 2. Use the DNA-based probes species, and individual identification. to assay game animal tissue for use for sex, 3. To place on-line, at Colorado State University, Department of Microbiology, a laboratory for routine testing of game animal tissue samples for gender determinations, population analysis, and wildlife law enforcement. METHODS This work is a cooperative endeavor between the Colorado Division Dr. Robert P. Ellis of Colorado State University. of Wildlife and Wildlife forensics using DNA assays have been ana Lyz ed by several investigators. These include assays using PCR primers (short DNA sequences which initiate DNA amplification) for sex determination of deer, antelope, and other mammals; species specific RFLP (restriction fragment length polymorphism) probes for discrimination of antelope, elk, and mule deer; mitochondrial DNA probes to distinguish mule and white-tailed deer (Cronin 1986, Blackett and Keim 1992, Menke 1992). Other commercial probes by AGTC (Analytical Genetic Testing Center, Inc.) are available for possible use in big game DNA analysis. Non-commercial probes are already available for use in objective 2. In November 1992, two papers by J. LeMay, S. Fain, and J. presented at the Northwest Association of Forensic Scientists Fall Portland, Oregon on the use of DNA assays for game animals (LeMay and Ruth 1992). These papers demonstrated successful development and use probes to identify, from meat and blood, the gender of most ruminant well as routine "individualization" of white-tailed deer. Ruth were Meeting in Fain 1992, of the DNA species as Development of individual-specific RFLP banding patterns from elk genomic DNA using a non-radioactive protocol has allowed for the elimination of radioisotope handling in DNA analysis (Stern 1992). These patterns arise by a conjugated antibody-enzyme mediated reaction. The use of non-radioactively labeled probes allow for reduction of safety concerns associated with radioactively labeled probes, easier di~osal of used probes and longer probe storage life. The non-isotopically labeled probes have at least equal resolution of the radioactive banding patterns. Although previous work has demonstrated adequate polymorphism using RFLP analysis, many forensic samples contain insufficient DNA to use RFLP methodology. �12 The third objective was to use RAPD-PCR or similar techniques in differentiating between individuals of the same species. This technique would be used as a "fingerprinting" scheme to determine the identity of tissues collected from the same animal. Likewise, tissues collected from different animals of the same species would have a different DNA "fingerprint" (Orita et al. 1989, Hayaski 1991, Kambhampati et a1. 1991, Welsh and McClelland 1991). In total, we would be able, in a matter of a few days, to produce irrefutable evidence of gender, species, and individual origin of tissue samples. RESULTS AND DISCUSSION Research was initiated to develop the ability in our Wildlife Forensics Laboratory to "DNA fingerprint" biological specimens and to determine whether such specimens are from the same animal (identical DNA fingerprint) or different animals (nonidentical DNA fingerprint). In many in~tances each year, it is necessary to establish such identity or nonidentity in;suspected illegal hunting activities. In some cases, physical examination and analysis of the evidence is sufficient to establish identity or nonidentity. In instances where physical techniques are not successful, DNA fingerprinting is the method of choice. DNA fingerprinting techniques are currently in use at the National Wildlife Forensics Laboratory (NWFL) at Ashland, Oregon. A workshop on DNA fingerprinting was presented by personnel at the NWFL in conjunction with the Northwestern Regional Forensics Meeting in October 1995. At this workshop instruction was presented regarding most aspects of DNA fingerprinting procedures. Specimens, from elk and grizzly bear, were collected and stored at -20C until DNA extractions were performed. Each specimen included in the development of our DNA fingerprint technique was accurately identified as to species, age, gender, and location of animal when the specimen was collected. Small portions of tissue were extracted and the resulting genomic DNA was purified and quantified spectrophotometrically. Purified genomic DNA, within the recommended DNA concentration range, was selected for further analysis. Each genomic DNA specimen was digested with the restriction enzyme Hinf I to yield different sized fragments of DNA. These fragments were electrophoresed to separate the fragments by size in base pairs. Intermediate sized fragments, from 500 - 2000bp, were blotted onto nitrocellulose membranes. The blots were immobilized so that they would not elute from the nitrocellulose membrane. Two different alkaline phosphatase-labeled DNA oligonucleotide probes were used to obtain a "DNA fingerprint" for each of the specimens with each probe. Alkaline phosphatase labeling replaces radioactive labeling and is not radioactively nor biologically hazardous. The commercially available probes used were the 33.15 and 33.6 probes described by Jefferys. Following annealing of the probes to their complementary sequences, the membranes were rinsed thoroughly to remove nonannealed probe and then placed next to X-ray film in X-ray film cassettes for development to visualize the positions where each probe annealed, and thus visualize the DNA fingerprint for that individual specimen as defined for each probe. Our results indicated that we were successful in differentiating individuals by the above techniques. Clearly discernable probe positional differences and, thus, different DNA fingerprints, were visualized for the elk �13 and grizzly bear specimens processed in our research. When the technique described above is applied to forensic analysis, more than one restriction enzyme should be used, so that fragments of DNA cut at different restriction sites are obtained. In addition to increasing sensitivity by using more than one restriction enzyme, more than two probes would be used. Commonly, 3 to 5 probes would be used, again increasing sensitivity and discriminatory power. For the DNA fingerprinting technique to be used for evidence analysis and presentation, a minimum of 25 specimens of each species to be included in our DNA fingerprint database should be collected, cataloged, and processed to establish the odds of two different animals of the same species having the same DNA fingerprint. In Colorado, I suggest that the species include Elk, Mule Deer, White-tailed Deer, Moose, Bighorn Sheep, Pronghorn, and Rocky Mountain Goat. Such analysis by the NWFL has indicated that the probability of the same fingerprint originating from two different animals of the same species is less than 1 chance in 1 million. Prepared �14 LITERATURE Blackett, R.S., deoxyribonucleic CITED and P. Keim 1992. Big game species identification (DNA) probes. Jour. Forensic Sci. 37:590-596. by Cronin, M.A. 1986. Genetic relationships between white-tailed deer, mule deer, and other large mammals inferred from mitochondrial DNA analysis. M.S. thesis, Montana State Univ., Bozeman. Hayaski, K. 1991. PCR-SSCP: a simple and sensitive method for detection of mutations in the genomic DNA in PCR methods and applications, Cold Spring Harbor LaB. Press, pp.34-38. Kambhampati, S., W.C. Black, IV, and K.S. Rai. 1991. RAPD-PCR for identification and differentiation of mosquito species and populations techniques and statistical analysis. J. Med. Entomol. 29:939-945. . LeMay, J.P. and S.R. Fain 1992. Gender determination of mammalian species from PCR amplified sex-linked genes. Forensic Scientist·Fall Portland, OR. wildlife Meeting. Menke, S.D. 1992. Protein synthesis during development of normal and mutant flight muscle in Drosophila melanogaster and isolation and characterization of sex specific DNA in deer and antelope. Ph.D. thesis, Univ. of Wyoming, Laramie. Orita, M., H. Iwahana, H. Kanayawa. K. Hayaski and T. Sekiya. 1989. Detection of polymorphisms of human DNA gel electrophoresis as single-strand conformation polymorphisms. Proc. Natl. Acad. Sci. USA. 86:2766-2770. Ruth, J.L. 1992. The forensic identification of individual deer using DNA probes. Forensic Scientist Fall Meeting. Portland, OR. Stern, C.M. 1992. Non-isotopic Fort Collins. DNA fingerprinting of elk. Colorado State Univ., Welsh, J., and M. McClelland. 1991. Genomic fingerprinting using primed PCR and a matrix of pairwise combinations of primers. Nucleic Aci~s Res. 19:5275-5279. �15 JOB PROGRESS REPORT State of __~C~o~l~o~r~a~d~o~ Project Work Plan Job Title: W-166-R-5 ___1 : Job Migratory Game Bird Investigations 22 Harvest distribution of mallards and pintails banded preseason in western Colorado Period Covered: Author: _ 01 April 1995 through 31 March 1996 Michael R. Szymczak Personnel: J. Gamble and staff, Brown's Park National Wildlife Refuge; J. Broderick, R. Caskey, D. Coven: P. Creeden, R. Del Piccolo, J. Ellenberger, V. Graham, J. Gray, J. Gumber, T. Mathieson, J. Miller, J. alterman, R. alterman, N. Smith, M. Szymczak, K. Wagner, S. Wait, and S. Yamashita, C~lorado Division of Wildlife ABSTRACT Ducks were trapped in modified Salt Plains bait traps and banded on 14 different wetlands in 4 areas across western Colorado in August and September 1995. About 1,850 mallards (Anas platyrhnchos) were banded, with the number captured generally well distributed between trapping areas. Only 24 northern pintail (Anas acuta) were banded in 1995. ��17 HARVEST DISTRIBUTION OF MALLARDS AND PINTAILS BANDED PRESEASON IN WESTERN COLORADO P. N. OBJECTIVE 1. Document the distribution of band recoveries captured preseason in western Colorado. of mallards and pintails 2. Determine if the geographic location of recovery of mallards pintails is dependent on the area of banding in Colorado. 3. Determine the relationship between the recovery distribution of western Colorado banded mallards and pintails and the distribution of recovery of those species banded ~n other areas of the Pacific Flyway. 4. Cooperate in analysis of Pacific Flyway-wide preparation of reports. band recovery and data and SEGMENT OBJECTIVES 1. Trap and band mallards and pintail in 4-5 areas of western Colorado in late August-early September using salt plains bait traps (Szymczak and Corey 1976). Recommended areas are: (1) Browns' Park National Wildlife Refuge, (2) Yampa River Valley below Craig, (3) Colorado River Valley below Glenwood Springs, (5) Uncompahgre-lower Gunnison River Valley, and (6) the Cortez - Mancos area. 2. Submit banding schedules and recapture reports to the U. S. Fish and Wildlife Services' Bird Banding Laboratory. Summarize and file band return reports. 3. Contribute Alberta. manpower and equipment to cooperative duck banding crews in INTRODUCTION In 1990, the Pacific Flyway Study Committee formulated a 5-year cooperative mallard and pintail preseason banding program that was endorsed by the Pacific Flyway Council. This program was designed to address banding needs throughout the western U. S., including Alaska, and in the provinces of British Columbia and Alberta. Through the first 4 years, about 7,000 ducks were banded under this program. This report covers the fifth year of banding during the preseason period in western Colorado. Background information can be found in Szymczak (1992). �18 METHODS Trap Area Selection Most breeding mallards in western Colorado are associated with small wetlands that are widely distributed throughout high elevation, mountainous areas. Only in Browns' Park, in the extreme NY corner of Colorado, are there extensive wetland complexes capable of supporting breeding ducks. Trapping on widely distributed wetlands with low densities of breeding ducks ,was assumed to be an inefficient method for banding western Colorado mallards. Therefore, strategically located areas, to which post-breeding and fledged Colorado mallards would move, were selected as primary trapping areas. In 1995, trapping occurred on the Browns' Park National Wildlife Refuge (BPNWR) , along the Colorado River Valley (CRV) from Debeque to near Fruita, in the Uncomphagre River Valley (URV) ,from Montrose to Delta including some locations east of Delta in the Gunnison River Valley, and in th~ Cortez-Mancos area (CM). Trapping Period Ducks were trapped about 10 days beginning banding periods in 1995 thru 8 September; URV - Trapping and Recording and banded in each area for a consecutive period of on 18 August and ending on 18 September. The actual were: BPNWR - 18 thru 9 September; CRV - 31 August 27 August thru 6 September; CM - 2 thru 11 September. Technique All birds were trapped in modified Salt Plains bait traps (Szymczak and Corey 1976) using whole shelled corn for bait. Traps were visited daily. Mallards and pintails were the target species, but green-winged teal (Anas carolinensis) were also banded in all areas, blue-winged teal (Anas discors) and/or cinnamon teal (Anas cyanoptera) were banded in Browns' Park and along the lower Colorado River, and gadwall (A. strepera), redheads (Aythya americana) and canvasback (Aythya valisineria) were banded in Browns' Park. Banded birds were recorded by wetland site. Band numbers of all birds captured that were banded in previous years or outside the specific area of trapping were recorded. Records were also maintained in some areas on the number of traps operated by wetland to evaluate capture/unit of effort at each trap site. Alberta Banding A volunteer was not obtained from the Colorado Division of Wildlife's permanent staff to work on a cooperative duck banding crew in southern Alberta. In lieu of obtaining a volunteer, $2,600 was forwarded to the Pacific Flyway banding coordinator through The Wildlife Management Institude to hire temporary personnel for banding in southern Alberta. �19 RESULTS Trap Locations Trapping was distributed over a total of 14 different wetlands in the 4 areas (Table 1). Hog Lake (BPNWR) and Hall Pond (URV) were reinstated as a trap sites in 1995, while Fravert Pond (CRV), Sanders' Pond (URV) and both trap sites near Yampa were dropped. Nearly all trap sites were in Palustrine Emergent Persistent Wetlands, but some sites in the Colorado River Valley were in Riverine Upper Perennial Rock Bottom wetlands (Cowardin et al. 1979). Banding and trapping efficiency About 1,850 mallards were banded during trapping.in western Colorado in 1995 (Table 2) bringing the 5-year total to slightly ~ver 9,000 mallards. The number of mallards banded increased in Browns' Park and in the URV but decreased in CRV and CM. The number of immatures in the banded ~ample on the BPNWR remained comparatively low at 24%. However, immature mallards comprised 74, 71, and 74% of the CRV, URV, and CM samples, respectively. Throughout the trapping area, 63.9% of the mallards banded we!e immatures compared to 64.8% in 1994, 61.9% in 1993, 62.6% in 1992, and '68.3% in 1991. Only 25 northern pintail, the secondary target species, were banded (Table 3). Addition species banded were green-winged teal, redhead, gadwall, canvasback, and blue-winged teal and/or cinnamon teal (Table 3). Band Reporting and Record Keeping All new banded birds and recaptures were submitted to the U. S. Fish and Wildlife Service's Bird Banding Laboratory on standard forms. Computer files containing the number of birds banded by area, site, day, age, and sex were constructed at the Colorado Division of Wildlife's Research Center. �20 Table 1. Trapping locations during preseason banding Colorado. August- September 1995. in western Wetland Area Browns' Park Natl. Wildl. Ref. Colorado R. Valley Name Location Flynn Marsh TlON, Rl03W, Sec 16, SE~ Spitzie Slough TlON, Rl03W, Sec 15, S~' Hog Lake TlON, Rl03W, Sec 9, SW~ Latham's Slough Morse's 'Pond TIS, RlE, S~c 34, NE~ Ute Meridian Walker Wildl Area North Uncompahgre Cortez/Mancos R. Valley Markley's T8S, R97W, Sec 27, S~ Pond TIN, R2W, Sec 36, SW~ Ute Meridian T50N, R9W, Sec 30, NW~ Sweitzer Lake T15S, R95W, Sec 28, S~ Hall Pond T14S, R94W, Sec 21, SE~ Totten Res. T36N, R15W, Sec 20, NW~ Weber Res. T36N, R13W, Sec 12, NE~ Summit Lake T37N, R14W, Sec 33, SW~ Williamson's Nolan's Pond Pond T36N, R15W, Sec 3l 4 SE~ T37N, R16W, Sec 8, N~ �21 Table 2. trapping Area Brown's Park Colorado River Uncomp. River CortezMancos TOTAL Number in of mallards banded by area, site, age, and sex during Colorado , 1995 western Site Flynn Marsh Hog Lake Spitzie Slough Subtotal N. Walker Morse's Pond Latham Slough Subtotal Markley's Sweitzer Lake Hall Pond Subtotal Nolan's Pond Totten Res. Weber Res. Summit Lake Williamson's Pd. Subtotal AM AF 137 47 _l Agelsex IF Totals 1M 49 36 28 24 11 7 _l _Q 185 74 39 56 11 17 5 90 34 .i _.2. _1]_ 74 31 151 83 54 26 30 166 90 250 89 Q _1 43 341 94 21 40 155, 257 71 __§J_ 448 220 48716 411 .: -2 _n. 144 61 278 17j 46 14 233 13 27 2 0 16 58 17 7 1 2 14 41 29 67 24 8 43 171 25 52 7 4 21 109 84 153 34 14 94 379 461 207 639 540 1847 pre-season �22 Table 3. Number of northern and sex in western Colorado, Species Northern pintail Area pintail and other species banded by area, site, age, preseason 1995. Site AM AF Latham Slough 0 1 0 0 1 Uncomp. River Markley's Pd. Sweitzer L. Hall Pd. 6 1 1 4 0 0 7 1 0 2 0 0 19 2 1 _Q Q 1 1 _2. 8 5 9 3 25 0 0 0 2 1 2 I Flynn Marsh Total Brown's Park Hog Lake Flynn Marsh 1 0 '0 0 Colo. River Walker Wildl. Latham Slough Morse's Pd 0 1 0 0 1 1 0 0 0 0 0 1 2 1 Uncomp. River Hall Pond Sweitzer L Markley's Pd. 46 0 10 14 3 2 7 3 20 3 3 6 70 9 38 CortezMancos Totten Res. Weber Williamson Pd. 1 2 19 0 0 0 0 2 3 _1 1 1 15 _..§. _!!..2. 80 24 47 23 174 1 Total Blue-winged/ Cinn. Teal Total Colo. R. Brown's Park Green-winged Teal AgeLSex IF 1M - Brown's Park Flynn Marsh Hog Lake 0 1 0 0 I 0 0 1 2 Colo. River Morse Pd. 1 Q 1 Q 2. 2 0 3 0 5 Total Redhead Brown's Park Hog Lake 0 2 9 25 36 Gadwall Brown's Park Hog Lake 0 0 0 I I Hog Lake 0 0 I 0 I Canvasback Brown's �23 LITERATURE CITED Szymczak, M. R. 1992. Harvest distribution of mallards and pintails banded preseason in western Colorado. Job Prog. Rep., Colorado Div. Wildl., Oct. 49-53. Pp. ________ , and J. F. Corey. 1976. Construction and use of the Salt Plains duck trap in Colorado. Colorado Div. Wildl., Div. Rep. 6. l3pp. Prepared by: Michael R. Szymczak LSSR IV ��25 JOB PROGRESS State of Colorado Project W-166-R-5 Work Plan 1__ Job Title: Author: : Job Integrated Period Covered: Migratory REPORT Game Bird Investigations 24 Waterbird 1 April Manage~ent 1995 through Studies 31 March 1996 James H. Gammonley Personnel: J. H. Gammonley, J. K. Ringelman, M. R. Szymczak, A. Callinan, R. Dieboll, M. Phipps, A. polansky, B. st. George, R. Sanders, Colorado Division of Wildlife; M. K. Laubhan, National Biological Service. ABSTRACT We studied foraging and nesting ecology of breeding waterbirds from 19 April to 4 July 1995 at Russell Lakes state Wildlife Area (RLSWA). During 4 15-day sampling periods, we measured habitat variables (water gepth, conductivity, and temperature; vegetation height, density, and species composition) in ~30 randomly-located plots within short emergent (SE), tall emergent (TE), shallow open water (SW), semipermanent open water (SPOW), saltgrass (SG), and upland shrub (US) cover types. During each sampling period, we collected invertebrate and seed samples (n = 1,620) from benthic, water column, and vegetation substrates in SE, SG, SW, and SP~W plots, to examine temporal and habitat-related changes in food availability. We collected mallards (n = 18), redheads (30), cinnamon teal (30), American avocets (27), killdeer (15), and Wilson's phalaropes (22) for analysis of food habits and body condition in relation to reproductive status; food samples were also taken at the site where each bird was collected to compare food use to availability at the scale of individual foraging locations. We also collected time-activity data for each species, and measured habitat variables at sites where birds foraged during each time budget bout. We conducted nest searches on all habitat plots and in .other portions of RLSWA; habitat variables were measured at 268 nest sites and nests were revisited to determine nest fates. Processing of food samples and data entry continued from July 1995 through March 1996. Habitat measurements, time-activity data collections, line-transect counts, waterbird and invertebrate collections, and nest searches and monitoring will continue from April through July in 1996. ��27 INTEGRATED WATERBIRD MANAGEMENT STUDIES P. N. OBJECTIVES 1. Map the location of wetlands and wetland 2. Document the hydrologic regime and water, characteristics of each wetland type. 3. Identify the aquatic invertebrates associated with each wetland community, and document seasonal trends in invertebrate diversity, abundance and biomass. 4. Quantify the abundance, spatial and temporal use patterns, behaviors, and food habits of waterbirds in different wetland types. Relate the dynamics of endogenous lipid and protein reserves to food habits and migration and breeding e~ology. 5. Determine the seasonal wetland habitat requ Lxement s for all waterbirds, and consolidate these needs into a conceptual design for an optimum wetland community. 6. Determine the water management protocol and wetland development guidelines needed to produce the optimum wetland community. Prepare wetland development and water management plan for the RLSWA. SEGMENT 1a. communities on the RLSWA. soil and vegetation a OBJECTIVES Map cover types (short emergent, tall emergent, shallow open water, semi-permanent open water, saltgrass, and upland shrub) and other landscape features (roads, ditches, gravel pits, and parking areas) RLSWA, and digitize into a GIS format. at lb. Randomly overlay a grid representing O.2S-ha plots on the GIS map. Select a stratified random sample of plots in each cover type (total 330) • 2a. Randomly select a subset (n = 15) of plots in each cover type for invertebrate and seed sampling. During a one-week period, collect seed and invertebrate samples and determine density and biomass of food items at random sites in each plot. Collect 3 replicate~amples each from the benthos, water column, and vegetation at each site.. Repeat sampling every 3 weeks from April to July. 2b. During a 2-week period, use focal sampling (Altmann 1974, Tacha et al. 1985) to determine activities of mallards, redheads, cinnamon teal, American avocets, killdeer, and Wilson's phalaropes. Repeat sampling with 1-week breaks between sampling periods, 15 April to 5 July. 2c. During each 2-week sampling period, search each selected plot (lb) for waterbird nests. Map the location of each nest on aerial photos and return at the expected hatch date to determine nest success. Obtain GPS coordinates of each nest and enter on GIS map (la). 3. On the first and last day of each 2-week sampling period, use linetransect sampling (Buckland et al. 1993, Laake et al. 1994) to estimate densities of each species in 2b within each cover type. Fourteen northsouth transects spaced at 440-m intervals across RLSWA will be walked during each count; total transect length is 38.4 km. n �28 4. During each 2-week sampling period, determine food habits of species in 2b by collecting actively foraging birds and measuring the numbers and biomass of food in esophageal contents. Determine reproductive status (pre-laying, laying, incubating, post-breeding), molt intensity, and nutrient composition of each collected bird. Collect food samples from collection sites using procedures in 2a. 5. At collection sites (4), nest sites (2c), focal time-budget sites (2b), food sampling sites (2a), and random sites in each selected O.25-ha plot (lb), measure the following habitat variables, where appropriate, during each sampling period: water depth, conductivity, and temperature; cover type; plant species; and vegetation height and visual obstruction (Robel et ale 1970). Use habitat variable measures to estimate the availability of each cover type for a) foraging and b) nesting waterbirds of each species listed in 2b. 6a. Determine food selection by species listed in 2b by comparing (density, biomass) esophageal contents to food contents at collection sites (4), and at random sites in e~ch cover type (2a). 6b. Determine foraging habitat selection by species;listed in 2b by comparing habitat variables at collection sites and focal observation sites to habitat variables at random sites (5). Also co~pare food density and biomass between collection sites and random~sampling sites, and among cover types (2a and 4). 6c. Determine nesting habitat selection by species listed in 2b by comparing habitat variables at nest sites to habitat variables at random sites (5), and by comparing nest densities in each cover type to the availability of each cover type (2c and 5). Also compare nest success among cover types (2c), and in relation to habitat variables (2c and 5). INTRODUCTION The San Luis Valley (SLV) is one of the most important breeding areas for waterbirds in Colorado (Ryder et al. 1979, CDOW 1989, Nelson and Carter 1990, Gilbert et ale 1996). A wetland ecosystem can be managed for habitats that maximize requirements for a narrow group of avian species, or for more diverse habitats that optimize resources for a variety of avian species. The latter approach of integrated waterbird management better fits the philosophy of increased emphasis on managing landscapes for species diversity. One goal of the Colorado State Waterfowl Management Plan is to provide habitat of sufficient quality to maintain duck and goose populations at desired levels for maximum recreational opportunities (CDOW 1989). In addition, the SLY draft management plan for waterbirds recommends maintena~ce of diverse wetland habitats with 25% of the actively managed habitat on public lands managed for nongame waterbirds (Olterman 1993). In 1994, CDOW, in cooperation with NBS, initiated a study to examine resource use by both game and nongame waterbird species breeding at Russell Lakes State Wildlife Area (RLSWA). STUDY AREA We studied waterbird ecology at RLSWA, a wetland complex in the SLY in Saguache County. We categorized habitats (cover types) at RLSWA according to hydrology (flooding depth and duration) and vegetation structure as follows: short emergent (SE), tall emergent (TE), shallowly flooded open sites with no emergent vegetation (SW), semi-permanently flooded sites with no emergent vegetation (SP), saltgrass (SG), and upland shrub (US). We created a GIS map of RLSWA based on these cover types, using aerial photos (scale 1:4,000) taken on 5 May 1994 (cover type designations were ground-truthed during summer of 1994). The 1,733 ha included in the GIS map were apportioned as follows: SE, 622 ha (36%); TE, 224 ha (13%); SW, 66 ha (4%); SPOW, 231 ha (13%); SG, 68 ha (4%), and US, 522 ha (30%). = �29 Our study focused on mallards, redheads, cinnamon teal, American avocets, killdeer, and Wilson's phalaropes. Other waterbird species were included in line-transect counts and nest monitoring. METHODS Cover types were delineated and mapped in a GIS. A grid of 0.25-ha square plots was randomly placed over the GIS map. Each plot was categorized according to its dominant (>50%) cover type, and a random sample of plots classified as SE, TE, SW, SPOW, SG, and US were selected for habitat sampling (total n = 330). Field work was conducted during 4 15-day sampling periods; a week separated the last day of a sampling period and the first day of the next sampling period. During eacb sampling period, we measured water depth, conductivity, and temperature, and vegetation height, density (Robel et ale 1970), and species composition (hereafter referred to as habitat variables). Aquatic invertebrates and seeds were collected from 15 plots each in SE, SG, SW, and SPOW cover types during each sampling period. In each plot, 3 samples were collected in the water column, vegetatioh, and benthos. We also measured habitat variables at each site. In the laboratory, invertebrates and seeds in each sample were identified to the lowest taxonomic status and counted. ~ On the first and last day of each sampling period, we used 14 line transects to census waterbirds on RLSWA. As observers walked each transect line, they recorded the species, sex (when possible), flock size, perpendicular distance from the transect line, and the cover type from which waterbirds flushed. We shot foraging waterbirds during each sampling period. We observed foraging birds for >20 minutes prior to collection, to ensure that individuals contained food items obtained at the collection site, and to determine the pair status of collected birds. Immediately following collection, the esophagus of each bird was removed and stored in 95% ethanol. Birds were weighed and stored in a freezer for later processing. In the laboratory, we identified and sorted food items contained in the esophageal contents of each bird. Each food item was dried (60 C) and measured to the nearest 0.1 mg. We also collected food samples and measured habitat variables at the site where each bird was collected to compare food and foraging habitat use to availability at the scale of individual foraging locations. We used focal sampling (Altmann 1974, Tacha et ale 1985) to determine the diurnal activities of selected waterbird species in ~ach cover type. During each 10-minute focal session, observers recorded each time the subject changed its activity (resting, preening, locomotion, foraging, alert, or aggression) or cover type. We measured habitat variables at locations where birds foraged during time-budget bouts. During each sampling period, we searched all selected plots for waterbird nests. We also searched other portions of RLSWA, to increase sample sizes. Habitat variables were measured, and the location of each nest was marked on an aerial photo and later digitized on the GIS map. We revisited each nest near its expected hatch date to determine its fate. A nest was classified as successful if >1 egg hatched. Unsuccessful nests were categorized as deserted, avian predation, mammalian predation, unknown predation, or other. �30 RESULTS Waterbirds varied interspecifically in their use of different cover types; percentages of each species recorded in different cover types also varied among sampling periods (Fig. 1). The probability of detecting waterbirds likely varies among cover types. Therefore, we will use linetransect analysis techniques (Buckland et al. 1993) to estimate detection probabilities and mean densities of species with adequate sample sizes using program DISTANCE (Laake et al. 1994). Invertebrate biomass (mg/L) was highly variable within and among cover types and over time; invertebrate biomass was usually higher in SE plots than in other cover types (Table 1). Invertebrate biomass was generally highest in the benthos (Table 1). Over 95% of the seed biomass in food samples occurred in benthic samples; seeds of spikerush (Eleocharis), bulrush (Scirpus), and Amaranthus dominated the seed biomass. Seed biomass in SE plots was much greater than in other cover types (Table 2). In SE benthic samples where mallards and cinnamon teal typically forage, seeds accounted for 88-95% of the total food (seeds and inverteb~ates combined) biomass., We collected 18 mallards, 30 redheads, 30 cinnamon teal, 27 American avocets, 15 killdeer, and 22 Wilson's phalaropes in 1995. In general, gut contents varied widely among individuals of each species. Pergent dry weight of esophageal contents comprised of invertebrates was (mean [SO]): mallard, 38.4 (30.8); redhead, 10.4 (15.6); cinnamon teal, 53.4 (39.4); American avocet, 95.8 (10.9); killdeer, 99.3 (3.5); Wilson's phalarope, 95.8 (18.2). Waterbird dissections and carcass analyses will be completed in 1997. Food habits will be analyzed in relation to species, sex, reproductive status, and body condition. Carcass lipid, protein, and mineral content will be compared among birds in different reproductive categories for each species. External body measurements will be used to correct for variation in nutrient composition due to individual differences in structural size. We collected 682 focal sessions (568.3 hr) of time-activity data in 1995. Time budget data will be analyzed to determine the effects of cover type, month, time of day, and social status on activities of each species. Percent time spent foraging in each cover type will be combined with linetransect data to rank the relative use of each cover type as foraging habitat by each species. We monitored 268 nests of 13 species in 1995 (Table 3). Mayfield estimates of nest success and daily survival rates of nests will be calculated. Habitat measurements were recorded and nest locations have been plotted on the GIS map. PLANS FOR 1996-97 Line-transect counts, habitat measurements in randomly located plots, collections of foraging waterbirds, time-activity budgets, invertebrate collections, and nest searches will continue in April-July in 1996. Analyses of data collected in 1994 and 1995 will continue. LITERATURE Altmann, J. 1974. Observational Behaviour 49:227-267. CITED study of behaviour: sampling methods. Buckland, S. T., D. R. Anderson, K. P. Burnham, and J. L. Laake. 1993. Distance sampling: estimating abundance of biological populations. Chapman and Hall, London. �31 Colorado Division of Wildlife. 1989. Colorado statewide waterfowl management plan 1989 - 2003. Colorado Div. Wildl., Terrestrial Wildl. Sect., Migratory Game Bird Program. 97pp. Gilbert, D. W., D. R. Anderson, J. K. Ringelman, and M. R. Szymczak. 1996. Response of nesting ducks to habitat management on the Monte Vista National Wildlife Refuge. Wildl. Monogr. 131. 44pp. Laake, J. L., S.T. Buckland, D. R. Anderson, and K. P. Burnham. 1994. DISTANCE user's guide, version 2.1. Colorado Cooperative Fish and Wildlife Research Unit, Colorado State Univ., Fort Collins, co. 84pp. Nelson, D. L., and M. F. Carter. 1990. Birds Luis Valley. Colo. Div. Wildl., Unpubl. Olterman, J., ed. Wildl. Rep. 1993. The San Luis Valley of selected wetlands Rep. 40pp. waterbird Ryder, R. A., W. D. Graul, and G. C. Miller. movements of ciconiforms 'in Colorado. Conf. 3:49-58. Tacha, T. C., P. A. Vohs, and G. C. Iverson. 1985. and continuous sampling methods for behavioral Ornithol. 56:258-264. Prepared by: ;9::~ LSSR III plan. of the San Colorado Div. 1979. Status, distribution, and Proc. Colonial Waterbird Group A compari~on of interval observations. J. Field �32 Table 1. Mean (SD) biomass (mg/L) of invertebrates in samples (n = 15 for each value) collected from 3 substrates in 4 cover types (vegetation was not present in SW sites) used by foraging waterbirds at RLSWA during 4 sampling periods in 1995. Substrate Cover type Benthos SP~W SE SG SW 25-29 APR 16-20 MAY 2.27 5.42 10.27 (4.10) (9.77) (29.01) (17.10) 19.05 33.07 15.01 31.58 (42.01) (74.76) (39.29) (61.56) 2.43 6.58 6.08 5.36 (6.76) (11.32) (8.86) (9.29) 2.28 4.88 11. 56 14.47 (5.81) (18.75) (26.58) 0.22 0.09 0.12 0.26 (0.68) (0.29) (0.43) (0.67) 12.49 2.50 1.95 3.50 (28.26) (4.24) (2.78) (8.83) 3.77 2.39 2.49 0.85 (7.72) (3.15) (6.04) (1. 34) 0.10 0.14 0.23 0.07 (0.30) (0.29) (0.45) (0.11) 1.26 0.89 1.48 0.16 (4.04) (1. 74) (3.62) (0.26) 1.26 0.63 4.75 0.24 (2.97) (0.92) (13.28) (0.26) 0.29 1. 74 0.95 0.44 (0.32) (6.75) (2.33) (0.71) (4.57 ) Vegetation SP~W SE SG Water column SP~W SE SG sw 6-9 JUN 24-28 JUN 7.44 �33 Table 2. Mean (SO) biomass (mg/L) of seeds in benthic samples (n each value) collected in 4 cover types used by foraging waterbirds during 4 sampling periods in 1995. Cover type SP~W SE SG SW 16-20 MAY 6-9 JUN 53.49 38.26 51. 73 61.96 (83.13) (86.59) (114.60) (99.59) 248.24 591.14 113.19 357.22 (507.97) (691.86) (142.54) (474.82) 12.90 25.09 10.02 6.33 (31.56) (57.31) (15.81) (11.39) 20.92 10.01 25.25 (42.41) (28.33) (47.85) (39.10) Fates (percent of total on RLSWA, 1995. Species number of nests) Unsuccessful Successful 24-28 of waterbird nests Unknown n Mallard 39.3 57.1 3.6 56 Redhead 36.4 45.5 18.2 11 Cinnamon teal 30.6 63.3 6.1 49 American avocet 69.6 0 30.4 23 40.0 33.3 26.7 15 Killdeer Wilson's phalarope 50.0 12.5 37.5 8 American coot 89.5 7.0 3.5 57 19.0 76.2 4.8 21 72 .2 27.8 -o 18 Gadwall Canada goose American bittern 100.0 0 0 1 Northern pintail 0 100.0 0 1 Northern shoveler 0 100.0 0 1 sandpiper 100.0 0 0 1 Spotted 15 for at RLSWA 25-29 APR 18.76' Table 3. monitored = JUN �w .p. Fig.. 1. during 4 Percent of individuals counted in different sampling periods in 1995. cover types on line-transect surveys at RLSWA Percent 100 • US II TE [Jl] 80 SG II SPOW o SW [ill] 60 40 20 o 1 2 3 American avocet 4 1 234 Cinnamon teal 1 2 3 4 123 4 123 4 1 2 3 Period Killdeer Mallard Redhead Wilson's phalarope 4 SE �35 JOB PROGRESS REPORT State of __~C~o~l~o~r~a~d~o~ Project W-166-R-5 Work Plan 10 Job Title: : Job Migratory Game Birds Investigations 1 _ Cooperative Management Programs Period Covered: Author: _ 01 April 1995 through 31 March 1996 Michael R. Szymczak Personnel: Michael R. Szymczak, Colorado Division of Wildlife ABSTRACT Recommendations for wetland habitat improvements and/or .anagement were provided for public and private land managers across the state. Proposals for funding projects with Duck Stamp monies were evaluated and rated. Presentations on wetland ecology in relation to waterfowl were given at workshops. Seven Wetland Focus Area Committees were formed in Colorado within the geographic range of the Intermountain West Joint Venture of the North American Waterfowl Management Plan. Chairmen for 3 additional Focus Areas ~ere recruited. Responsibilities as Colorado's representative on Pacific Flyway Study Committee and Council, including chairman of subcommittees for Four Corners band-tailed pigeons and Pacific Flyway consultant to the U. S. Fish and Wildlife Service's Regulation Committee were fulfilled. A 3-5 year cooperative post-breeding goose banding program was continued in the CortezMancos and Durango areas and initiated in the upper Gunnison Basin. ��37 COOPERATIVE MIGRATORY BIRD MANAGEMENT PROGRAMS Michael R. Szymczak In 1988, the Colorado Division of Wildlife (CDOW) created the Migratory Game Bird Program (MBP) within the Terrestrial Wildlife Section. This administrative change combined all individuals having statewide responsibilities for research and management of migratory game birds. Members of the MBP work in concert to improve migratory bird management in Colorado. This job was created to allow team members to participate in these management programs. In November 1993, project personnel assumed additional responsibility for leading and administering the Duck Stamp wetland development program. In July 1994, the author became Colorado State Coordinator for the Intermountain West Joint Venture of the North American Waterfowl Management Plan. P. N. OBJECTIVES 1. Participate in developing and implementing habitat-based waterfowl management plans on 'a statewide, habitat region, and p roj ect; basis. 2. Advise state and federal land managers on beneficial habitat acquisitions and/or developments and provide expertise in preparation of development and/or management plans. Advise private land managers in developing habitat management plans and assessing impacts on waterbird populations '. 3. Present information workshop attendees, 4. Participate in migratory flyway levels. 5. Cooperate in developing surveys and techniques impact of migratory bird management programs. on the principles of waterfowl management to educational classes, and conservation organizations. bird management meetings at the state and that will assess the SEGMENT OBJECTIVES 1. Serve as Colorado coordinator and on the technical review committee for the Intermountain West Joint Venture of the North American Waterfowl Management Plan. Obtain chairmen for the state's 7 geographic Focus Areas and assist in formulating implementation plans for each Focus Area. 2. Serve as chairman of the Waterfowl Habitat Project Review Committee (WHPRC). Provide biological expertise on waterfowl biology and wetland development programs on public and private areas when requested. 3. Prepare and present requested. lectures on migratory game bird management when �38 4. Compile appropriate migratory bird population status information and represent Colorado at Western Migratory Upland Game Bird meetings and Pacific Flyway Study Committee and Council meetings. Attend migratory game bird program and biologist meetings in Colorado, when requested. 5. Provide methodology for wetland'habitat population surveys when requested. 6. Cooperate in Canada goose trapping and banding operations Durango-Cortez area and the upper Gunnison Basin. 7. Conduct experimental surveys of waterfowl breeding pairs o~ wetland development units under management by the U. S. Fish and Wildlife through Partners in Wildlife program. and migratory game bird in the RESULTS Wetland Developments. and WHPRC and IWJV Activities Potential sites for wetland developments and existing wetlands were visited and recommendations for development and/or managemen~were made for: the Sandsage State Wildlife Area(SWA) near Wray, a potential acquisition near Barr Lake SWA, the Hotchkiss SWA near Hotchkiss, Gunnison SWA near Gunnison, Partners For Wildlife projects in North Park (Private/U. S. Fish and Wildlife Service), potential Bureau of Land Managment (BLM) development sites near Dotsero and near Finger Rock, Hebron Sloughs in North Park (BLM), wetlands on the Browns Park National Wildlife Refuge (USFWS), a Colorado Department of Transportation proposed wetland near Limon, a wetlands mitigation project near Cortez (Private), wetlands along the Uncomphagre River between Ridgway and Delta (Private), Markley's Pond near Olathe (Private, potential purchase or easement), the Arikaree River bottom near Wray (Private), playa wetlands near Haxton (Private), the Van Tyl ranch near Gunnison (private), and the Evans Ranch near Pueblo. As chairman of the WHPRC, I: chaired 1 committee meeting for ranking and funding proposals submitted for the 1995-96 funding year; informed proposal submittees of the outcome of their funding request; periodically monitored progress of project planning, construction, and money of new and previous years funded projects; coordinated Site Specifi~ Agreements and fund reimbursement with the Ducks Unlimited MARSH program; solicited, reviewed, and obtained additional information needed to complete project proposals submitted for 1996-97 funding; distributed proposal packets to committee members, and scheduled the 1996 meeting to review proposals. As State Coordinator for the IWJV, I solicited potential partners in wetland acquisition and development in the IMJV area in Colorado (all of Colorado west of the eastern foothills), chaired a meeting of the partners in which 7 geographic Focus Areas were selected, recruited chairman for each Focus Areas Committee, attended 5 Focus Area Committee meetings, and attended 2 meetings of State Coordinators of the IWJV. In addition, I prepared and forwarded background information on preparing Implementation Plans to Focus Area chairman and provided background information on waterfowl populations for some Focus Areas. �39 In addition, I began the formation northeast Colorado (South Platte River), and along the Front Range of Colorado. Informational of Wetland Focus Committees in southeast Colorado (Arkansas.River), Programs A program on waterfowl use of wetlands, designing wetlands for waterfowl, and applying for Colorado Duck Stamp and Ducks Unlimited MARSH monies was presented to the CDOW Wildlife Technicians group. Waterfowl Technical Committee and Council Meetings I attended the July 1995 Pacific Flyway Study Committee (PFSC) and Council meetings. Waterfowl population status was reviewed along with characteristics of the 1994-95 waterfowl hunting season harvest, and proposed 1995-96 hunting season recommendations were formulated,and forwarded through the Council to the USFWS Regulation Committee. Populations of specific interest to Colorado whose status was reviewed in July were (1) breeding and wintering mallards inhabiting western Colorado and (2) the Rocky Mountain Canada goose population. I chaired the Rocky Mountain Canada -goose population subcommittee. The winter meeting of the PFSC featured discussion and decisions concerning the content of objective function for mallards under Adaptive Harvest Management (ARM). The SC decided to give equal weight in the function to: (1) the value of hunting and (2) achieving the North American Waterfowl Management Plan goal for breeding mallards in the mid-continent region (8.1 million). The initial function completely de-valued hunting when the mallard breeding population was reduced to 4 million. The March 1996 meeting of the Pacific Flyway Study Committee allowed committee members to exchange general information on migratory game bird populations and formulate regulatory recommendations for the Flyway Council, for species hunted before October 1, including mourning doves, band-tailed pigeons, snipe, rails, and cranes. I chaired the Four Corners band-tailed Pigeon subcommittee. Early season special Canada goose seasons were also considered. Of special interest was the review of duck regulation packages proposed for the 1996-97 hunting seasons. Three packages (restrictive, moderate, liberal) were offered, and the one selected would be based on a combination of the (1) mallard breeding populations in prairie U. S. and Canada, and (2) number of ponds in prarie Canada, both as measured during the May 1996 breeding pair survey. For all meetings reports containing topics pertinent to Colorado were written, compiled and distributed to appropriate CDOW personnel. In addition to PFSC duties, I was assigned as the Colorado representative on the Pacific Flyway Council. I attended the March Council as Colorado's representative. Colorado's representative was also responsible for representing the Pacific Flyway Council as a consultant to the USFWS's Regulation Committee. Council consultants also serve on the ARM Task Force and I attended 1 Task Force Meeting, and filed a report of that meeting to the Council. �40 Population Survey Methodology The survey of nesting and brood-rearing Canada geese on Walden Reservoir in North Park was conducted in 1995 but few nests were found because low water levels resulted in the conversion of islands (preferred nesting habitat) into penisu1as. Cooperative Canada Goose Banding Canada geese were banded in the Cortez-Mancos area for the fourth consecutive year, in the Durango-Bayfield area for the second consecutive year and in the upper Gunnison basin for the first year. The trapping operations were conducted with the cooperation and personnel of the CDOW Southwest Region. A total of 126 goslings and 35 adults was banded at 6 locations in the Cortez-Mancos area, and 129 goslings and 63 adults were banded in the Durango-Bayfield area at 3 locations in late June 1995. (Appendix A). The total number banded in 4 years'in the Cortez-Mancos ~rea has been 742 with 398 in the Durango-Bayfield area. A total of 61 goslings and 34 adults was banded near Gunnison a~ 1 location. DISCUSSION Project personnel provide useful information in planning and evaluating waterfowl management and habitat enhancement programs in Colorado and educating land management agency personnel about the habitat requirements of waterfowl. With increasing emphasis on wetland habitat in Colorado, and the initiation of new programs with expanded responsibilities for project personnel, wetland related objectives of this job will receive more emphasis in the near future. Colorado will shortly have 10 Wetland Focus Area Committees functioning in the state that will require coordination and expertise in wetland project planning. As in past years the resources provided by project personnel will insure that the money raised through the Colorado Duck Stamp program or any other funding initiative will be spent in accordance with the objectives of the program. Conducting and/or formulating surveys and banding efforts and informing management agency personnel. about aspects of waterfowl and wetland ecology provides a valuable service to management agencies, the waterfowl resource and, in some cases, the hunting public. Continued participation on Pacific Flyway committees ensures that Colorado will remain informed on migratory bird matters, have recommendations for migratory bird hunting regulations, and influence on habitat programs affecting migratory game birds. Prepared by: Michael R. Szymczak LSSR IV �41 Table 1. Age, sex, number, and band numbers of Canada geese banded wetlands in the Cortez-Mancos area, June 1995 Wetland Loc. M Age and sex Loc , F Ad. M Ad. F Totals on Band numbers 19 23 7 9 59- 858-11213 - 271 Colbert's 8 11 3 3 25 858-11188 - 212 La Verde Pds 9 9 5 8 31 858-11157 - 187 Cortez Golf. Course 24 18 4 5 51 858-11106 - 156 7 10 2 3 22 858-11084 - 105 0 1 8 858-11272 - 279 16 19 161 Baike Thomas's Dudd1eson Pds. Totals "Tnc Lude 4 62 s 1 bird of unknown 3' 64 age and unknown sex. Table 2. Age, sex, number, and band numbers of Canada geese banded wetlands in the Durango-Bayfield area, June 1995 Wetland Loc. M Age and sex Loc. F Ad. M Ad. F Totals on Band numbers - 335 James Ranch 18 23 4 12 57 858-11280 858-11337 O'Neal 17 31 16 18 83- 858-11336 858-11338 858-11413 - 411 - 420 Vallecito 4 5 2 0 411 858-11421 - 430 Tay-Col Cattle Co. 9 14 6 5 34 858-11431 - 464 Dove Ridge Subdivision 6 2 0 0 8 858-11465 - 472 54 75 28 35 193 Totals "Tnc Lude s 1 bird of unknown age and unknown sex. �42 Table 3. Age, sex, number, and banding years of Canada geese banded wetlands in the Cortez-Mancos area. Wetland Thomas's Loc. M Pd. Age and sex Loc. F Ad. M Ad. F on Years Banded Totals 27 41 12 12 92 92,93,95 15 22 7 12 56 92,94,95 12 24 3 6 45 92,93 Baikie Pds. 49 55 12 15 131 92,94,95 Colbert's Pd. 50 48 15 17 130 92,93,95 Dudd1eson Pds. 58 '44 9 16 127 92,93,94,95 Pd. 5 5 2 5 17 93 Forest's 10 7 0 2 19 94 Cortez Golf Course 44 45 13 13 115 McPhee Res 1 3 3 3 10 Totals 271 294 76 101 742 La. Verde' Dolores' Pds. Hat. Browning's Table 4. Age, sex, number, and banding years of Canada geese banded area, June 1995 wetlands in the Durango-Bayfield Wetland Loc. M Age and sex Loc. F Ad. M Ad. F Totals 94,95 92 on Years banded James Ranch 30 43 16 23 1J_2 94,95 O'Neal 31 41 29 27 128 94,95 Vallecito 11 15 19 25 70 94,95 Tay-Co1 Cattle Co. 19 27 11 17 74 94,95 Dove Ridge Subdivision 7 5 1 1 14 94 95 98 131 76 93 398 Totals �43 JOB PROGRESS REPORT State of __~C~o~l~o~r~a~d~o~ Project W-166-R-5 Work Plan 22 Job Title: Migratory Period Covered: Author: _ Migratory : Job GameBirds Investigations _2_ Game Bird Publications 01 April 1995 through 31 March 1996 Michael R. Szymczak Personnel: James Garnmonley, James K. Ringelman, Colorado Divisron of , Wildlife and Michael R. Szymczak, ABSTRACT The following list submitted for publication contains those articles or published during this Gammonley, J .H. 1995. Nutrient teal. Condor 97:985-992. reserve that were prepared segment: and organ dynamics of breeding and/or cinnamon 1996. Cinnamon teal (Anas cyanoptera). The birds of North America, No. 209 (A. Poole and F. Gill, eds.). The Academy of Natural Sciences, Philadelphia, PA, and The American Ornithologists' Union, Washington, D.C. Gilbert, D. W., D. R. Anderson, J. K. Ringelman, and M. R. Szymczak. 1996. Response of nesting ducks to habitat management on the Monte Vista National Wildlife Refuge . Wildlife Monograph l~l : 44pp. Pierce, C. L., J. K. 'Ringelman, M. R. Szymczak, andvH, J. Manfredo. An investigation of factors aff~ctirtg wa,te:dowl' hunt.Lng participation, in Colorado. Colorado Div. Wildl. and Colorado State Univ., HumanDdmensLons in Natural Resour. Unit., Fort Collins. Proj. Rep. 10. 86pp. Prepared by: i\ ",t-aWW<00 b 'iJ~ch(l1~ , Michael R. Szymczak LSSR IV l e1.l!) ��45 JOB PROGRESS State of: Project: Work Plan: Job Title: Colorado W-167-R-5 1 -=-- : Job Evaluation in Eastern Period Covered: Authors: REPORT Thomas Upland Bird Research 24 of Habitat Colorado 01 January through E. Remington Developments 31 December and Warren for Ring-necked Pheasants 1995 D. Snyder Personnel: C. E. Braun, T. J. Davis, M. A. Etl, K. M. Giesen, E. T. Gorman, L. K. Haynes, R. W. Hoffman, D. D. Johnson, E. N. Landes, J. L. Mekelburg, M. A. Porter, T. E. Remington" B. J. Rosenbach, W. D. snyder, M. L. Trujillo, J. D. Weiland, B. T. Weinrneister, J. A. Yost, D. J. Younkin, Colorado Division of Wildlife; G. A. Peterson, D. Poss, D. G. Westfall, Colorado State University. ABSTRACT Expenditures under the Pheasant Habitat Improvement Program (PHIP) increased from about $278,000 in 1994 to $299,000 in 1995. Most habitat developments were sorghum plantings (295 of 552), although, as in past years, most PHIP expenditures went toward establishment of plum thickets. PHIP has now resulted in the establishment of 538 such thickets in 4 years. Above average spring moisture throughout the area improved plum seedling survival but delayed planting of sorghum. As a result, cover value of sorghum plantings for ringnecked pheasants (Phasianus colchicus) was only fair, with an average height of only 5.1 drn, and a vertical obstruction reading (VOR)of 2.2 drn among 50 plots measured. Average counts of crowing males did not differ (P = 0.38) between treatment and control blocks. Counts averaged about 14 calls per station in 1993, 17 in 1994, and 13 in 1995. Hunter pressure declined about 25% from 1994, while harvest rates increased about 50% to 0.08 birds per hour in 1995. One hundred and seventy-seven hens were captured and radiomarked, bringing the sample of birds available for survival estimates to 203 (including 26 survivors from the 1993 and 1994 trapping efforts). Annual survival of radio-marked hens (1 Oct - 31 Sep) declined to 15% from 41% in 1993-94. The decline in, survival was attributed to poor cove rd.n wheat stubble and Conservation Reserve Program (CRP) fields following drought condd.t.Loris during spring of 1994. No differences in survival between treatment and 'control blocks were apparent. Wheat stubble resulting from combining wheat using a stripper header had an average VOR 'of 5,5, dm and an average height of 8.0 drn. These values were significantly greater (P < 0.05) than the VOR and height of 2.4 and 3.6 drn, respectively, for wheat stubble resulting from combining wheat with traditional headers. Stripped wheat , stubble contained less waste grain than conventionally-cut wheat stubble (P < 0.05; 12.0 vs. 7.6 gms/square m). Both stubble types contained enough waste grain to provide an adequate food resource for pheasants over winter •. ��47 EVALUATION OF HABITAT DEVELOPMENT IN EASTERN Thomas E. Remington FOR RING-NECKED PHEASANTS COLORADO and Warren D. Snyder INTRODUCTION Pheasants are pursued by more hunters than any other small game species in Colorado (83-88% of small game license buyers). In a recent survey, 74% of pheasant hunters rated their hunting trips in Colorado as poor (45%) or fair (29%), while only 10% rated their trips as very good or excellent. Lack of birds and places to hunt were identified as the most significant reasons why some hunters did not hunt pheasants in Colorado. Small game license sales in Colorado have declined by about 90,000 (45%) in the last 10 years. It is apparent that if the Division of Wildlife is going to turn this decline around, pheasants will be a key species. Presumably, recruitment and retention of hunters will increase i~ the quality of pheasant hunting is improved, i.e., increases in pheasant numb'ers and places to hunt. Previous research has indicated that over-winter survival of ph~asants is the most critical factor limiting pheasant populations. The Pheasant Habitat Improvement Program (PHIP) was created to establish overwinter survival cover within historically good pheasant range in eastern Colorado. The program was conceptually designed to overcome significant obstacles to developing habitat, mainly a lack of manpower and a burdensome contractual system (costs of administering contracts exceeded costs of developments). Under PHIP, the Division of Wildlife contracts with individual Pheasants Forever chapters in eastern Colorado to contact landowners and develop habitat on private lands following specific guidelines. Each chapter develops contracts with individual landowners and pays them when the habitat work is completed and verified. Division of Wildlife personnel inspect habitat developments and verify completion and compliance with guidelines. A new method of combining wheat has potential to increase pheasant survival. Stripper headers strip the grain from the stalks rather than cutting and thrashing the stalks to separate the grain, which leaves much taller stubble. Pheasant survival from July through April is largely dependent upon the height and cover value of wheat stubble (Snyder 1985). We measured the height and cover value of paired blocks of stripped and conventionally-cut wheat to assess the value of stripped wheat for pheasants. Stripper headers are potentially more efficient than conventional headers, i.~. less waste grain. While this is positive for farmers, it could be negative for-pheasants and other granivorous animals if food in the form of waste grain is unavailable as a result. We measured the amount of waste grain in stripped and conventionally-cut fields to determine food availability. Farmers typically make decisions about equipment purchases on an economic rather than a wildlife basis. Because stripper headers are expensive (-$100,000), farmers are unlikely to purchase them unless some economic benefit can be demonstrated. One potential benefit is increased soil moisture storage over winter because taller stubble should trap and hold more blowing snow, and evaporation will be reduced by the shading effect of taller stubble. A significant increase in soil moisture would allow farmers to grow 2 crops in 3 years, rather than the traditional crop every other year under a wheat-fallow rotation. We measured the fall soil moisture profile of stripped and conventionally-cut wheat stubble fields as a baseline to compare relative moisture storage over winter. �48 P. N. OBJECTIVES To determine if habitat developments offered through the Pheasant Improvement Program increase pheasant survival, breeding density, harvest within selected northeast Colorado study areas. SEGMENT 1. Habitat and pheasant OBJECTIVES Work with Pheasant Forever Chapters, management personnel, and landowners to develop habitat within treatment sites and elsewhere the primary pheasant range in northeast Colorado. in 2. Monitor hen pheasants previously radiomarked with mortality-sensing transmitters within treatment and control blocks to compare survival, nesting success, and use of habitats through 1995. Trap and radiomark additional hens as necessary to reach a sample size of 100 hens each in the treatment and control blocks in fall 1995. 3. Conduct pheasant during April-May 4. Monitor control hunting sites. 5. Conduct cover. evaluations 6. Prepare an annual crowing'counts 1995. pressure within and pheasant of the quality progress all treatment harvest and control within of annual plantings t~tment blocks and as survival report. METHODS Procedures used during this work segment were described by Remington and Snyder (1995). The PHIP Habitat Project Guidelines for participating Pheasants Forever chapters were modified slightly from 1994 and are attached (Appendix A). Paired stripped and cut blocks were selected from within wheat fields or occasionally from adjacent fields. Cover value of stripped and cut wheat fields was evaluated using Robel poles to measure height and visual obstruction similar to how cover quality was measured in sorghum plantings. Twenty measurements of VOR and height were obtained in each field. Waste grain was vacuumed by shop vacuum from within 0.25 x 1.0 m aluminum sample frames. Two sets of 6 samples were taken from each field. Each set consisted of samples taken from 1, 2, and 3-m left and right of the center of combine wheel tracks. The contents of the shop vacuum were placed in paper bags. Wheat seeds were separated from other material using different size screens, dried in a convection oven, and then weighed. Soil cores were taken using hydraulically driven stainless-steel soil corers and placed in numbered aluminum tins. Soil moisture was determined as the difference in weight of samples before and after drying in a convection oven. RESULTS The Pheasant Habitat Improvement Program continued to be a popular program with landowners in northeastern Colorado and with members of participating Pheasants Forever chapters. The Frenchman Creek Chapter of Pheasants Forever initiated its first contract for PHIP funds and began habitat work in the Fleming area. The 6 cooperating Pheasants Forever chapters began the 1995 growing season with $357,239.18 in PHIP funds. Of this total, $320,000 was obtained through new contracts with the Colorado Division of Wildlife and the balance was carried over from the previous year. Total PHIP expenditures were $298,679.91 in 1995 (Table 1). �49 Pheasants Forever chapters also spent their own chapter money on purchase and maintenance of equipment, fuel, food for volunteers, etc., which increased the amount of habitat developed. None of the habitat development under PHIP would be possible without Pheasants Forever members and landowners donating their time. Pheasants Forever chapters continued to use a variety of approaches to planting plum thickets; this flexibility was facilitated by payment of a reasonable labor rate. The Phillips county Chapter of Pheasants Forever planted 42 plum thickets (most with juniper windbreaks) and subcontracted an additional 21 thickets to a professional planter. The labor portion of the PHIP payment for the thickets planted by the chapter was invested in additional shrub plantings (not paid under PHIP) , purchase of switchgrass seed, and major overhaul on 2 grass drills owned by the local Natural Resource Conservation service office but used by the Phillips County Chapter under a cooperative agreement. They, and 3 other chapters hired a temporary employee to contact landowners and layout planting sites. The Washington County Chapter of Pheasants Forever subcontracted all of their 50 woody plantings to local Future Farmers and Young' Farmers chapters whose members donated their time and equipment to planting as fund raising proje~ts. The Pheasants Forever chapter retained funds to cover administrative costs, but most of the planting costs for labor went to the subcontracted groups. ~The Northeast Colorado Chapter in Sterling received assistance in their plantings from local Explorer Scouts and their group received some reimbursement for their labor from the Pheasants Forever chapter. The Yuma and Phillips County chapters received some volunteer assistance in planting from members of Front Range Pheasants Forever chapters. A large group of local people donated their efforts toward the first-year planting efforts by the Frenchman Creek Chapter. All plantings completed by the Eastcentral Chapter in Kit Carson county were contracted to a professional tree planter. In 1995, 552 plantings were completed on 1,672.9 acres (Table 1). Woody plantings continued to be the most popular habitat option among Pheasants Forever chapters. Their efforts resulted in 191 thicket plantings in 1995, which brought the total to 538 in the past four years (Table 2). Nearly all consisted of a 0.1 to 0.3-acre shrub thicket accompanied to the north and west by a small 3-row juniper windbreak. They were also the most costly item, but when costs are prorated over their expected life (> 30 years) they may be the most cost-effective habitat component. Seedling survival for most plantings was good to excellent, the poorest occurring in southeastern Sedgwick County where less rainfall was received. Wet field conditions delayed completion of planting by some groups, and a few thickets were planted late using weakened planting stock with resulting low survival. Laying of fabric around the seedlings could not be completed on a few sites in Washington county which were subsequently abandoned for the year. The Washington County Pheasants Forever Chapter absorbed the cost of the seedlings and donated their labor for these thickets. Precipitation was deficient throughout northeastern Colorado during 1994 and averaged 9.76 cm (3.84 inches) below average for the Holyoke, Sterling and Akron weather reporting stations. In contrast, April, May, and June 1995 were exceptionally wet and cool which halted field preparation and planting until after mid-June. Although data for December 1995 were not available, annual precipitation in the study area was above average for 1995 and totaled 48.9 cm when averaged for Holyoke, Sterling, Yuma, and Akron weather stations for January through November. Greater than 0.25 cm (0.10 in.) of precipitation was received on 12 days of May, and total precipitation was greater than average at all reporting stations. This facilitated establishment of switchgrass that had previously been seeded but markedly delayed site preparation and planting of sorghum plots. Nearly all sorghum was planted during the last 2 weeks in June and satisfactory stands were established. �50 Table 1. Pheasant Habitat Improvement Program expenditures by Pheasants Forever chapters during 1995 in northeastern and east-central Colorado, by habitat type. Number Plantings Acres Habitat/Contract Sorghum Plantings Northeast Colorado Phillips County Yuma County Washington County Subtotal (sterling) 5 ~5 Switchgrass Plantings Northeast Colorado Phillips County Yuma County Washington County Subtotal 4 22 1 2 ~ Disturbance Tillage (Annual Forbs) Northeast Colorado (sterling) Phillips County Yuma County Washington County subtotal Tall Wheat Yuma Stubble 64 55 171 5 22 1 ~ $8,856.00 8,218.00 29,377,00 560.00 $47,011.00 17.8 113.1 3.0 14.0 147.9 887.50 5,160.00 150.00 700.00 $6,897.50 26.8 9.0 166.0 4.5 206.3 $1,072.00 360.00 3,520.00 180.00 $5,132.00 18.0 $360.00 Retention Shrub Thickets and Windbreaks Northeast Colorado Phillips County Yuma County Washington County Frenchman Creek (Fleming) Kit Carson Subtotal 15 63 38 50 8 17 191 552 Plantings/acres Custom Tillage (Contracted Northeast Colorado Phillips County Custom Yuma County Washington County Subtotal Replacement of Seedlings Yuma County Washington county Subtotal Total 221.4 216.7 740.8 14.0 1,192.9 . 8 1 Total Payment Expenditures 7.5 34.4 22.4 29.2 5.0 9.3 107.8 $16,641. 80 77,865.87 47,083.05 65,512.80 9,888.76 17,737.80 $234,730.08 1,672.9 Site Preparation) 4 2 24 50 --so 30 9 ~ 16.0 6.0 72.0 103.0 197.0 163.50 90.00 504.00 2,707.50 3,465.00 $809.00 275.33 $1,084.33 $298,679.91 �51 Table 2. Pheasant Program, 1992-95. Woody plantings Sorghum habitat a food/cover plots switchgrass plantings Disturbance tillage Tall wheat stubble Totals created through the Pheasant Habitat Improvement 1992 1993 1994 1995 Total 49 136 162 191 538 52 304 333 295 984 0 24 73 29 126 11 47 37 36 131 13 6 8 1 28 125 517 613 552 a Woody plantings consist of 0.1 to 0.3-acre plum thickets; accompanied by 3-row juniper or juniper/plum windbreaks.: However, precipitation was well below normal from July through combination of a short growing season (caused by late planting and insufficient summer moisture stunted growth of both sorghum plots. Wet snows in late-september and October accompanied by over many of the annual weeds but most of the sorghum remained relatively dry winter. 1,807 most were mid-september. The and an early frost) and annual weed high winds broke standing through the Over half of the 295 sorghum plantings were planted within treatment areas. No additional sorghum plots were broken out within CRP fields. A few plots, previously planted to sorghum in 1994, were retained in annual weeds (disturbance tillage) in 1995 because of wet field conditions that severely restricted time for site preparation and planting. Sorghum plots averaged 5.1 dm in height with an average visual obstruction reading (VOR) of 2.2 (Table 3). While this was considerably better than cover values in 1994, plots were still below the VOR of about 5 needed to influence pheasant survival. Ninety percent of plots had at least 1 VOR reading> 5, indicating there were small clumps of adequate cover available in most plots. Pheasant use was rated as high in 6 of 50 (12%) plots, low in 25, while no pheasant use was detected in 19 plots (38%). We have been unsuccessful in establishing consistently good stands of sorghum over the 3 years of this program. Extreme weather, both wet and dry, has contributed to variable and poor stands, but it is also becoming clear that normal ~infall won't support successful growth of sorghum if annual food plots are planted year after year in the same location. This problem is compounded in CRP fields where moisture is typically depleted when the plot is broken out of grass, and competition from grass persists. We could pay landowners to fallow the plot every other year, or every second year, but this would increase the cost per acre of standing sorghum. This is probably cost effective only in ideal situations, such as center-pivot irrigation corners adjacent to winter roosting cover. Switchgrass plantings were given additional emphasis as permanent survival cover that does not need annual planting or maintenance. Twenty-nine switchgrass plantings were completed in 1995, most of these (22) were planted by the Phillips County Chapter. In addition, a number of sites planted in 1994 that failed to establish due to drought were replanted in 1995 and these appear to have become established. �52 Table 3. Vegetation characteristics of sorghum plots planted within treatment blocks in 1995 and sampled in January-March 1996, northeastern Colorado. VOR Height (%) CanoE:t cover Treatment block N (dm) S.D. (dm) S.D. Sorghum S.D. Kurtzer Pauli Fleming Clarkville Y-W 10 10 10 10 10 2.70 2.23 2.08 2.69 1. 07 0.72 0.58 0.87 0.89 0.36 6.84 4.93 5.01 5.45 3.46 1.07 0.82 1. 37 1.17 0.70 50 44 34 40 36 11 16 9 11 9 20 45 31 29 12 5 18 16 8 4 2.15 0.67 5.14 1.21 41 7 27 12 Means Pheasant Crowing Forbs S.D. Census Average courrt.sof crowing males did not differ between tf'eatment and control blocks (Table 4; P = 0.50 and 0.95 using average and high count of replicate counts, respectively). Average counts declined from about 17 calls per station in 1994 to about 13 in spring 1995. The lack of a difference in breeding density between treatment and control blocks is not surprising given the generally poor quality of the sorghum plantings as survival cover. Average counts may have been reduced since we were unable to get replicate counts for many areas. Table 4. Pheasant crowing census data among treatment northeastern Colorado, spring 1995. and control blocks, Count Block 1 2 3 Average of count.s? Highest count High count/ s t at.Lon- Treatments Holyoke SE Mailander Kurtzer Clarkville Pauli Y-W Co. Line Fleming Kuntz otis Curve 19.7 7.4 14.0 12.6 10.2 6.8 8.5 11.8 4.8 10.7 6.9 4.3 14.8 Means 19.7 7.9 12.9 12.6 7.5 10.7 6.9 4.3 14.8 19.7 8.5 14.0 12.6 10.2 10.7 6.9 4.3 14.8 19.7 9.5 16.8 12.6 10.4 10.7 8.0 4.3 14.8 10.8 + 4.7 11. 3 + 4.6 11. 7 + 4.5 Controls 18.6 Paoli NE 14.1 Haxtun NE Paoli South st. Pete Kelly Yuma Co. Lonestar Platner Washington W. 8.8 16.0 37.0 10.0 5.2 using 16.3 13.3 8.2 16.0 37.0 18.6 14.1 8.8 16.0 37.0 19.9 15.1 10.9 16.0 37.0 13.3 11. 6 5.2 7.4 13.3 5.2 7.4 14.5 5.2 7.4 14.4 + 10.0 15.1 + 10.0 15.8 + 9.8 7.4 Means ~ Obtained 14.1 12.5 7.7 the highest count per station among counts before averaging. �53 Hunter Pressure and Success Hunter contacts during the opening weekend (397) declined by 25% from 1994 levels, but this was still 53% above 1993 levels (Table 5). Weather on Saturday and Sunday was similar to last year; warm, sunny, and conducive to hunting. Harvest rates improved from 0.05 birds per hour in 1994 to 0.08. Although this represented a greater than 50% increase, on average hunters would still have had to hunt about 13 hours to kill a rooster. This compares to an overall harvest rate of about 0.19 birds per hour (5.2 hours to harvest a rooster) in 1993 when snow on Sunday boosted hunter success. Harvest rates were, on average, similar within treatment and control blocks (0.06 birds/hour). Sorghum and disturbance tillage plots continued to be popular with hunters where these provided adequate cover. Many hunting parties did not feel they had enough hunters to effectively hunt quarter sections of wheat or CRP effectively. The value of PHIP sorghum and disturbance tillage plots as accessible places to hunt cannot be discounted. Pheasant Trapping and Survival Pheasant trapping began on 2 October and continued until 11 December. We placed radios on 177 new hens. Twenty-six transmitters were still functioning (as of 1 Oct) on surviving hens trapped and marked in 1993 and 1994, bringing the total of hens available for assessment of survival in 1994·;:"95 to 203. Annua'L survival of hens radio-marked between October and December 1994, was low compared to the 1993 cohort, or compared to what is needed to increase populations. Of 206 hens radiomarked between October and December 1994, 26 survived to 31 September, 1995, while 144 died,S lost their transmitters, and we lost contact with 31 during this interval. This represents a survival rate of about 15% (26 of 170 birds not lost or missing), compared to 41% survival the previous year. Mortality was relatively high from November through June, then declined from July through September (Fig. 1). Most mortality was due to mammalian or avian predation, although a few hens were shot by hunters, hit by vehicles, or died from some other accident. There were no weather-related mortalities as snowfall was light and snow cover did not persist. High fall, winter, and early spring mortality can be attributed to poor cover in wheat stubble and CRP which resulted from drought conditions during the 1994 growing season. This pattern of persistent mortality contrasts with that reported for hen pheasants by Snyder 1985)in northeastern Colorado from 1979 to 1981 when taller varieties of wheat predominated. 25 101993-19941 ~ 1994-1995 20 4 (J. r+ 15 - >.~ r- ~ ~ ro .•.... •... ~ gX 0 10 ~ e- 5 o r - - r- ~ ~ I Nov Dec Jan Feb Mar ~ ~ ~ Apr - I ~ May Jun Jul Aug Sep Oct Month Fig. 1. Percent of radio-marked hen pheasants that died each month. �Table 5. Pheasant November 1995. hunters Hunters Block contacted, Flush/ Hunter hunter Bag/ Hunter Inside effort, and hunter Birds/ Hour success Hunters in Treatment Flush/ Hunter Block and control Bag/ Hunter outside blocks, Birds/ Hour Block 12-13 Birds/ Hour combined Treatments 6 1. 00 0.67 0.102 - - - - 0.102 Mailander 13 0.38 0 0 10 1.5 0.3 0.08 0.043 Kurtzer 28 0.93 0.11 0.068 16 0.81 0.06 0.022 6 4.17 0.50 0.074 10 2.80 0.30 0.082 Holyoke SE Clarkville 32 Pauli Y-W County 2 0.044 0.078 0.50 0.16 0.056 45 1.33 0.47 0.078 0.073 0 0 0 20 1. 95 0.40 0.124 0.120 Fleming 10 0.90 0.10 0.065 2 3.00 0.50 0.333 0.109 Kuntz 40 0.15 0.08 0.090 36 1.25 0.33 0.093 0.092 otis Curve 13 0.31 0.15 0.094 10 1.50 0.3 0.059 0.070 0.63 0.12 0.063 Means Controls Paoli NE 26 0.35 0.12 0.067 19 3.58.. 0.63 0.162 0.126 Haxtun 23 1. 00 0.30 0.149 20 1.35 0.20 0.075 0.109 - - - - 0.108 '1.00 NE • , Paoli south 2 1. 50 0 0 st. Pete 9 0.33 0.11 0.108 Kelly 10 1. 30 0.10 0.125 8 0.13 0.034 0.054 Yuma Co. 42 3.24 0.52 0.073 16 3.25 0.75 0.20 0.094 Lonestar 32 0.56 0.03 0.012 3 6.67 0.33 0.067 0.020 Platner 23 0.13 0 0 9 1. 67 0.11 0.049 0.015 0 0 0 6 0 0 0 0 1.20 0.20 0.064 Wash-West Means 6 I - 0 V1 ..,.. �55 stripped Wheat Cover Value Wheat harvested with a stripper header provided over twice the cover value as wheat cut with a conventional header (Table 6). The VOR for stripped wheat ranged from 4.3 to 7.7 and averaged 5.5 dm, higher (paired t-test, ~ < 0.05) than the average of 2.4 dm for cut wheat. stripped wheat ranged from 6.9 to 9.3 dm in height, and averaged 7.9 dm. Cut wheat averaged 3.6 dm tall. VOR's and height of stripped wheat were substantially greater than those of sorghum plantings. Heights of both stripped and cut wheat stubble were greater than normal because spring rainfall was much greater than normal. stripped wheat was so tall that much of it lodged when several inches of heavy, wet snow, driven by high winds, fell in early October. This reduced the cover value of many fields for the rest of the winter. Waste grain was abundant in both stripped and cut wheat fields. Cut wheat contained more (P < 0.05) waste grain than stripped wheat; 12.0 versus 7.6 g/m2 (Table 6). Samples taken directly between combine tire tracks contained more (~< 0.05) waste grain than samples taken from either side of the combine track, indicating waste grain is not randomly distributed throughout wheat fields. stratifying sample collection ,by distance from the center of the combine track was a useful way to reduce variance. Fall soil moisture profiles were similar between stripped and cut wheat fields (Fig. 2), although soil moisture in depth stratum 2 (1-2 ft)~~as higher in stripped than cut fields (11.9 to 9.3%; P = 0.03). It is not clear whether this is an anomalous result or due to increased shading and wind protection afforded by the taller stripped stubble. Soil moisture profiles will be measured again in spring 1996 to ascertain whether stripped wheat fields trap and hold more snow and consequently retain more soil moisture than cut fields. 14 10 stripper 12 - .~:=:~; ~ conventional I - - r-- - 10 r ~ 0 Q) ':J 8 r +-' C/) r-4 0 E 6 - 4 - r-: 0 (/) 2 r ~ 0 1 2 3 4 5 6 Depth (feet) Fig. 2. Soil moisture profiles in stripped and cut wheat fields, Aug. 1995. �56 Table 6. Height, available vertical in wheat fields harvested VOR Field 1 Height (VOR) , and waste 1.9 7.5 grain and stripper (drn) stripped cut 4.3 reading with conventional (drn) stripped Schlacter obstruction Waste (g/m2) headers, 1995. grain cut stripped cut 3.0 2.9 1.9 2.0 Schlacter 2 6.2 2.8 8.8 3.6 1.8 Schlacter 3 5.1 2.7 7.7 4.0 3.7 2.0 Schlacter 4 5.6 2.8 8.2 4.2 2.8 2.0 Schlacter 5 4.9 2.2 8.1 3.6 2.5 3.0 Schlacter 6 7.0 2.5 7.8 3.7 5.1 1.5 8.1 2.8 3.0 2.0 Rafert 1 Rafert 2 4.6 1.7 7.0 3.5 Rafert 3 7.7 1.7 9.3 2.9 3.6 7.0 Rafert 4 5.6 1.7 8.1 2.7 2.1 Rafert 5 5.7 3.2 8.3 4.7 1.1 4.6 - 5.7 . 0.6 Heerrnan 1 6.7 3.4 9.1 4.4 1.5 8.7 Heerrnan 2 4.7 2.4 7.9 3.5 1.0 3.7 Heerrnan 3 4.3 2.4 7.3 3.2 1.1 2.4 Heerrnan 4 4.4 1.9 7.0 3.1 0.9 1.9 Heerrnan 5 5.6 2.9 8.1 4.1 0.9 4.4 Gerk 4.9 1.9 7.0 3.1 0.9 1.6 5.8 2.1 7.8 3.2 7.1 3.7 9.0 4.9 0.9 1.0 5.4 2.6 7.6 4.0 1.9 1.6 5.5 + 1.0 2.4 + 0.6 8.0 + 0.7 - 1.9 + 2.5 - 3.0 + 4.9 1 Gerk 2 Gerk/Einsp. Gerk 3 4 Means LITERATURE Remington, T. E., and W. D. Snyder. necked pheasants Fed. Aid snyder, W. D. 1985. Wildl. Prepared by Proj. W-167-R. Survival Manage. LSSR IV CITED Evaluation Colorado. Apr. of habitat Colorado development Div. Wildl., for ring- Prog. Rep., 1-19. of radio-marked hen ring-necked pheasants in Colorado. 49:1044-1050. --rt~"'-'tf.-d {_ Thomas 1994. in eastern 3.6 + 0.6 E. Remington Prepa red by \ >JOIUt91\ Warren D. Snyder LSSR IV 1), JruV-tt!\ 1 { .RI\~ J. �57 Appendix A. PHIP specifications, SHRUB THICKETS 1994 AND SUPPLEMENTAL WINDBREAKS Shrub (plum) thickets are the priority item. Small windbreaks, if planted, must be associated with a thicket and will not be funded if planted alone. Plantings will be eligible for funding only in farmed areas and must be within 0.1 mile of cultivated cropland. Plantings must remain undisturbed for at least 10 years. Maximum Funded: No more than 1 thicket (with or without wind barrier) can be planted per 80 acres. Each thicket/windbreak must at least 1/4 mile from another thicket/windbreak. be Size: Shrub thickets must be at least 1/10th acre (4,300 ftz) and no larger than 2/10ths acre (8,800 ft2) in size, and must include at least 8 rows (excluding windbreak rows). Twelve hundred (1,200) feet is the maximum linear feet of fabric funded per thicket. Supplemental windbreaks, if planted, must be placed on the north and west side of the thicket and must include no more than 900 linear feet total if straight and 1,200 linear feet total if L-shaped. They must include at least 3 rows, one of which must be juniper or cedar. Spacing between the thicket and the windbreak should approximate 100 feet (range 60 ft. minimum; 120 ft. maximum). Payment Rate: Payment will be at $0.55 per linear foot of fabric (6 ft. wide) to the maximums listed above if completed by PF Chapters. Actual costs not to exceed $.60/ft. if contracted to the State Forest Service or a local SCD. The maximum payment rate for approved private contractors will be $0.64 per linear foot of fabric. Supplemental payment rates/linear foot will be: $0.03 for application of polymer, $0.05 if 8 ft. wide fabric is used, and $0.01 for band application of an approved herbicide along the exterior edge of the fabric to reduce weed competition. Use of fertilizer will not be funded. Labor for landowners planting their own plots will not be funded. Planting Pre-Plant Dates: Between March 20 and May 15. Treatment: sites must be tilled, preferably the fall prior �58 to planting. Tillage must be to bare soil with little residue remaining and must be deep enough to kill existing vegetation. Approved species: American Red Cedar (potted). Plum (bare root), Rocky Mt. Juniper, Between-row spacing: A maximum of 10 feet will be permitted feet spacing is recommended) for shrub thickets. A maximum feet will be permitted for wind barriers. E. (6 to 8 of 15 In-row Spacing: A maximum of 8 feet will be permitted for shrub thickets (6 feet is recommenqed for plums) and 8-12 feet will be permitted for evergreens within wind barriers. : Mulching: Woven polypropylene fabric is required for-all plantings. Minimum fabric width is 6 ft. (3 ft. on each side of the row). PERENNIAL GRASS AND GRASS-LEGUME PLANTINGS switchgrass provides tall cover that stands well over winter and has high value for pheasants. Small unfarmed tracts, currently in short, sodded grasses, are recommended for revegetation to switchgrass. other shorter, cool-season grass-legume mixtures may be used in roadsides where snowdrift is a problem. This practice is funded only in farmland (not rangeland) settings. Payment Rate: $50.00 per acre as a one-time payment for sites up to 10 acres. For each additional acre (in sites larger than 10 acres [40 acres maximum]) the rate is $35.00. An additipnal $15.00 per acre will be paid for breaking out sod- in heavily sodded sites and supplemental discing prior to planting switchgrass (this does not apply to roadsides). Preplant Soil Preparation: Adequate tillage to completely destroy existing perennial vegetation and to establish a moist, weed-free, firm seed bed is required. Interseeding is not approved. A pre emergent herbicide (e.g. Ally at 1/10 oz/acre) is recommended when planting switchgrass. Planting a tall-sorghum mix (for which payment is available) is recommended the first year. switchgrass can be seeded into the residual sorghum without tillage during the subsequent spring but application of Ally herbicide is recommended. �59 P1antina Procedures: Planting procedures outlined in the Division's Game Information Leaflet #113 should be considered when planting switchgrass. In general, about 20 pure live seeds/ft2 (2 - 3 lbs/acre) should be planted using a drill with double-disk furrow openers, I-inch depth bands, and packer wheels. If a herbicide is not used, up to 1 lb/acre of an adapted dry land alfalfa and up to 1/2 lb of sweet clover should be added. Approved Species: In plots, switchgrass should comprise at least 75% of the live seed (alfalfa and sweet clover are approved additions). within roadsides, switchgrass is the priority species where snowdrift is not a problem. Other approved warm-season grasses include bluestems and Indian grass. Where these can not be used the tallest wheatgrasses (tall, intermediate, or standard crested) the roadside site will allow, should be used in combination with alfalfa (1 to 2 lbs/acre). Plantina Dates: Warm-season grasses including switchgrass: March 15 to May 15: Cool-Season Grass-legume Mixtures: March 15 - July 15. Plot Duration: Grass and grass-legume plantings must remain ungrazed and undisturbed for at least 7 years. Roadsides should remain unmowed unless essential to reduce snowdrift. If essential, mowing should be delayed until after 1 August and restricted to the road shoulder. Prescribed burning, thinning tillage, or other renovation treatments to rejuvenate grass stands may be applied after 7 years. Grass stands that are relatively thin provide talTer, better cover for pheasants. Legumes provide nitrogen and increase growth and quality when added to mixtures. DISTURBANCE TILLAGE AND TALL WILD ANNUALS wild sunflowers, kochia, pigweed, and other tall annuals which attain 4 - 6 ft. height stand better through winter than other herbaceous vegetation, and provide excellent cover for broods, protection from blizzards and predators, and supplemental food. This is the most effective and least expensive approach for increasing pheasants and other upland game birds. Fallow land that is left idle usually converts to annual grasses or dog-hair stands of weeds by the 2nd �60 year following tillage. Thus, at least one tillage each spring usually needed to promote growth of tall annuals and a second thinning tillage is sometimes needed. Maximum Funded: 14 acres/0.25 section, 28 acres/section. larger than 3 ac. should be at least 0.25 mi. apart. FUnding is Plots Rate: $30.00/year for patches 0.1 to 0.5 acres in size, patches larger than 0.5 acres are considered 1 acre. $40.00/acre/year for sites up to 5 acres (7'ac. in pivot corners). $30.00/acre/year for additional acres up to 10 (6th10th acres). Seeding wild sunflower or other approved wild annuals at 2 to 4 lbs. per acre will be funded at direct seed costs (see seed sources below. Plot Dimensions: easily inundated Short, relatively wide patches, which will not be by drifting snow, are preferred. Placement: Adjacent to woody cover when possible. Draw bottoms that already contain weeds and above average moisture are ideal. sites containing noxious perennials should be avoided. specifications: Initial tillage with a disk plow or mold-board plow is needed in sites containing perennial grass to destroy all perennial cover, preferably, immediately after th~ground has thawed in early March. Large clod size is preferred to retain thin stands of annual forbs. Initial tillage in subsequent years should be conducted prior to May 1 A second thinning tillage may be used prior to the 1st of June. Spring tillage is needed'each year to retain tall annuals. Annual grasses usually dominate if tillage is not used each spring. wild sunflowers and annual ragweeds can be drilled or broadcast and harrowed at low rates to help establish tall annuals, if they are not already present. Known sources in Colorado include the Arkansas Valley Seed Company - Denver & Longmont, and Sharpe Bros. Seed Company - Greeley. �61 Retention: Tall annuals must remain undisturbed through March of the following year. Sites should be prepared for the next year's growth during. early to mid April if weedy cover exists. �62 ANNUAL SURVIVAL PLANTINGS - SORGHUMS & DRY LAND CORN APPLICATION: On CRP, Annual Set Aside, and other cropland or tilled wasteland. When applied within CRP fields, SCS specifications for CP-12 must be used (see supplement). Dryland corn is primarily applicable in center-pivot corners next to irrigated corn. MAXIMUM FUNDED: 1 plot/80-acre field, 2 plots/160 acres, 4 plots ac.)/section. Plots must be at least 1/4 mile apart. (28 PAYMENT RATE: $40.00/acre/year for 1-5 acres (7'acres within center pivot corners) and $25.00/acre/year for additiohal acreage~ in tracts larger than 5 acres (12-acre maximum). $30.00/acre/year if planted after June 15th. $6.00/acre for application of 30 lbs of nitrogen/acre. $15.00 per acre will be paid for breaking out sod in CRP or heavily sodded sites and supplemental discing prior to planting. Landowners can not be paid for labor/tillage on their own land other than for breaking out sod. PLACEMENT: Plots should be placed within placed crosswise to prevailing winds. SPECIFICATIONS: Preplant Soil Preparation: Initial to destroy existing perennial vegetation or near cropland and treatment: Adequate tillage in early spring prior to annual growth. Subsequent years: Preferably minimum tillage shredding of old materials as needed prior to April 25. Annual application of nitrogen at 30-40 lbs./ac. is recommended. Plot Dimensions: Minimum total plot width shall be 150 feet. Wider strips are preferred to reduce impacts of drifting snow. (See restrictions on dimensions in CRP) . Row Spacing: 36. Sorghums - 15 to 30 inches; Dryland corn - 30 to �63 Seed Specifications: Sorghum Patches - At least 60% (75% preferred) of an adapted tall forage sorghum that will stand well with minimal lodging and will mature before frost. Up to 40% can be adapted varieties of grain sorghum. These can be mixed or planted in separate rows (i.e., 2 rows of grain sorghum to 6 rows of forage sorghum. These sorghums should equal a minimum of 75% of the total weight. Maximum amounts for other grains include: Dryland corn (25%), sunflowers (10%) and proso millet (10%). Addition of 1 to 2 lbs./ac. of wild sunflower seed is recommended (Source: Arkansas Valley Seed Company - Denver). Dryland Corn Plots - Early maturing dryland varieties adapted to NE Colorado. Seed from these varieties that is one y~gr· removed from purchased hybrid can be used to reduce seed cost. Planting Dates & Rates: Sorghums - Between April 25 and June 15; Mid to late May is recommended. Plantings conducted after June 15 will be assessed a $10/acre payment reduction. Sorghums should be planted at 4-8 lbs./acre (30-inch rows) and at higher rates if drilled. Dryland corn - Between April 25 and May 15. Plantings after June 1 will not be accepted for payment. Seeding for dryland varieties should be from 10,000 to 13,000 seeds/acre. At least one CUltivation is needed for corn and fertilizer should also be used. Plot Duration: 1 year. Sorghum plantings must remain undisturbed through March of the following year. Dryland cor~must be left standing through March unless harvested. Harvesting may be conducted after March 15 of the following year. SUPPLEMENT FOR SORGHUM PLANTINGS WITHIN CRP sca Notification: The CRP contract must be amended at the local SCS office prior to implementing CP-12 and breaking out food plots within CRP. This requires filling out a one-page form at your SCS office. The ASCS must be advised of the change for their records. Dryland corn is not approved for plots within CRP. Once a winter cover-food plot is broken out within CRP it must remain �64 as such until the end of the CRP contract. Payments will be made annually based on seeded acres. If the farmer wishes to discontinue this practice he must reestablish grass (required by the ASCS). Reimbursement will be at $40.00/acre to cover reseeding grass. Maximum Funded: 1 plot/80-acre be at least 1/4 mile apart. Maximum contain field, 2 plots/160 acres. Plots must Size: The maximum size is 3 acres per site. CRP fields at least 40 acres to be eligible for a CP-12 food plot. must Plot Dimensions: Plantings may be up to 200 feet wide (100 ft in sandy soils). Typical 3-acre plots measure 198 x 660 feet." Where a 100 ft maximum is required a 30 ft wide buffer of untilled grass is left between two 99 x 660 ft parallel strips to obtain a 3 acre plot. Smaller plots should have reduced length to retain at least the 150 ft. minimum width. For example, a plot 99 ft wide x 440 ft. long equals 1 acre and two adjacent plots will exceed the minimum width requirement. Placement: Preferably within 50-100 yds of edge and near cropland, but location can vary depending on soil, wind, and moisture, and location of other winter covers if they occur. Sorghum plantings are not permitted in soils containing free lime (shows effervescence), or soils that are deep sands or choppy sands. �65 RETENTION OF STANDING WHEAT! TALL STUBBLE Application: Where winter wheat on set Aside (ACR) acres is left standing under the ASCS wildlife food plot option. This option must be approved at your ASCS office. The objective is to provide taller, more secure cover for night roosting, feeding, loafing, and escape by pheasants through summer, fall, and winter. A primary concern with respect to pheasants is that unharvested wheat often does not stand well over winter. If the heads can be clipped (with ASCS approval) the resulting cover will be much more valuable to wintering wildlife. Payment Rate & Maximum Funded: (3) Funding to retain uncut wheat under the ASCS Wildlife Option on set Aside tracts ~ill be at $10.00 per acre up to 10 acres and $5.00 per acre for each additional acre up to 20 acres maximum. If the ASCS will permit clipping of heads to retain tall (>20 inch) stubble within Set Aside tracts payment rates will increase to $20.00 per acre for up to 10 acres and $10.00 for each additional acre to 20 acres. Maximum funded is 20 acres per quarter section and 40 acres per section. Soecifications & Retention: Wheat or clipped stubble must remain standing through winter (ungrazed) until March 31 of the following year. Tall annual weeds, if present, must be left standing and can not be treated with herbicides. Treatments will not be funded unless the entire stubble field is to be left undisturbed through the subsequent fall and winter. Placement: Standing wheat patches should be near corn, sorghum, or other cropland preferably within the southeast part of the stubble fields. SUPPLEMENTAL Purpose: To prepare the proper equipment PAYMENTS FOR CUSTOM SITE PREPARATION planting sites when the landowner does not have or does not have time to prepare the site. �66 Treatment: Breaking out small tracts within CRP or sodded waste areas with a mold-board plow or heavy discing to completely destroy existing vegetation for reseeding to switchgrass or planting sorghum patches. Tillage must be to a depth of at least 6 inches. Payment Rate: Payment rate will be $15.00/acre for adequate may involve two to three treatments. Equipment transportation to and between payment will be $15.00/hour. ' site preparation small tracts. which Supplemental �67 JOB FINAL REPORT State of: Colorado Project: W-167-R-5 Upland Bird Research _22_ Work Plan: _-:1,,--_: Job Job Title: Farming for Ring-necked Pheasants - Program Development and Evaluation Period Covered: 01 January 1991 through 31 December 1995 Author: Thomas E, Remington Personnel: L. L, Bixler, C. E. Braun, S. M. DeMasso, J. W. Moore, T. E. Remington, and W. D. Snyder. Colorado Division of Wildlife; K. J. Manfredo, and J. J. Vaske. Colorado St~te University. ABSTRACT A telephone survey was conducted of Colorado small game license buyers to investigate their experience, success. and satisfaction with ring-necked pheasant (Phasianus colchicus) hunting in Colorado. Respondents were also asked several questions designed to ascertain their experience with, and willingness to participate in, fee hunting for pheasants. While 92.4% of respondents had hunted pheasants in the past, only 63% hunted pheasants in Colorado in 1991. Colorado pheasant hunters generally hunted 3 'days or less, bagged 5 or fewer pheasants (31% bagged none). and rated hunting as poor or' fair. Many hunters (23%) had hunted pheasants on shooting preserves, clubs, on land they had leased. or had paid landoltmers for access. Twenty-two percent of respondents reported hunting pheasants out-of-state in 1991. Barriers to participation in pheasant hunting identified by hunters included lack of access to hunting areas, lack of birds, and distance to hunting areas. Only 9% of respondents had no interest in pheasant hunting the following year. Hunter will~ngness to participate in a hypothetical fee hunt of wild pheasants ranged from 75 to 20% as daily fee increased' from $15 to $70. Flush rate, a purported measure of quality, did not influence participation. These data demonstrate substantial interest in pheasant hunting ••and willingness to pay for;,it, by C~lorado .saa'l.L, game hunters. A conceptual model of a communitybased fee 'hunting program was developed to link pheasant hunters willing to pay with interested landowners. ��69 INTRODUCTION Small game license sales have declined markedly in Colorado over the past 10 years, from a high of about 200,000 in 1982 to a low of about 111,000 in 1995. Declines in participation in small game hunting are even more striking when considered as a percentage of Colorado's population. A recurring theme in surveys conducted to identify barriers to participation in hunting has been access to places to hunt, particularly places that have reasonably good hunting (Peterson and Manfredo 1993). Pheasants are the most commonly hunted small game species in Colorado (Braun et ale 1994). Declines in small game hunter numbers may be attributable, in part, to difficulties in acquiring access to places to hunt (pheasants) (Rounds 1975) and/or hunter dissatisfaction because of declines in pheasant populations (Farris and Cole 1981). Thus, pheasants were identified as a pivotal species in any strategy to reverse declines in small game hunter participation, and access to quality pheasant hunting was identified as a key management goal (Braun et al. 1994). Pheasant populations have declined in eastern Colorado because of a lack of survival cover and secure nesting cover (Snyder 1984, 1985, 19~1) caused by intensive farming. Landowners are unlikely to alter agricultural practices to' benefit pheasants or other wildlife, or in some cases allow hunting access, without significant financial incentives (Matulich and Bagwell 1979, Bishop 1981, Rasker 1989). P. N. OBJECTIVES Develop a program to link hunters willing to pay for pheasant hunting opportunity with landowners willing to: 1) provide access for a fee, 2) develop habitat for pheasants within the program area, and 3) amend farming practices to make them more compatible with production and survival of pheasants. RESULTS A conceptual model for a.Pheasant Cooperative Program was developed as a means to develop habitat, increase local pheasant populations, and provide hunting access to interested hunters for a fee (Appendix A). As a prelude to implementation of this program, we conducted a survey to ascertain small game hunter experience, satisfaction, and future interest in pheasant hunting in Colorado. The survey was designed to measure hunter willingness to pay to hunt wild pheasants, and to predict the impact that fee rate and quality of hunting would have on rates of participation in a fee hunting program for pheasants. The results of this survey have been reported previously (Remington 1993), and have been published in a peer-reviewed Journal (Remington, T. E., M. J. Manfredo, J. J. Vaske, and S. M. DeMasso. 1996. Fee hunting pheasants in Colorado: experimental evidence. Human Dim. of Wildl. 1:51-59.). 4 This research has established that small game hunters will support a feehunting program by small game hunters. A work package was submitted to Division administrators to implement a program of this type but has been rated as low priority. If, and when priorities change and a fee hunting program is developed for pheasants we will develop a study plan to evaluate the success of the program. �70 LITERATURE CITED Bishop, R. C. 1981. Economic considerations affecting landowner behavior. Pages 73-87 in R. T. Dumke, G. V. Burger, and J. R. March, eds. Wildlife management on private lands. Wisconsin Chapt. The Wi1d1. Soc., Madison. Braun, C. E., K. M. Giesen, R. W. Hoffman, T. E. Remington, and W. D. Snyder. 1994. Upland Bird Management Analysis Guide, 1994-1998. Colorado Div. Wi1d1., Denver 48 pp. . Farris, A. L., and S. H. Cole. 1981. Strategies and goals for wildlife habitat restoration on private agricultural lands._ Trans. North Am. Wildl. and Nat. Resour. Conf. 46:130-135. , Matulich, G. C., and G. Bagwell. 1979. On-farm pheasants enhancement potentials in irrigated agriculture. West J. Agric. Econ ..4:99-l09. Peterson, M. R., and M. J. Manfredo. 1993. -Pub l.Lc access in Colorado: what are recreationists' perceived problems and preferred solutions? Human Dimensions Persp. 15. 6 pp. Rasker, R. 1989. Agriculture and wildlife: an economic analysis of waterfowl habitat management on farms in western Oregon. Ph.D. Diss., Oregon State Univ., Corvalis. 24lpp. Remington, T. E. 1993. Farming for ring-necked pheasants - program development and evaluation. Colorado Div. Wild1., Prog. Rep., Fed. Aid Proj: W-167-R. Apr.:'25-36. Rounds, R. C. 1975. Public access to priyate lands for hunting. Div. Wildl. Spec. Rep. 2. l79pp. Colorado Snyder, W. D. 1984. Ring-necked pheasant nesting ecology and wheat farming on the high plains. J. Wildl. Manage. 48:878-888. 1985. Survival of radio-marked hen ring-necked pheasants in Colorado. J. Wild1. Manage. 49:1044-1050. 1991. Wheat stubble as nesting cover for ring-necked pheasants in northeastern Colorado. Wi1dl. Soc. Bull. 19:469-474. Prepared by _~\'fl\<}O C;. ~Vn~ Thomas E. Remingt~"'" LSSR IV �71 APPENDIX A PHEASANT COOPERATIVE PROGRAM The Pheasant Cooperative Program is designed to develop habitat and increase pheasant populations by overcoming several significant barriers to habitat improvement for pheasants and other farm wildlife in eastern Colorado; namely providing financial and other incentives for landowners to develop habitat, offering technical assistance, and generating funding. The Division, landowners, and sportsman will be equal partners in improving ph~asant populations and hunting in Colorado. In addition, this program has educational benefits, increases access for hunting, and should significantly increase pheasant harvest and decrease landowner-sportsman conflicts over hunting. The program minimizes CDOW FTE commitments because sportsman do much of the legwork and habitat developments are completed by landowners and/or sportsmen. At least two communities in eastern Colorado have organized programs to match hunters willing to pay to hunt pheasants with landowners willing to provide access for a fee. These are Ringneck Raiders in Yuma, run b~the Future Farmers of America, and Rooster Roundup, rUn by the Burlington Rotary Club. These programs provide and control access but do not improve habitat and consequently have not increased pheasant popUlations. Landowners do not receive any proceeds from these programs because they are used as fund-raisers for the sponsoring groups. Hunter satisfaction is poor and organizers may be tempted to stock pen-raised pheasants to the gun to retain hunter interest and dollars. We propose that the Division assist in the formation of Community Pheasant Cooperatives consisting of individual farmers and sportsman groups. Hunters would pay a fee to hunt lands within the Cooperative. Some of this money would be paid to the landowners, some would pay for habitat developments within the Cooperative, and a small portion would pay expenses of Cooperative administration. The Division of Wildlife would match hunter contributions for habitat development. Community groups, particularly those which benefit from hunting revenue such as gas stations, restaurants, motels, Chambers of Commerce, etc., would be encouraged to join the Cooperative as sponsors by donating $100, $250, $500, etc. for habitat development .• The Division would match these contributions as well. The Division would contract with the Cooperatives while the Cooperatives would contract with the individual farmers. This greatly reduces CDOW FTE requirements to process contracts. Habitat developments would follow prescriptions described in the Pheasant Habitat Improvement Program (PHIP) and would be detailed in individual management plans prepared for each farm in the Cooperative at the outset. It is anticipated that this habitat development money can be used to leverage significant Federal Farm Bill money by paying the farmer's cost share on developments through the Conservation Reserve Program, Forest Stewardship Program, and other Farm Bill programs. Thus, the Division contribution may represent only 25% of total costs. The Cooperative would contract with individual farmers to provide required amounts and quality of habitat, and agree to alter specified farming practices to benefit pheasants (such as timing of stubble tillage in spring to prevent pheasant nest destruction). The Cooperative would also serve as a forum for educational efforts to �72 increase landowner and sportsman knowledge about farming and habitat development practices beneficial to pheasants and other farm wildlife. Many details about how this program would be administered remain to be developed. We believe most details should be left to individual cooperatives since it will be their program. For the Division to participate and cost share developments, habitat plans would have to be developed and habitat quantity and quality goals would have to be met. Cooperatives would be run by a Board consisting of landowner representatives, sportsman representatives, and a CDOW representative. This board would make decisions about.program implementation, monitor compliance, and make cash distributions (of hunter fee money). Examples of questions/answers about program implementation details are illustrated in Table 1. Actual programs may differ from this example. Table 1. Questions/answers about Pheasant Cooperative ,Program implementation. Q. What are significant advantage a of thia program? <, A. Provides monetary return to the farmer for access for hunting and burden of hunter control is shifted from the landowner. Farmers are compensated for developing habitat on productive farm ground and this compensation is from groups directly benefitting, i.e., hunters, motels, restaurants, etc. Will significantly increase pheasant populations and harvest at minimal cost to the Division in dollars and personnel. Access and hunter participation should increase, albeit at a fee. Q. Where will money coae from to develop habita·t? A. From hunter fees (25-50% to habitat, remainder to farmers after costs), and community sponsors (motels, restaurants, Chambers of Commerce, Pheasants Forever, etc.). The Division of Wildlife would match these contributions, and the entire pool would be used to leverage Federal Farm Bill program money. Q. How much will huntera be charged? A. Rates will be set by individual cooperatives and influenced by supply and demand, but $15/day, $25/weekend, $75/season seem reasonable. Possibilities exist for embellishments such as reduced fees or a sliding scale for youngsters, discounts for mid-week, or surcharges for opening weekend. Q. What about enforcement? A. Additional enforcement to prevent trespass will likely be necessary, at least initially. It is anticipated this program will be self-policing to some extent, because both landowners and hunter participants have a vested interest to prevent trespass. Par t LcLpatrl.ng hunters will be identifiable by windshield/dashboard cards and by some form of identification worn on their backs or on their head to facilitate enforcement. Acreage enrolled will be �73 intensively signed (provided by CDOW and erected by cooperative) and participants will be furnished maps. Q. What about liability to landowners and other cooperative members? A. They may be covered under state's general liability immunity (true for participants in South Dakota's pheasant program). If not. liability insurance would be obtained by the Cooperative and paid from fees generated. Q. How will money be paid to landowners equitably. and yet in a manner that encourages habitat and farming practices to benefit pheasants? A. We suggest half of the money to be paid to farmers be allocated based on the amount of land enrolled. The other half would be allocated based on what proportion of pheasants harvested came from that farmers' land. or what proportion of hunters hunted on his land. Evaluation The type, quantity, and rlistribution of habitat developments will parallel those in PHIP; thus. intensive evaluation of pheasant population responses will not be necessary as they can be inferred from evaluation of PHIP. Evaluation will focus on hunter participation, harvest per unit area and effort. and hunter· and landowner satisfaction relative to pre-Cooperative levels and/or similarly farmed areas not in the Program. Acreage of habitat improvements and income generated for farmers will also be documented. ��75 JOB PROGRESS REPORT State of: Colorado Project: W-167-R-5 Upland Bird Research Work Plan: _----'1~2=___: Job _--:l....,8::,...__ Job Title: Genetic Variability In Merriam's Wild Turkeys and Its Relationship to Reproductive Performance Period Covered: 01 January through 31 December 1995 Author: Richard C. Dujay Personnel: Richard W. Hoffman; Colorado Division of Wildlife; Richard C. Dujay, Colorado State University ABSTRACT Blood samples were collected from 230 Merriam's wild turkeys (Meleagris gallopavo merriami) trapped (n - 225) and/or harvested (n - 5) in Colorado between September 1995 and February 1996. Samples were obtained from 3 East Slope and 5 West Slope populations. In addition, samples were obtained from the Sacramento Mountain population in New Mexico (n - 5) and the Mogollon Rim population in Arizona (n - 32). DNA was successfully extracted from 260 of the 267 samples processed. Five randomly-selected samples assayed using spectrophotometry showed concentrations of DNA ranging from 0.50 to 1.01 ~g/~l at a wave length of 260 nm. Once purification and quantification of the extracted samples are completed, each sample will be subjected to restriction length polymorphism and/or microsatellite analysis using polymerase chain reaction techniques. Collection and analysis of samples will continue in 1996-97. ��77 GENETIC VARIABILITY IN MERRIAM'S WILD TURKEYS AND ITS RELATIONSHIP REPRODUCTIVE PERFORMANCE Richard TO C. Dujay INTRODUCTION Genetic considerations have been largely ignored in wild turkey (Meleagris gallopavo) restoration and range expansion programs due to the lack of cost-effective techniques for examining genetic attributes of populations (Leberg 1990, Stangel et al. 1992). Establishing high genetic diversity in introduced populations is considered desirable because.the level of genetic diversity influences the population's growth rate and ability to adapt to new environmental conditions (Leberg 1990, Vrijenhoek and:Leberg 1991). Practices common in wild turkey transplant programs that may contribute to. low genetic diversity in new populations include releasing more females than males, releasing birds captured from the same flock, and obtaining release stock from the same population (Leberg 1990, 1991). This lack of genetic diversity in the released stock may explain why some wild turkey populations experience consistently low reproductive success and fail to increase even when habitat conditions appear suitable. Supplemental releases into genetically stressed populations may be one way to enhance genetic vigor and stimulate reproductive performance. However, before any further manipulations are attempted, as much genetic information as possible must be gathered from existing populations (Leberg et al. 1994). With advancements in DNA technology, large scale surveys can now be economically conducted over broad geographic areas to assess the current genetic condition of populations and to evaluate the genetic consequences of past management activities. These genetic data are essential in prescribing supplemental releases and in directing management efforts towards other factors, such as habitat quality, if the data reveal no genetic problems. P. N. OBJECTIVES The objectives of this study are to (1) compile a database of genetic structure and variation among populations of Merriam's wild turkeys (M. g. merriami) in Colorado, (2) compare genetic diversity of introduced/reestablished populations with the original source population, (3) compare genetic diversity of Merriam's wild turkey populations in Colorado with nonmanipulated (pure) populations of Merriam's turkeys in Arizona and New Mexico, and (4) identify populations that are genetically stressed and develop management recommendations to enhance the genetic attributes of these populations. �78 SEGMENT OBJECTIVES 1. Review literature pertinent to the objectives of this study. 2. Collect blood samples, extract DNA and electrophoretically survey for genetic diversity (i.e., heterozygosity, number of alleles per locus, and percent polymorphic loci) within an introduced population (Larimer County) suspected of experiencing a genetic bottleneck. 3. Collect blood samples and determine genetic diversity within the source population (Spanish Peaks Wildlife Area). 4. Collect blood samples and determine genetic diversity within potentially unrelated populations in Arizona, New Mexico, South Dakota, and elsewhere in Colorado. 5. Depending upon results of the DNA analyses, develop a study plan to investigate the effects on reproductive performance of supplemental releases (i.e., introduction of new genetic material) into the Larimer County population. 6. Trap and radiomark 40 female wild turkeys within the Larimer County population to ascertain habitat use, movement patterns, and reproductive performance prior to conducting any supplemental releases. 7. Compile data, analyze results, and prepare progress report. STUDY AREA AND METHODS The following populations/geographic areas were selected for sampling: 1. Larimer County - between the Poudre and Big Thompson drainages. This area is classified as nonhistoric turkey range. The population originated from the release of 15 birds (8 males, 7 females) captured in 1957 at what is now the Spanish Peaks State Wildlife Area. 2. Boulder County - between Lefthand and Saint Vrain creeks. There are no records of any releases in Boulder County. This population occurs at the northern periphery of the native distribution of Merriam's wild turkeys and probably originated from the natural expansion of native populations northward. However, it is also possible that birds from the introduced population in Larimer County expanded southward into Boulder County. 3. Las Animas County - on and surrounding the Spanish Peaks State Wildlife Area. Spanish Peaks and Devil's Creek State Wildlife Area in Archuleta County have been the primary sources of birds for transplant programs elsewhere in the state. Situated within the core of the native distribution of Merriam's turkeys, Las Animas County supports some of the highest densities of turkeys in Colorado and is the leading harvest area in the state. No recent transplants have been made into Las Animas County. However, in the 1940's and 1950's, male turkeys were �79 interchanged between Las Animas breeder exchange program. and Archuleta counties as part of a 4. Archuleta County - including Devil's Creek State Wildlife Area and Cat Creek (Southern Ute Indian Reservation). This area also is within the core of the native distribution of Merriam's wild turkeys. Supplemental releases were made into Archuleta County (specifically at Devil's Creek State Wildlife Area) during the 1940's and 1950's as part of a breeder exchange program. 5. Montezuma County - including Boggy Draw and Hartman Draw. This area has a history of population declines followed by reintroduction programs. The first decline occurred prior to 1940. Reintroductions were made during the early 1940's using birds trapped at the Devil's Creek State Wildlife Area. By the early 1960's, turkeys were presumed extirpated from the area. Reintroduction efforts were init~ated in 1983 using birds trapped from Las Animas and Pueblo counties. The popUlation has substantially increased in recent years and has: provided a source of transplant stock for .other areas in southwestern Colorado .. 6. Montrose County - between the Dave Wood and Delta-Nucla roads. Historical records suggest wild turkeys are native to Montrose County, but disappeared from the area by 1900. Restoration attempts were first made in 1934 using birds obtained from private sources in Texas, Oklahoma, and New Mexico. These may have been captive birds, and some were eastern (M. g. silvestris) or more likely Rio Grande (M. g. intermedia) wild turkeys. Few, if any birds, remained by 1940. New attempts to restore the population began in 1944 and continued through 1949 using birds trapped in Archuleta County. The reintroduced population increased and expanded its range through the early 1960's, when another crash occurred. A third restoration attempt was begun during the early 1980's using source stock from Las Animas and Pueblo counties. Some turkeys still remained in the area when the third restoration program was initiated. The restored popUlation has increased and expanded into formerly occupied habitats. 7. Mesa County - within the Plateau Valley. There are no records of turkey releases in the Plateau Valley. However, releases .were made in the 1950's and 1980's near Rifle along Divide and Beaver creeks. Turkeys also were released near Cedaredge on the south side of Grand Mesa. It is possible birds from these releases expanded their range into Plateau Valley. 8. Garfield County - including Beaver Creek, Divide Creek, and Rifle Creek. This area, along with Plateau Valley, is considered non-historic turkey range. Releases made in the 1950's were comprised of birds trapped in southwestern Colorado, primarily from Archuleta County. Releases in the 1980's were from birds trapped in Las Animas County. More recently, birds have been trapped and moved within the Garfield County area. 9. Mogollon Rim, Chevelon Ranger District, Apache-Sitgreaves National Forest, southwest of Flagstaff, Arizona. This area was selected because it supports one of the few remaining unmanipulated populations of Merriam's wild turkeys within the native distribution of the subspecies. �80 Turkeys have been trapped and moved from the Mogollon Rim, but no birds from other populations have been released into the area. 10. Sacramento Mountains, Cloudcroft Ranger District, Lincoln National Forest, southeastern New Mexico. This is another unmanipulated population of Merriam's wild turkeys that is isolated from other populations. Turkeys were baited with oat hay and corn and live-trapped using cannon nets, drop nets, or box traps. Captured turkeys were classified ,as to age and sex, and banded with serially-numbered aluminum leg bands. Ages were recorded as subadult (8-10 months) or adult (> 18 months). Blood was collected by jugular venipuncture. At least 1.5 mls of blood were collected from each bird and placed in EDTA purple top vials for DNA analysis. Another 4-6 mls of blood were collected from a random sample of birds from each flock for disease testing. Samples collected for DNA analysis were s.to red in a -20 C freezer until extractions were performed. DNA extractions were done using Analytical Genetic Testing Center's Quik Gene Kit with adapted protocol f:or avian blood. The whole blood samples were thawed and a volUme of 0.250 mls was drawn from each vial for DNA extraction. The unused portion of whole blood was re-frozen and stored as a backup sample for later extraction. The extraction protocol was: 1. 0.250 ~l of whole avian blood was placed into a graduated centrifuge tube. conical 2. 2.5 ml of ice-cold red blood cell (RBC) lysis buffer (IX) was added to the whole blood, shaken to mix, and vortexed for 30 seconds. 3. The sample was placed on ice 10 minutes minutes at 3500 rpm. 4. The supernatant was decanted and discarded. still lumpy, steps 2 and 3 were repeated. 5. 1 ml of RBC lysis buffer (IX) was added and the pellet was rinsed. The pellet was not rinsed if it was necessary to pipette the supernatant. 6. 4 ml of white blood cell (WBC) lysis buffer (IX) was added to the sample and the mixture was shaken until viscous, then vortexed for 5-10 seconds. 7. The sample was incubated water bath. 8. 200 ~l 10% SDS, 500 ~l protein precipitant solution, and 1 ml of ultra pure water were added, ,and the mixture was vortexed for 1 minute. 9. The sample was again incubated and then centrifuged for 30-45 minutes 25 If the sample was at 55 C in a shaker in S5 C water bath for 20 minutes �81 and then centrifuged at 3500 rpms for 25 minutes. If the sample was not clear after spin, 1/2 volume of ultra pure water and 200 ~l of SDS were added, then the sample waS vortexed 30 seconds, incubated in a 55 C water bath for another 20 minutes and centrifuged at 3500 rpms for 25 minutes. 10. The clear supernatant was pipetted into a clean test tube and 2-3 volumes of ice-cold absolute alcohol were added to precipitate the DNA. 11. The DNA was removed with a sterile glass hook, dried slightly, and dissolved in a microcentrifuge tube containing 700 ~l of TE buffer (EDTA TRIS at 7.4 pH). 12. The sample was refrigerated at 4 C for 24-48 hours, then placed into a -20 C freezer for short term storag~ (3-6 months) or a -70 C freezer for long'term storage. The washing and purification procedure involved thawing the extracted DNA samples and placing the solution into a 15 ml conical centrifuge tube. The volume of solution was determined, and 10% SDS and protein precipitant were added at 50 ~l and 90 ~l per 1.0 ml of DNA solution, respectively. The samples were mixed thoroughly by vortexing and allowed to stand at room temperature for 10 minutes. The samples were then centrifuged at 3500 rpms for 25 minutes and the supernatant was decanted or pipetted into a clean centrifuge tube, where the DNA was precipitated using absolute ethanol. The DNA was collected and dissolved again in TE buffer and frozen at -20 C. Once the purification procedures are completed, restriction length polymorphism and/or microsatellite analysis using polymerase chain reaction techniques will be performed on several samples to determine which method produces the best results. These analyses will be performed at the Analytical Genetic Testing Laboratory in Denver. RESULTS AND DISCUSSION Blood samples were collected from 267 Merriam's wild turkeys trapped (n (n - 19) in Colorado, Arizona, and New.Mexico (Table 1). Within Colorado, samples were obtained from 3 East Slope and 5 West Slope populations. DNA was extracted from 260 of the 267 samples processed. All samples failing to yield any DNA were from hunter-harvested birds. Five randomly-selected samples subjected to spectrophotometric analysis showed moderate to high concentrations (0.50-1.01 ~g/~l) of high quality (absorbency 260 nm 0.100-0.202) DNA. -248) or harvested Power calculations were performed using SAS statistical software in a completely randomized design. Calculations that compared each of the 10 populations under investigation (n1 - nlO) to the pooled sample (N) indicated that a power> 0.80 was obtained when any n = 20 and N > 300. To assure confidence in statistical analysis, the minimum sample size from any population should be > 20. This indicates that additional samples should be obtained from Boulder County, Larimer County, Las Animas County, Garfield County, and the Sacramento Mountains in New Mexico. �82 Trapping required more time than expected because of the statewide scope of the project and because the lack of snow made it difficult to attract birds to prebaited trap sites. The extraction of DNA from the samples also required more time than expected because of the tedious nature of the extraction protocol and the large number of samples that needed to be processed. For these reasons, it was decided not to prepare a study plan and proceed with the radiomarking segment objectives until the analyses of the DNA samples were completed. Table 1. Blood samples collected Arizona, and New Mexico, 1995-96. from Merriam's wild turkeys in Colorado, Location Boulder County Larimer County Las Animas County Montezuma County Archuleta County Montrose County Mesa County Garfield County Sacramento Mountains, Mogollon Rim, AZ Total 14 11 4 61 33 38 51 18 5 32 NM (N) 267 LITERATURE CITED Leberg, P. L. 1990. Genetic considerations in the design of introduction programs. Trans. North Am. Wildl. and Nat. Resour. Conf. 55:609-619. 1991. Influence of fragmentation and bottlenecks on genetic divergence of wild turkey populations. Conserv. Biol. 5:522-530. _____ , P. W. Stangel, H. O. Hillestad, R. L. Marchinton, and M. H. Smith. 1994. Genetic structure of reintroduced wild turkey and white-tailed deer populations. J. Wildl. Manage. 58:698-711. Stangel, P. W., P. L. Leberg, and J. I. Smith. 1992. Systematics and popUlation genetics. Pages 18-28 in J. G. Dickson, ed. The wild turkey �83 biology and management. Stackpole Books, Harrisburg, Pa. Vrijenhoek, R. C., and P. L. Leberg. 1991. Lets not throw the baby out with the bath water: a comment on management for MHC diversity in captive populations. Conserv. BioI. 5:252-254. Prepared by: R"chard C" Dujay Research Te h. I Approved by: : ��85 JOB PROGRESS REPORT State of: Colorado Project: W-167-R-5 Work Plan: Job Title: 13 Upland Bird Research Job: 10 Movements. Reproductive Success. and Habitat Use by Introduced -, Plains Sharp-tailed Grouse Period Covered: 01 January through 31 December 1995 Author: Kenneth M. Giesen Personnel: Jim Aragon, Clait E'.Braun, Kenneth M. Giesen, Chuck Loeffler, Colorado Division of Wildlife ABSTRACT A total of 43 plains sharp-tailed grouse (Tyrnpanuchus phasianellus jamesi) was trapped in southeastern Wyoming and transplanted onto Raton Mesa in Las Animas County, Colorado in April 1995. Ten of 20 males and 12 of 23 females were fitted with radio tra~mitters prior to release. Weather conditions at the time of release included blizzard conditions with cold temperatures, high winds, and> 1.0 m of snow. Mortality of radio-marked birds was 55.5% within 60 days postrelease with raptor predation a major cause. Dispersal from the release site ranged from 1.2 to 11.1 km with 8 of 18 birds moving ~ 6.0 km. Four radio-marked birds were not relocated following release. Home ranges of 3 birds surviving 60 days ranged from 0.48 to 1.61 km2• Height density indices within grasslands used by radio-marked birds ranged from 1.56 ± 0.77 dm to 5.52 ± 1.62 dm. Sightings of unmarked birds and survival of some radiomarked birds through December 1995 indicates that spring through fall habitat on Raton Mesa meets the minimum requirements for this species. ��87 MOVEMENTS, REPRODUCTIVE SUCCESS, AND HABITAT USE BY INTRODUCED PLAINS SHARP-TAILED GROUSE Kenneth M. Giesen INTRODUCTION Plains sharp-tailed grouse historically occurred in suitable foothill and riparian habitats along the Front Range of Colorado. Sharp-tailed grouse populations declined with human settlement and were extirpated from most of their range in eastern Colorado by the late 1800's. Although the historical breeding population of sharp-tailed grouse in Douglas County continue to decline, small breeding populations and winter migrants or transients have been reported in recent years from Yuma, Logan, and Weld counties (Hoag and Braun 1990; C.E. Braun unpubl. data). However, the total breeding population of plains sharp-tailed grouse in Colorado remains small. «300 birds) and most populations are associated with' privately-owned lands and, therefore, subject to land management activities which may have detrimental consequences. Plans to increase distribution and populations of plains sharp=tailed grouse in Colorado will rely primarily on transplants (Braun et al. 1992). While numerous transplants of prairie grouse have been attempted in North America, few have been successful (Toepfer et al. 1990, Hoffman et al. 1992, Rodgers 1992). A previous transplant of sharp-tailed grouse into Las Animas County near Raton Mesa was attempted with a total of 85 males and 83 hens being released over a 3-year period (1987-89). While this transplant was not thought to be successful in establishing a breeding population, little followup of released birds was conducted and important data on survival and movements are lacking. Thus, it is desirable to document responses of sharptailed grouse to experimental transplants and evaluate parameters potentially aff~cting success including movements, habitat use, mortality, and reproduction. P. N. OBJECTIVES The objectives of this project are to assist with trapping and transplanting of plains sharp-tailed grouse into selected sites along the Front Range of Colorado and evaluate transplant success. Population characteristics of the transplanted population including movements and home range size, timing and causes of mortality, habitat use, and nest success will be compared to those described in the literature for native and transplanted prairie grouse. Results of this study will assist in developing transplant protocols for future transplants of prairie grouse. SEGMENT OBJECTIVES 1. Review literature habitat use. on prairie grouse introductions, movements, and 2. Coordinate efforts with Wyoming Game and Fish personnel and affected landowners in southeastern Wyoming to locate potential trapping sites for plains sharp-tailed grouse. �88 3. Transplant up to 50 plains sharp-tailed grouse from southeastern into suitable habitats along the Front Range of Colorado. 4. Radiomark up to 25 sharp-tailed grouse in the transplanted population and monitor movements, habitat use, reproduction, and mortality. 5. Conduct site. 6. Prepare annual progress a pre-release evaluation of the habitat at the selected Wyoming release report. METHODS Contact was made with personnel of the Wyoming Game and Fish Department to obtain permits for trapping plains sharp-tailed grouse jn southeastern Wyoming for transplant into Colorado. Active dancing grounds in southeastern Wyoming were located as potential trapping sites and permissiQn from affected landowners was obtained. Walk-in funnel traps (Toepfer et al. 1988, Schroeder and Braun 1991) were used to capture male and female sharp-tailed grouse on dancing grounds. Captured birds were classified to age and sex and fitted with serially-numbered aluminum bands on the right leg and yellow colored plastic bandettes on both legs. Battery-powered transmitters (weight 12-13 gms) were attached with a necklace (Amstrup 1980) to selected birds to facilitate monitoring of movements and survival after release on Raton Mesa. A portable Global Positioning Satellite (GPS) receiver was used to record locations of birds and minimum convex polygon home ranges were calculated using the McPaal software package (M. Stuwe and E. E. Blohowiak, Conserv. Res. Cent., Natl. Zool. Park, Smithsonian lnst., Front Royal, Virginia, 1985). Vegetative cover (height-density indices) in the release area was measured using a Robel pole (Robel et al. 1970). RESULTS Transplant of sharp-tailed grouse A total of 43 (20 males, 23 females) sharp-tailed grouse was captured in Platte and Goshen counties, Wyoming and released onto Raton Mesa in Las Animas County, Colorado in April 1995. These birds were trapped on 4 dancing grounds (Baker Swale = 6 males, 4 females; Gladys'- 4 males, 1 female; Thomas Jeffersons - 3 males, 12 females; Grange - 7 males, 6 females). Captured birds were held in captivity up to 4 days prior to release. Birds were transported by truck and helicopter to the release site. At the release site birds were placed in holding boxes for 5-10 minutes before the doors were opened to allow escape. Twenty-one birds (20 males, 1 female) were released on 11 April, 5 hens were released on 15 April, and 17 hens were released on 21 April. Ten male and 12 female sharp-tailed grouse were fitted with radio transmitters prior to release. Weather conditions at the time of release were less than ideal. Unusually cold and wet weather in March and April resulted in 100% snow cover (depth> 1.0 m) at the release site and adjacent areas on Raton Mesa when birds were released. These weather conditions persisted into late May. Snow conditions �89 and restricted access on Raton Mesa hampered weeks following release. monitoring birds for several Mortality Mortalities of 12 released birds (7 males,S females) were documented from 11 to 60 days post-release (Table 1). Mortalities were documented from 1.24 to 11.16 km from the release site with most occurring within 30 days of release. Four birds (1 male, 3 females) were not located (no radio signals received) following their release and it is suspected that most dispersed from the study area and likely died soon after. Several attempts to locate these radio signals from aircraft were unsuccessful. The high initial mortality (minimum 60%) was likely caused by the severe weather conditions at time of release and the total snow cover at the release site. Most of the known causes of mortality were attr~buted to raptor depredation and may have been influenced by the condition of the birds following capture, captivity (1-4 days), and lack of food availability following release. Table 1. Fates of radio-marked sharp-tailed Las Animas County, Colorado, 1995. Bird II Age Sex Max Dist" (m) 0925 1045 1184 1254 1275 1334 1534 1655 1684 1745 1765 1775 1805 1835 1845 1875 1885 1904 1945 1955 1965 1994 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 1+ 2+ 2+ 2+ 2+ F M M M F F F M F M M F M M n d ." i 7,530 8,410 8,000 (est.) 1,400 1,240 8,200 1,230 n.d. F M F F n.d. n.d. F 1,750 M on Raton Mesa, Home Range (km2) 2,530 2,250 11,160 7,000 (est.) 6,000 (est.) 3,580 6,024 2,220 2,120 3,560 F F grouse released 1.607 0.476 0.137 Fate Raptor kill, 3 May Raptor kill, 2 May Raptor kill, ? May Mortality signal, May Mortality signal, Jun No signal postrelease Last signal 29 Jun Radio fell off, 13 Sep Mortality signal, May Raptor kill, 2 May Raptor kill, 2 May Alive in Sep Radio fell off, 14 Jun No signal postrelease Last signal 2 May Morality, 18 May Mortality signal, 3 May Mortality, 2 May Raptor kill, 3 May No signal postrelease No signal postrelease Last signal 2 May " Maximum distance from release site. No data, bird not located after release onto Raton Mesa. b �90 Movements and Home Range The distribution of movements from the release site was bimodal with one group of birds staying relatively close to the release site while another group established ranges ~ 6.0 km from the release site (Table 1). High mortality of some birds soon after release may have skewed innate movement patterns, although some birds dispersed> 6.0 km and died away from Raton Mesa within 13 weeks after release. Some of these birds were not located or the radios recovered although mortality signals were received for >30 days. Although sample sizes were small, it did not appear that post-release move~ents were related to sex of the grouse, with both males and hens showing long dispersal movements. Minimum convex polygon home ranges were calculated for 3 grouse (2 males, 1 female) which were located periodically for> 60 days after release. Home ranges were relatively small (0.137 - 1.697 km2, Table l) and may have been affected by the few locations of each bird. However, consecutive locations at 1-2 week intervals showed that birds were usually within 200-400 m of their previous location. Habitat Height-density of vegetation was measured during June-August at 150 points within 3 pastures (50 measurements/pasture) where radio-marked sharp-tailed grouse were regularly located. The greatest height-density was measured at the release site pasture (5.52 ± 1.62 dm). Two grazed pastures in New Mexico where sharp-tailed grouse occurred during May-September had mean heightdensity measurements of 1.56 ± 0.77 dm and 1.95 ± 1.17 dm. However, heightdensity was quite variable in these pastures with 33% having height-density measurements ~ 2.25 dm, and 15% ~ 3.0 dm. The height-density measured in 1995 likely reflected the high precipitation the areas received during March-June. DISCUSSION Results from the initial year of this transplant indicate high mortality of released birds and dispersal from the release site. The long movements may have been related to the snow cover and weather conditions at time of release. However, Hoffman et al. (1992) recorded post-release movements up to 29 km for greater prairie-chickens (Tympanuchus cupido) transplanted in spring. High winds and the location of the release site relative to the edge of the mesa likely resulted in some birds flushing from the mesa top and landing in forested habitats. Because the elevation on top of Raton Mesa is 300-600 m above the surrounding terrain, any birds leaving the mesa top may have perished before finding their way back. No birds were known to leave the top of the mesa and return. The west, north, and east sides of Raton Mesa are surrounded by forests and steep cliffs; conditions which may hinder return movements to the mesa top. Furthermore, cold weather and persistent snow cover at the time of release may have increased energetic costs for the birds at a time when food resources were most restricted. These conditions, combined with several days of captivity, may have weakened the birds and increased their susceptibility to predation. �91 It is not known whether the radio-marked birds suffered higher mortality than those not marked as reported in other studies (Marks and Marks 1987). Sharptailed grouse without radios were observed regularly following the release indicating actual survival may have been higher than indicated by radio-marked birds. While no nesting attempts were documented, it is possible that reproduction may have occurred by unmarked birds and was not detected. The success of this project may depend upon the successful release of additional sharp-tailed grouse in 1996. If weather conditions are favorable, mortality and dispersal movements of released birds may be reduce~ and chances for reproduction enhanced. One factor not studied is the availability of winter food for this population. Waste grain from wheat fields was available in Wyoming where these grouse were captured but was lacking on Raton Mesa and adjacent areas. If the grouse released onto Raton Mesa need to disperse into New Mexico to find winter food, they may not return to breed near their release site. LITERATURE Amstrup, S. C. 1980. A radio-collar 217. Braun, CITED for game birds. J. Wildl. C. E., R. B. Davies, J. R. Dennis, K. A. Green, 1992. Plains sharp-tailed grouse recovery plan. Denver. 33 pp. Manage. 44:214- and J. L. Sheppard. Colorado Div. Wildl., Hoag, A. W., and C. E. Braun. 1990. Status and distribution tailed grouse in Colorado. Prairie Nat. 22:97-102. of plains sharp- Hoffman, R. W., W. D. Snyder, G. C. Miller, and C. E. Braun. 1992. Reintroduction of greater prairie-chickens in northeastern Colorado. Prairie Nat. 24:197-204. Marks, J. S., and V. S. Marks. 1987. Influence of radio-collars sharp-tailed grouse. J. Wildl. Manage. 51:468-471. Robel, R. J., J.N. Briggs, A.D. Dayton, and L.C. Hulbert .•1970. Relationships between visual obstruction measurements and weight of grassland vegetation. J. Range Manage. 23:295-297. Rodgers, R. D. 1992. A technique for establishing sharp-tailed unoccupied range. Wildl. Soc. Bull. 20:101-106. on survival grouse in of Schroeder, M. A., and C. E. Braun. 1991. Walk-in traps for capturing prairie-chickens on leks. J. Field Ornithol. 62:378-385. Toepfer, J. E., R. L. Eng, and R. K. Anderson. 1990. Transplanting pra1r1e grouse: what have we learned? Trans. North Am. Wildl. and Nat. Resour. Conf. 55:569-579. �92 _________ , J. A. Newell, and J. Monarch. 1988. A method for trapping prairie grouse hens on display grounds. Pages 21-23 in A. D. Bjugstad, Tech. Coord. Prairie chickens on the Sheyenne National Grasslands. U.S. Dep. Agric., For. Servo Gen. Tech. Rep. RM-159. Prepared by Kenneth M. Giesen LSSR IV �93 JOB FINAL REPORT State of: Colorado Project: Upland Bird Research W-167-R-5 : Job _7_ 'Work Plan: 17 Job Title: Population Period Covered: Dynamics 01 January of White-tailed Ptarmigan 1991 through 31 December 1995 Clai"t E. Braun and Kenneth M. Giesen Author: Personnel: Kathy Martin, University of British Columbia; Clait E. Braun Kenneth M. Giesen, 'Colorado Division of 'Wildlife and ABSTRACT Long-term studies of populations of white-tailed ptarmigan (Lagopus leucurus) at hunted (Mt. Evans) and unhunted (Rocky Mountain National Parkr-areas in Colorado that started in 1966, ceased after the 1995 field season. Data analysis and manuscript preparation will continue under Work Plan 22, Job 1. Major publications in the 1991-95 period resulting from this study were: Braun, C.E., D.R. Stevens, K.M. Giesen, and C.P. Melcher. 1991. Elk, whitetailed ptarmigan and willow relationships: a management dilemma in Rocky Mountain National Park. Trans. North Am. Wildl. and Nat. Resour. Conf. 56:74-85. _________ , K. Martin, and L.A. Robb. 1993. White-tailed ptarmigan (Lagopus leucurus) in A. Poole and F. Gill, eds. The Birds of North America. Acad. Nat. Sci., Philadelphia, Pa. and Am. Ornithol. Union, Washington, D.C. 68. 24 pp. Giesen, K.M., and C.E. Braun. 1992. characteristics of white-tailed 104:263~272: ___________ , and juvenile white-tailed Prepared by Winter home range and habitat ptarmigan in Colorado. Wilson Bu LL; 1993. Natal dispersal and recruitment of ptarmigan in Colorado. J. Wildl. Manage. 57:72-77. ~E.~ Clait E. Braun Wildlife Research Leader ��95 JOB PROGRESS REPORT State of: Project: Colorado W-167-R-5 Upland Bird Research Work Plan: 22 Job Title: Upland Bird Research Publications Period Covered: Author: Job _1_ 01 January through 31 December 1995 Clait E. Braun , Personnel: Clait E. Braun, K. M. Giesen, R. W. Hoffman, T. E. Remington, and W. D. Snyder, Colorado Division of Wildlife ABSTRACT The following articles were published in 1995: Braun, C.E. 1995. Distribution and status of sage grouse in Colorado. Prairie Nat. 27:1-9. Giesen, K.M. 1995. Re-establishment of plains sharp-tailed grouse in Colorado. Proc. Prairie Grouse Tech. Counc. 2l:Abstract. Remington, T.E. 1995. Reaping what you sow. Colorado Outdoors 44(5):26-28. Schulz, J.H., S.L. Sheriff, Z. He, C.E. Braun, R.D. Drobney, R.E. Tomlinson, D.O. Dolton, and R.A. Montgomery. 1995. Accuracy of techniques used to assign mourning dove age and gender. J. Wildl. Manage. 59:759-765. Prepared by __ ~~=~'.:..__.....;2=---._ . ..!..~~==r....=:..._. _ Clait E. Braun Wildlife Research Leader ��97 JOB State of: PROGRESS REPORT Colorado Project: Upland Bird Research W-167-R-5 : Job _1_ Work Plan: 26 Job Title: Analysis of Upland Bird Population Trends Period Covered: Author: 01 January through 31 December 1995 Clait E. Braun Personnel: C1ait E. Braun, Kenneth M. Giesen, Richard ·W. Hoffman, Thomas E. Remington, and Warren D. Snyder, Colorado Division of Wildlife ABSTRACT The following reports were prepared in 1995. Braun, C. E. 1995. Blue Mountain sage grouse harvest data, 1976-95. 1995. Cold Spring Mountain harvest data, 1976-95. 1995. Eagle County sage grouse harvest data, 1977-95. 1995. Sage grouse harvest data, Eastern Moffat and Western Routt counties, Colorado, 1976-95. 1995. Gunnison Basin sage grouse harvest data, 1977-95. 1995. Middle Park sage grouse harvest data, 1975-95. 1995. Northcentral Moffat County sage grouse haryest data, .1976-95. 1995. No rtih Park sage grouse harves~ dat.a.,1973-95 .. 1995. Piceance Basin sage grouse harvest data, 1977-95. 1995. Yampa Area sage grouse harvest data, 1995. Commons, M. L., C.A. Brigham, and C. E. Braun. 1995. Sage grouse investigations, Dry Creek Basin/Miramonte Reservoir, San Miguel County, Colorado, March-August 1995. Giesen, K. M. 1995. Columbian sharp-tailed grouse harvest data, northwest Colorado, 1976-95. �98 1995. Hatching chronology of Columbian sharp-tailed grouse in northwestern Colorado, 1980-1995. Hoffman, R. W. 1995. 1995. Analysis of statewide blue grouse wing collections for _____ , and M. T. Tucker. 1995. Analysis of ruffed grouse range expansion program. Joint Rep., Colorado Div. Wi1d1. and U. S. For. Serv., Region 2. Remington, T.E. 1995. The quail alternative. 1995 Colorado hunting guide. Colorado Div. Wi1d1., Denver. pp. 8-11. Woods, C.P., and C. E. Braun. 1995. Sage grouse investigations, Glade Park and Pinon Mesa, Mesa County, Colorado, April-September 1995. Prepared by »» .L2.~~ ~~ ~~~~ C1ait E. Braun Wildlife Research Leader _ �99 JOB FINAL REPORT State of: Colorado Project: W-l67-R-5 Work Plan: Job Title: 27 Upland Bird Research 1 : Job Experimental Range Period Covered: 01 January Author: W. Hoffman Personnel: Richard Expansion through of Ruffed Grouse 31 December in Colorado 1995 Clait E. Braun, Richard W. Hoffman, Richard M. Lopez, Jim Olterman. Robin Olterman, Colorado Division of Wildl~fe; Kathy Peckham, Mark Tucker, U. S. Forest Service. A. ABSTRACT No ruffed grouse (Bonasa umbellus) were trapped and transplanted into Colorado. Attempts by the Colorado Wildlife Commission to obtain a compromise position between opposing sides of the issue were unsuccessful. Thus, due to public opposition, lack of support from the U. S. Forest Service, disagreements internally, and concern about a prolonged legal process, the Colorado Division of Wildlife has indefinitely delayed activity on this project. ��101 JOB PROGRESS REPORT State of Colorado Project: W-171-R-I: Work Plan __l_ : Job __l_ Bald Eagle Nest Site Protection Job Title: Bald Eagle Nest Site Protection Period Covered: and and Enhancement Program Enhancement Program I July 1995 - 30 June 1996 Personnel: G.R. Craig, Colorado Division of Wildlife ABSTRACT Bald eagles occupied 26 Colorado nesting territories in 1996. Five new territories was discovered and one that bad been vacant in 1995 was reoccupied. Nineteen territories hatched young and all pairs fledged 32 young. Productivity -averaged 1.23 young per occupied territory. / This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the author. ��103 BALD EAGLE NEST SITE PROTECTION AND ENHANCEMENT PROGRAM Gerald R. Craig SEGMENT OBJECTNES 1. Monitor nest site occupancy and reproductive success. 2. Document survival rates and mortality factors. 3. Determine migration and wintering areas. 4. Determine ifpbilopatry occurs in breeding eagles. 5. Determine nest site tenacity by individual breeding eagles. 6. Quantify nesting habitats and associated foraging areas in an effort to document nest site parameters conducive to improved reproduction. 7. Document pesticide contamination through eggshell measurement and chemical analysis of nonviable eggs. 8. Where necessary, implement actions to stabilize nests and maintain occupancy. METHODS 1. Ammally visit all documented breeding sites to determine the presence of bald eagles. Pairs at territories will be documented by DWMs and other fiekl personnel. Previously unrecorded pairs will be revealed in the course of aerial eagle and waterfowl flights. DWM's will confirm actual incubation from ground visits. 2. Occupied territories will be visited by DWM's periodically throughout the breeding season to determine hatch of young, nesting failures, etc. 3. In May and June, a Utility Worker will observe breeding eagles from a distance and endeavor to follow their movements to locate important foraging areas. Responses of eagles to various human activities and land uses will be recorded. 4. In June, when the young are determined to be old enough to band, sites will be' visited by Craig and Knight to place a federal band on one leg and a colored, alpha numeric marker on the other. The color markers will permit identification if the young return in subsequent years. During the same nest visit the following will be recorded: Physical parameters such as tree species, height, DBH, condition, and dominance. Nest condition, size, and location. Vegetative community and land use practices. In addition, prey remains, nonviable eggs, and eggshell fragments will be collected. 5. Approximately Sec's of blood will be collected from each nestling. The blood will be analyzed at the Savannah River Ecology Lab in Aiken, South Carolina. Electrophoretic examination will permit genetic comparison with samples collected from other populations in Saskatchewan, the Lake States and Arizona, as well as determine the heterogeneity of the Colorado birds. 6. When necessary, remedial actions will be taken to stabilize nests that are threatened by wind throw. Should the tree be decadent and in danger of falling, an artificial nest base may be placed in a suitable, adjacent tree. Action will be taken only after it bas been deemed desirable to encourage the eagles to nest at the same location. �104 RESULTS AND DISCUSSION Territory Occupancy Bald eagle nesting activities in Colorado have varied (Fig. 1 am Table 1).. In 1996, 26 territories were occupied of which 5 (Jackson, Moffat #5, Rio Blanco #6 am #7, am Routt #2) were discovered in 1996. Nest size suggests that Rio Blanco #7 and Moffat #5 were probably first nesting attempts. Although the Jackson site was discovered in 1996, the ranch caretaker indicated that it had been occupied at least the 2 previous years. Presence of an adult at Routt #2 in 1995 suggests that site was active the previous year. The Fremont site was occupied by adults early in the breeding season, but did not produce young. Although the Grand site nest tree blew down shortly after the young fledged in 1995, the pair relocated to an artificial osprey nest platform and fledged 1 young. After the Rio Blanco #4 nest slipped out of the tree in 1994, the pair relocated upstream approximately 0.5 miles. The new nest was discovered in the course of a helicopter inventory of big game. It is likely that the pair used the nest in 1995. Montezuma #2 and #3 were occupied by Canada geese. Fig. 1. BREEDING PAIRS AND PRODUCTIVIIT 35 -r--------------------------------------------------------------, 30 - ------. ----.- -------- ----------- ---- ------- ------------- ----------. ------ -------------- --- Years Reproduction Reproductive efforts for 1996 varied (Table 2 and Fig. 1).. In 1996, 21 pairs laid eggs and 32 young were produced by 19 successful pairs (1.68 young per successful pair) which yielded an overall productivity of 1.23 young per territory occupying pair. Seven pairs either did not produce eggs" or failed during incubation. Single unhatched eggs were encountered during banding visits at 2 nests (Moffat #1 and #4). Sixteen young were banded and color marked at 11 locations (Adams, Jefferson, Grand, La Plata #1, Mineral, Moffat #1, #2, #3 and #4, Morgan and Routt #2). Fish and Wildlife Service bands were affixed to the nestlings' right legs and red alpha-numeric bands with yellow vinyl flags were affixed to their left legs. Culmen length and foot pad length measurements were obtained from eaglets that were banded. Eaglets at 4 other sites were too old to band, and landowner permission could not be obtained at Rio Blanco #4. Land Status The 5 new territories «Jackson, Moffat #5, Rio Blanco #6 and #7, and Routt #2) were on private property and livestock �105 grazing is the primary land use. Banded Adults The adult male at Rio Blanco #5 was banded and color marked. He was produced at Rio Blanco #3 in 1991 which is approximately 16 miles downstream. The female was unbanded. The female at the Adams site, which had been banded on the Rocky Mountain Arsenal during the 1986 winter was replaced by an unbanded female in 1996. She was banded as an adult, so given at least 4 additional years to achieve adult plumage, she was at least 13 years old when she disappeared. Nest StabjJjzation Efforts No nest stabilization efforts were undertaken in 1996. The nest at Rio Blanco #5 which blew out killing the chick in 1995 year, was reconstructed in a more substantial position in 1996. The Rio Blanco # 1 nest tree continued to remain upright in spite of its dead condition. The pair did not frequent an artificial nest that had been constructed in an adjacent tree a nwnber of years previously. After being vacant in 1995, Rio Blanco #1 was reoccupied and was successful despite the fire damage that killed the tree. The Archuleta site successfully fledged 2 young, although at time of fledging, the nest fell to the ground and the fledglings had to perch on nearby limbs. An artificial nest base will be placed at the site in the fall. The Jefferson nest is vulnerable to wind throw due to placement in the crotch of a dead limb. Since the limbs cannot be stabilized, consideration should be given to placement of an artificial base in an adjacent tree. The dead nest tree at the Adams site had its life extended when it was guyed up by REA. Prepared by: G .R. ~ Gerald R. C Life Science Researcher N �Table 1. Bald Eagle Nesting Efforts in Colorado I--' o 0\ Site La Plata Co. #1 Moffat Co. #1 La Plata Co. #2 Grand Co. Montezuma Co. #1 Rio Blanco Co. #1 Rio Blanco Co. #3 Weld Co. #1 Montezuma Co. #2 Moffat Co. #2 Moffat Co. #3 Adams Co. Archuleta Co. Montezuma Co. #3 Weld Co. #2 La Plata Co. #3 Rio Blanco Co. #4 Morgan Co. #1 Mesa Co. #1 Fremont Co. Routt Co. #1 Gunnison Co. Mineral Co. Weld Co. #3 Montezuma Co. #4 Jefferson Co. Mesa Co. #2 Moffat Co. #4 Jackson Co. Rio Blanco #5 Moffat CO.#5 Routt Co. #2 Rio Blanco #6 Rio Blanco #7 Total Young Total Pairs Young/Gcc. Terr. IA = Inactive A 1974 1975 1976 legg IA IA lyng 2yng 2yng 1977 IA 2yng 2yng 1978 IA 2yng 2yng Oyng 1979 IA lyng Oyng Oyng 1980 ? -IA IA A 1981 ? 2yng IA IA A 1yng 1982 ? 2yng IA IA A 1yng 3yng 1983 ? -IA IA A ? 2yng 1984 ? 2yng IA IA IA eggs 2yng 2yng 1985 ? Oyng IA IA IA Oyng 2yng 2yng 2yng 1986 eggs 1yng IA IA IA 2yng Oyng eggs 1yng 1yng legg eggs eggs 1987 2yng 2yng A IA IA 2yng lyng IA lyng Oyng IA 1egg 2yng 1996 lyng 1yng IA 1yng IA 3yng 2yng IA IA 2yng 2yng 1yng 2yng IA IA Oyng 2ng 3yng IA Oyng Oyng LAD 2yng Oyng IA 1yng 2yng lyng 2yng -lyng Oyng lyng 2yng Oyng o 1 4 4 4 1 0 3 6 2 6 6 5 10 8 16 13 19 20 24 19 25 32 1 1 2 2 3 3 1 3 4 2 4 5 10 9 8 10 10 13 14 20 17 21 26 0.00 1.00 2.00 2.00 1.33 0.33 0.00 1.00 1.50 1.00 1.50 1.20 0.50 1.11 1.00 1.60 1.30 1.46 1.43 1.20 1.12 1.19 1.23 = Active LAD = Lone Adult . \ 1988 lyng lyng IA IA IA 2yng A IA lyng 2yng ? eggs IA 1yng 1989 2yng 3yng IA IA IA 2yng 2yng IA 1yng 3yng eggs 2yng I.A 1yng eggs 1990 Oyng 2yng IA IA IA 2yng lyng Oyng 1yng 2yng IA 2yng IA 1yng IA 2yng 2yng 1991 1992 lyng lyng 2yng 2yng IA IA IA IA IA IA 2yng 1yng 3yng 3yng IA IA Oyng 1yng 2yng 3yng Oyng Oyng 3yng 3yng Oyng Oyng 2yng lyng IA IA 2yng? 1yng 2yng 2yng 3yng Oyng eggs 1993 2yng 2yng IA IA IA 2yng 2yng IA IA Oyng 2yng 2yng eggs 2yng IA ? 3yng 2yng eggs 2yng 2yng 1yng A A Oyng Oyng 1994 3yng 2yng IA IA IA 2yng 1yng IA IA Oyng 2yng 3yng IA lyng IA IA Oyng Oyng Oyng IA Oyng 2yng ? Oyng IA Oyng 2yng lyng ?yng 1995 Oyng 2yng IA 1yng IA IA 2yng IA IA 3yng 1yng 3yng 1yng 2yng IA 1yng Oyng 3yng IA Oyng 2yng Oyng Oyng Oyng IA Oyng 2yng lyng 2yng Oyng �107 Table 2. Colorado Bald Eagle Nesting Efforts - 1996 Site Age of Birds Male Female Young Produced Comments ~£D!Iru Rcgion Adams Co. Jefferson Co. Adult Adult Adult Adult I I Nortl}ell,<;tRegion Jackson Co. Morgan Co. Weld County #3 Adult Adult Adult Adult Adult Adult 2 3 0 Adults present early, eggs not confirmed. ' Southeast Regio!} Fremont Co. Adult Adult ? Adult perched by nest early in season. Southwest Region Archuleta Co. Gunnison Co. La Plata Co # I La Plata Co. #3 Mineral Co. Montezuma Co. #2 Montezuma Co. #3 Northwest Region Grand Co. Mesa Co. #2 Moffat Co.#1 Moffat Co #2 Moffat Co. #3 Moffat Co. #4 Moffat Co. #5 Rio Blanco Co.#1 Rio Blanco Co. #3 Rio Blanco Co. #4 Rio Blanco Co. #5 Rio Blanco Co. #6 Rio Blanco Co. #7 Routt Co. #1 Routt Co. #2 Total Total pairs: 26 Egg laying pairs: 21 Productive pairs: 19 IA = Inactive Adult Adult Adult ? SubadAdult Adult Adult Adult Adult .2 > 0 Fledged, nest fell out at that time . Adult female only bird OJ>SCIVed. I 0 2 Goose in nest Goose in nest Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult SubadAdult Adult Adult 26 23 I 2 I 2 2 I 0 3 '2 2 I 2 0 0 I 32 Unhatched egg in nest. Unhatched egg in nest. Female in incubating posture, failed. Relocated to new nest, probably used in 95. Adult male was hatched at Rio Blanco #3 in 1991. ObSCIVedfrom aircraft. Female in incubating posture, failed. � https://cpw.cvlcollections.org/files/original/e34bfb31befb42c7512bbe95e365d6f4.pdf c361260c8a38ad72a638df4e656ede07 PDF Text Text Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS state of Project Colorado No. W-153-R-9 Mammals Work Plan No. Covered: Author: Research Hu1tispecies Inyestigations Consulting Services for Mark-Recapture Analysis Job No. Period REPORT July 1, 1995 - June 30, 1996 G. C. White Personnel: R. M. Bartmann, R. B. Gill, T. D. I. Beck, D. J. Freddy ABSTRACT Progress towards the objectives of this job include: 1. A computer program (Program NOREMARK) developed to operate on personal computers for the computation of population estimates based on resightings of marked animals was published in the Wildlife Society Bulletin. 2. A study of compensatory effects of harvest on the Piceance Basin mule deer population was continued as part of Federal Aid Project W-153-R Work Plan 2 Job 15, entitled Compensatory Effects of Harvest in a Mule Deer Population. Experimental harvests have been conducted in December, 1989, 1990 and 1991. Radio collars to monitor over-winter survival of fawns were placed on the animals during November, 1989-95, and results are currently being summarized and written. 3. Preliminary cooperation 4. A World-Wide-Web page at http://www.cnr.colostate.edu/-gwhite/software.html has been established to distribute software developed contract. analysis of elk sightability with David Freddy. models was performed under in this ��3 CONSULTING SERVICES FOR MARK-RECAPTURE ANALYSES G. C. White P. N. OBJECTIVES Evaluate density dependence ·in the Piceance SEGMENT 1. deer populations. OBJECTIVES Summarize evidence of for density fawns in the Piceance Basin. RESULTS Basin mule dependence in survival of mule deer AND DISCUSSION Introduction. Density dependence in a deer population has major implications for management because the concepts of maximum sustained yield, compensatory mortality, quality versus quantity assume density-dependent feedback in the population. Density dependence is generally defined as a negative relationship between population growth rate and population size (McCullough 1990). As McCullough (1990) pointed out, growth rate is a misnomer in the sense that its value can be positive, zero, or negative; the latter resulting in a decreasing population. A positive growth rate is often associated with populations below carrying capacity, whereas a negative rate is more typical with populations above carrying capacity. A concept closely tied to density dependence is compensatory mortality, or a change in the rate of the remaining sources of mortality in response to a change in the rate of 1 mortality source. As Kautz (1990) stated, the only reasonable mechanism to explain compensatory mortality is density-dependent mortality. Objectives of this paper are 1) to present models of how density dependence can operate in a population'S dynamics, 2) to review evidence in the literature of density dependence in deer populations, and 3) to discuss some potential problems in experimental design and statistical analysis procedures when investigating density dependence in a deer population. Acknowledgments. We thank the numerous personnel from the Colorado Division of Wildlife, Los Alamos National Laboratory, and Colorado State University, and numerous other volunteers that helped in conducting the 15 years of mule deer research in the Piceance Basin. STUDY AREA The study area on mule deer winter range in Piceance Basin in northwest Colorado was previously described in a similar investigation of compensatory mortality by Bartmann et al. (1992). The area was a northwest-southeast oriented ridge, about 10-km long x 5-km wide, containing 48 km2• Most of the study area was on the north side of the ridge, but a variety of exposures were created by drainage patterns. Elevations ranged from 1,770 to 2,170 m with steep topography and elevational changes ~350 m within 2 km common. Climate was semi-arid with warm summers and cold winters. Mean annual precipitation �4 was 33 cm with about half occurring october through April. as snow. Deer were usually present from Pinyon pine (finy edulis) and Utah juniper (Juniperus osteosperma) were the main overstory species. A shrub understory included big sagebrush (Artemisia tridentata), Utah serviceberry (Amelanchier utahensis), true mountainmahogany (Cercocarpus montanus), antelope bitterbrush (Purshia tridentata), mountain snowberrry (Symphoricarpos oreophilus), rabbitbrushes (Chrysothamnus ~.) and Gambeloak (Quercus gambelii). Additional shrubs, forbs, and grasses were described by Bartmann (1983). The study area was split into a treatment unit on the west (21.4 km2) where the deer population was reduced by >50%, and a control unit on the east (26.4 km2) with no reduction other than harvest during regular hunting seasons. Estimates of vegetation biomass on the study area during summer 1988 indicated more total vegetation and more grass on the control unit (£ 50.045), but shrub and forb biomass did not differ (£ ~ 0.126) (Bartmann et al. 1992). During the study, the!e were several fires with the most area burned on the treatment unit (get BLM data). METHODS Population Density and Reduction Aerial line transect surveys described by Bartmann et al. (1992) and White et al. (1989) were continued during this study to estimate deer density on control and treatment units. The transects were first flown in December 1985 and were continued in mid-December or early January every winter, except 1993-94, through 1994-95. For 1995-96, deer densities were estimated with an aerial mark-recapture procedure utilizing radiocollared fawns as the marked population (Bartmann et al. 1987). To enable quick identification of collared fawns, a 5 x 10-cm vinyl tab was riveted to the top of each collar. Tabs were color-coded by unit: orange for control and white for treatment. Thirty-three collars retrieved 1-4 months post-survey all had tabs intact supporting the assumption that no marks were lost. All collared fawns were located immediately prior to the surveys to verify the number on each unit. Three complete-coverage helicopter flights (morning, midday, and afternoon) were made on each unit 6-7 January 1996 with 2 observers plus the pilot. The mule deer population on the. treatment unit was reduced by >50% over a 4year period by public hunting during late seasons in December. The original treatment unit included 21.4 km2, but unit boundaries were altered by adding 6 km2 on the west side and excluding a 1 x 6-km strip along the south side to make them align with roads. Boundary roads were signed at -0.5-km intervals to identify the side open to hunting. Hunting occurred during all of December in 1989-91, and for 9 days in 1992. There were 375 licenses issued in 1989, 400 in 1990, 350 in 1991, and 100 in 1992 with each hunter allowed 2 antlerless deer. Prior to each season, hunters were mailed a packet with information about the season, a detailed map of the hunt area, and a postage-paid tooth envelope to record kill data and to enclose 2 lower incisors. �5 Check stations were operated during daylight hours on weekends at the 2 main access locations to the hunt area. During the week, hunters were checked as encountered in the field during daily radio-tracking activities. Each checked deer was weighed (kg) and its age estimated. The lower jaw was taken from deer older than yearlings. Hunters not checked could mail tooth envelopes or deposit them in a collection box at either main access point. A sample (percent???) of hunters was also included in the CDOW's regular harvest survey with mailed questionnaires. These data were used to estimate hunter participation, success rate, and harvest. Deer harvest on the 2 study units could not be derived from CDOW survey data for regular hunting seasons. Therefore, an estimate of the minimum harvest of antlerless deer during regular seasons was obtained from field checks of hunters. Data for these deer were the same as collected during the late season. Survival From 1989 through 1992, deer were captured with dropnets (Ramsey 1968, Schmidt et al. 1978) during the -3-4-week period between the end of the regular hunting season in November and the start of late seasons in early or midDecember. Dropnets were used in 1989 through 1991. In 1992, about half the deer were captured with dropnets and half with helicopter netguns (White and Bartmann 1994). Netguns were used exclusively the next 3 years except .1994 when 26 fawns and 13 adult does were also captured with dropnets. From 1989 through 1991, the goal was to radiocollar 40 fawns on each study unit. This was increased to 80/unit the following 4 years. Goals for radiocollared adult does were 3D/unit in 1989, 40/unit in 1990 and 1991, and SO/unit the last 4 years. Does were radiocollared to allow detecting any changes in survival patterns documented over the previous 7 years and to detect movements between or off units as the population reduction progressed. Adult collars were permanently attached so only enough new does were collared each year to meet annual goals. No does were radiocollared in 1995 for the final year of the study. For dropnetting, 26 to 38 sites, distributed over both study units, were used each year except 1994 when only 10 sites were used. For netgunning, there were 2 to 4 processing sites/unit each year to minimize flight time and stress on animals. Fawns captured less than -0.5 km from the processing site were released at the site with the rest flown back to the capture location and released. Adult does were released at the processing site. From 1989 through 1993, female fawns were fitted with expandable collars so survivors could be monitored as adults. Female fawn collars in 1994 and 1995 and male fawn collars all years were designed to drop off after -8 months. Each fawn and adult transmitter had a motion switch set with a 3-4 hour delay to allow quick detection of mortalities. Fawn survival estimates were based on the -6-1/2 to 7-month period from capture in mid-November/early December to the following 15 June when they were considered adults. Annual survival for adult does was estimated for the 1 December-3D November year. All deer were monitored from the ground 5~7 days/week from trapping until migration in late April or early May. Aerial monitoring was then done about every 2 weeks until 15 June. Aerial tracking �6 continued 1-2 times/month through the summer and fall to monitor survival and locate drop-off collars for reuse. adult All radiocollared deer were located 3 times each winter during the mid portions of January, February, and March. This allowed detecting movements of deer between units, or out of either unit, as the treatment population was reduced. Deer that moved permanently off either study unit were censored for survival estimates. Weight (kg), total body length (cm), and left hind leg length (cm) were recorded for each fawn and doe (Anderson et al. 1974). Deer captured by netgunning averaged -2 cm longer than those caught with dropnets. This was attributed to different methods of restraint (White and Bartmann 1994). This had little effect on comparisons between units within years as all deer were ~ither captured with the same method, or similar proportions on both units were captured with each method. Between years??? statistical Methods Deer density was computed with line transect estimators from helicopters (White et al. 1989, Buckland et al. 1993) using Program DISTANCE. Distance data were taken in January, 1985-1993, and again in 1995, for each of the 2 areas. Detection distances were grouped into 10 intervals: 0-15 m, 10-m increments from 15 to 95 m, and 95 - 155 m; and truncated after 155 m. From past work (White et al. 1989), the negative exponential model was known to generally fit distance data taken on mule deer from helicopters. Thus, we included the negative exponential key function in the list of candidate models considered, along with the 3 adjustments available in DISTANCE: cosine, polynomial, and Hermite polynomial. The half-normal key with the Hermite polynomial adjustment and the hazard key with the polynomial adjustment were also included. For each estimator, up to 5 adjustment terms were allowed.· Models were fit separately to each year*area combination. Models were also fitted with detection distances pooled across all year*area combinations, and pooled within years across the 2 areas. AIC was used for model selection, both within key functions and among key functions. Density in Jan, 1996, was estimated based on the joint hypergeometric estimator (Bartmann et al. 1987) using Program NOREMARK (White 1996). Radiocollared fawns were used as the sample of marked animals, with 3 surveys of each area completed. Statistical analyses of physical condition variables were performed with PROC GLM of SAS (1987) and of survival rates with PROC GENMOD of SAS (1996). A multivariate model predicting body weight, total body length, and left hind leg length was constructed. Independent variables were area (control, treatment), gender, areaxgender, period (before or after population reduction), area x period, gender x period, area x gender x period, year within period, area x year within period, genderXyear within period, and areaxgenderXyear within period. Tests of effects were constructed from a mixed model analysis of variance using the RANDOM / TEST statement of PROC GLM. Year within period and all interactions containing this term were considered random effects. F-tests of area and areaxperiod were constructed with area X year within period in the denominator, gender and gender x period with gender x year within period, period with year within period, area x gender x period with area x gender x year within period, year within period. with area x year within period, area x year within period and gender x year within �7 period period with area x gender x year withing period, with the within cell error term. and area x gender x year within Kaplan-Meier estimates of survival curves were computed with the staggered entry method of Pollock et al. (1989). Overwinter fawn survival was modeled as a function of area '(contro1, treatment), period (before or after population reduction) area x period, year within period, gender of fawn, and weight of fawn. Because the areaxperiod effect explained any differences in area, this interaction was modeled directly with 2 different approaches. The effect of the treatment is expected to increase as the number of deer removed increases, and density is lowered. Thus, a linear trend (1, 2, ••• ) was fit, starting with 1989. The treatment effect is delayed because of incremental removals, so we also delayed the start of the trend from 1989 to 1990, up to 1993. We also considered a constant treatment effect, i.e., a dummy variable consisting of either 0 or 1. This variable was also started at different years, from 1989 to 1993. The best fitting model was ·selected from the list of candidate models generated by combinations of these variables with Ale (Burnham and Anderson 1990). RESULTS Reduction of the treatment area was accomplished with hunter harvest (Table 1), with a corresponding decrease in numbers of animals on the treatment area compared to the control area (Fig. 1). Table 1. Hunter participation and success and deer harvest estimates for the December late season on the treatment unit of the Ridge study area, 1989-92. 1989 No. Licensesa Survey respondents Did not hunt Unsuccessful Harvested 1 deer Harvested 2 deer Harvest-Does Fawns Total a Hunters were (SE) 375 60 81 38 56 200 342 114 456 allowed % 1990 No. % 21.7 10.0 15.0 53.3 400 92 48 109 104 139 75.0 25.0 (47.5) 263 .120 383 1991 No. % 12.0 27.2 26.0 34.8 350 69 41 76 122 112 11. 6 21.7 34.8 31. 9 68.6 31. 4 (40.0) 214 131 345 62.1 37.9 (39.4) to take 2 antlerless 1992 % No. 100 37 deer on a license. 32 30 38 83 22 105 32.4 29.7 37.8 78.9 21.1 (15.8) �8 DENSITY 120 110 100 90 80 70 60 50 40 30 20 10 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 YEAR AREA - - Control Remoyal Fig. 1. Density estimates (deer/km2) on control and treatment areas, Little Hills Wildlife Area, Meeker, Colorado, 1985-96. The 1996 estimates were computed from resighting data, with the remaining years computed with line transect distance estimators. Separate models for each year*area combination provided the smallest AlC: 13213.85. Pooling the areas for each year increased the AlC to 13221.22, and pooling across all year*area combinations resulted in an AlC of 13286.21. The model with the smallest AlC for each of the 18 year*area combinations were negative exponential: with no adjustments, 6; with cosine, 3; and with polynomial, 1; half normal with no adjustments, 5; and hazard with no adjustments, 1, and with cosine adjustments, 1. For all data pooled across the 18 area*year surveys, the model selected was a negative exponential with a cosine adjustment. Thus, the negative exponential model tends to be selected with the AlC model selection criterion. Because the detection distances had to be modeled for each year*area combination separately, the confidence intervals on the estimates (Fig. 1) were wide. Nov-Dec fawn body size increased (interaction of areaxperiod, ~ = 0.079) the removal area after population reduction (Fig. 3), with most of the increase contributed by hind foot length (~ = 0.007) and body weight (~ = 0.052), but little by total body length (~ = 0.191). on Overwinter fawn survival improved on the treatment area after density was lowered (Fig. 2). The model including gender (~< 0.001), weight (~< 0.001), year (~ < 0.001), and with areaxperiod interaction modeled as a constant treatment effect starting in 1993 (~ = 0.001) was the best-fitting model based on AlC. However, this model is somewhat suggested by the data in Fig. 4, because the treatment effect seemed to increase after the discontinue of removals in 1992. The model including gender (~< 0.001), weight (~< 0.001), year (~ < 0.001), and with area x period interaction modeled as a linear trend starting in 1989 (~ = 0.003) was within 0.71 AlC units of the best AlC model found. All the models with the area x period interaction effect modeled as a �9 linear trend or a constant effect showed the effect to be important (~ < 0.005) in explaining the observed overwinter fawn survival rates. For the model with area X period interaction modeled as a effect for 7 years, the increase in survival, assuming an average year with a female fawn weighing 32 kg, would be from 0.43 with no removals to 0.55 for the level of removals in this experiment. For the model with area X period interaction modeled as a linear trend starting in 1989, and assuming an average year with a female fawn weighing 32 kg, survival would increase from 0.44 with no removals to 0.51 after 3 years, to 0.62 after 7 years. The treatment effect increases with time, but as shown in Fig. 3, year-to-year variation produces even greater fluctuations in survival. Thus, the compensatory effect of the population reduction is disguised by temporal variation. The cumulative harvest on the removal area from Table 4, with 0 used for the control area and prior to harvest on the removal area, provided a predictor (~ = 0.005) of fawn survival, with weight, gender, and year included in the model. This model was within 1.72 AlC units of the best model found. Density (Fig. 1) did provide some predictive capability (~ = 0.096) when weight, gender, and year were in the model. Density is likely not a good predictor of survival because of the lag in the response of survival to the reduction in density. By the time we observed differences in survival, the densities on the 2 areas were similar, whereas, during and immediately following the removals, when density differences were the greatest, we did not observe differences in survival. �10 37 36 W 35 ."I 34 Q 33 h t 32 ( 31 k 30 ~ 29 28 27 1981 1982 19831984 L e n Q t 19851986 19871988 19891990 19851986 1987 1988 19891990 1991 1992 19931994 19951996 43 42 h h I 41 n d 40 e Q ( c m 39 1981 1982 19831984 1991 1992 1993 1994 19951996 135 B 134 o d y 133 132 131 130 129 120 127. CI 126 t. 125 h 124 e n ( 123 ~ g~ ) ~~~ 1981 198219831984 1985 1986 1987 198819891990 1991 19921993 1994 1995 1996 YEAR REMAREA - - Control Remoyal . Fig. 2. Body size of fawns (sexes combined) measured during Nov-Dec on control and treatment areas, Little Hills Wildlife Area, Meeker, Colorado, 1982-1995. �1981 1982 19831984 19851986 19871988 19891990 1991 1992 1993 1994 19951996 YEAR REI'IAREA D i f f e ,. e n c e n Control --- Removal 0.5 0.4 0.3 0.2 ; o. 1 ; / ••••• - 0.0 -0.1 S -0.2 ~ -0.3 v v a 1 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 YEAR Fig. 3. Staggered-entry Kaplan-Meier estimates of overwinter fawn survival (sexes combined) and 95% confidence intervals (upper graph) measured Nov-Jun on control and treatment areas, Little Hills Wildlife Area, Meeker, Colorado, 1982-1996. Hunting mortalities were censored. Bottom graph shows the differences of estimates (treatment minus control) and their 95% confidence interval. DISCUSSION �12 One reason that fawn survival did not immediately respond to decreased density is the stress caused by the December hunts. Based on our field observations, deer on the control area were commonly feeding during day-time hours, whereas deer on the removal area were not out in the open feeding because of heavy hunting pressure. Thus, the stress of the harvest affected deer behavior and likely had an effect on body condition. The lag in response of vegetation to lowered deer densities provides another explanation for why we did not immediately observe a response in fawn survival to lower densities. LITERATURE CITED Bartmann, R. M., G. C. White, L. H. Carpenter, Aerial mark-recapture estimation of confined woodland. J. Wildl. Manage. 51:41-46. Buckland, S. T., D. R. Anderson, sampling: estimating abundance London. 446 pp. and R. A. Garrott. 1987. mule deer in pinyon-juniper K. P. Burnham, and J. L. Laake. Distance of biological populations. Chapman and Hall, Pollock, K. H., S. R. Winterstein, C. M. Bunck, and P. D. Curtis. 1989. Survival analysis in telemetry studies: the staggered entry design. J. Wildl. Manage. 53:7-15. SAS Institute Inc. 1987. edition. SAS Institute SAS/STATm Guide for personal Inc., Cary, NC. 1028pp. computers, version 6 SAS/STAT® Software: Changes and enhancements SAS Institute Inc. 1996. through release 6.11. SAS Institute Inc., Cary, NC. 1104pp. white, G. C., R. M. Bartmann, L. H. Carpenter and R. A. Garrott. 1989. Evaluation of aerial line transects for estimating mule deer densities. Wildl. Manage. 53:625-635. white, G. C. 1996. NOREMARK: population estimation surveys. Wildl. Soc. Bulletin. 24:50-52. from mark-resighting J. �13 Colorado Division Wildlife R~search July 1996 of Wildlife Report ADDENDUM JOB FINAL REPORT state of Project Colorado No. W-153-R-9 Mammals Research Work Plan No. Multispecies Job No. Consulting Research Period Covered: Authors: July G. C. White, Personnel: Inyestigations Services for Mammals 1, 1995 to June 30, 1996 R. B. Gill, G. C. White, and T. D. I. Beck R. B. Gill, and T. D. I. Beck ABSTRACT Harvest strategies for a south-central Colorado black bear (Ursus americanus) population (Beck 1991) were evaluated with a Monte Carlo simulation model. The model included binomial variation for birth and death rates, plus two types of annual environmental variability: 1 year in 10 heavy harvest because of optimal hunting conditions, and 1 year in 10 failure of the berry crop, causing increased natural mortality and decreased reproduction. No densitydependent responses were included. Six harvest scenarios were simulated: no harvest, 2 spring seasons with 30 and 35% of the harvest consisting of females, and 3 fall seasons with 35, 40, and 45% of the harvest consisting of females. Harvest rates were 5, 10, 15, and 20% of bears ~2 year old. Because spring harvest is disproportionately males, our simulations suggest more bears can be harvested during spring seasons. Without density-dependent compensation in the population, -15% of the population ~2 years old can be removed annually based on observed reproductive and survival rates, and the population would not decline. Increased environmental variation caused by berry crop failures and occasional heavy harvest can significantly reduce population growth rate, and thus affect long-term management. ��15 Addendum to: Consulting Gary C. White, Services R. Bruce Gill, for Mammals and Thomas D. Research I. Beck P.N. OBJECTIVE Evaluate compensatory effects of harvest SEGMENT 1. on the Piceance mule deer population. OBJECTIVE Prepare a manuscript for submission to a peer-reviewed scientific journal on the results of simulations of black bear population dynamics with revised estimates of survival. RESULTS A copy of the manuscript follows. ��17 July 15, 1997 Gary C. White Dept. Fishery and Wildlife Colorado State University Fort Collins, CO 80523 970-491-6678 gwhite@cnr.colostate.edu RH: Black Bear SIMULATION Simulation MODELING Model OF BLACK • White et al. BEAR HARVEST STRATEGIES GARY C. WHITE, Department of Fishery and wildlife University, Fort Collins, CO 80523 R. BRUCE GILL, Research Section, Colorado Prospect, Fort Collins, CO 80526 THOMAS D. I. BECK, Research Section, Prospect, Fort Collins, CO 80526 Biology, Division Colorado Colorado of Wildlife, Division State 317 West of Wildlife, 317 West Abstract: Harvest strategies for a south-central Colorado black bear (~ americanus) population (Beck 1991) were evaluated with a Monte Carlo simulation model. The model included binomial variation for birth and death rates, plus two types of annual environmental variability: 1 year in 10 heavy harvest because of optimal hunting conditions, and 1 year in 10 failure of the berry crop, causing increased natural mortality and decreased reproduction. No density-dependent responses were included. Six harvest scenarios were simulated: no harvest, 2 spring seasons with 30 and 35% of the harvest consisting of females, and 3 fall seasons with 35, 40, and 45% of the harvest consisting of females. Harvest rates were 5, 10, 15, and 20% of bears ~2 year old. Because spring harvest is disproportionately males, our simulations suggest more bears can be harvested during spring seasons. Without densitydependent compensation in the population, -15% of the population ~2 years old can be removed annually based on observed reproductive and survival rates, and the population would not decline. Increased environmental .variation caused by berry crop failures and occasional heavy harvest can significantly reduce population growth rate, and thus affect long-term management. ~ Words: Binomial distribution, environmental variation, Monte Carlo simulations, population growth rate, spring harvest, temporal variation, americanus. ~. Hildl. Manage. ~ 00:000-000. No where in their range can black bears be considered numerous. The species evolved as long-lived with exceptionally low reproductive rates and low natural mortality rates. Unlike many other species hunted for sport, rate of population increase for black bear populations is low. In addition, wildlife managers have had difficulty in developing harvest strategies for black bear populations because populations are difficult to monitor. Population estimation methods are expensive, and often ineffective, due to the secretive nature of the animal. Individuals are difficult to capture, and' thus to obtain an adequate sample of marked animals for mark-recapture methods. Further, revenue from the sale of licenses is not adequate to justify extensive management. As of a result of these limitations, little �18 research has been conducted on harvest strategies. Public controversies have revolved around when to hunt, how to hunt, how many bears should be harvested, how many hunters should be allowed, how nuisance bears should be controlled, and how illegal hunting should be eliminated. We developed a Monte Carlo computer simulation model of a Colorado black bear population to evaluate possible harvest strategies. The stochastic model is patterned after the grizzly bear (~. arctos horribilis) model of Knight and Eberhardt (1985). structure of their model was modified to be appropriate for Colorado black bears, with parameter estimates taken from research of Beck (1991) in south-central Colorado. Six harvest regimes were simulated: no harvest, 2 spring seasons with 30 and 35% of the harvest consisting of females, and 3 fall seasons with 35, 40, and 45% of the harvest consisting of females. In addition, two types of additional environmental variability were considered: occasional heavy harvest because of optimal hunting conditions, and occasional failure of the berry crop, causing increased natural mortality and decreased reproduction. Acknowledgments. Funding was provided by Colorado Federal Aid in Wildlife Restoration Project W-153-R. Division of Wildlife, METHODS The black bear population model (available from the senior author) was developed using the Statistical Analysis System (SAS 1985). Structure of the model is based on 3 arrays. The subadult array holds numbers of cubs and yearlings of both sexes whose mothers have died. The male array holds numbers of males of ages 2 to 20 (the maximum age attainable). The 2-dimensional female array holds numbers of females of ages 2 to 20, and also their reproductive status (current number of offspring). ~eproductive status 0 means no cubs or yearlings, 1 means i cub, 2 means 2 cubs, 3 means 3 cubs, 4 means 1 yearling, 5 means 2 yearlings, and 6 means 3 yearlings. Although the yearlings are not actually with the mother, the mother is not allowed to breed in 2 consecutive years (unless the cubs are lost). Thus categories 4-6 provide a mechanism to store females for one year until they are ready to breed again. Parameter values for survival and reproduction rates (from Beck 1990) are presented in Tables 1 and 2. These rates represent a non-hunted population, but with some illegal harvest plus some emigration to areas where harvest is allowed. Stochasticity in the model is in the form of binomial processes for birt'hs and deaths. For example, a 71% survival rate for adult males does not multiply the current population size by 0.71 to obtain the number of survivors. Rather, the N animals in the adult male population are assigned a fate (lived or died) according to the probability 0.71, using the RANBIN function of SAS. This binomial variation increases the reality of model simulations. Two extensions to the grizzly bear model developed by Knight and Eberhardt (1985) were made. Beck (1991) showed that survival of cubs and yearlings and reproductive rates are affected by berry crop. Late spring frosts curtail mast and berry production. Therefore, some simulations incorporated 2 sets of values for birth and death rates, depending on whether the current year.was a "good" berry crop or a "bad" berry crop. Bad berry crops occurred at random with a frequency of 1 year out of 10 using the RANUNI function (SAS 1985). �19 In addition to effeqts from bad berry years shown in Tables 1 and 2, all cubs born during bad years had reproductive rates as if it were a bad year until they were 6 years old. After a bad berry year, all barren adults had 0.90 reproduction instead of 0.80, compensation for the previous bad year (unless the second year was also a bad berry year). The second modification of the model by Eberhardt and Knight was to include harvest. A major consideration of the various experimental harvests was the proportion of males versus females in the harvest. The proportion of females in the harvest can be affected by the timing of the season, plus the method of take (Gill and Beck 1990). Six harvest regimes were simulated: no harvest; spring seasons with the expected harvest consisting of 30 (labeled S30) and 35% (labeled S35) females; and fall seasons with the expected harvest consisting of 35 (labeled F35), 40 (labeled F40), and 45% (labeled F45) females. Because of the differential survival of males and females, an unhunted population would consist of 72% females for animals ~2 years old. Therefore, experimental harvests incorporated differential vulnerability of females and males. Harvest vulnerability was determined based on the expected harvest and the relative proportion of the population consisting of males and females ~ 2 years old. For example, to obtain a harvest with 30% females, the proportion of females to harvest is equal to the expected number of females in the harvest divided by the number of females in the population, and similarly for males. The only exception to this procedure was when the proportion of the population segment to be harvested exceeded 90%, the maximum allowable harvest was set to 90%. Actual number of animals .harvested was determined stochastically with the RANBIN function (SAS 1985). Cubs of any females harvested in a spring season (April-June) are all assumed to die, whereas cubs of females harvested in the fall (September-october) are assumed to survive at the rate of cubs without mothers. In addition, an increase in vulnerability to harvest of the entire population was included in the model using the RANUNI function (SAS 1985). On average, 1 year out of 10 had a double success rate (e.g., 20% harvest instead of 10%). This doubling of harvest was included in the model to represent the effects of variable weather on harvest (Gill and Beck 1990). Initial population sizes were constructed by making 100 runs of the model for 30 years with no harvest, and using the resulting relative proportions of the various age and sex classes. Values for each of the age and sex classes are initialized the same in each simulation. A summary of these. initial conditions are: cubs (both sexes) - 2499, yearlings (both sexes) - 1296, adult males - 1740, and adult females - 4465, for a total population at time 0 (No) of 10,000. These initial conditions represent a large population, approximately the state-wide population of Colorado (Beck 1991). All simulations were performed for 30 years, Le., the population was projected forward for 30 years. This process was replicated 100 times to provide a mean and standard deviation for each of the parameters measured on the simulated population. Here, we report the X population level at the end of the 3D-year simulation (}Lo), plus X annual rate of increase ~, where ~ (}Lo/No)(1/30) - 1. All simulations. were repeated for 100 times to estimate means and SDs. Statistical tests between pairs of treatments were performed with a.-test using a pooled variance computed as I12 = [(n1 - 1)I112 + (n2 - 1)I1/]/ (n1 + lli 2) • �20 RESULTS Annual rate of increase for simulations with constant annual survival and reproduction, berry crop failures 1 year in 10, double the normal harvest 1 year in 10, and both berry crop failures 1 year in 10 and double harvest 1 year in 10 are presented in Fig. 1. Harvest rate is 10%. Effect of berry crop failures on the model population was to increase variability of population size after 30 years and to lower expected !Lo. As an example, 35% females in the spring harvest under no berry failures had a 3.7% X annual increase, whereas with berry failures, the X annual growth rate was 2.8% (£ < 0.001). Under the fall harvest scenario with 35% females, no berry failures generated a X population growth rate of 4.2%, berry failures 3.2% (£ < 0.001). 50's of ~ for the no annual variation simulations were 0.085, 0.071, 0.083, 0.089, and 0.095, for 530, 535, F35, F40, and F45 seasons, respectively. Corresponding 50's for the berry failure simulations were 0.540, 0.571, 0.664, 0.592, and 0.614. A 10% chance of double harvest reduces (£ < 0.001) the rate of population growth compared to where harvest is always the same from year to year. Another impact of the occasional double harvest is an increase in variability of the final populations. For example, 50's of ~ for the occasional double harvest simulations were 0.188, 0.236, 0.220, 0.245, and 0.274 for 530, 535, F35, F40, and F45 seasons, respectively. These values are intermediate between the no annual variation and occasional berry failure simulations presente~ above. Impact of both berry crop failures and occasional double harvest is to further increase variation, and to further lower the average annual rate of increase. However, impacts of increased annual variation does not change the relative impact of the 5 seasons (Fig. 1). The ordering of the annual rate of increase for 530 through F45 is the same for all 4 scenarios shown in Fig. 1, and absolute differences appear to be constant across the 4 scenarios. For reproductive and survival rates measured in south-central Colorado, approximately 15% of the population can be harvested annually if no densitydependent compensation is assumed (Fig. 2). Occasional double harvest and berry crop failures are included in the simulations in Fig. 2. This annual variation would decrease the proportion of the population that can be harvested annually. Under constant annual conditions, the proportion of the population that could be harvested would be >15%. DISCUSSION Additional stochasticity included in the model by incorporating survival and reproduction as binomial processes makes the results more realistic for small populations. Typically black bear populations are small in a numerical sense, even though large areas may be inhabited. Chance variation should be included in a realistic model. Additional variation from berry crop failures and occasional double harvest also makes the model's predictions more realistic. Also, this annual variation makes the model more conservative in its predictions of the permissible annual harvest. Responses of black bear populations to hunting removals are not well known. Some studies suggest that black bear populations respond to hunting removals with increased survival of subadults and juveniles. Others suggest that increased survival occurs only among subadult and juvenile males. Still �21 others suggest that populations do not respond with increased survival at all, rather dispersing subadults colonize home ranges vacated by the harvest removals of resident bears. This controversy is more than academic because allowable harvests depend upon the degree of compensation. Simulations reported here portray harvest as additive mortality, with no compensation assumed to result from removal of harvested animals. Thus, our results magnify effects of harvest -- any compensation would reduce differences between no harvest and harvest scenarios. We chose to not simulate a compensatory mechanism because adequate data are lacking to perform a realistic simulation. We know A priori that any compensation occurring in the population dynamics of black bears means that a greater harvest can be exerted, and less effect observed in the resulting population levels. Hence, the question of compensation is of interest only to the degree that compensation results. However, no reliable data exist to justify a level of compensation for our simulations. Thus, any results pertaining to compensation would only reflect our potentially unrealistic assumptions, and not provide insight into bear population dynamics. Our results provide the maximum effect expected from the harvest scenarios we simulated. If compensation occurs, then the effect of the harvest will be less than portrayed here. MANAGEMENT IMPLICATIONS The 15% harvest rate estimated with this model is too liberal if we consider that some illegal kill takes place, plus loss of depredating bears from the population. This harvest rate represents the total allowable removals from the population, because our simulation model did not include illegal harvest or removal of problem bears. Thus, to maintain existing population levels, we recommend that black bear populations similar to the one simulated should be harvested at a rate <15% to permit additional removals from illegal harvest and problem bears. LITERATURE CITED Beck, T. D. I. 1991. Black bears in west-central Colorado. Tech. Publ. 39, DOW-R-T-39-91. Colorado Div. of Wildl., Fort Collins. Gill, R. B., and T. D. I. Beck. 1990. Black bear management plan. Colorado Div. Wildl. Rep. No. 14, Fort Collins. 44pp. Knight, R. R. and L. L. Eberhardt. 1985. population dynamics of Yellowstone grizzly bears. Ecology 66:323-334. SAS Institute Inc. 1985. SAS Language Guide for Personal Computers, Version 6 Edition. SAS Institute Inc., Cary, NC. 429pp. Receiyed Accepted Associate Editor �22 Table 1. Survival rates for the various age and sex classes in the black bear simulation model, derived from Beck (1991). Only cub survival was affected by annual variation in berry crops. All other age and sex classes were modeled as shown, regardless of variation in berry crops. Age Class Males All years, regardless of berry crops Adults, > 4 years old 0.70 4-year olds 0.70 3-year olds 0.76 2-year olds 0.76 Yearlings (with mother) 0.94 Yearlings (without mother) 0.94 Females 0.96 0.96 0.94 0.94 0.94 0.94 Constant environment -- no variation in berry crops Cubs (with mother) 0.56 0.56 Cubs (without mother) 0.56 0.56 variable environment -- good berry crop years Cubs (with mother) 0.56 0.56 Cubs (without mother) 0.56 0 56 e . variable environment -- bad berry crop years Cubs (with mother) 0.33 0.33 Cubs (without mother) 0.33 0.33 Table 2. Reproductive rates for females in the black bear simulation model, derived from Beck (1991). For simulations with no variation in years, the parameters listed for good berry years were used • .. ' Parameter Definition Good Berry Years Bad Berry Years 0.001 0.22 0.29 0.60 0.50 0.80 0.20 0.60 0.20 50:50 0.001 0.001 0.001 0.001 0.20 0.20 0.20 0.60 0.20 50:50 Prop. of 2 year olds producing first cubs Prop. of virgin 3 year olds producing first cubs Prop. of virgin 4 year olds producing first cubs Prop. of virgin 5 year olds producing first cubs Prop. of virgin 6+ year olds producing first cubs Prop. of bar ren" females having cubs Prop. of litters with 1 cub Prop. of litters with 2 cubs Prop. of litters with 3 cubs Sex ratio at time of 2nd birthday a Barren means cubs born more than 1 year ago, or cubs lost during first year so that female actually breeds in 2 consecutive years. �23 Annual Rate of Increase (%) 5 r-------------------------------------------, 4 r+ 3 r + + + l] +-1- 2r -I- + Maximum Mean Minimum -I- 1 I- o r---------------------------------------~r-~ -1 ~----------------------------------------~ Population after 30 Years 40,000 30,000 20,000 10,000 S30 F35 F45 S35 F40 S30 F35 F45 S35 F40 S35 F40 S30 F35 F45 S35 F40 S30 F35 F45 Constant Berry Double F~il11rp.~ H~1VP.~t o Both Figure 1. Annual rate of increase (%) and final population size (initial population size 10,000) for simulations with constant parameters (no annual variation), berry crop failures 1 year in 10, double the .normal harvest rate 1 year in 10, and a combination of berry crop failures 1 year in 10 and double the normal harvest 1 year in 10. Five different harvest scenarios were simulated: 530 - spring season with 30% of harvest females, 535 - spring season with 35% of harvest females, F35 - fall season with 35% of harvest females, F40 - fall season with 40% of harvest females, and F45 - fall season with 45% of harvest females. Minimum and maximum values were obtained from 100 repetitions. �24 Annual Rate of Increase (% ) t 10 ~------------------------------------------~ 8t : ttttt t tt o Maximum Mean Minimum ~------------------+-~~~4-4-;-4-~~----~ -2 -4 -6 ~ ~------------------------------------------~ Population after 30 Years ,--------------------------, 100,000 80,000 60,000 40,000 20,000 0 ~~~~~~~~~~~~~~~~~~~~=u~~ None S35 F40 S30 F35 F45 S35 F40 S30 F35 F45 S30 F35 F45 S35 F40 S30 F35 F45 S35 F40 5% 10% 15% 20% Figure 2. Annual rate of increase (%) and final population size (initial population size 10,000) for simulations of 5 harvest levels (0, 5, 10, 15, and 20%) for 5 seasons (spring 30 and 35% females in harvest and fall 35,40, and 45% females in harvest, see Fig. 1). The model includes annual variation of berry crop failures 1 year in 10 and double the normal harvest 1 year in 10. Minimum and maximum values were obtained from 100 repetitions. �25 Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS State of Project Colorado No. W-153-R-9 Mammals Work Plan No. 7 Covered: Author: July Jacqueline Personnel: Research Hultispecies Job No. Period REPORT Mammals Library Inyestigations Publication, Services Editing, and 1, 1995 - June 30, 1996 A. Boss Jacqueline A. Boss and Nancy W. McEwen ABSTRACT During the Segment the following were accomplished: * 2 publications were purchased at the request of Mammals Researcher personnel and placed into the Colorado Division of Wildlife Research Center Library collection. * 12 free reports and short publications from state or federal agencies or from private sources were located, ordered, and obtained for use by mammals Research personnel. * 40 theses or books were obtained on Interlibrary for use by Mammals Research personnel. * 1,243 individual articles were located for use by Mammals Research personnel. * 15 manuscripts by Mammals accepted for publication. * 8 manuscripts were prepared personnel for peer review. Research Loan or as gifts and delivered personnel and submitted on request were published by Mammals or Research ��27 MAMMALS PUBLICATION, EDITING Jacqueline AND LIBRARY SERVICES A. Boss P. N. OBJECTIVE To provide a centralized support program for manuscript editing services to facilitate publishing results of research conducted Federal Aid project W-153-R. SEGMENT and library by staff of OBJECTIVE To provide a centralized support program for Mammals Research editing, library, and publishing services so that Mammals Research personnel can be most efficient in publishing results of their research. SUMMARY OF SERVICES Publications Purchased with Mammals Research Funds and Placed in the Research Center Library Gray, G. G. 1993. Wildlife and people: the human dimensions of wildlife ecology. Chicago, IL : University of Illinois Press. 260pp. Grenfell, B. T. and A. P. Dobson. 1995. Ecology of infectious diseases in natural populations. Melbourne: Cambridge University Press. 521pp. Theses and Books Loan or as Gifts Obtained on Interlibrary for Use by Researchers Archon, M. 1992. Ecology of the San Joaquin kit fox in western Merced County, California. M.S. Thesis, California State University Fresno, CA. 45pp. Alberta Forestry, Lands and Wildlife. Fish and Wildlife Division. 1992. Management plan for cougar in Alberta. Wildlife management planning series; No.5. Edmonton, Alberta: Alberta Forestry, Lands & Wildlife. Fish & Wildlife Div. 91 leaves. Atkinson, D. E. 1976. Population dynamics and predator-prey relationships of the Carmen Mountains white-tailed deer. M.S. Thesis, Texas A&M University, College Station, TX. 89pp. Bailey, T. N. 1993. The African leopard: ecology and behavior of a solitary felix. New York : Columbia University Press. 429pp. Barlow, J. C. 1965. Land mammals from Uruguay: ecology and zoogeography. Ph.D. Dissertation, University of Kansas, Lawrence, KS. 346 leaves. Broomier, A. I., and G. M. Van Dyne, eds. 1980. Grasslands, systems analysis, and man. New York : Cambridge University Press. 950pp. Brown, F. ed. 1993. International meeting on transmissible spongiform encephalopathies ~ impact on animal and human health. New York : Karger. 233pp. Craig, D. L. 1986. The seasonal food habits in sympatric populations of puma (Puma concolor), coyote (Canis latrans), and bobcat (Lynx rufus) in the Diablo Range of California. M.A. San Jose State University, San Jose, CA. 61pp. Dowd, M. 1993. Social influences on declining number of hunters in Texas. M.S. Thesis, Texas A&M University, College Station, TX. 133p. Drake, J. A., [et a1.]. eds. 1989. Biological invasions: a global perspective. New York: J. Wiley. 525p. �28 Dubey, J. P., C. A. Speer, and R. Fayer. 1989. Sarcocystosis of animals and man. Boca Raton, FL : CRC Press, Inc. 215pp. Ehrlich, P., and A. Ehrlich. 1981. Extinction: the causes and consequences of the disappearance of species. New York : Random House. 305pp. Firchow, K. M. 1986. Ecology of pronghorns on the Pinon Canyon Maneuver Site, Colorado. M.S. Thesis, Virginia polytechnic Institute and State University, Blacksburg, VA. 83pp. Fischer, C. L. 1993. Anti-predator behavior of Gunnison's prairie dogs in response to aerial predators. M.S. Thesis, Northern Arizona University, Flagstaff, AZ. 32pp. Fuller, D. P. 1993. Black bear population dynamics in western Massachusetts. M.S. Thesis, University of Massachusetts, Amherst, MA. 136pp. Haltenorth, Th. 1953. Die wildkatzen der alten welt : eine ubersicht uber die untergattung Felis. Leipzig: Akademishe Verlags Gesellschaft Geest & Portig K.-G. 166pp. Hengeveld, R. 1989. Dynamics of biological invasions. New York: Chapman & Hall. 160p. Hudson, W. L. 1993. Density estimation, population dynamics, and dispersal· of red and gray foxes in central west Virginia. M.S. Thesis, West Virginia University, Morgantown, WV. 142pp. Iriarte, J. A. 1988. Feeding ecology of the patagonian puma (Felis concolor patagonia) in Torres del Paine National Park, Chile. M.S. Thesis, University of Florida, Gainesville, FL. 80pp. Leroux, N. 1993. What are biological species? The impact of the current debate in taxonomy on the species problem. M.S. Thesis, McGill University, Montreal, Quebec, Canada. 133pp. Markgren, G. 1975. Winter studies on orphaned moose calves in Sweden. Sweden : Svenska Jagareforbundet. Viltrevy: Swedish Wildlife; Vol. 9(4)193-219. Mooney, H. A., and J. A. Drake, eds. 1986. Ecology of biological invasions of North America and Hawaii. New York: Springer-Verlag. 321p. Musgrave, R. s., and M. A. stein. 1993. State wildlife laws handbook. Rockville, MD : Government Institutes, Inc. 840pp. New Mexico Dept. of Game and Fish. 1967. New Mexico wildlife management. Santa Fe, NM : New Mexico Dept. of Fish & Game. 250pp. Noble, W. o. 1994. Characteristics of spring foraging ecology among black bears in the Central Coast Ranges of Oregon. M.S. Thesis, Oregon State University, Corvallis, OR. 96pp. Okel10, M. M. 1993. Pocket gopher (Thomomys talpoides) food preferences, habitat relationships, and damage prevention. M.S. Thesis, University of Idaho, Moscow, ID. 64pp. Redford, K. H., and J. F. Eisenberg, eds. 1989. Advances in neotropical mammalogy. Gainesville, FL : The Sandhill Crane Press, Inc. 614pp. Robinson, J. G., and K. H. Redford. 1991. Neotropical wildlife use and conservation. Chicago: University of Chicago Press. 520pp. Shaw, H. G. 1989. Soul among lions: the cougar as peaceful adversary. Boulder, CO : Johnson Printing Company. 140pp. Schmidly, D. J. 1977. The mammals of Trans-Pecos Texas: including Big Bend National Park and Guadalupe Mountains National Park. College Station, TX : Texas A&M University Press. 225pp. Schulte, B. A. 1993. Chemical communication and ecology of the North American beaver (Castor canadensis). Ph.D. Thesis, State University of New York, Syracuse, NY. 194pp. Schwab, F. E. 1985. Moose habitat selection in relation to forest cutting practices in northcentral British columbia. Ph.D. Thesis, University of British Columbia, Vancouver, BC. 176pp. �29 Shull, Scott D. 1994. Management of nuisance black bears (Ursus americanus) in the interior highlands of Arkansas. M.S. Thesis, University of Arkansas, Fayetteville, AR. 101pp. Sigman, M. J. 1977. The importance of the cow-calf bond to overwinter moose calf survival. M.S. Thesis, University of Alaska, Fairbanks, AK. 185pp. U.S. Dept. of Ag. Animal and Plant Health Inspection Service, U.S. Forest Service, and U.S. Dept. of Interior. Bureau of Land Management. 1993. Animal damage control program : supplement to the draft environmental impact statement: Volume 1 of 2. Washington, D.C. : US Dept. of Ag. & US Dept. of Interior. various pagination. Wagner, F. H. 1988. Predator control and the sheep industry. Claremont, CA : Regina Books. Contemporary issues in natural resources and environmental policy; #1. 230pp. Waller, J. S. 1992. Grizzly bear use of habitats modified by timber management. M.S. Thesis, Montana State University, Bozeman, MT. 64pp. Wauer, R. H. 1973. Naturalist's Big Bend: an introduction to the trees and shrubs, wildflowers, cacti, mammals, birds, reptiles and amphibians, fish, and insects. College Station, TX. : Texas A&M University Press. 149pp. Ecology of mountain lions in the Sun River area of Williams, J. S. 1992. northern Montana. M.S. Thesis, Montana State University, Bozeman, MT. 1l0pp. Zumbaugh, D. M. 1984. Natural history of foxes in Kansas. M.S. Thesis, Fort Hays State University, Hays, KS. 37pp. Reference Document Location and Deliyery The Research Center Library staff also located and delivered approximately 1,255 individual articles or free documents on request for Mammals Researcher personnel during this segment. Manuscripts Job Progress Published Reports; FY 1995-96 Federal Aid. All studies. Fitzhugh, E. L., and A. E. Anderson. 1996. Trends in lion mortality in the western United states. In: Fifth mountain lion workshop : 27 February 1 March, 1996 : San Diego, California: abstracts. Organized by: Calif. Dept. of Fish & Game, South. Calif. Chapt. of the Wildl. Society. p.8. Franklin, A. B., and T. M. Shenk. 1995. Meta-analysis as a tool for monitoring wildlife populations. In: Integrating people and wildlife for a sustainable future : proceedings of the First International Wildlife Management Congress. eds. Bissonette, J. A. and P. R. Krausman. Bethesda, MD : The Wildlife Society. 484-487pp. Gill, R. B. 1996. The wildlife professional subculture : the case of the crazy aunt. Hum. Dim. Wildl. 1(1):60-69. Jessup, D. A., E. T. Thorne, M. W. Miller, and D. L. Hunter. 1995. Health implications in the translocation of wildlife. In: Integrating people and wildlife for a sustainable future : proceedings of the First International Wildlife Management Congress. eds. Bissonette, J. A. and P. R. Krausman. Bethesda, MD : The Wildlife Society. 381-385pp. Kraabel, B. J., M. W. Miller, D. M. Getzy, and J. K. Ringelman. 1996. Effects of embedded tungsten-bismuth-tin shot and steel shot on mallards (Anas platyrhynchos). J. of Wildl. Diseases 32(1):1-8. Kufeld, R. C. 1996. Status and management of moose in the Colorado State Forest and adjacent area on North Park. In: Colorado State Forest Ecosystem Planning Project : strategic plan. Colorado State Board of Land Commissioners. Colo. Dept. of Natural Resources. App. J. �30 Miller, M. W., and C. W. Mccarty. [1996] Models: potential applications in managing brucellosis in the Greater Yellowstone area. In: National Brucellosis Symposium, Jackson Hole, Wyoming, September, 1994. Thorne, E. T., P. Nicoletti, M. Boyce, and T. J. Kreeger (eds.). (in press) Miller, M. W., M. A. Wild, and W. R. Lance. 1996. Efficacy and safety of naltrexone hydrochloride in antagonizing carfentanil citrate immobilization in captive Rocky Mountain elk (Cervus elaphus nelsoni). J. Wildl. Dis. 32: 234-239. Pojar, T. M. 1996. Colorado pronghorn: compatibility and conflicts with agriculture. Wildlife Information Leaflet # 116. Fort Collins, Co Colo. Div. of Wildl. 7pp. Pojar, T. M. 1996. Helicopter net-gun capture of pronghorn. In: 17th biennial pronghorn antelope workshop. (in press) Reed, D. F. 1995. Preservation of small reserves. BioSci. 45(8):516. Reed, D. F. 1996. Corridors for wildlife. Sci. 271(5246):132. Shenk, T. M., and G. C. White. 1995. Detecting density dependence from temporal trends in demographic parameters using logistic regression. Bull. Ecol. Soc. Am. (Suppl.) 76(2): 95 (abstract) Shenk, T. M., and G. C. White. 1995. Detecting density dependence from temporal trends in demographic parameters using logistic regression. The Wildlife Society, Bethesda, MD (abstract) Shenk, T. M., A. B. Franklin, and K. R. Wilson. 1995. A model to estimate the annual rate of golden eagle population change at the Altamont Pass Wind Resource Area. In: Proceedings of National Avian - Wind Power Planning Meeting II. Richardson, W. J. (ed.) King City, Ontario: LGL Ltd., Environmental Research Associates. (in press) Manuscripts in Review FY 1995-96 Andelt, W. F., R. L. Phillips, R. H. Schmidt, and R. B. Gill. 1996. Foothold traps : an overview of the biological, social, and political issues surrounding a public policy controversy. Wildl. Soc. Bull. (in review) Baker, D. L., G. W. Stout, and M. W. Miller. 1996. A diet supplement for captive wild ruminants. -J. Zoo Wildl. Med. (in review) Kraabel, B. J., and M. W. Miller. 1996. Effect of simulated stress on susceptibility of bighorn sheep neutrophils Pasteurella haemolytica leukotoxin. J. Wildl. Dis. (in review) Kufeld, R. C. 1996. Design and evaluation of an expandable radio-collar for male Deer. Wildl. Soc. Bull. (in review) Kufeld, R. C., and D. C. Bowden. 1996. Movements and habitat selection of Colorado moose (Alces alces shirasi). Alces (in review) Manfredo, M. J., D. Fulton, J. Pate, and R. B. Gill. 1996. Public acceptance of wildlife trapping in Colorado. Hum. Dim. Wildl. (in review) Miller, M. W., J. A. Conlon, H. J. McNeil, J. M. Bulgin, and A. C. S. Ward. 1996. Experimental evaluation of a multivalent Pasteurella haemolytica vaccine in captive bighorn sheep (Ovis canadensis). J. wildl. Dis. (in review) Spraker, T. R., M. W. Miller, E. S. Williams, D. M. Getzy, W. J. Adrian, G. G. Schoonveld, R. A. Spowart, K. I. O'Rourke, J. M. Miller, and P. A. Merz. 1996. Spongiform encephalopathy in free-ranging mule deer (Odocoileus hemionus), white-tailed deer (Odocoileus virginianus), and Rocky Mountain elk (Cervus elaphus nelsoni) in northcentral Colorado. J• .Wildl. Dis., (in review) Prepared by Jacqueline Librarian A. Boss �31 Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS state of Project REPORT Colorado No. W-153-R-9 Mammals Research Work Plan No. Multispecies Job No. Mammals 1 Research Administration Period Author: Covered: Inyestigations July 1, 1995 - June 30, 1996 R. Bruce Gill Personnel: R. Bruce Gill and Diane K. Haerter ABSTRACT Six federally funded and 1 independently funded jobs were planned, budgeted, supervised, and administered during the Segment. Three jobs involved multiple species dimensions, 2 invoved deer research projects, 1 involved elk research, and 1 involved moose research. All research objectives were accomplished within the allocated time limits and budget constraints. Forty reports, short publications, and books were added to the Research Library holdings at the request of Division research staff members. Two manuscripts/books were prepared by Mammals 1 staff members in professional journals and/or symposia proceedings. and published ��33 Mammals Research Administration R. Bruce Gill P.M. OBJECTIVE Administer research studies within productivity and the lowest cost. the Mammals Segment 1. 1 Research Unit for the highest Objectives Supervise and administer research Mammals Research Section. on deer, elk, and moose within the RESULTS Seven projects were active during the segment. segment objectives completed successfully for all 7 projects. Highlights include: were { Approximately 35% of the time of Mammals Program Leader allocated to Mammals 1 Research was consumed by reorganization and Management Review implementation tasks. { Forty reports, short publications, and books were added to the Research Library holdings at the request of Division research staff members. { Three articles/books authored by Mammals 1 Research accepted for publication during the segment. { Two professional/technicai manuscripts were in final stages of preparation by members of the Mammals 1 Research staff or were in the peer review process of professional journals. { All program projects and activities were achieved allotted time and allocated fiscal resources. Prepared by: R. Bruce Gill Mammals Program Leader staff were within the ��35 Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS state of project REPORT Colorado No. W-153-R-9 Mammals Research Work Plan No. Deer Job No. Compensatory Effects of Harvest in a Mule Deer Population Period Covered: Author: Personnel: July 1, 1995 - June 30, 1996. R. M. Bartmann, E. White Investigations and G. C. White. o. Younkin. ABSTRACT ~ Deer density estimates from aerial mark-recapture surveys indicated similar densities on the 2 study units (control 26.0/km2 and treatment 27.6/km2). Fawn survival on the treatment unit (0.766, SD 0.051) was higher than on the control unit (0.624, SO 0.057) as it has been for the previous 5 years, but the difference was not significant (~ = 0.079). There were no differences in fawn weight, total body length, left hind foot length, or weight/length ratio between the 2 study units (~~ 0.181). Adult doe survival remained high on both units (control 0.974, SO 0.026 and treatment 0.893, SO 0.051). Field data collection for this study was terminated 30 June 1996. Work during the next segment will involve data analysis and preparation of at least 1 manuscript for publication. . ��37 COMPENSATORY EFFECTS OF HARVEST IN A MULE DEER POPULATION Richard M. Bartmann and Gary C. White P. H. OBJECTIVES 1. Increase the winter survival rate of mule deer fawns by lowering deer density to reduce competition for forage during winter. 2. Increase the harvest rate of deer through increased productivity of adult does and decreased natural mortality of fawns resulting from closer alignment of population size with carrying capacity. SEGMENT total OBJECTIVES 1. Maintain the winter population of mule deer on the Ridge a density <40/km2 for 5 years. 2. Estimate winter survival rates of fawns on control 3. Estimate units. annual survival rates of adult 5. Estimate condition of fawns on control females treatment and treatment on control and treatment unit at units •. and treatment units. METHODS Methods, except for population estimation, remained the same as previously reported by Bartmann (1990) and Bartmann and White (1991) with modifications by Bartmann and White (1992). All deer were captured with helicopter net guns by Helicopter wildlife Management, Inc. Previous deer population estimates from aerial line transects showed the expected decline in the population on the treatment unit due to late season harvests. However, population estimates on the control unit also started declining until they were nearly the same as on the treatment unit. To see if there may have been a problem with line transects, an aerial mark-recapture survey was used for the 1995-96 winter (Bartmann et al. 1987). Radiocollared fawns were used as the marked population. A 5 x 10-cm vinyl tab was riveted to the top of each radiocollar to enable quick identification of marked fawns. Tabs were color-coded by unit: orange for control and white for treatment. All fawns were located immediately prior to the surveys to verify the number on each unit. Three complete-coverage helicopter flights (morning, midday, and afternoon) were made on each unit with 2 observers and a pilot. Field data collection for this study was terminated as of 30 June 1996. Work during the final segment will consist of data analysis and preparation of at least 1 major publication. �38 RESULTS AND DISCUSSION Maintain Population Aerial mark-recapture surveys were flown on the Ridge study area 6-7 January 1996. As for the previous winter, density estimates were similar for the 2 units: control 26.0 deer/km2 and treatment 27.6 deer/km2• This supports the close estimates for 1994-95 from line transects. The reason for the population decline on the control unit is unknown. It is not supported by fawn or doe survival estimates. This leaves decreased productivity as a possible explanation, but we did not collect these data. ~ CONTROL •• ~••• TREATMENT 120r----~---------------------------------------~ YEAR Fig. 1. Deer density estimates (w/95% confidence intervals) from aerial line transects and mark-recapture surveys (1995-96 only) on control and treatment units during early to mid-winter. Fawn Survival Helicopter netgunning was done 14 November-20 December 1995 with effort alternated daily between control and treatment units. Little snow cover and reduced populations made fawn captures difficult and were the reasons for the prolonged capture period and our failure to capture the desired 80 fawns on each unit. We radioco11ared 77 fawns on the control unit and 74 on the treatment unit. For the Kaplan-Meier survival estimates, 1 fawn was censored on the control unit and 3 on the treatment unit. Snow cover remained scant during the first half of the 1995-96 winter, but started increasing in mid-January. As during the past 5 winters, fawn survival on the treatment unit (0.766, SE 0.051) was higher than on the control (0.624, SE 0.057), but the difference was not significant.(£ = 0.079) (Table 1). Predation was the leading There were few starvation 2 years. Adult mortality cause for fawns during 1995-96 (Table 2). losses on either unit as there has been the previous Poe Survival No new adult does were radiocollared in 1995. Estimated adult doe survival from 1 December to 30 June was 0.974 (SE 0.026) on the control unit and 0.893 (SE 0.051) on the treatment unit (Table 3). The difference was not significant (£ = 0.190) Condition of Fawns All 3 fawn body measurements and the weight/length ratio (control 0.261, SD 0.019 and treatment 0.265, SD 0.023) on the treatment unit exceeded those on the control unit but none of the differences were significant (£ ~ 0.181) (Table 4). �39 Table 1. Kaplan-Meier estimates of survival rates (S) for radio-collared mule deer fawns on control and treatment units of the Ridge study area in Piceance Basin, Colorado, from time of collaring in November and December until the following 15 June 1982-83 through 1995-96. Hunting mortalities are censored. l:t:~atm~nt ynit CQntt:Ql Ynit n Winter 1982-83 1983-84 1984-85 1985-.86 1986-87 1987-88 1988-89 1989-90 1990-91 1991-92 1992-93 1993-94 1994-95 1995-96 28 28 34 59 60 32 34 38 34 28 80 80 82 76 .s. SE(.s.) n 0.321 0.071 0.196 0.537 0.431 0.241 0.270 0.779 0.320 0.456 0.112 0.550 0.701 0.624 0.088 0.049 0.078 0.070 0.064 0.077 0.083 0.078 0.090 0.107 0.036 0.056 0.051 0.057 31 32 26 58 58 28 28 44 36 42 74 80 80 71 LITERATURE .s. SE(.s.) 0.387 0.033 0.431 0.439 0.471 0.107 0.445 0.745 0.339 0.548 0.148 0.767 0.825 0.766 0.087 0.033 0.105 0.070 0.067 0.058 0.096 0.070 0.106 0.098 0.043 0.048 0.042 0.051 E. of equal .s.ct.) 0.578 0.774 0.075 0.157 0.565 0.006 0.509 0.659 0.909 0.481 0.102 0.006 0.079 0.079 CITED Bartmann, R. M. 1990. Compensatory effects population. Colo. Div. Wildl., Wildl. of harvest Res. Rep. in a mule deer July: 187-196. _____ , and G. C. White. population. colo. 1991. Compensatory Div. Wildl., Wildl. effects of harvest in a mule Res. Rep. July:27-40. deer _____ , and G. C. White. population. colo. 1992. Compensatory Div. Wildl., Wildl. effects of harvest in a mule Res. Rep. July:27-37. deer _____ , , L. H. Carpenter, and R. A. Garrott. 1987. Aerial markrecapture estimation of confined mule deer in pinyon-juniper woodland. J. Wildl. Manage. 51:41-46. Prepared by _ Richard LSSR M. Bartmann III Dr. Gary C. White Professor �40 Table 2. Cause of mortality for radiocollared mule deer fawns on control and treatment units of the Ridge study area in Piceance Basin, Colorado, from time of collaring in November and December until the following 15 June 1982-83 through 1995-96. Percentages are of total uncensoreda fawns. Winter n C§iln~Qt:§ilg Hunting other stat:::latiQn % No. MQt:talit~ ~5:U.UHil ~t:§ilgatiQn Qtb§ilt: No. % No. % CQntt:Ql unit 1982-83 1983-84 1984-85 1985-86 1986-87 1987-88 1988-89 1989-90 1990-91 1991-92 1992-93 1993-94 1994-95 1995-96 28 28 34 59 60 32 34 38 34 28 80 80 82 77 1 7 11 6 2 5 14 9 7 1 4 5 3 2 1 i5 22 16 10 14 22 10 3 6 7 12 3 56 79 59 21 26 73 34 14 24 33 15 4 6 8 4 4 5 7 17 15 14 19 15 31 5 17 10 3 50 22 15 19 40 14 64 29 19 26 2 3 2 11 13 1 1 7 10 9 22 25 4 4 3 2 26 8 9 9 15 8 39 11 11 14 1 7 3 2 7 3 3 3 7 11 9 2 4 15 6 7 24 14 12 14 9 14 12 3 2 7 3 1 1 5 5 4 4 4 7 8 4 3 14 2 2 18 20 13 20 17 10 11 5 5 Tt:eatment unit ., 1982-83 1983-84 1984-85 1985-86 1986-87 1987-88 1988-89 1989-90 1990-91 1991-92 1992-93 1993-94 1994-95 1995-96 31 32 26 58 58 28 28 44 36 42 74 80 80 74 1 1 4 9 5 5 9 9 6 3 9 7 9 1 10 3 11 15 27 8 17 16 19 9 6 8 7 23 3 1 4 50 87 36 35 30 68 36 20 40 29 34 4 1 6 a Uncensored fawns are those that were not killed by hunters, that had nonfailing radios, or that had collars that did no.t drop off prematurely. �41 Table 3. Kaplan-Meier estimates of annual (1 Dec-30 Nov) survival rates (A) for radiocollared adult female mule deer on control and treatment units of the Ridge study area in Piceance Basin, Colorado, 1982-83 through 1995-96. Hunting mortalities are censored. Winter n Control unit Hunting ~ SE(~) n 1982-83 1983-84 1984-85 1985-86 1986-87 1987-88 1988-89 1989-90 1990-91 1991-92 1992-93 1993-94 1994-95 1995-96a 10 15 9 25 27 14 7 23 39 41 46 49 47 38 0.800 0.779 1.000 0.917 0.756 0.818 0.857 1.000 0.969 0.758 0.716 ·0.917 0.872 0.974 11 15 10 21 18 10 5 28 41 43 52 46 44 39 a Survival 1 3 ·2 1 rate estimates 0.126 0.113 0.056 0.087 0.116 0.132 0.031 0.071 0.067 0.040 0.049 0.026 for 1995-96 Treatment Hunting ~ 1 9 12 6 7 unit SE(~) 0.909 0.929 1.000 0.900 0.878 1.000 0.800 1.000 0.906 0.809 0.719 0.804 0.908 0.893 are only through 0.087 0.069 0.067 0.081· 0.179 0.065 0.071 0.066 0.058 0.044 0.051 30 June £ of equal ~ 0.448 0.271 1.000 0.821 0.432 0.329 0.854 1.000 0.474 0.608 0.910 0.123 0.513 0.190 1996. �42 Table 4. Weights (kg) and body measurements (cm) of mule deer fawns trapped on control and treatment units of the Ridge study area in Piceance Basin, Colorado, 1982-95. Total t!Q!l~l~ngtb H~igbt n Year it so CQntrQl 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 28 28 34 60 60 33 34 40 35 28 82 85 83 76 34.6 31.7 32.2 32.6 31.9" 29.9 29.5 32.7 30.8 30.7 30.4 30.7 33.6 34.7 3.10 4.40 4.65 4.02 3.89 3.60 3.10 3.31 4.29 3.58 3.80 3.91 3.59 3.49 Treatment 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 31 32 26 60 61 28 30 47 36 43 83 80 82 74 "n n b c n 58 27 30 32.8c 32.3 32.3 32.3 31.7 30.2 28.8 30.6 30.7 32.6 30.3 32.9 33.9 35.5 4.18 3.12 5.07 4.62 4.13 5.34 4.13 3.33 4.42 3.54 4.68 3.96 3.90 3.86 it so Left hind fQQt l~ngtb it so unit 124.0 124.2· 123.9 124.4 128.1 127.3 123.8 131.0 126.5 128.2 126.8 129.1 132.9 132.8 4.64 5.65 7.25 6.26 6.53 6.12 7.83 5.81 9.02 5.47 6.67 6.17 7.13 5.47 41.1 40.6 40.8 41.1 41.0 40.8 41.0 41.8 40.8 40.6b 40.6 40.5 41.7 42.0 1.08 1.73 1.53 1.48 1.95 1.72 1.37 2.16 1.75 1.32 1.58 1.68 1.41 1.31 5.45 5.53 7.25 6.28 6.58 8.86 7.07 5.33 6.54 6.11 7.94 5.76 6.26 5.04 41.1 40.6 40.8 40.8 41.0 41.2 40.6 40.7 41.1 40.8 40.5 41.3 41.9 42.2 1.65 1.34 1.89 1.77 2.11 1.66 1.88 1.41 1.69 1.29 1.77 1.43 1.29 1.53 unit 121.7 123.6 124.7 124.4 125.9 127.5 124.9 126.6 127.8 129.8 126.7 131.6 131.0 133.4 �Colorado Division Wildlife Research July 1996 of Wildlife Report JOB FINAL REPORT state of Project ~C~o~l~our~a~d~o~ _ No. W-153-R-9 Mammals Research Work Plan No. Elk Inyestigations Job No. Effects Seasons Period Covered: Author: Personnel: of Early Hunting on Elk Distribution July 1, 1995 - June 30, 1996 D. J. Freddy M. M. Conner, G. C. White, J. Madison, J. Ellenberger Abstract A final study plan for the experimental evaluation of the effects of archery and muzzleloading hunting on elk movements during early fall hunting seasons was completed by Mary M. Conner, Ph.D. candidate. A copy of the final study plan is included in this report. During fall 1995, radio collared elk were monitored to document generalized movements of elk during early fall hunting seasons. This monitoring concluded the pilot study, and analysis of preliminary data collected during 1992-1995 is included in the final study plan. Additionally, radio collars were purchased so that 80 adult female elk could be captured and marked during July 1996 when field work for the experimental portion of this project is scheduled to begin. ��45 EFFECTS OF EARLY JOB FINAL REPORT HUNTING SEASONS ON ELK DISTRIBUTION David J. Freddy P, N. OBJECTIVE Evaluate the effects of early big game hunting seasons muzzleloading deer and elk seasons) on the distribution River Data Analysis Unit. SEGMENT (archery and of elk in the White OBJECTIVES 1. Complete during final study plan and analysis 1992-95. of preliminary data 2. Purchase and prepare radio collars for marking 80 adult July 1997 when field work is scheduled to begin • collected female elk during . INTRODUCTION Preliminary data collected from 1992 through 1995 suggested elk were seeking refuge areas, often private lands, in response to disturbance associated with early hunting seasons. Based on these results, and the varied interests and often opposing concerns of several user groups, we opted to design and implement an 2-year experiment to evaluate the effects of early hunting seasons on elk movements from public to private lands, or non-refuge to refuge areas. RESULTS We designed an experiment whereby opening dates of early fall hunting seasons will be 3 weeks apart on 2 adjacent hunting units. These opening dates will be 'early' and 'late' and will be crossed-over between hunting units during the second year of this project. Movements of elk will be monitored intensively for 30 days prior to and after opening dates of seasons. A final study plan is attached and serves as the primary result for this segment's work. Radio collars for 80 adult female elk were purchased during that collars would be available for placement on elk during Prepared by: David J. Freddy Life Science Researcher IV this July segment 1996. so ��47 ELK MOVEMENTS IN RESPONSE TO EARLY-SEASON HUNTING IN mE WHITE RIVER AREA STUDY PLAN Mary Conner Department of Fishery and Wildlife Biology Colorado State University Fort Collins, Colorado 80523 Spring 1996 �48 EXECUTIVE SUMMARY The White River elk herd (Cervus elaphus) has been growing since its re-establishment to the area in 1929 and its numbers are now at the upper bounds of the desired management objectives. The number of hunters using the area has grown along with the elk herd, with especially significant increases in the number of early-season archery hunters, during the past ten years. Also over the past ten years, there have been increasing observations and complaints that elk have been moving off easily accessible public lands to lower elevation private lands or to remote and inaccessible areas during the early-season hunting period. The increasing disturbance of early-season hunting may be causing the elk to move off their summer ranges before fall migration. Early movement has lead to complaints by local landowners onto whose land the elk are moving, by resource managers, and by early-season hunters. All parties indicate that it is not the number of elk in the area causing the problem, but the distribution of the elk. Documented responses of elk to hunting from previous studies includes increased movement, movements away from hunted areas, movement into inaccessible areas, and movement into no-hunting areas. There is some:evidence that elk response may be dependent on the hunter density and the amount of escape cover in the vicinity. Further, there is an indication that the responses of elk to hunters is of short duration. All the previous elk studies with respect to hunting have been observational in nature and have not tested for a cause and effect relationship between hunting activity and elk movement. A preliminary study on the proposed study area was conducted by George Bear and Jeff Madison from the Colorado Division of Wildlife. They trapped and radio-collared 20 elk cows in 1992. From 1992 to 1995 the radio-collared elk were intensively monitored during August and September, approximately one month before and one month after the opening day of archery hunting. Corresponding to the opening of archery hunting season, all elk located on non-refuge areas (areas easily accessible to hunters) moved to refuge areas (private land or wilderness area not easily accessible to hunters), while all elk located in refuge areas stayed in refuge areas. The mean date of movement was not significantly different from the opening date of early-season hunting. Although the proportion of elk moving from non-refuge to refuge areas and the date of movement indicated that elk movement correlates with the opening of early-season hunting, there are several alternative hypotheses that could also explain these movements. To infer a casual relationship between elk movement and early-season archery hunting, the study needs to be continued with a manipulative experiment. The objective of this study is to evaluate the impacts of early-season hunters on the movement and distribution of elk with the basic premise or null hypothesis that early-season hunting does not effect elk movements. A manipulative experiment will be conducted to determine if the presence of hunters is contributing to the movement of elk off hunting accessible areas, to relatively hunting inaccessible areas. The principle objectives to answer this question are: 1. Test the hypothesis that early-season hunting disturbance of elk does not result in movement of elk from heavily hunted areas to lightly hunted areas. 2. Evaluate alternative hypotheses about causes of elk movement such as livestock grazing, woodcutting, recreationalists, weather, or forage quality. 3. Compare the movements of this heavily hunted elk population with movements of elk from concurrent studies in other parts of Colorado. Elk will be captured and radio-collared in Game Management Units (GMUs) 12,23,24, and 33, and treatment and control groups established. A total sample size of 80 radio-collared elk are recommended based on power analyses. Telemetered elk will be located at least twice a weekfor the month preceding and the month following the opening date of archery hunting. Locations will be labeled as either refuge or nonrefuge for the analysis. The experimental design is a crossover design with the study area split into two experimental units of the GMU areas north and south of the White River. In the first year of the study, one area would be treated with hunting moved forward one week, while the remaining area would be treated with �49 a hunting delayed two weeks from traditional opening day, yielding three week difference in opening dates. The treatments would be reversed in the second year of the study. Data from the United States Forest Service on livestock grazing areas, woodcutting permits, and recreational use will be used to evaluate the effects of these disturbances on elk movements during the study period. Livestock effects on elk movement will be further evaluated by locating livestock herds in thevicinity of radio-collared elk. Analysis of the elevation shifts and movements of the control and treatment elk will be used to evaluate the effects of weather and forage quality on elk movements. Results from this study will provide managers with scientifically defensible and publicly credible information that can be used to defme management alternatives for managing early-season elk distribution problems. The conclusions will have direct inference to the White River area and will also contribute to the body of scientific knowledge on elk in the western United States. Outputs from this research will include written reports, scientific publications and spatial use maps. �50 TABLE OF CONTENTS INTRODUCTION BACKGROUND '.' Elk Movement in Response to Hwnan Activities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Elk Movement in Response to Hunting Conclusions OBJECTNES STUDY AREA PILOT STUDY METHODS , Study Design '. . . . . . . . . . . . . . . .. Open hunting season one week earlier on one area (chosen randomly) and two weeks later on the remaining area. Reverse treatments for year two. . . . . . . . . . . . . . . . . . . . . . . . . .. Capture and Collaring . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Radio-Telemetry Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Hunter Sruveys Application of Results '. . . . . . . . . . . . . . . . . . . . .. LITERATURE CITED APPENDIX A. Logistic regression analysis for 1992 - 1995 : APPENDIX B. Alternative Study Plans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Alternative two - Close hunting one year on two GMUs(chosen randomly) and issue the full nwnber of permits for the other two GMUs. Reverse the treatments the following year. . Alternative three - Move the early-season hunting forward for two weeks on non-refuge areas only, then close hunting on public land and open it for the following two weeks on refuge areas only '.' APPENDIX C. Study Design Under Ideal Conditions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 52 53 53 54 56 56 57 58 61 63 63 68 69 69 70 70 73 82 82 83 84 FIGURES Figure 1. Nwnber of early-season archery hunters for DAU E-6 in the White River area, Colorado, 1984 - 1994 ~ Figure 2. White River study area. Figure 3. Example of a logistic regression curve to estimate the date of movement for an elk moving from a non-refuge area to a refuge area (A), and elk staying in a refuge area (B). Figure 4. Proportion of elk on refuge areas for elk in non-refuge and refuge areas before opening of early-season hunting in 1992 and 1993. Figure 5. Study design layout showing nested levels of experimental units. Figure 6. Hypothetical dates of elk movement to refuge areas if movement is caused by a factor other . than hunting that is common across the 2 areas (woodcutting, weather, forage quality etc.). . .. Figure 7. Hypothetical dates of elk movement to refuge areas if movement is caused by hunting Figure 8. Hypothetical dates of elk movement to refuge areas if movement is a response to hunting and has a learned behavior component. ,........................... 52 57 59 61 64 67 67 68 �51 TABLES Table 1. Locations of radio-collared elk before and after opening day of early-season hunting, White River elk herd, 1992 - 1994 Table 2. Mean dates of movement of elk from non-refuge to refuge areas, opening date of early-season hunting, and p-values from a t-test for differences for the White River elk herd, 1992 - 1994 ... Table 3. Example of crossover design blocking out effects of sheep grazing activity with a hunting and no-hunting treatment. Table 4. Layout of Latin square crossover design with corresponding model for alternative one Table 5. Sample size (number of collared elk) per half for treatment groups based on estimates of effect size and 90% power for primary response variables. Table 6. Layout of Latin square crossover design with corresponding model for alternative two Table 7. Layout of Latin square crossover design for alternative three 59 60 62 65 66 82 83 �52 INTRODUCTION The White River elk (Cervus elaphus) herd of Data Analysis Unit (DAU) E-6 has been one of the most heavily hunted, managed, and documented elk herds in Colorado (Boyd 1970, Freddy 1987, Gray et al. 1994). The herd has been growing in number since the year hunting resumed in the White River area in 1929 and is now at the upper bounds for the desired management objectives (Grayet al. 1994). The number of hunters using the area has grown along with the elk herd, with especially significant increases in the number of early-season archery hunters between 1985 and 1992 (Figure 1). ~ ~ 3,000 :::J ::I: ~ m 5 2,500 !G en Q) ~ 2,000 (II w '0 •... ..8 E :::J 1,500 Z 1,000 -t----+---+---r----+----t----+---+------i----l 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 Year Figure 1. Number of early-season archery hunters for DAU E-6 in the White River area, Colorado, 1984 1994. Along with the increase in early-season archery hunters has been an increase in the use of four-wheel drive (4WD) and all-terrain vehicles (ATV's) (Grayet al. 1994). Disturbance caused by an increasing number of early-season hunters and/or hunters using 4WO vehicles and ATV's may be causing elk to move off summer range areas early. Historically, the main migration from summer to winter ranges took place between late November and early January (Boyd 1970). However, Freddy (1987) noted that elk were commonly found in lower elevations, typical winter range, during the September and October hunting seasons, which was uncommon during the 1960's. Now there is evidence that elk are moving during August and early September during early-season archery hunting (Gray et al. 1994), from areas easily accessed by hunters to areas not easily accessed. Refuge areas are defmed as easily accessed areas that are primarily public forest lands with abundant road access, while non-refuge areas are difficult access areas; either private lands or wilderness areas with sparse or no public road access. Private lands have not typically been hunted by archery hunters, but have been hunted later during rifle season, thus providing elk a refuge from hunting during archery season. These movements and resulting redistribution of elk has lead to complaints by local private landholders, resource managers, and hunters (Grayet al. 1994). As the number of archery hunters continues to increase, early-season hunter induced elk movements could lead to elk redistribution with corresponding overpopulation problems in localized areas. Also, early fall movements may leave nutritious summer forage uneaten at the cost of over-grazing winter range. There is evidence that poor winter range correlates with low elk condition (Weber et al. 1984), possibly leading to �53 lower reproductive performance of female elk. In the White River area, there have been complaints by private landholders bordering public areas of range damage, complaints by resource managers that riparian areas are being damaged by this redistribution, and complaints by early-season hunters of lower success rates in the public areas they hunt. All parties indicate that it is not the number of elk in the area causing the problem, but the distribution of the elk (Gray et al. 1994). Although the phenomena of elk movements from public to private lands has been discussed, it is not well documented. A three-year pilot study from 1992 to 1994 was conducted on 14 to 20 radio-collared White River elk from Game Management Units (GMU) 12 ,23,24, and 33. During the study, all radiocollared elk located on non-refuge areas moved to refuge areas. For all years combined, the mean day of movement was not different than opening day of early-season hunting. Although a causal relationship was not established by this study, it provided strong evidence of a correlation between elk movements and the opening of early-season hunting. To more thoroughly understand elk response to hunting, this study describes a manipulative experiment to evaluate the cause and effect relationship between the movement of elk and the opening of early-season archery hunting in the White River DAU. The null hypothesis is that elk movements are the same on hunted and non-hunted areas, or that early-season hunting does not cause elk to move. Hypotheses on the timing and extent of other factors that may explain the elk movements will also be examined. BACKGROUND Elk Movement in Response to Human Activities Elk response to recreational activities such as hiking, cross-country skiing and driving seems to vary depending on how habituated the elk are and whether the population is hunted. In Rocky Mountain National Park, where elk were accustomed to people and were not hunted, elk showed little response to approaches of automobiles and people on foot, day or night (Schultz and Baily 1978). Another study of an unhunted elk population in Yellowstone National Park found variation in elk response to cross country skiers (Cassirer et al. 1992) depending on how accustomed the elk were to people. In areas of typically low human activity during the winter, elk responded with flight distances between 125 - 1,700 m, while in an area of high human activity, the flight distance was much less, ranging between 0 - 300 m. Interestingly, elk in the area of high human activity showed a three-fold increase in flight distance when they were disturbed just outside the 100ha area where people were present 24 hours each day (Cassirer et al. 1992). Thus, even habituated elk appear to fluctuate in their response to human activity patterns depending on their habituation. Acceptance of human activity may be a learned response of unhunted elk. Movements with respect to roads seems to be related to the habituation of elk to vehicles and to the level of road use. In Rocky Mountain National Park, where elk were not hunted and there was high traffic volume, Schultz and Bailey (1978) found that none of their 14 delineated elk behaviors changed with traffic volume and there was little or no avoidance of the roads in the winter. In contrast, on the Roosevelt National Forest, which is adjacent to the Rocky Mountain National Park, Rost (1975) found that a population of hunted elk avoided roads in winter ranges. Wright (1983) found a variable response; mean distance to jeep trails more than doubled (800 m to 2,100 m) during fall from just before and during hunting season. Elk may learn to accept human activity that poses no threat. However, behavior of elk in areas with respect to recreational activity has not been examined in hunted areas for comparison. Elk may also respond to the level of activity on a road. Wright (1983) found that elk had double the average distance to heavily traveled paved roads than to lightly traveled jeep trails. Similarly, Hershey and Legee (1982) found elk crossed secondary roads more frequently than primary roads. Czech (1991) found a significant increase in mean elk distances to a road after it was opened to the general public with a corresponding increase in traffic levels. If elk respond to different levels of activity on a road they likely also respond to different levels of hunting activity. �54 Effects of logging and mining disturbances on elk movements depend on the cover available near the disturbances. Edge and Marcum (1985) found normal elk movements were changed by logging disturbances; elk moved significantly longer distances away from logging areas than towards them. Typically, a buffer zone of between 500 - 1,000 m separated areas of high elk use from areas of disturbance depending on the cover in the area (Edge and Marcum 1985). Czech (1991) found that elk tolerance oflogging operations was correlated positively with proximity to hiding cover. In one of the few manipulative experiments on elk movements, elk calves were subjected to simulated surface-mining activities (Kuck et al. 1985). Compared to undisturbed calves, disturbed calves moved greater distances, used larger areas, showed greater use of coniferous forest, and lacked selection for favorable physiographic elements (Kuck et al. 1985). A strong case for a cause and effect relationship between elk movement and mining disturbance was built with this manipulative experiment. All studies that evaluated elk movements after the disturbance ended found that displacement appears to be temporary in most situations. The Yellowstone elk displaced by cross-country skiers typically returned close to their original locations after people left the area (Cassirer et al. 1992). Road proximity returned to pre-hunting distances after hunting season closed (Wright 1983) and mean distance to roads decreased soon after the roads were closed for the season (Hershey and Legee 1982). During weekends, and immediately following the end oflogging activities, elk moved back into logged areas (Ward 1976, Edge and Marcum 1985). Elk Movement in Response to Hunting How and to what extent hunting affects elk is not clearly understood (Adams 1982). Elk response to hunting pressure has been examined in several studies (Martinka 1969, Knight 1970, Craighead et al. 1972, Lemke 1975, Morgantini and Hudson 1979, Wright 1983) and two theories about elk response have been tendered. The first theory is that elk in recent years have learned to move to unhunted refuges and choose migration routes through lightly hunted areas (Altmann 1956, Martinka 1969) (learned behavior theory), while the second theory is that elk populations were decimated if they migrated through or lived in heavily hunted areas, whereas elk living in lightly hunted areas persisted (Rudd et al. 1983, Boyce 1991) (population redistribution by differential hunting pressure theory). Neither theory has been experimentally tested. Documented responses of elk to hunting includes movement away from hunters or heavily hunted areas, increased movement, movement to thick cover, shifts in circadian patterns, and elevation shifts in migration patterns. Most responses seem to occur before winter migration, but some studies have noted changes in migration patterns that were attributed to hunting pressures. Responses of the elk may depend on density of hunters, roughness of terrain, or density of cover. Altmann (1956) described an evasive "migration" of elk hunted adjacent to Yellowstone National Park. With opening of hunting season, movements of these elk became relatively long (5 - 13 km), and they stopped moving only when they reached the sanctuary of the Park. Similarly, Martinka (1969) found that all of 12 marked elk located in the National Elk Refuge just prior to hunting season moved between 8 and 22 km to areas closed to hunting in Grand Teton National Park. Other studies have found less dramatic movement distances to refuge areas. With opening of hunting season, elk moved to densely forested (Irwin and Peek 1979) or shrubby (Morgantini and Hudson 1979) areas adjacent to their typical areas of activity, which provided refuge from hunting. In a study of elk responses to hunting pressure, elk moved toward winter range earlier than normal fall migration time, and the number of locations on private land increased during rifle season (Wright 1983). Additionally, Wright (1983) found that mean straight-line distances traveled by radio-collared elk tripled during hunting season versus the months before hunting season. Hunting disturbances may also affect the timing of cropland use; a hunted elk population limited open cropland grazing to early morning hours (roughly 0200-0400), while an unhunted population would enter cropland during daylight hours (Strohmeyer and Peek in press). A reversal in downward elevation migratory movement of elk, which coincided with the opening of the hunting season, was found by Knight (1970) and Morgantini and Hudson (1979). While some elk appear to have learned to move to refuge areas, elk in refuge areas may learn to stay there in response to hunting pressure. In Jackson Hole, both resident and migratory elk use the National Elk �55 Refuge for their winter range (Martinka 1969). Of the 183 marked elk in the study, resident elk were defmed as elk located July through August within 15 km of the winter range, while migratory elk were not located in . the area. In the fall, both resident and migratory elk had to traverse areas where hunting was permitted while moving to the National Elk Refuge winter range. Resident elk were conditioned to hunters presence and tended to remain on areas closed to hunting while migratory elk moved through them to the winter range. These studies of elk moving to, or staying in, refuge areas support the theory that elk are responding, possibly as a learned behavior, to hunting pressure. Other studies found evidence to indicate elk are not responding to hunting pressure, but are being differentially removed from heavily hunted areas. In Wyoming, a resident population of elk just east of Yellowstone National Park were being detrimentally reduced by differential hunting pressure on them (Rudd et al. 1983). Like the Jackson Hole herd, migratory elk that summered in protected areas of Yellowstone wintered with resident elk in the unprotected range just east of the park. Hunting seasons occurred before normal elk migrations from the park, placing heavy hunting pressure on the resident elk. The proportion of migratory elk from refuge areas had increased proportionally to the resident elk from no apparent behavior shifts due to hunting. Boyce (1991) noted that migration routes of elk wintering on the National Elk Preserve in Jackson Hole have changed dramatically between 1950-1954 and 1980-1984. The primary migration route shifted from being predominately on open hunting national forest lands to routes through Grand Teton National Park, where hunting is more restricted. In the 1950-1954 period, 22.1 % of the migrating elk were estimated to have moved through Grand Teton National Park, growing to 51.4% by 1980-1984. Boyce hypothesized that this may be to differential removal of the migrating animals from the population by hunting and not due to individuals shifting to alternative routes. However, the mechanism of these movement shifts in response to . hunting has not been identified or tested. . Elk responses to hunting pressure may depend on density of hunters. Wright (1983) examined elk response to different hunting seasons: archery/muzzleloading, rifle special elk. rifle special deer, and deer and elk rifle. The greatest density of elk hunters was during the rifle special elk, with distances traveled by elk the greatest during that season. Elk traveled distances three to four times greater during this season than during archery and muzzleloading season. Wright (1983) concluded that radio-collared cow elk were not adversely affected by archery hunters or muzzleloaders. However, it is possible that the lack of effect by archery hunters and muzzleloaders could be due to their low densities (0.13 hunters/knr' ) compared to the higher densities of rifle hunters (1.42 rifle hunters/knr'). In a previous study on the proposed study area in 1985, Consolidation Coal Company had 23 elk collared to evaluate ·the perceived movement of elk onto their protected land during early-season hunting. In that study, 87% of the elk were still found on National Forest lands mid-way through archery and muzzleloading seasons (Camp, Dresser and McKee Inc. 1986). However, due to a change in hunting regulations in 1985, only 37% as many archery hunters were afield during the study as the previous year (Gray et al. 1994) perhaps explaining the lack of movement that year. Also, Zahn (1974), Lemke (1975), and Hershey and Legee (1982) noticed an increase in movements during the first 1012 days of hunting season which decreased to normal during the remaining of the hunting season. These authors noted that hunting pressure was heavy during that time and relatively light for the remainder of the season. These studies suggest that there is perhaps a critical density of hunters that must be reached before elk begin to move in response to the hunting disturbance. An interesting observation from the Zahn (1974), Lemke (1975) and Hershey and Legee (1982) studies is that elk response to hunter disturbance was short lived. All studies noted that elk movements returned to normal after the initial 10-12 days of opening day of hunting, and all studies noted that the initial 10-12 days was when hunting activity was the greatest. Elk not only discontinued their erratic and increased movements but returned to their pre-season activity areas, indicating that hunting activity was a temporary disturbance. Elk movements during hunting may depend on the amount, location and quality of hiding cover, as well as the topography of the area. Elk either greatly increased their daily movements (Altmann 1956, Martinka 1969) or were found in thick vegetation inaccessible to most hunters (Lemke 1975, Irwin and Peek �56 1979, Morgantini and Hudson 1979). Also, preliminary data from George Bear's study (unpubl. data) on the elk on the proposed study area found that elk located in rugged and relatively inaccessible areas (n = 9) moved short distances (mean = 2.5 Ian) to dark timber and rough terrain when early-season hunting opened (Gray et al. 1994). In contrast, elk located in accessible areas (n = 11) moved an average of 13 km to private land. There were no areas of dense timber or rough terrain for elk on the accessible areas; therefore they needed to move to private land for a refuge. Areas of rough topography and dense timber may serve as refuge; if it is available, elk may not move far even in the face of high hunting pressure. Conclusions Elk are extremely plastic in response to human activity, habituating to people in areas where there is no hunting, and actively avoiding hunters in areas where they are hunted. Elk response to human activities appears to depend on the amount of contact and the danger of contact with humans. Avoidance of roads changes with the amount of use and the danger level. The presence of dense cover for hiding seems to attenuate movement responses to human disturbance. Finally, elk responses to human disturbances do not seem to persist after the disturbance is removed. Whether hunting is causing learned responses of avoidance and refuge seeking, or whether populations with certain behaviors are differentially removed are two theories of elk responses to hunting. Most hunter induced movements appear to occur before migration. As with other human activities, the level of activity, hunter density, and the amount of cover in the vicinity seem to affect elk responses to hunters. The literature indicates that elk responses to disturbance is short-lived. All of the studies of elk responses to hunting have been observational and it remains untested whether there is a causal relationship between hunting and elk inovements. This study will examine this issue with an experiment designed to determine if elk move from non-refuge to refuge areas in response to early-season hunting. OBJECfIVES The objective of this study is to evaluate the impacts of early-season hunters on the movement and distribution of elk in the White River area with the null hypothesis that elk movement is not affected by earlyseason archery hunting. The study will focus around a manipulative experiment to determine if the presence of the hunters is contributing to the movement of elk off areas easily accessible to hunting (public land I nonrefuge areas), to areas not readily accessible to hunting (private land or wilderness I refuge areas), but will also examine alternative hypotheses. Basic study objectives are: l. 2. 3. Test the hypothesis that early-season hunting disturbance of elk does not result in movement of elk from non-refuge to refuge areas. Evaluate hypotheses about other possible causes of elk movement such as livestock grazing, woodcutting, recreationalists, or weather. Compare the movements of this heavily hunted elk population with movements of elk from concurrent studies in other parts of Colorado. Early-season archery hunters are the focus of the study because they are the first to begin hunting and they are alleged to be causing the premature movement of elk off summer ranges. Although later-season rifle hunters may also cause movement of elk to refuge areas or keep elk in refuge areas, timing of the rifle season movements are more acceptable to resource managers and private landholders. To eliminate confounding factors from rifle hunting pressure, the manipulations will be limited to the early-season hunting. �57 STUDY AREA The study area consists of the GMUs 12,23,24, and 33, a subset ofDAU-E6, located in and adjacent to the White River and Routt National Forests (Figure 2). The area is approximately 4,540 knr' and is bounded on the north by the William's Fork of the Yampa River, on the west by state highway 13 (west side of the Grand Hogback), on the south by 170 between Rifle and Canyon Creek (12 Ian east of New Castle), and on the east by Canyon Creek in the south, through the eastern side of the Flattops, joining up with the East Fork of the William's Fork in the north. Hamilton t N I . LEGEND [II Flat Tops Wilderness Area D GMU Boundary _ State Wildlife Area D Study Area Boundary II White River National Forest Km r o 20 Figure 2. White River Study Area. White River 40 • Towns �58 Topography, climate, and vegetation vary widely throughout the study area. Elevation ranges from 1,629 m along the Colorado River to 3,700 m on the Flattops. Higher elevations have severe winters with heavy snowfall, while lower elevations have comparatively mild winters. Mean annual precipitation at 3,000 m in the Routt National Forest is about 100 em, while at Rifle (1,629 m) and Craig (1,856 m) mean annual precipitation is about 30 cm. Vegetation types range from the montane/subalpine zone in the higher elevations (>2,600 m), to the transitional zone in the middle elevations, to the Great Basin zone at the lower elevations «1,980 m) in the southern and northern parts of the study area. The area is described in detail by Gray et al. (1994). Higher elevations of the Flattops, Sleepy Cat Ridge and the White River/Colorado River divide areas provide good summer and fall forage for elk. Lower elevations are typically used as winter range by elk. Hunting pressure is relatively light on the Flattops, due to limited road access, and relatively heavy on the Sleepy Cat Ridge and the White/Colorado River divide areas due to abundant road access. The study area is 34% private land and 66% public land, with most public land being USFS (54%). Unit 24 is mostly (92%) public land, while the other three units average 57% public land. Private land is comprised mainly of ranches and coal mines. Much of the public land is grazed in the summer by sheep and cattle. Hunting for large and small game is economically important to local residents in the study area (Gray et al. 1994). Guides, outfitters, local service sectors, and private landowners (who sell trespass permits) make a large proportion of their annual income during the elk and deer hunting seasons, with most hunting revenues made during rifle hunting season. PILOT STUDY George Bear and Jeff Madison from the Colorado Division of Wildlife (CDOW) trapped and radiocollared 20 elk cows in the proposed study area in 1992. In 1995, an additional 11 cows were radio-collared in the study area, and the number of early-season hunter permits was reduced by 65%. From 1992 to 1995 the radio-collared elk were intensively monitored during August and September, from approximately one month before the opening day of early-season hunting to one month after opening day. There were 20, 16, 14, and 13 elk tracked in 1992, 1993, 1994, and 1995 respectively. A minimum of 18 locations was collected for each animal used in the analysis. Each location was classified as either being in a "refuge" or "non-refuge" area. Refuge areas were defined as having limited or no vehicle access, typically privately owned land or wilderness areas. Non-refuge areas were accessible by motor vehicles, typically USFS or BLMland. For each location of a telemetered cow, a zero was assigned to non-refuge locations, and a one was assigned to all refuge locations. All elk located on refuge areas remained on refuge areas throughout hunting season, while elk located in non-refuge areas moved to refuge areas with the advent of hunting season (Table I). . From these data, a logistic regression was done for each animal to estimate the date of its movement from non-refuge to refuge area (Appendix A). The date of movement was defined as the date at which the logistic curve crossed 0.5 on the y-axis, indicating an equal probability of being on a non-refuge or refuge area (Figure 3). From this analysis, a mean date of movement for all animals was computed. A two-tailed t-test was computed to determine if the mean date of movement was different from the opening date of early-season hunting for each year (Table 2). The analysis was not applied to the refuge elk since none of them moved to a non-refuge area during the study period. �59 Table 1. Locations ofradio-collared elk before and after opening day of early-season hunting, White River elk herd, 1992 - 1994. Movement pre-opening YEAR Combination.: -> p oat-op en in g 1992 1993 1994 1995 refuge -> refuge 10 10 6 7 refuge -> non-refuge 0 0 1 0 non-refug·e -> refuge 10 6 5 7 non-refuge -> non-refuge 0 0 2 2 20 16 14 16 sam pie size 1.5 (A) Refuge 1 ( & Estimated date of movement at 0.5 probabi&tyof being on.refugE 0 •• IIOlH'Bfuge 1- refuge -FItted logistic cwve 0.5 NOIH'efuge ) 0 712M13 eI5I93 8115/93 8125.'93 • estimated date of .movement er2!SI93 . 11(4193 11t14193 . Q'24193 (B) 1.5T""""------'--------:--------, Refuge 1 No movement & 0 = non-refuge. 1 = refuge -Fitted logistic curve 0.5 Non-refugeo 7f2!J1f1J7~ 8/993 8119/93 8I'2M13 918193 9118/93 9I2BI93 1G'8/93 Figure 3. Example of a logistic regression curve to estimate the date of movement for an elk moving from a non-refuge area to a refuge area (A), and elk staying in a refuge area (B). �60 Table 2. Mean dates of movement of elk from non-refuge to refuge areas, opening date of early-season hunting, and p-values from a t-test for differences for the White River elk herd, 1992 - 1994. 1992 1993 Mean date of movement 27 August, n= 10 28 August, n=6 8 September n=7 1 September n=6 95% Confidence Interval 24August31 August 23 August2 September 27 August20 Sept. 21 August13 Sept Opening date of early-season hunting (da) 29 August 28 August 27 August 26 August Ho: d m = opening date p=0.28 P = l.00 p=O.06 p=O.25 - 1994 1995 These data were also used to evaluate the proportion of elk that began on a non-refuge area and moved to a refuge area, before and after opening of early-season hunting (Figure 4). Because only the elk in non-refuge areas moved, the proportion of elk beginning in refuge areas remained constant (upper line in Figure 4) in the analysis. These data were appropriate for a Before-After-Control-Impact-Paired-Sampling (BACIPS) analysis with a t-test for the difference in proportions (Stewert-Oaten and Murdoch 1986, Osenberg et al. 1994). Instead of comparing the mean difference in a response, such as proportion moved from non-refuge to refuge areas between a controland impact site, the BACIPS approach compares the mean difference (control impact) in the before and after periods, where samples are temporally paired, thereby controlling for naturally occurring spatial and temporal variation (Osenberg et al. 1992, 1994). The average of the proportional differences Before and After the opening of early-season hunting was tested with the null hypothesis; Ho: db = da and Ha: db < da. The null hypothesis was rejected for 1992 (p < 0.0004),1993 (p < 0.0002), and 1995 (p = 0.012), which supports the alternative hypothesis. The difference before was less than the difference after, that is, the proportion of elk that moved to refuge areas increased for the non-refuge group after the opening of early-season hunting. This suggests that early-season hunting opening had an influence on elk locations, specifically, elk moved to refuge areas after the opening of early-season hunting. Although mean date of movement, and difference in proportion of elk found on refuge or non-refuge areas before and after the opening of early-season hunting indicates that elk movement correlates with the opening of early-season hunting, several alternative hypotheses, such as woodcutting activity or weather, could also explain the movements. . There are three main criteria required to strongly infer a cause-and-effect relationship between two factors: 1) there is a correlation between factor A and factor B, 2) the presence of factor A by itself causes factor B, and 3) factor B does not occur when factor A is removed. Criteria one has been satisfied by the pilot study. Criteria two, everything else being the same, a change in early-season hunting causes a change in elk movement, and criteria 3, there is no elk movement when there is no early-season hunting, can only be tested by a manipulative experiment. An effect consistently seen in a replication of a well-designed experiment can only reasonably be explained as being caused by the manipulation of the experimental variables (Manly 1992). With an observational study the same consistency of results may occur because all of the data are affected in the same way by some unknown and unmeasured variable (Manly 1992). Thus, a manipulative experiment, that isolates the effect of early-season hunting, and controls for, or blocks for, other potentially confounding variables (weather, woodcutting activity etc.) is required to infer that early-season hunting is causing elk to move from refuge to non-refuge areas in the White River area. �61 1992 i::hm~ "! differences before, db 0.8 differences after, d. Cl ~ ,:,e. iii '0 0.4 c ., o 0.2 V 0.0 -17 -12 Days before opening day 1993 -Begin in refuge V a. £. --Begin in non-refuge II V._ ~ 0.6 o -4 o 7 Archery Season Opening Date 12 18 22 28 Days after opening day ~Tff --~ i::: "& 0.8 _ •• noes before, d, ~~"", renoes after, d. •....• ~ § 0.6 I~ ~ ----eegin in nonrefuge / ,:,e. iii '0 0.4 / g ., V -,,- •..... Begin in refuge £0.2 / 0.0 ._ •.....• H-J.-+-+--+-+--JH-+-+-+--+-f--iI---f-+-+4 -26 Days before opening day -16 -9 o 9 Archery Season Opening Date 18 26 Days after opening day Figure 4. Proportion of elk on refuge areas for elk in non-refuge and refuge areas before opening of earlyseason hunting in 1992 and 1993. METHODS There are several alternative methods that could answer the question of whether the elk movement patterns in the White River area in late August are related to the presence of hunters. Study designs considered in the development of this plan are presented in Appendix B. An optimal study design, one with no limitations on money, resources, or time is presented for a comparison with this study plan and alternatives in Appendix C. �62 Although elk may respond in many ways to disturbance, effects of early-season hunter-induced movement will be tested by four response variables: mean date of movement, proportion of elk that move from one classification to the other, mean elevation changes, and mean distance moved between successive locations. The primary hypothesis tested by this study is: Ho: Elk movement from non-refuge to refuge areas is not different between hunted and non-hunted groups. Ha: Elk movement from non-refuge to refuge areas is greater for hunted groups. To build a strong case inferring a causal relationship between elk movement and the early-season hunting pressure we need to: 1) devise and test alternative hypotheses, and nearly as possible, exclude one or more of the hypotheses (platt 1964). Elk have been shown to move in response to recreational activity, logging activity, and road activity, so other activities that occur in the area should be considered. The following alternative hypothesis could also explain elk non-migratory movements in the White River area: Hal: Ha2: Ha3: Ha4: HaS: Elk are moving in response to livestock grazing activity/forage quality Elk are moving in response to presence or activity of woodcutters Elk are moving in response to recreationalists Elk are moving in response to weather/forage quality Elk have a learned, for an unknown reason, to move at this time of year. To assess the distribution of resource activity on the study area, data from the forest service on livestock grazing allotments, woodcutting permits, and recreational use will be evaluated. Additional data will be collected to evaluate livestock effects on elk movement; coordinates of livestock in the vicinity of radio-collared elk will be recorded every time elk are located. Distances between elk and livestock will be tested for elk avoidance of livestock. Hypotheses generation and testing of these factors outside of the crossover design framework will be observational in nature. If the distribution of these activities remains uniform from year to year, then the crossover designs presented in this proposal will block out for these effects and only hunting will vary. What makes the crossover designs good for isolating for hunter effects . alone is that each hunting treatment is used on each area. As a result, the effects of resource use averages out for each treatment. For example, say there were 10 bands of sheep grazing on one half of the study area, and only 5 bands grazing on the other half. As long as this was consistent from year to year each treatment would experience each level of grazing, averaging them out when comparing between treatment (Table 3). Table 3. Example of crossover design blocking out effects of sheep grazing activity with a hunting and nohunting treatment. Year I Year 2 North half of study area no-hunting treatment 5 bands of sheep South half of study area hunting treatment 10 bands of sheep hunting treatment 5 bands of sheep no-hunting treatment 10 bands of sheep Any persistent learned behavior will cause a bias toward acceptance of the null hypothesis, that elk movement is not different between hunted and non-hunted groups. For example, if elk are hunted one year, and the next year still move as a learned behavior even when they are not hunted, there will be less difference �63 between the movement patterns of hunted and unhunted groups the second year. Thus, the learned behavior could confound hunting induced movements. Because the literature indicates that elk responses to disturbance are short lived (Zahn 1974, Lemke 1975, Ward 1976, Hershey and Legee 1982, Edge and Marcum 1985, Cassirer et al. 1992), the study is designed on the assumption that there will be little persistent behavior from year to year. Study Design Open hunting season one week earlier on one area (chosen randomly) and two weeks later on the remaining area. Reverse treatments for year two. Blocking the confounding effects of space (area effects) and time (year effects), and relative ease of implementation were key factors in selection of this design. The study area to the north or south of the White River will recieve treatment one, opening hunting season one week earlier than the typical opening date of the last weekend in August. Treatment two would consist of opening hunting season two weeks later than opening date on the remaining half of the study area. Number of hunting permits issued is based on the mean number calculated from 1991- 1994 (2,545); 1,300 permits will be issiued on the north half of the study area and 1,245 will be issued on the south half of the study area. Hunters would choose and hunt on the study area GMUs to the north or south of the White River. Hunters choosing the early opening area would have an extra week of hunting. Hunters choosing the later opening season area would loose one week of hunting because the season will be extended one week later than usual, but they will have more time during the elk rut period. The study area will be split roughly east-west by the White River and North Fork of the White River into two halves for application of early or late opening treatments. GMU 12 and part of GMUs 23 and 24 will comprise the north half, while GMU 33 and part of GMUs 23 and 24 will comprise the south half. The primary response variables will be mean date of movement and proportion of elk that change classification (refuge -> non-refuge or non-refuge-e-refuge). Elk movement responses of elevation changes and distance between successive locations will also be analyzed to examine effects of hunting activity, as well as the effects oflivestock grazing, woodcutting, and recreational activity. This is a geographically nested design, with two levels of analysis. One level of analysis is within half, where samples (collared elk) will be randomly selected from refuge or non-refuge areas. Refuge or nonrefuge areas are the experimental unit, elk are the sampling unit, and hunting is the treatment (Figure 5). This design is not a true experiment since hunting (treatment) is not randomly allocated to experimental unit (refuge/non-refuge area). If hunting has no effect on elk movements, then date of movement to refuge areas should not necessarily be the same as opening date, date of movement should not differ between non-refuge or refuge areas, and proportion of elk on refuge and non-refuge areas should not change with the opening of hunting. The hypothesis tested would be: Ho: date of movement = opening date Ha: date of movement « opening date Ho: date of movement of elk on non-refuge areas Ha: date of movement of elk on non-refuge areas = date of movement of elk on refuge areas of elk on refuge areas * date of movement Ho: proportional difference of elk on non-refuge and refuge areas difference of elk on non-refuge and refuge areas after hunting Ha: proportional difference of elk on non-refuge and refuge areas difference of elk on non-refuge and refuge areas after hunting before hunting opens = proportional opens before hunting opens '" proportional opens Differences in time of movement for the fist two hypotheses will be tested with a one sample t-test, and proportion of elk moving will be tested as was the 1992-1993 pilot data. There would be two spatial replicates and two temporal replicates of each test. \ �64 Figure 5. Study design layout showing nested levels of experimental units. On the next level, this is a two-period crossover design (Manly 1992, Ott 1993) which is a essentially a 2 x 2 Latin squares design (Table 4). In this design, the experimental unit is north or south half of the study area, the sampling unit is individual elk, and the treatment is early or late opening of early-season hunting (Figure 5). Here, the responses between the differently treated halves would be compared. A key issue in this wider analysis is that only elk on non-refuge areas will be compared between halves in statistical tests since response in movement of hunted elk is the primary concern of this elk study. This is not to say that elk on refuge areas are not important - they serve as a crucial behavioral control in the within-half analysis. Ifhunting has no effect on elk movements, then there should be no difference between the date of movement to refuge areas or the proportion of elk on non-refuge areas for either treatment. The null hypothesis tested at this level would be: Ho: mean date of elk movement on early hunting GMU = mean date of elk movement on late hunting GMU Ha: mean date of elk movement on early hunting GMU GMU '# mean date of elk movement on late hunting On Early and/or Late Opening Date Ho: proportion of elk on non-refuge areas in early hunted GMUs in late hunted GMUs Ha: proportion of elk on non-refuge areas before hunting opens after hunting opens. '# = proportion of elk on non-refuge areas proportion of elk on non-refuge areas �65 Table 4. Layout of Latin square crossover design with corresponding model for alternative one. ce North half of study area South half of ~ area Collared Elk n n 1996 Early opening Late 1997 Late opening Early This analysis is based on an assumption of minimal interaction between the blocked sources of variation (GMU and year). The corresponding analysis of variance for the model is: where: 4. = ~(kr fixed effects for kth sequence random effects for individual animal a, = fixed effects due to treatment /3.i = fixed effects due to period (year) A power analysis to determine the number of cows needed from each half of the study area was done for: 1. date of movement compared to opening date within half 2. proportion of elk that move from non-refuge to refuge between halves on each opening date 3. date of movement between halves Power of at least 90% was specified for strong confidence that elk movement was not attributable to early-season hunting. Variance was estimated from the pilot study data of 1992 and 1993 as 4 days for mean date of movement tests. Previous proportions of elk on refuge and non-refuge areas before and after hunting season from pilot data were used in the estimation of variance and effect sizes for change in proportion tests. Because the hunted elk, or elk on non-refuge areas, are the key issue in this study, emphasis in sample size calculations was given to the comparisons between halves. Sample sizes were calculated (Chapman 1995) at different effect sizes for all three variables (Table 5). Between half analysis showed response variable 2 to require the largest sample size (Table 5). Based on an expected effect size of ~0.5 (i.e., at least 50% difference in proportion of elk on non-refuge area between treatments, at an opening date) a sample size of22 cows per half are required for non-refuge areas. For refuge areas, the behavioral control, collaring of 16 cows per half would allow for detecting a 9 day difference in movement date. This also allows for testing the difference of proportion of elk on refuge and non-refuge areas for likely scenarios (slightly more extreme differences), although this leaves slightly less power to detect more subtle differences. Because some cows may move from area to area, or move from non-refuge to refuge before the study period, some extra cows should be collared. If two extra cows are collared on nonrefuge areas for contingency during the study, this totals to 40 cows per half (22 non-refuge + 16 refuge + 2 contingency). Therefore, 80 total collars are required for the study. �66 Table 5. Sample size (number of collared elk) per half for treatment groups based on estimates of effect size and 90% power for primary response variables. Primary response variable 1- Nwnber of days between date of movement and opening date within half FffectSize Nun:Der of days between rreen date of movement date and Sample Size 16 10 7 10 14 8 Primary response variable 2 - difference in proportion of elk on non-refuge areas before and after an opening date (early or late) between early and late hunting treatments. Also can be used to evaluate proportion of elk on refuge and nonrefuge (within halves) areas before and after an opening date (early or late). Proportion of elk on non-refuge areas for early hunting treatment area Proportim of elk on non-refuge areas fur late hunting treatment area 0.8 0.7 0.9 11 15 8 16 22 11 22 36 15 0.1 0.2 0.3 Primary response variable 3 - date of movement between non-refuge and refuge. Effect Size Nun:Der of days between rreen date of movement for early and late hooting treatments Sarrple Size 7 10 14 22 12 8 Testing of alternative hypotheses would be straightforward under this alternative. The forest service data of livestock grazing allotments, woodcutting permits, and recreational use will be evaluated for each half of the study area. If these activities are relatively constant from year to year they will be averaged out. Weather will be assumed to be relatively constant by elevation band between the halves. Any factor that is common across the 2 areas, such as grazing, woodcutting, weather, forage phenology etc., that could be causing elk to move, would result in movement patterns closer than the minimum 7 day difference detectable in this design (Figure 6). In this case we would conclude that early season hunting was not the cause of elk movement, but would not be able to assign a cause. If there is a marked difference in activity on the two halves between year one and year two, then there a slightly different analysis would be performed. Only elk at distance greater than 1 km from a disturbance would be used in the analysis. A distance of 1 km was chosen since most studies found that elk farther away from disturbances ranging from hikers to logging did not respond to the disturbance (Ward 1976, Hershey and Legee 1982, Wright 1983, Edge and Marcum 1985, Cassirer et a1. 1992). The strength of a crossover design that it blocks for two sources of external variation (GMU and year), allowing for much stronger inference of cause and effect than the same design without a strict crossover (Ratti and Garton 1994), and is more efficient (smaller sample sizes needed) than a randomized design (Ott 1993). Further, the second two factors required to establish a causal relationship of factor A on B would be established with this design. That is, if elk move to refuge areas in the GMUs where hunting is moved forward one week, while remaining in non-refuge areas on the other GMUs, the presence of factor A (hunting) by itself can be inferred to be causing factor B (elk moment to refuge areas) (Figure 7). Conversely, if the elk on the late opening GMUs do not move to refuge areas before hunting begins, then it can be inferred that factor B (elk movement to refuge areas) did not occur in the absence of factor A (hunting). �67 Early Opening ~ ~10 ~ •• .!E '0 5 ::J Z Opening Day Late 0 pening Opening Day Figure 6. Hypothetical dates of elk movement to refuge areas if movement is caused by a factor other than hunting that is common across the 2 areas (woodcutting, weather, forage quality etc.). Early Opening '" c: '5' ~ 10 "'" ~ '0 .2l 5 E ::J Z o Opening Day Late Opening Opening Day Figure 7. Hypothetical dates of elk movement to refuge areas if movement is caused by hunting. �68 Early Opening Late Opening g .i5 E 10 .>< Qj '0 .2l E :::I Z 5 Opening Day Opening Day Figure 8. Hypothetical dates of elk movement to refuge areas if movement is a response to hunting and has a learned behavior component. Two weaknesses of this design apply to all alternatives. There may be a learned effect between years that the design cannot determine (Figure 8). For example, elk on non-refuge areas that were hunted early the first year may move early the second year, well before late hunting begins, as a random effect or a learned response. Similarly, elk hunted late the first year may move late the second year, after the opening date. In this situation the effect size is diminished; we will not be to detect effects due to early-season hunting on elk movements because of low power. However, because all studies indicate that disturbances have a short term effect on elk responses, this error is assumed to be minimal. The second weakness is the lack of spatial and temporal replication. There is only one spatial replicate, and it borders on a pseudoreplicate because it is not randomly chosen. The two years of the study are a fixed effect, restricting inferences to only the two years of this study. Thus, all inferences and results will only be applicable for the GMUs in the study during the time of the study. Capture and Collaring All capture and constraint procedures will be in accordance with Colorado Division of Wildlife and Colorado State University Animal Care and Use Protocals approved for this study. Helicopter netgunning will be the primary method of elk capture and will be contracted out to Helicopter Wildlife Management from Salt Lake City, Utah. Once a group of elk are located and an individual is "randomly" selected from the group, it is perused (typically < 30 sec) until the netgunner can fire a net over the elk. Once the elk becomes entangled in the net, it is blindfolded, hobbled and the net is removed to allow for installation of the telemetry collar (Phillips 1994). Collars will be in the 148 - 151 MHz transmitting range with a mortality sensor. Only cows will be collared for this study because bulls move with cows, so collaring either sex is representative of movement patterns. Also, bulls are lost at a high percentage to hunting, hence the loss of collared bulls during hunting season could reduce sample size below levels required for powerful hypotheses testing. Because only cows will be collared, the collars will not be of an expanding design. �69 Preferably, elk will be captured on their swnmer range in early to mid July. Of the seven elk radiocollared by netgunning in August of 1995, the average distance between their capture sites and location four days after capture was 6.4 Ian. The elk moved < 2.5 Ian between their first and second location after capture (3 days between first and second locations after capture). Therefore, capturing elk in early July will be early enough to avoid confounding capture movements with early-season hunter effects, but late enough so that the elk will be in their swnmer ranges to help ensure the desired distribution for the samples. Once the number of cows from each GMU is decided, capture sites will be randomly selected until an appropriate number of sites are located within the refuge or non-refuge areas. The pilot will go to the randomly selected location, then capture the first cow found from that location. In 1997, cows will be captured to replace losses from the sample due to mortality and transmitter failure. The cows will be replaced to balance the number of elk in the GMUs and classifications (refuge or non-refuge) to compensate for elk shifting location as well as lost elk. Radio-Telemetry Methods All elk locations will be collected using aerial telemetry. An H-antenna will be mounted to each strut of an airplane, and the signal received using a four-band scanning receiver. Location will be determined using a Global Positioning System (GPS). Telemetry system error is a combination of observer error and GPS error. The error will be tested by selecting a minimum of 40 locations from a variety of topographical and vegetative types represented on the study area (stratifIed random sample). These selected sites will be located to ±50 m using the GPS and a beacon will be left at the site to be located by air. To ensure the error contains the component of observer error as well as location error, this will be a blind test with the selected locations unknown to the observer. Elk locations will be collected at least two times a week at regular intervals with two to three days between collection. The main hypothesis that this study addresses is the relationship between elk movements and early-season hunting, especially around the opening day of early-season hunting. Therefore, the time frame of the 3-month study period of July 15th - October 15th; the 3-month interval allows valid comparison between treatments for the entire herd. Elk locations before and after this time interval will be collected and may yield information about elk responses to other variables (livestock grazing, woodcutting etc.), but confining the intensive data collection to around the opening of early-season hunting will eliminate confounding factors from the analysis and focus on the disturbance of interest. I will attempt to collect a minimum of 15 locations on each animal the month before and after the opening date of early-season hunting. Locations of livestock in the vicinity of radio-collared elk will also be collected during these flights. A Geographical Information System (GIS) map will be used to record all spatial information, such as refuge and non-refuge polygons, and elk locations. The map will be a 1:100,000 scale with 30 m elevation intervals, and will contain hydrography, property ownership, boundaries, and roads. Universal Transverse Mercator (UTM) 1983 will be the coordinate system used for all spatial analysis. Hunter Surveys To determine when and where hunters were afield during the study, hunters in the study area will be surveyed as soon as archery season closes. Hunters will be asked; 1) if they hunted opening day, and 2) what days they were in the study area. To improve the precision, sampling rate of hunter surveys will be increased for the study area. �70 Application of Results The results from this study will reject or establish a causal relationship between elk movements and early-season hunting activity. These results will provide managers of the White River elk herd with scientifically defensible and publicly credible information that can be used to design strategies for managing early-season elk distribution problems. All inferences and results will only be applicable for the GMUs in the study during the time of the study. In a meta-analysis framework, conclusions will have direct applicability to the White River area but will also contribute to the body of scientific knowledge on elk in the western United States. Outputs from this research will include written progress reports, scientific publications and spatial use maps. The primary scientific publication would be an article in the Journal of Wildlife Management on 'Elk movements in response to early-season hunting in the White River area'. Offshoots from this project may include an article in the Wildlife Society Bulletin on 'Video methods for viewing animal movement', or an article about large-scale field experimentation. LITERATURE OTED Adams, A. W. 1982. Migration. Pages 301-322 in J. W. Thomas and D.E. Toweill, eds., Elk of North America: ecology and management. Stackpole Books, Harrisburg, PA. Altmann, M. 1956. Patterns of herd behavior in free-ranging elk of Wyoming, Cervus canadensis nelsoni. Zoologica. 41:65-71. Boyce, M.S. 1991. Migratory behavior and management of elk (Cervus elaphus). App. Animal Beh. Sc. 29:239-250. Boyd, RJ. 1970. Elk of the White River Plateau, Colorado. Colorado Div. of Game, Fish and Parks. Tech. Publ. No. 25. 126pp. Camp, Dresser and McKee Inc. 1986. Meeker PRLA elk mitigation study: monitoring report Volume 2. Prepared for Consolidation Coal Company. Camp Dresser and McKee Inc., Denver, Colorado. Cassirer, E.F., DJ. Freddy, and E.D. Ables. 1992. Elk responses to disturbances by cross-country skiers in Yellowstone National Park. Wildl. Soc. Bull. 20:375-381. Chapman, P. 1995. SAS power calculation programs. Statistics Department, Colorado State University, Ft. Collins, CO. Craighead, JJ., G. Atwell, andRW. Wildl. Monogr. 29. 49pp. O'Gara. 1972. Elk migrations in and near Yellowstone National Park. Czech, B. 1991. Elk behavior in response to human disturbance at Mount St. Helens National Volcanic Monument. App. Animal Beh. Sc. 29:269-277. Doncaster, C. D. 1990. Non-parametric estimates of interaction from radio-tracking data. J. Theor. BioI. 143:431-443. Edge, W.D., and C.L. Marcum. 1985. Movements of elk in reaction to logging disturbances. J. Wildl. Manage. 49:926-930. Freddy, DJ. 1987. The White River elk herd: A perspective, 1960-85. Colorado Div. of Wild I. Tech. Publ. No. 37. 64pp~ Gray, J. P., G. Byrne, and J. Madison. 1994. White River elk data analysis unit plan: game management units: 11,211,12,13,131,231,23,24,25,26,33. Colorado Div. ofWildl. 45pp. �71 Gurevitch, L, L. L. Morrow, A. Wallace, and 1S. Walsh. 1992. A meta-analysis of competition in field experiments. Am. Nat. 140:539-572. Hershey, TJ., and T. A. Leege. 1982. Elk movements and habitat use on a managed forest in north-central Idaho. Idaho Dept. ofFish and Game. Wildt. Bull. No. 10. 23pp. Irwin, L. L., and J. M. Peek. 1979. Relationships between road closures and elk behavior in northern Idaho. Pages 199-204 in M. S. Boyce and L. D. Hayden-Wing eds., North American elk: ecology, behavior, and management. University of Wyoming, Laramie. Knight, R.R. 1970. The Sun River elk herd. Wildt. Monogr. 23. 66pp. Kuck, L., G. L. Hompland, and E.H. Merrill. 1985. Elk calf response to simulated disturbance in southeast Idaho. 1Wildl. Manage. 49:751-757. Lemke, T.O. 1975. Movement and seasonal ranges of the Burdette Creek elk herd, and an investigation of sport hunting. Montana Fish and Game Dept. Job Final Rept., Study No. 32.01, Job No. BG-3.15. I 27pp. Manly, B. F. J. 1992. The design and analysis of research studies. Cambridge University Press, New York. 353pp. Martinka, lC. 1969. Population ecology of summer resident elk in Jackson Hole, Wyoming. J. Wildl. Manage. 33:465-481. Minta, S. C. 1992. Tests of spatial and temporal interaction among animals. Ecol. App. 2:178-188. Morgantini, L. E., and R. 1Hudson. 1979. Human distribution and habitat selection by elk. Pages 132-139 in M. S. Boyce and L. D. Hayden-Wing eds., North American elk: ecology, behavior, and management. University of Wyoming, Laramie. Osenberg, C. W., S.l Holbrook, and R. 1Schmitt. 1992. Implications for the design of environmental assessment studies. Pages 75-89 in P. M. Grifman and S. E. Yoder eds., Perspectives on the Marine Environment. Proc. on the Marine Env. of Southern Calif., Los Angeles, CA. Osenberg, C.W., R. 1. Schmitt, S. 1. Holdbrook, K. E. Abu-Saba, and A. R Flegal. 1994. Detection of environmental impacts: natural variability, effect size, and power analysis. Ecol App. 4: 16-30. Ott, R L. 1993. An introduction to statistical methods and data analysis. Fourth ed. Wadsworth, Inc., Belmont, CA. 1051pp. Philips, G.E. 1994 Upper Eagle River Valley elk study: draft study plan. Dept. Fishery and Wildt. Bio. Colorado State Univ., Ft. Collins. 26pp. Platt, lR. 1964. Strong inference. SCience. 146: 347-353. Ratti.J. T., and E. O. Garton. 1994. Research and experimental design. Pages 1-23 in S. Hieb, ed., Research and management techniques for wildtife and habitats, Fifth ed. The Wildl. Soc., Bethesda, Maryland. Rost, G. R. 1975. Responses of deer and elk to roads. M.S. Thesis, Colorado State Univ., Ft. Collins. Rudd, W. r, A. L. Ward, and L. L. Irwin. 1983. Do split hunting seasons influence elk migrations from Yellowstone National Park? Wildt. Soc. Bull. 11:328-331. Schultz, R.D., and 1. A. Bailey. 1978. Responses of national park elk to human activity. J. Wildt. Manage. 42:91-100. �72 Stewart-Oaten, A. and W. W. Murdoch. 1986. Environmental impact assessment: "pseudoreplication" in time? Ecology. 67:929-940. Strohmeyer, D.C., and 1. M. Peek in press. Wapiti home range and movement patterns in a sagebrush desert. Northwest Sci. Ward, A. L. 1976. Elk behavior in relation to timber harvest operations and traffic on Medicine Bow Range in south-central Wyoming. Pages 32-43 in S. Hieb, ed., Proc. of the Elk-Logging-Roads Symposium. Univ. ofIdaho, Moscow. Weber, B. 1., M. L. Wolfe, G. C. White, and M. M. Rowland. 1984. Physiologic response of elk to differences in winter range quality. 1. Wildt. Manage. 48:248-253. Wright, K.L. 1983. Elk movements, habitat use, and the effects of hunting activity on elk behavior near Gunnison, Colorado. M.S. Thesis. Colorado State Univ., Ft. Collins. 20Opp. Zahn, H. M. 1974. Seasonal movements of the Burdette Creek elk herd. Montana Fish and Game Dept. Job Final Rept., Study No. 32.01, Job No. BG-3.13. 68pp. �73 Appendix A Logistic regression analysis for 1992 - 1995. 1992 --------------------------- YEAR-1992 Ie-1I8.010 ------------------~------Plot of GRilFUGE·DATE. Legend: A - lobs, B-2 ----------------.------------ YEAR-1992 Ie-1I8.070 --------------- obs , etc. Plot of GRilFUGE·DATE. Legend: A - lobs, B-2 _ obs, etc. R Limited Access + 1.1. e f Limited Access A A A A A .1. A A A 1.1. u g e S t a Motor Veh. + A A A A t u A A + s Motor Yah. ---+----------+---------..:+----------+----------+----------+-08/10/92 08/20/92 08/30/92 09/09/92 09/19/92 A. A A A 1.1.1. I ---+----------+----------+----------+----------+----------+-08/10/92 . 08/20/92 08/30/92 09/09/92 09/19/92 09/29/92 09/29/92 DATE --------------------------- YEAR-1992 Ie-1C8.020 --------------------------Plot of GREFUGB*DATB. Legend: A - 1 01>3, B.- 2 01>3, --------------------------- etc. DATE YEAR-1992 Ie-lC8.190 --------------------------- Plot of GREFUGB*DATB. Legend: 1.- 1 01>3, B-2 01>3, etc. R Limited Access + A A A A 1.1.1. A 1.1. 1.1. A A 1.1. 1.1. I I I I u g e I. I S t a t I I Motor Yah. •f Limited I + u " ---+----------+----------+--:--------+----------+----------+-08/10/92 08/20/92 08/30/92 09/09/92 09/19/92 09/29/92 Access + A 1.1. 1.1. A A 1.1. 1.1. I I I I I I I I I 1.1. A A 1.1.1. + Motor Veh. I ---+-------_..:._+----------+----------+----------+----------+-08/10/92 08/20/92 08/30/92 09/09/92 09/19/92 09/29/92 DATE DATB --------------------------- YEAR-1992 Ie-1C8.0S0 ------------------------_-Plot of GRilFUGB*DATB. Legend: A - 1 01>3, B-2 01>3, etc. I I LiJoitedAccess + A 1.1.1. A AIAAABA 1.1. I I I I I I I I I Motor Veh. A A + I ---+----------+---------+---------+----------+----------+-08/10/92 08/20/92 08/30/92 09/09/92 09/19/92 09/29/92 DATE --------------------------- YEAR-1992 Ie-1C8.060 ---------------------------Plot of GRilFUGg*DATE. Legend: A - lobs, B-2 obs, etc. I I Limited ACCe3,5 + ---------_--------------- YEAR-1992 Ie-lC8.310 --------------------------- Plot of GREFUGE*DATB. Legend: A-lobs, B-2 cbs, etc. R •f Limited Access + A 1.1. 1.1. A A 1.1. 1.1. I g I e I I S I t I I t I u I s Motor Veh; 1.1.1. A B A A + I ---+----------+----------+----------+----------+----------+-08/10/92 08/20/92 08/30/92 09/09/92 09/19/92 09/29/92 DATE ---------------------------- YEAR-1992 Ie-1C8.330 ---------------------------Plot of GRilFUGg·DATE. Legend: A - lobs, B-2 obs , etc. u •• R A 11 A 1.1.1. A 1.1. 1.1. A A 1.1. A A e f Limited Acce.5.5 A A A 1.1. 1.1.1.1.1.1.1.1.1. 1.1. u g e s a t Motor Veh. u oS ---+----------+----------+----------+----------+----------+-08/10/92 08/20/92 08/30/92 09/09/92 09/19/92 09/29/92 Motor Veh. + I ---+:..._--------+----------+----------+----------+----------+-08/10/92 08/20/92 08/30/92 09/09/92 09/19/92 '09/29/92 �74 ------------------- YKAR-1992ID-148.39~ ,Plot of GRBFUGStDATE.lAgend: A-I ----____________ cbs, B-2 cbs, --------------.,---- .te. YKAR-1992ID-148.600 Plot of GRBI'UGStDATB.Leg.nd: A-I _ cbs, B-2 obs, .te. R Liaited Access Motor Veh. + I A AA A AA A AAAAAAAA AA •f Liaited Access u I I I I I I I I 9 e + :J S t a t u Motor Veh. ---+----------+----------+----------+----------+----------+-08/10/92 08/20/92 08/30/92 09/09/92 09/19/92 + I I I I I I I I I + I AA A AA A Plot Liaited YKAR-1992IO-U8.tOO I I + I 08/10/92 08/20/92 08/30/92 ----------------------_----cbs, --------------------------- etc. Plot 09/09/92 YKAR-199210-148.750 09/19/92 09/29/92 ----------------------- of GREFUGEtDATB.Legend: A-lobs, B ,- 2 obs, _ etc. R A A A A AAA A AAAAAliAA AA •t Liaited Aceess u s Motor Veh. + g I I I I I I I + I ---+----------+----------+----------+----------+----------+-08/10/92 08/20/92 08/30/92 09/09/92 09/19/92, 09/29/92 + I I I I I I I I I u • s t a t A, AA A AAA ---------------------- YKAR-199210-148.450 AA AA A A AA 09/29/92 DATB ------------------------- of GRBFUGBtDATE.Legend: A-lobs, A I ---+----------+----------+----------+----------+---------+-08/10/92 08/20/92 08/30/92 09/09/92 ,09/19/92 DATE Plot AA ---+----------+----------+----------+----------+----------+-- 09/29/92 I Motor Veh. AA DATK of GRBFUGEtDATB.Legend: A - 1 oce , B-2 Access A' A A A A A A DATK ___________________________ A B-2 obs, --------------------- YKAR-199210-148.810 -,--------------------- , Plot 'of GREFUGEtDATE.Legend: A - 1 cbs, ate. B-2 obs, ate. R a Liaited Access + I I I A AA A AAA A AA AA A A AA AA Access • s I I I I + I ---+----------+----------+----------+--------+----------+-08/10/92 08/20/92 08/30/92 09/09/92 09/19/92 09/29/92 + AAA t a t u Plot YEAR-1992 10-148.460 of GREFUGKtDATE. ,Legend: A - lobs, A A A'A A A AA AA A AA A + I ---+----------+----------+----------+-:... •.. _-------+----------+-08/10/92 08/20/92 08/30/92 09/09/92 09/19/92 09/29/92 s Motor Yah. DATE --------------------------- A I I I I I I I I I g I I Motor Veh. t Liaited u DATB ---------------------------B-2 ~--------------------------- obs , etc. Plot YKAR-199210-149.011 of GREFUGEtDATE.Legend: A - lobs, ---------------------------B-2 obs , etc. R e Limited Acce.5.5 + A AAAAAAAA AA f Limited Access AAA + A AAAAAAAA AA u g e I s • t u Motor Veh. + A A A A A A A ---+----------+----------+----------+----------+----------+-08/10/92 08/20/92 08/30/92 09/0~/92 09/19/92 09/29/92· DATE !I Motor Veh. + A A A A ---+----------+----------+----------+----------+----------+-08/10/92 08/20/92 08/30/92 09/09/92 09/19/92 09/29/92 �75 __________________________ - YKAIl-1992II>-U 9.060 ---------------------------- -------------------------Plot Plot of GRBPUGK·IlIITK. Legend: A - 1 obs, 8 - 2 obs, etc. YKAIl-1993II>-UB.OI0 -------------- of GRBPUGK·IlIITIl. Legend: A - 1 obs, _ 8 - 2 obs, etc. R Liaited ,, ,, ,, ,, , , Acce,," + Motor Veh. A A A· A AI. AI. e f Limited Acce"" + AI. u 1.1.1. 08/20/92 , S , a , , s Motor Veh. 08/30/92 09/09/92 09/19/92 , + A AAMAA 07/26/93 08/15/93 09/04/93 09/24/93 10/14/93 illiTE YKAIl-1992I[)OU 9. 080 ---------------------------- Plot of GRBFUGE·OATIi. Legend: A - lobs, MAl. --+-------------+-------------+-------------+-------------+-- 09/29/92 illiTE --------------------------~ AMAMA , e ---+----------+----------+----------+----------+----------+-- 08/10/92 M , u AA A + AI. . , 9 B-2 ---------------------------Plot obs, etc. YEAR-199311>-148.020 of GRliFUGE·OATIi.Legend: A - lob", B-2 ob", etc. R Liaited ,, ,, ,, Acc."" + e A AA A 1.1.1. A 1.1.1.1.1.1.1.1. u s t a t , u , .s Motor + 08/10/92 08/20/92 + + .L 08/30/92 09/09/92 Veh• +__ 09/19/92 09/29/92. + I __+ 07/26/93 + 08/15/93 IlIITi! ------------------- YEAR-1992II>-U9.100 AM AM A ,, ,, , S + __ + A A A M A A M A A A A .M I e ,, , Motor Veh. ,, f Limited Aeee.s.s + AI. + + 09/01/93 09/21/93 +__ 10/14/93 IlIITIl ---------------------------- Plot of GREFUGII·IlIITIl.Legend: A - 1 db", B-2 ---------------------------Plot db", etc. YKAIl-199311>-148.060 ---------------------------- of GRliFUGIi·OATIi.Legend: A - 1 obs, B-2 ob:s, etc. R Liaited ,, ,, ,, ,, Acce"" + A AA A 1.1.1. A .1.1. AI. A A AI. AI. I + I Motor.veh. e· f Liaited Acce"" + u , g e , , S , t a t u I I " Motor Veh. + I 08/20/92 08/30/92 09/09/92 09/19/92 Plot Limited Access ,, + A A A A AAA A B-2 obs , 07/26/93 ---------------------------Plot .etc. AAAAAAAA AI. ,, , ,,, , Motor Veh. 08/15/93 09/01/93 09/2'/93 10/11/93 R e f u Li.m..ited Access YEAR-199311>-148.190 ---------------------------- of GRIiFUGE'OATIi.Legend: A - lobs, + A AAMAA MAAAA B-2 M obs , etc. "MAMA 9 e S a u + .'---+----------+----------+----------+----------+----------+-- 08/10/92 AMAMA IlIITIi Y&AR-199211>-149.120 --- ..------------------------ of GRliFUGIi·OATIi •. Legend: A - lob", M --+-------------+-------------+-------------+-------------+-09/29/92 OATIi --------------------------- MAAAA , ---+----------+----------+----------+----------+----------+-08/10/92 A AAMAA 08/20/92. 08/30/92 1993· 09/09/92 09/19/92 09/29/92 s Motor Veh. --+-------------+-------------+-------------+--------~----+-- 07/26/93 08/15/93 09/04/93 09/2'/93 10/14/93 �76 --------------------------Plot YBAR-1993100U 8.810 ---------------------------- of GRBPUGB·DATB.Legend: A-lobs, B-2 obs, ---------------------------- etc. Plot Lilli ted Access + MAAAA M AMAMA 1 1 1 1 1 1 1 1 1 A AAMAA Motor Yah. + 1 --+-------------+-------:------+-------------+---------.:..---+.:..07/26/93 08/15/93 09/04/93 09/24/93 10/14/93 R e f Lillited of GRBPUGE·DATE.Legend: A-I u Access + 1 9 1 e 1 1 5 ·1 t 1 a t u 1 1 S Motor YBAR-1994100U 8.050 A AA A Plot 01>3, B-2 A A 08/20/94 08/30/94 ---------------------------Plot •f LiAited Acces5 u 9 e 5 t a t u .5 Motor YEAR-199410-148.060 --~--------------------B-2 obs, Veh. ---------------------------- etc. Plot ---------------------- _ B - i obs , etc. YEAR-199110-148.190 ----------------------- of GRBPUGE·DATE.Legend: A - lobs, R e f Lilli ted Access + u 1 1 9 e 1 1 5 1 t 1 a 1 t 1 u I s Motor Veh. + A A A AA B-2 A A A 01>3, etc. A A A ---+----------+----------+----------+----------+----------+-08/10/94 08/20/91 08/30/94 09/09/94 09/19/94 09/29/94 DATE YBAR-199310-14 9.100 ---------------------------- of GREFUGS·DATE.Legend: A - lobs, 09/29/94 I· 1 A AA A A A + A A A A A A I 1 1 1 1 1 I I I + I ---+----------+----------+----------+----------+----------+-08/10/94 08/20/94 08/30/94 09/09/94 09/19/94 09/29/94 DATE Plot 09/19/94 DATE LiAit..:! Access + A AAMAA MAA1t.A M AMAMA I 1 1 1 1 1 I 1 I Motor ;Veh. + I --+------------+-------------+-------------+-------------+-07/26/93 08/15/93 ·09/04/93 09/24/93 10/14/93 --------------------------- 09/09/94 of GREPUGE·DATE.Legend: A-lobs, DATE - YBAR-199310-119.080 A ---+----------+----------+----------+----------+----------+-08/10/94 R of GRBPUGB·DATB.Legend: A - 1 obs, A Veh. 01>3, etc. LiAit..:! Access + 1 1 1 1 1 1 1 1 1 Motor V.h. A AAMAA MAA + I -+---------+-------------+-------------+-------------+-07/26/93 08/15/93 09/04/93 09/24/93 10/14/93 Plot A A oos , etc. DATE YBAR-199310-14 9.·0·60 ---------------------------- of GREPUGB·DATE.Legend: A-I 01>3, B-2 A A DATE --------------------------- _ B-2 obs , ---------------------------Plot etc. YEAR-1994 10-14 8.310 ---------------------------- of GIUiFUGE'DATS.Legend: A - lobs, B-2 obs , etc. R Lillited Access + I A AAMAA MAAAA M e f Limited AMAMA Acce~~ + A A A A A A A A A u 1 9 1 e 1 1 5 1 1 Motor Veh. a t 1 1 + 1 u ~ Hotor --+-------------+-------------+-------------+-------------+-07/26/93 08/15/93 09/01/93 1994 09/21/93 10/11/93 Veh. + AA A ---+----------+----------+----------+----------+----------+-·08/10/94 08/20/91 08/30/91 09/09/91 OATS 09/19/94 09/29/91 �77 ___________________________ YEAR-~99110.118.330 Plot -----_______________________ of GREFUGS'DATS.lAgend: A - 1 01>0, B-2 01>0, ---------------------------- YEAR-199110.118.600 ---------------------------- iI - 2 cbs, 1'lot of GREFUGS*DATS.lAgend: A - 1 cbs, etc. etc. R LiJaited Access + A AI. A A A A A A A A A •f Limited I u I s I e Acee!!!! I 5 I I I t a t I u I I I I I I I I I + I s Motor Veh. + I Motor Vah.. + A AI. A Plot A A A A A 08/20/94 08/30/91 09/09/91 09/19/91 ---+----------+----------+----------t----------+----OS/10/91 09/29/91 OS/20/91 OS/30/91 YEAR-1991Io.U8.390 of GREFUGE'DATE.Legend: A - t --.-------------------------- obs, .09/09/91 +__ 09/19/91 09/29/91 DATE DATl> --------------------------- A A I ---+---~------+----------+----------+-~--------+----------+-- OS/10/94 A B-2 obs, ---------------------------Plot etc. YEAR-1991Io.US.750 ---------------------------- of GREFUGE'DATE.Legend: A - l.obs, B-2 obs , etc. R Lillited Aceess e + f Limited A I I u g I e I I I I I I a t u s Motor + A AI. A A A Veh. + A A A A ---+----------+----------+----------+----------+----------+-OS/10/91 08/20/9. 08/30/9. 09/09/9. 09/19/9. 09/29/91 DATE Plot YEAR-1991Io.IU.IOO DATE -----------:----------------- of GREFUGS*DATE. Legend: A - 1 01>0, A I I -----,---------------------- A I I I I I I I I I 5 A A A A + A AI. A A A A ---+----------+----------+----------+----------+----------+-08/10/91 08/20/91 08/30/91 09/09/91 09/19/91 09/29/91 Motor Veh. Acee.ss B-2 obs, ---------------------------Plot etc. YEAR-1991Io.US.810 ---------------------------- of GREFUGE*DATE. Legend: A - lobs, B-2 cbs, etc. R Lilli ted Acc."s + A 1.1. A A A· A A A A A A I I I I u g I 5 t Access + u s Motor Veh. 08/20/91 08/30/9.. 09/09/91 09/19/91 + I A AI. A A OS/10/94 09/29/91 08/20/91 OS/30/91 .DATE· A A A 09/09/91 09/19/91 09/29/91 DATE YEAR-1991Io.U 8.450 ---------------------------- of GREFUGE*DATE. Legend: A - lobs, A --~+----------+----------+----------+----------+----------+-- ---+----------t----------+----------+----------+----------+-- Plot A t I --------------------------- A a + 08/10/91 A e I I I I Kbtor Veh. e f Limited B-2 obs, ---------------------------Plot etc. YEAR-199110.119.060 ----,------------------------. of GREFUGE·DAT.E_ Legend: A - lobs .• B-2 obs , etc_ R e Limited. Access + A A A A A Limi ted A Access A + u 9 e s a t u Motor Veh. + I A AA A .1. A .5 ---+----------+----------+----------+----------+----------+-OS/20/91 08/30/94 09/09/91 09/19/91 09/29/91 . OS/10/91 DATE ..... ' .. ' Hotor Veh. + A AA .1. A A A A A A A ---+----------+----------+----------+----------+----------+-- 08/10/91 08/20/91 OS/30/9' 09/09/91 OATS 09/19/94 09/29/91 �78 --.:.------------;..------Plot LilIited YBAIl-,199' II>-U 8. 330 -------------------------- of GRBFUGS·DATI. Legend: •••- 1 ees , B-2 Access Motor Yah. + .•. .•..•. oba, .•. .•. .•. .•. .•. .•. ---------------------:------ etc. Plot .•. .•. .•. u S + s Motor Yah. t a t u 08/20/9' 08/30/9' 09/09/91 09/19/91 Plot LilIited Access B-2 obs, A loA 08/10/94 ---------------------------- etc. A I I I A A·••• A A ••• A A A· A A R e f LUaited Access + u I I 9 e I I S I t I a I t I u I s Motor Veh .• + I ---+----------+----------+----------~---------+----------+-08/20/94 OS/30/94 09/09/9' 09/19/94 09/29/94 08/20/94 08/30/94 - 1 obs. A loA A Access + I I I A AA A A A B-2 obs. ---------------------------- A A A A A A A A A f u Limited YBAIl-1994·11>-148.810 S t a t u s Motor Veh. ---+----------+----------+----------+----------+----------+-09/09/91 09/19/91 I I I I I + I A A ••• A A Plot etc. A A A A A 08/10/91 08/20/94 08/30/94 09/09/91 09/19/91 09/29/91 DATE YEAR-1991 11>-148.450 ---------------------------- of GRBFUGE·DATE.Legend: A - lobs, A obs. ---+----------+----------+-----.;..----+----------+----------+-- 09/29/91 DATE --------------------------- ---------------------------- I I I e 08/30/91 A A B-2 .•. Access + I 9 08/20/94 A A Plot of GRBFUGE·DATE.Legend:. A-lobs. etc. I I I I I + I 08/10/91 ~-------- obs , etc. ---+----------+----------+----------+----------+----------+-08/20/94 08/30/91 09/09/91 09/19/91 09/29/91 I Motor Veh. B-2 A R ·e LilIited 09/29/91 DATI! YBAIl-199.111>-148.400 ---------------------------- of GRBFUGS·DATB.Legend:'" 09/19/94 YBAIl-1994 11>-118.750 ---_--------- DATE Plot 09/09/94 08/10/94 OSi10/94 ----.--------------------- .•. A Plot of GRBFUGS·DATE.Legend: A - lobs, A + I I .+ .•. .•. OATS I I I I Motor Veh. etc. ---+----------+----------+----------+----------+----------+-- 09/29/94 YBAIl-1994 11>-148.390 ---------------------------- of GRBFUGS·DATS.Legend: A - lobs. obs, .•. .•. .•. .•. + DATE --------------------------- B-2 I I I I I I I I I 9 e ---+--------~-+----------+----------+----------+----------+-- 08/10/91 _ of GRBFUGS·DATI. Legend: •••- 1 obs. R e cceas f Lilli ted ••• I I I I I I I I I YBAIl-199' I.I>-U8.600 ---------------------------- YEAR-1991 10-119.060 ---------------------------- Plot of GREFUGS·DATE.Legend: A - lobs, B •. 2 obs , etc. B-2 obs, etc. R Limited Acce!5!5 + A A A A A A e f u Limited Access + A 9 e S a t u Motor Yah. + A loA A A A ---+----------+----------+----------+----------+----------+-- 08/10/94 08/20/91 08/30/94 A AA s Motor veb • 09/09/94 DATE 09/19/91· 09/29/94 A A A A A A A A ---+----------+----------+----------+-------~--+----------+-- 08/10/94 08/20/94 08/30/94 09/09/91 DATE 09/19/91 09/29/91 �79 --Plot YBAR-199' 11>-119.080 -..;--------- of GRBFUGB·DATB.lAgend: A - 1 ob:s, B-2 LiJoit<od Acc."" + A AA A A A A A --------------------------- ob:s, etc.. A A Plot A A R e f LiJaited YKAR-1991;11>-118.210 ------------------ of GRBFUGB.DATB.Legend: A - 1 eea, B-2 _ obs , etc. I I· Access A u I I I 9 e I Hotor veh , I 5 I I I I t a + s Motor Veh. t u A A A A A A A I ---+----------+----------+----------+----------+----------+-- 08/10/94 08/20/94 08/30/94 09/09/94 09/19/94 ---+----------+----------+----------+----------+----------+-- 09/29/94 08/15/95 08/25/95 09/04195 Plot 09/24/95 10/04/95 OATS DATE ----------------.----------- 09/14/95 YKAR-199411>-119.100 --- ..• ----------------------- of GRBFUGS·DATS.Legend: A - 1 ebs , B-2 ---------------------------Plot ebs , etc. YKAR-199510-148.230 of GRBFUGS·DATE.Legend: A - 1 obs , B-2 obs , etc. R LiJoited Aceess Motor Veh. + I I I I A AA A A A A A A A A A e f Limited Access + A A A A A A u 9 e I I I I I 5 + I s Motor Veh. t a t u + I A A· ---+----------+----------+--------+----------+----------+-- ---+------~-+----------+----------+----------+----------+-08/20/94 08/30/94 09/09/94 09/19/91 09/29/94 08/15/95 08/10/94 08/25/95 09/04/95 09/11/95 09/21/91; 10/04/91; DATS -------------------------Plot ---------------------------- YKAR-199411>-149.190 --------------------------- of GRBFUGS·DATE.Legend: A - lobs, B-2. Plot obs , etc. OATS YKAR-1991;10-14 8.270 ---------------------------- of GRBFUGS·DATS.Legend: A - lobs, B-2 obs , etc. R LiJoited Acc.ss Motor Yah. + I I I I I I I I I + I •f LiJoited A u 9 • S t a t Access + I I I I I A A A I I I + u Veh. -+---------------------------+-----------------------~---+09/21/91 DATS A A I .s Motor 01/01/60 A A ---+----------+----------+----------+----------+----------+-08/15/95 06/11/29 08/25/9~ 09/04/91; 09/14/95 09/21/95 10/01/95 DATE 1995 --------------------------Plot YKAR-199511>-148.190 ---------------------------- of GRBFUGS·DATB.Legend: A - lobs, B-2 -------------------------~-Plot obs , etc. YKAR-1995·10-148.290 .---------------------------- of GRBFUGS·DATB.Legend: A - 1 obs , B-2 obs , etc. R Limited Access + A A A A e f u Limited Access + A A A A A 9 e 5 a .t u Kotor Veh. + A A A A A AA A S Motor Veh. A --+-------------+-------------+-------------+--~----------+-- 07/16/95 08/05/95 08/25/95 DATB 09/11/95 10/04/95 A A A ---+----------+----------+----------+----------+----------+-- 08/15/95 08/25/95 09/04/95 09/11/95 DATS 09/24/95. 10/04/95 �80 ------------------------. Plot Liaited YEAR-J995 100U 8.312 ---------------------------- of GRllFUGE·DATE.Legend: A - lobs, ,, ,, .',, ,, , B-2 ®s, etc. Plot of GRllFUGE·DATE.Legend: A-lobs, R a f u Access + Motor Veh. + Limited Accass + AAA AAA 9 AA Plot Liaited A A A A A A s Hotor Veh. A 08/25/95 09/01/95 09/11/95 09/21/95 - - +- - - - -- - ---- -- + - --07/16/95 08/05/95 10/01/95 --- YEAR-1995IO-U8.330 ---------------------------B-2 obs , et c , A A A A A f u Liaited S t a t u A :J ,, ,, , ,,, , , Legend: A - lobs, Access + g a A YEAR-199510-118.450 ---------------------------- Plot of. GREFUGS·OATE. R a A Hotor Veh. + A A A A 8 - 2 obs, etc. A A A A A A ---+----------+----------+----------+----------+----------+-- ---+----------+----------+----------+----------+----------+-08/25/95 09/01/95 09/11/95 09/24/95 10/01/95 08/05/95 08/15/95 08/25/95 DATE 09/0l/95 09/14/95 09/2l/95 DATE ---------------------------- YEAR-199510-118.390 of GRllFUGE·DATE.Legend: A-lobs, -- - - -+---- --- - - - - -- +-09/14195 10/04/95 - -- --- + -- - - - --08/25/95 DATE 08/15/95 Plot A u A ,, ,, ,, , ------------------------ A a t ,, , AAA A e Access + + etc. S t of GRllFUGE·DATE.Legend: A-lobs, Motor Veh. ®S, , DATE ----------_---------------- B-2 , ---+----------+----------+----------+----------+----------+-- 08/15/95 YEAR-1995IO-U8.130 B·- 2 ®S, etc. YEAR-199510-148.460 Plot of GRllFUGB·DATE.Legend: A-I ®S, B-2 ®S, etc. R Liaited ,, Access + A.A A A A A AA AAA A A ,, ,, ,, Motor ..Veh. e f Limited Access + u , g e , A A A A A A A S t a , , t u + s Motor Yah. , + A ---+----------+----------+----------t.,;..---------+----------+-- --+-------------+-------------+-------------+-------------+-08/05/95 08/25/95 09/11/95 10/01/95 08/15/95 07/16/95 08/25/95 09/01/95 09/14/95 09/21195 10/01l95 DATE -------------------------Plot ------------------.----'------ YEAR-199510-14 8.100 ---------------------------- of GRllFUGE·DATE. ·Legend: A-lobs, B-2 obs, YBl\R-1995 IO-H8. 810 ---'------------------------- Plot of GRllFUGE·DATE.Legend: A-I etc. Ob3, B-2 obs , A AAA etc. R e Limited Access + AAA AAA AA A A A A A A t Limited ·Access + A A u 9 e ·1 s. t. u s Motor Veh. Motor Veh. --+-------------+-------------+-------------+-------------+-- 07/16/95 08/05/95 08/25/95 DATE 09/11/95 10/01/95 + A A A A A AA --+-------------+-------------+-------------+-------------+-07/16/95 08/05/95 08/25/95 DATE 09/H/95 10/04/95 �81 ___________________________ YBAR-~99S10-119.100 Plot ~-------------------------- of GRBPUGS·DATI.Legend: A - 1 01>5, B-2 01>5, ------------------~--------- etc. Plot ---------------------------- YEAR-199S10-150.310 of GRBFUGS·DATE.Legend: A - 1 ob.s, B-2 01>5, etc. R AAA LiAited Access + 1 1 1 1 1 1 1 1 1 Motor Veh. + 1 AAA AA A AAA A A e f Limited Access + A, A a t u Motor Veh. oS 08/05/95 08/25/95 09/14/95 + 1 ---+--------+--------+--------+--------+--------+--------+-08/30/95 10/01/95 09/04/95 09/09/95 --------------------------- YBAR-199SIO-l19.t12 ---------------------------- of GRBFUGS·DATS.Legend: A - lobs, LiAi ted Access + 1 1 1 1 1 1 1 1 1 Motor Yah. + I AAA AAA AA B-2 01>5, A AAA ---------------------------Plot etc. A' A '1 e s t a t u s Access + I 1 1 1 1 1 1 1 1 Motor Veh. + 1 09/29/95 YEAR-199SX0-1S0.320 ---------------------------- AA AAA A 07/16/95 08/05/95 oos , etc. A A A A 08/25/95 DATS 09/11/95 10/01/95 DATB ------------------------ of GRBFUGB·DATE.Legend: A - lobs, 09/24/95 --+-------------+-------------+-------------+-------------+-- --+-~-----------+-------------+-------------+-------------+-08/05/95 08/25/95 '09/11/95 10/01/95 07/16/95 YBAR-199S10-150.001 09/19/95 of GRBFUGS·DATS.Legend: A - 1 obs , B-2 R e f Limited u _'--- 09/14/95 DATE DATE Plot A S t --+-------------+-------------+--~----------+-------------+-- ------------, A '1 e 07/16/95 Plot A u B-2 ---------------------------Plot eee , etc. YEAR-l99S 10-150.920 ---------------------------- ot GRBFUGE·DATE.Legend: A - ,1 obs , B-2 ebs , etc. R e Limited Acc.ss Motor Yah. + 1 1 1 1 1 1 1 1 I AAA AA A AA AA A A t Limited Access + <} e S t a t u + A s Motor Yah. A AA '1 --+--------_._-+-----------+-------------+-------------+-07/16/95 08/05/95 08/25/95 09/11/95 10/01/95 A A.A A A A AA A A A + 1 --+-------------+-------------+-------------+-------------+-07/16/95 08/05/95 08/25/95 09/11/95 10104/95 DATE --------------------------Plot YEAR-199S10-150.240 A 1 1 1 1 1 1 1 1 1 u DATE ---------------------------- of GRBFUGS·DATE.Legend: A - 1 ebs , B-2 ---------------------------Plot ob.•, etc. YBAR-199S10-151.010 ---------------------------- of GRBFUGS·DATE.Le<}end: A - 1 ob.s, B-2 ob.s, etc. R e Limited Acce.5~ + 1 1 1 1 AA AAA A A A AAA A A f Limited Acce.•.• + A AAA A A u <} e 1 S 1 t a t u Motor Veh. '+ 1 s Motor Veh. --+-------------+-------------+-------------+-------------+-07/16/95 08l05/95 08/25/95 DATE 09/14/95 10/04/95 + AAA AAA AA --+-------------+-------------+-------------+-------------+07/16/95 08/05/95 08/25/95 09/14/95 10/04/9 �82 Plot Liaited Analysia DATI YEAR-1995 10-151.139 - ___________________________ of ·GRBFUGII·DATII.Legend: A-lobs, Access Motor Veh. of IUk Mov••• nt During Archery Season 1992 - 1995 Dat. SOl probablility for refuge _ Ii - 2 005, YEAR N Obs etc. Variable DAY50 DATlI50 A + I I I I I· I I I I + I ·A A A A A A A 08/05/95 08/15/95 A 08/25/95 6 DAYARClI days from Arch. DAY50 Julian day of 0.5 DATE50 Day of 0.5 6 6 6 0 240.0000000 12293.00 1994 7 DAYARClI days from Arch. DAY50 Julian day of 0.5 DATE50 Day of 0.5 7 7 7 12.1428571 251.1428571 12669.14 1995 11 DAYARClI days from Arch. DAY50 Julian day of 0.5 DATE50 Day of 0.5 11 11 11 5.0000000 213.0000000 13026.00 Plot Limited YEAR-1995 10-151.199 Variable 1992 10 DAYARCH days from Arch. DAY50 Julian day of 0.5 DATlI50 Day of 0.5 4.6439925 4.6439925 4.6439925 1.1685594 1.4685594 1.4685591 1993 6 DAYARCII days from Arch. Julian day of 0.5 DAY50 DATESO Day·of 0.5 5.2153619 5.2153619 5.2153619 2.1291626 2.1291626 2.1291626 1991 7 DAYARCII days frc-. Arch. DAYSO· Julian day of 0.5 DATlI50 Day of 0.•5 13.1960312 13.1960312 13.4960312 5.1010203 5.1010203 5.1010203 1995 11 DAYARClI days from Arch. DAY50 Julian day of 0.5 DATESO Day of 0.5 12.7671453 12.7671153 12.7671453 3.8191392 3.8191392 3.8194392 ---------------------- of GRBFiJGE·DATE. Legend: A-lobs, Access YEAR N Obs of lilt Movement During Archery Season - 1995 Data Plots of refuge status by date _______________________ B-2 A + etc. 005, A I I I Label Std Dev Std Error ------------------------------------------------------------------------- I I I I I I + Motor Veh. Julian day of 0.5 Day of 0.5 -1. 7000000 240.3000000 11927.30 09/04/95 DATE Analysis Mean N 10 10 10 1993 --+-'------------+-------------+-------------+-------------+-- 07/26/95 Label ----------~-------------------------------------------------------1992 10 DAYARCII days frOOl Arch. YEAR N Obs Variable Label DAY50 DATE50 Julian day of 0.5 Day of 0.5 231.0000000 11918.00 2.0000000 241.0000000 11931.00 Mina ••• Maxiaua --------------------------------------------------------------------------1992 10 DAYARCII days ·frOlll Arch. -11.0000000 A A A A A A A A A A I ---+--------+--------+--------+-------+--------+--------+-. 08/05/95 08/15/95 08/25/95 09/04/95 09/14/95. 09/24195 1993 6 DAYARCII days frOli Arch. DAYsO Julian day of 0.5 DATlI50 Day of 0.5 -7.0000000 233.0000000 12286.00 7.0000000 217.0000000 12300.00 1991 7 DAYARCII days frOla Arch. DAYSO Julian day of 0.5 DATESO Day of 0.5 -3.0000000 236.0000000 12651.00 31.0000000 270.0000000 12688.00 1995 11 DAYARCII days frora Arch. DAY50 Julian day of 0.5 DATESO Day of 0.5 -17.0000000 221. 0000000 13001.00 21.0000000 262.0000000 13015.00 10/04/95 DATE Analysis of lilt Movement During Arc-hery Season Totaled 1992-1995 501 probablility Variable Label for Data refuge Mean N Std Dev Std Error --'. ----------------------------------------------------- DAYARCII days frc-. Arch. DAYSO Julian day of 0.5 DATlI50 Day of 0.5 Variable 31 34 31 Label DAYARCH days frOlll Arch. DAY50 Julian day of 0.5 DATE50 Day of 0.5 Variable 3.6176471 243.3529412 12500.03 10.9323393 10.5021855 153.3916361 Minimum Maximwil -17.0000000 221. 0000000· 11918.00 Label DAYARCH days from Arch. DAY50 Julian day of 0.5 DATE50 Day of 0.5 31.0000000 270.0000000 13015.00 Variable YEAR N Obs 1.8718807 1.8011100 77.7565386 10 1992 Label T Prob>ITI --------------------1.1575970 DAYARCII days frOll Arch. DAYSO Julian day of 0.5 DATESO Day of 0.5 163.6297166 8121.77 0.2768 0.0001 0.0001 T 1993 6 1. 9295345 135.1127621 160.7585630 DAYARClI days from Arch. Julian day of 0.5 DAY50 Day of 0.5 DATE50 0 112.7203721 5773.63 1.0000 0.0001 0.0001 1994 7 DAYARClI days from Arch. DAY50 Julian day of 0.5 DATE50 Day of .0.5 2.3801761 19.2338478 2183.65 0.0517 0.0001 0.0001 1995 11 DAYARClI days from Arch. Juli~n d~y of 0.5 DAY50 DATE50 Day of 0.5 1.2988905 63.1260789 3383.87 0.2231 0.0001 0.0001 Prob>ITI 0.0623 0.0001 0.0001 -----------------------------------------------------------------------Analysis OBS 1 2 3 1 5 YEAR of Elk: Movement During Archery Season 1992 - 1995 Data 95' CIon days from archery opening day _TYPE_ 1992-1995 1992 1993 1994 1995 0 1 -FRBQ_ 31 10 6 7 11 MEAN 3.6176 -1. 7000 0.0000 12.1129 5.0000 STDERR 1. 8749 1.1686 2.1292 5.1010 3.8494 N 31 10 6 7 11 LCI -0.19683 -5.02211 -5.17319 -0.33889 -3.57708 UCI 7.1321 1.6221 5.1732 il.6216 13.5771 �83 Appendix B Alternative Study" Plans Alternative two - Close hunting one year on two GMUs(chosen randomly) and issue the full number of permits for the other two GMUs. Reverse the treatments the following year. This alternative is essentially identical to alternative one with different treatments on the north and south halves of the study area. Treatment one would be business as usual, with early-season hunting beginning on the last weekend in August on one half. Treatment two would consist of no early-season hunting permits (deer or elk) issued on the other half. There would be no limitation on the number of early-season hunting permits issued, thus the treatment could have high hunter densities during the study. While higher hunter densities would not be typical for the area, they would provide a strong treatment and prevent local business from suffering losses during the study. GMUs 12 and 23, and 24 and 33 would be paired as in alternative one, with the same response variables and testing of alternative hypothesis. Like alternative one, this would be a two-period crossover design (Manly 1992, Ott 1993) with different treatments (Table 6). Table 6. Layout of Latin square crossover design with corresponding model'for alternative two. FactcrB 1996 CdlaredFlk Sooth half cf area n Early-seasm hunting n No earl -seascn huntin No early-seasm hunting Statistical analysis, sample size calculation, and testing of alternative hypothesis would be the same as for alternative one. The weakness of alternative one are applicable to this alternative, but there are additional logistical and practical problems. Hunters may boycott the White River area because they can't hunt in their traditional areas, or because they fear the crowds on the two areas left open to hunting. There would be numerous hunter complaints about this alternative. Second, the guides and outfitters have their traditional hunting areas defined, Closing of GMUs could cause them time and money in the establishment of new camps. Further, guides, outfitters, and local businesses in general, may have a decrease in their overall early-season revenues, if hunters boycott the area. Alternative three - Move the early-season hunting forward for two weeks on non-refuge areas only, then close hunting on public land and open it for the following two weeks on refuge areas only. Alternative three is a crossover design similar to alternative one (Table 7), but with the experimental units being refuge and non-refuge areas, the sampling units being elk, and treatment being hunting or no hunting. However, the number of collared elk would change between the first and second periods, some to all of the elk in the non-refuge areas may move to refuge areas during the second period seriously unbalancing the sample sizes. Analysis of this alternative would be the same as for alternative one for both the main and alternative hypothesis. �84 Table 7. Layout of Latin square crossover design for alternative three. CdlarooElk First 2 \\efks ofseasn Nn-refuge n n ofseasm N>hmting Hntin The recommended sample size calculation is the same as for alternative one except area is refuge or non-refuge area instead of each half of study area. Therefore, 22 cows would be collared on refuge areas, and 22 cows would be collared on non-refuge areas (44 total collars). Alternative three suffers from all the drawbacks of alternative one and two, except for hunter displeasure. Additionally, alternative three would be more logistically difficult than either alternative one or two, because the opening and closing of areas would have to be coordinated with public and private landowners. Timing and coordination of closing and opening of areas to hunting is likely to be a logistic nightmare requiring many personnel out in the field coordinating the activity. Almost daily aerial radio tracking of the elk would be required to ensure enough samples for reaction to two perturbations so close in time. Finally, the analysis and inferences may be weakened by unbalanced sample sizes. �85 Appendix C Study Design Under Ideal Conditions Observational studies and small manipulative experiments cannot predict responses of elk to hunter pressure. This information is only attainable through large-scale field experiments, which due to economic, administrative, and biological factors, often are not adequately replicated. Meta-analysis is one alternative to assist in prediction of widespread or typical responses to hunting (Gurevitch et al. 1992), but the best alternative is to have adequate spatial and temporal controls. Before the experiment is designed, much thought should be given to defining the important responses of interest, that is, responses that managers would like to be able to understand and/or predict for management of an elk herd. To evaluate movement of elk to refuge areas in response to the opening of early-season hunting, I would use a spatially replicated crossover design, with four treatments on four adjacent areas for each study area. The four treatments would be: Area Area Area Area 1 - no hunting 2 - full hunting pressure at normal time 3 - full hunting pressure two weeks early 4 - full hunting pressure two weeks late. This design would take four years, as each treatment would be administered each season. The spatial replicate would be the study area (each with four sub-areas for treatment). The study areas would be chosen by random stratified sampling throughout the state where there is a reasonable amount of elk hunting. I would recommend at least three study areas to evaluate spatial heterogeneity in elk responses to early-season hunting and to get a good estimate of the error term. The sequences would have to be related in each study area so each row and column received each treatment as required by the Latin Block design, which the crossover design is based on. However, each study area could have a different sequence to eliminate possible sequence effects from the analysis. A weakness of this design is that it cannot separate out persistent learned behaviors from the error term. Allowing several years for each treatment or adding a treatment including the import of elk naive to hunting would allow measurement of learning effects. However, because the literature indicates that disturbance effects tend to be short lived, I would begin with this experiment on the assumption that there will be little persistent behavior year to year. This experiment would not address effects of density of hunters, cover etc.; it is specifically designed to test for a cause and effect relationship between opening of early-season hunting and movement of elk to refuge areas. Once the relationship between opening of early-season hunting and movement of elk was established, then the next logical step would be do design an experiment to evaluate hunter density effects or establish threshold densities that result in movement during early-season hunting. �86 Budget -1996 1 ElkColiars 10 collars x 80 elk x Item Costs $220 'collar shipping and handeling for 80 collars $400 'elk captured SubTotals $2.200 $50 $32,000 $34,250 2 Aighttime AUg-Sept July, Oct-Nov HPPSupport 2 dayslweek x 1 dayslweek x 9 weeks x 6 weeks x 8 hrsldayx 8 hrsldayx $185 Ihr $185 Ihr $26,640 $8,880 ($20,000) $15,520 3 Salary Ever-faithful graduate research asslstart salary and tuition Volunteer for 2 months Intensive data collection Equipment maintenance and office supplies Travel money for presentations $25,000 Total $74.770 Budget-1997 Item Costs SubTotals 1 Elk Collars 10 elkx $4,000 $400 'elk captured $4,000 2 Aighttime Aug-Sept July, Oct-Nov HPPSupport 2 dayslweek x 1 dayslweek x 9 weeks x 6 weeks x 8 hrsldayx 8 hrsldayx . $185 Ihr $185 Ihr $26,640 $8,880 ($20,000) $15,520 3 Salary Ever-faithful graduate research assistant salary and tuition Volunteer for 2 months Intensive data collection Equipment maintenance and office suppli~ Travel money for presentations $25,000 Total $44,620 �87 Colorado Division Wildlife Research July 1996 of Wildlife Report JOB state of Project Work No. W-153-R-9 Mammals Research Elk Inyestigations Job No. Author: REPORT Colorado Plan No. Period PROGRESS Estimating Developing Population Covered: July Survival Rates of Elk and Techniques to Estimate Size I, 1995 - June 30, 1996 D. J. Freddy Personnel: J. Broderick, G. Byrne, A. Coriell, D. Crane, J. Ellenberger, D. Fox, J. Frothingham, V. Graham, J. Gray, G. Loucks, D. Masden, C. Mehaffy, J. Ritchie, L. Stevens, P. Will, CDOW; D. Bowden, G. White, CSU; CSU Diagnostic Laboratory; K. Crane, C. McCarty, N. Miers, D. OUren, A. Ryel, volunteers; BLM Glenwood Springs, NBS Ft. Collins, Rocky Mountain Elk Foundation, USFS Rifle, cooperating. ABSTRACT We captured and radio-collared 71 calf elk (Cervus elaphus nelsoni) 6-months old in December 1995 to estimate survival rates during winter 1995-96 and to increase numbers of radiocollared elk available for experiments utilizing mark-resight models to estimate population density. Elk were captured using helicopter net-gunning and portable corral traps. Survival rates (± 95% CI) for 6-11 month old calves during winter-spring were 0.92 ± 0.06, 0.90 ± 0.07, and 0.88 0.08 in 1993-94, ,'1994-95, and 1995-96, respeotively. Survival was similar among years (P > 0.50) and sexes (P > 0.70). Suspected primary causes of death for calves were predation (57%) and malnutrition (33%). Survival rates for adult females (~ 12 months old) during winter-spring were 0.96 ± 0.05, 0.96 ± 0.04, and 0.95 ± 0.04 also during 1993-94, 1994-95, and 1995-96 and survival was similar among years (P > 0.70). Hunting was the primary cause of death (57%) for adult females during winter-spring. Annual survival rates (1 December-30 November) for adult females were 0.78 ± 0.10 and 0.84 ± 0.09 in 1993-94 and 1994-95, respectively. Survival was similar between years (P > 0.50) and hunting annually accounted for 88% of the adult female deaths. Hunting removed 79% of the adult 2-year old males. Survival of males from 6month old calf to 35-month old adult were 0.09 ± 0.10 compared to 0.86 ± 0.12 for females of the same cohort. Wounding loss on adult males was 9% and illegal loss of yearling males averaged 10%. Estimates of population size based on mark-resight estimators ranged from 3,272-3,618 elk or about 26 elk/mi2 of winter range. The IEJHE mark-resight model using data pooled from ± �4 aerial flights provided the best estimate of population size which was 3,415 elk with a 95% CI of 3,129-3,807. Estimates based on counts of elk on randomly selected 1 mi2-quadrats during 2 replicate flights were 2,165 and 3,175 elk. Sighting bias models to adjust for negative bias of quadrat counts are currently being evaluated. We believe stratified sampling systems using quadrats as sample plots and with counts corrected for sighting bias holds promise for providing reasonable estimates of population size. �B9 ESTIMATING SURVIVAL RATES OF ELK AND DEVELOPING ESTIMATE POPULATION SIZE David TECHNIQUES TO J. Freddy P. N. OBJECTIVE Estimate estimate survival rates of adult population size. female and calf elk and develop techniques to SEGMENT OBJECTIVES 1. Radio-collar 75 calf and 15 adult female elk during Game Management unit 42 south of Rifle, Colorado. 2. Estimate winter arid annual survival known fates of radioed elk. 3. Estimate density of elk in a portion of GMU-42 using 2 replicate flights of a quadrat sampling system. Apply sighting bias corrections to adjust numbers of elk counted during each replicate. Compare biase adjusted estimates with mark-resight estimates based on numbers of marked elk seen during the 2 replicate flights plus 2 nonrandom mark-resight flights. 4. Analyze 5. Prepare draft of manuscript on sighting collected in 1994 and 1995. 6. Continue elk. data and summarize to monitor annually locations December rates of calf and adult in Federal elk from Aid Job Progress bias models and movements 1995 in developed of selected reports from data radioco11ared INTRODUCTION OUr objectives are to provide reliable estimates of survival rates for calves and adult females during winter and tor adult females throughout the year for the period 1993-94 through 1996-97. Additionally, we will develop and test a system for estimating population size that will incorporate estimates of sighting bias in conjunction with a random sampling system using search quadrats as sample units. Our winter study area encompasses about 839 km2 2 (324 mi ) in the eastern half of Game Management Unit 42 south and east of Rifle, Colorado. Elk winter range vegetation types include juniper-pinyon woodland (Juniperus osteosperma-Pinus edulis), oakbrush-mountain shrub (Quercus gambelii-Amelanchier alnifolia), aspen (Populus tremuloides), sagebrush (Artemisia tridentata), and agricultural fields (Freddy 1993, 1994). �90 METHODS Marking We placed radio collars (172~176MHz) having mortality sensors on 71 calves (6 months old), of which 37 were males and 34 were females. Of these, 68 calves were trapped from 8-11 December 1995 using helicopter net-gunning and 3 calves were trapped on 21 December using portable corral traps. Helicopter capture occurred at 10 remote sites located primarily on public lands while corraltrapping occurred at 1 site on public lands. Trapping effort was allocated among 8 geographic trap zones to assure that radioed elk were representative of most if not all segments of the population (Table 1). Radio collars were of the same type used in 1993 and 1994 (Freddy 1994). Calves captured by net-gunning were ferried by helicopter to processing points usually within 1.6 km of capture sites. At processing points, body weight, total body length, hind foot length, and rectal body temperature (F) were measured and calves were then radio-collared and released. Similar measurements were also made on calves that were corral-trapped and then released at the trap site. Body measurements for calves were compared between sexes and years using Proc FREQ, GLM, and REG (SAS 1988). Survival We monitored life or death status of radioed elk during daily ground surveys and aerial surveys conducted at 2-4 week intervals from December 1995 through April 1996 and via monthly aerial surveys from May to November 1995 and May to June 1996. Survival rates (S) of radioed elk were calculated using the binomial estimator with a variance, VAR(S) = S(l-S)/n (White and Garrott 1990) (Proc FREQ, SAS 1988). Survival rates are expressed as the mean estimate ± the 95% confidence interval. We used x2 -contigency tests to compare survival rates. We chose not to use the staggered entry approach and did not use a Kaplan-Meier approach because elk were captured in a short time interval and few animals were censored (White and Garrott 1990). We defined 4 major time intervals for survival analyses: winter-spring was 1 December to 14 June, summer-fall was 15 June to 30 November, annual was 1 December to 30 November to coincide with timing of capture and radiocallaring, and yearly for yearling elk aged 12-23 months was 15 June to subsequent 14 June. Life or death status of all calves radioed in December 1995 (71) was known for the period 8 December 1995 through 15 June 1996 except for 1 male calf collar, 174.619/95 which apparently Slipped off the elk in May 1996. On 15 June, calves become yearlings for purposes of calculating rates of calf survival. Life or death status for adult females •.adult males, yearling females, and yearling males collared in December 19.93 or 1994 was known for 198 of 205 elk through 15 June 1996. Causes of death were estimated from multiple sources of evidence including: presence or absence of gunshot wounds, presence or absence of bite wounds on carcass and predator tracks or scat at carcass site, physical positioning of carcass remains whether buried, covered, scattered, or consolidated, relative amount of internal fat and marrow fat if present with carcass, and results of histopathology and marrow fat analyses (Wade and Browns 1982, Halfpenny and Biesiot 1986). Fat content (percent dry matter) of bone marrow and estimates of age based on dental cementum were obtained for dead elk by the Colorado Division of Wildlife Laboratory while histopathology analyses were provided by �91 the Colorado state University Veterinary Diagnostic Laboratory. Photographs were taken of nearly all mortalities so that physical evidence could be reviewed and judged by outside experts (pers. comm. A. Anderson, T. Beck, W. Andelt) • PQpulatiQn Estimates Elk population size in GMU 42 from East Alkali Creek west to Grass Mesa was estimated during mid-January and early March 1996. We counted elk using a Bell-SolQY helicopter on 4 flights. Two replicate flights involved counting elk on 53 quadrats about 1 mi2 (2.59 km2) in size representing a 40% stratified random sample of the 132 mi2 winter range. Each potential quadrat in 8 geograhic blocks were ranked as a high or low density quadrat resulting in 14 strata. Intensity of sampling was 60% in high density and 32% in low density strata with a minimum of 3 quadrats in anyone strata. Two flights involved counting elk encountered on a nonrandom route that traversed the same 132 mi2 area. One flight of each type was conducted during the time periods' 15-20 January and 29 February-3 March 1996. During helicopter surveys, fixedwinged flights were conducted to locate 219 radiocollared elk to determine whether these marked elk were within or outside the 132 mi2 sample area. During all flights, numbers of marked and 'unmarked elk were tallied by observers. We generated estimates of population size using several markresight estimators for each individual flight, pairs of flights, and all 4 flights pooled using program NOREMARK (White 1996). We considered estimates based on all 4 flights pooled to be our best estimate of population size and the benchmark against which individual flights, especially quadrats, would be compared. Estimates from quadrat sampling were computed using program DEAMAN (Colo. Div. Wildl. software). At this time, we have n2t. applied sighting bias correction factors to the counts of elk observed on quadrats (Freddy 1995). Moyements We continued to locate selected radioed elk at least once per month since capture to document seasonal movements via telemetry using a Cessna 185. These elk were selected at random from within trap zones and equalized by age class in Janaury 1994. Elk from the original sample that died were replaced each subsequent January primarily with randomly selected 6 month-old calves of the same sex from the same trap zone(s) as elk that died. As of 1 January 1996, these 44 elk were classified as 22 adult females, 4 yearling females, 6 female calves, 2 adult males, 4 yearling males, and 6 male calves. During June 1996, we again located 28 adult females that were selected at random in 1995 to document locations during the calving period. As needed, we located other elk to document unusual movements. RESULTS AND DISCUSSION ca,pture Two elk calves died during helicopter net-gunning activities. These have been the only deaths during capture of 235 elk using the helicopter. One calf died of a broken neck during capture and one calf (174.800/95) died within 10 days of capture due to the effects of capture myopathy (histopathology, CSU Diagnostic Lab) and was subsequently censored from survival analyses. �92 Survival Between 1 December 1993 and 14 June 1996, 86 radiocollared elk died (Table 2, Appendix 1). Hunting was apparently involved in 66% of the deaths. For adults ~1 year-old, hunting accounted for 87% of 65 deaths. There were 2 periods of mortality during the year. Calves died from February to May while adults died during fall and early winter when hunting seasons occurred (Fig. 1). Calves Survival rates for 6-11 month old calves during winter-spring were 0.92 ± 0.06, 0.90 ± 0.07, and 0.88 ± 0.08 in 1993-94, 1994-95, and 1995-96, respectively (Table 3). We failed to detect differences in survival of calves among years (X22 = 0.40, P >0.50), between sexes pooled among years (X21 = 0.58, P >0.70), and between sexes within each yearly cohort (x\ !£ 0.79, P > 0.70) (Tables 3, 5, 6). Sex of dead calves for all years was 12 male and 9 female (Table 2). These survival rates were associated with winters considered mild in temperature and having low or moderate snow depths. In 1994-95, considerable snow fell during March and April but usually melted rapidly at lower elevations. Causes of death for calves during winter-spring were suspected malnutrition (33%), mountain lion predation (33%), suspected predation (19%), bear predation (5%), and unknown cause (10%) (Tables 2, 4). Percent fat in bone marrow of calves suspected of dying from predation was 82 for males (range 6495, n = 6) and 48 for females (range 29-63, n = 3) and for calves dying from suspected malnutrition, 14 for males (range 0.2-41, n = 3) and 11 for females (range 8-15, n = 3) (Table 4). Yearlings Survival rates for elk 12-23 months of age were 0.86 ± 0.09 for males (n = 57) and 0.95 ± 0.05 for females (n = 66) when yearlings were pooled among years and hunting mortalities were included (Tables 5, 6). Excluding or censoring hunting mortalities increased survival rates to 0.98 ± 0.04 for males and 0.98 ± 0.03 for females. Males had lower survival rates when hunting mortalities were included (X21 = 3.38, P = 0.07) but not when hunting mortalities were excluded (x\ = 0.03, P > 0.50). Of the 11 deaths involving elk 12-23 months old, 9 (82%) were hunting related (Table 3). Adult Females Survival rates for adult females (~ 12 months of age) during winter-spring and inclusive of hunting mortalities were 0.96 ± 0.05, 0.96 ± 0.04, and 0.95 ± 0.04 in 1993-94, 1994-95, and 1995-96, respectively. Excluding hunting mortalities increased survival to 0.98 ± 0.03, 0.99 ± 0.02, and 0.97 ± 0.03 in the same 3 years (Table 7). We failed to detect differences in survival among years inclusive (X22 = 0.37, P >= 0.50) or exclusive (X22 = 1.52, P > 0.70) of hunting mortalities. Of 14 winter-spring deaths, 8 (57%) were related to late-season rifle hunting: 4 legal and 1 illegal harvest and 3 wounding losses (Table 2). Six natural deaths included 2 suspected predation, 3 of unknown cause, and 1 calvingrelated. Of the 3 elk lost to wounding, 2 were suspicious kills. Both of these elk died from 1 rifle shot to the upper neck or head in relatively plain �93 sight near or on an agricultural field where retrieval of the carcass was not difficult. These animals may have been accidentally shot, maliciously shot, or abandoned because of the radiocollar. Survival rates for adult females (~ 12 months of age) during summer-fall and inclusive of hunting mortalities were 0.87 ± 0.07 and 0.94 ± 0.04 in 1994 and 1995, respectively. Excluding hunting mortalities increased survival to 0.99 ± 0.02 and 1.00 in the same 2 years (Table 7). We failed to detect differences in survival among years inclusive (x\ = 3.13, P = 0.09) or exclusive (x\ = 1.38, P > 0'.70) of hunting mortalities. Hunting accounted for 20 (95%) of 21 summer-fall deaths. The one natural death was calvingrelated. During hunting seasons in Fall 1994 and 1995, 100 and 129 radiocollared adult females, respectively, were available to hunters (Table 7). Proportions of these marked elk lost to hunting-related mortalities were 12% in 1994 and 8% in 1995. Annual survival rates for adult females (~ 12 months of age) marked as a cohort in December 1993, inclusive of hunting mortalities, were 0.78 ± 0.10 and 0.84 ± 0.09 in 1993-94 and 1994-95, respectively. ' Excluding hunting mortalities increased survival to 0.96 ± 0.05 and 0.98 ± 0.04 , respectively (Table 8). We failed to detect differences in survival among years inclusive (X21 = 0.19, P > 0.50) or exclusive (x\ = 0.15, P > 0.50) of hunting mortalities. For this cohort of adult females, hunting accounted for 22 (88%) of 25 deaths during both years. Adult Males Survival of males from 6-month old calf to 35 months of age was 0.09 ± 0.10 compared to 0.86 ± 0.12 for females of the same calf cohort (Table 5). Hunting was the overwhelming cause of mortality among male elk. From the 1993-94 calf cohort of 36 male calves, 28 (78%) lived to become 2-year old legal branch-antlered bulls for the 1995 hunting seasons. Of these 28, 22 or 79% were harvested in the Fall of 1995 inclusive of 2 wounding losses (9% of 22). Rifle seasons accounted for 16 (73%) of the hunting mortalities (Table 2). Most of these 2-year old bulls were harvested in the Grand Mesa DAU or in adjacent GMU 43. One exception was a bull taken during archery season in GMU 63 on Black Mesa about 100 miles south of its capture site in GMU 42. One surviving bull was located in March 1996 in GNU 47 east of Basalt, CO about 35 miles southeast of its capture site in GMU 42. Yearling spike-antlered bulls were generally not legal quarry during hunting seasons. In 1994, 32 yearling bulls from the 1993-94 calf cohort entered the hunting seasons presumably as spike-antlered bulls and 4 (12.5%) were illegally taken with 3 taken during rifle seasons and 1 taken prior to rifle seasons. In 1995, 30 yearling bulls from the 1994-95 calf cohort entered the hunting season and 2 (7%) were illegally taken and 1 (3%) was fatally wounded during archery season in an area where the bull was legal quarry. Calf Bogy Size Body weights of male and females calves were largest in 1995 with the increase most pronounced for males (Tables 9, 10). Increases in weights were not unexpected as the locally moist summer of 1995 was favorable to forage production on summer ranges. However, we failed to detect differences in �94 body weights among years (P = 0.206) and any interaction between year and sex (P = 0.380). Overall, male calves had larger body weights (P = 0.0001), longer total body length (P = 0.0386), longer hind leg lengths (P = 0.0001), and higher condition indexes (P = 0.0001) than female calves (Table 10). Accurately predicting body weight from total body length, hind foot length, or a combination of these 2 measurements does not look promising at this time, although all regressions were significant (P < 0.0001). The multiple regression using body length and hindfoot length as independent variables provided the best correlation coefficients (~) of 0.63 for female calves, 0.69 for male calves, and 0.69 for sexes combined. Calves suspected of dying from predation (Table 4) averaged 121 kg in weight for males (range 100-140 kg, n = 6) and 91 kg for females (range 79-117, n = 5). Predators thus appeared to take larger than average males and smaller than average females (Table 10). Calves suspected of dying from malnutrition (Table 4) averaged 95 kg in weight for males (range 77-127, n = 3) and 107 kg for females (range 100-120, n = 3). Malnutrition thus appeared to affect smaller than average males and average sized females (Table 10). Population Estimates Our elk population was not geographically or demographically closed during the time interval encompassing flights. During our first pair of flights, some elk were still moving down in elevation in response to heavy snow in early January resulting in fewer than expected radiocollars on the sample area. By the March flights, all elk had reached elevations encompassed by the sample area but some elk moved lower in elevation onto private lands outside of but adjacent to the sample area and some elk moved to winter ranges adjacent to but also outside the sample area. These movements resulted in radiocollared elk moving into and out of a "pool" of elk that was different and larger than the elk population encompassed by the sample area. Between pairs of flights, 2 radiocollared elk died indicating some loss of elk was occurring during our flight time interval. Fortunately, both quadrat and nonrandom flights reflected proportionately similar and positive changes in total elk and total marked elk counted between pairs of flights indicating that movements of marked and unmarked elk into the sample area were similar (Table 11). Population estimates from 4 mark-resight estimators ranged from 3,272-3,618 elk or 24.8-27.4 elk/mi2 of winter range within the sample area (Table 12). At this time, the estimate of 3,415 elk generated by the Immigration/Emigration JHE mark-resight estimator is probably the best estimate of population size on the sample area because it accounts for movement of elk on and off the sample area. Adding the 165 elk directly counted on private lands in lower Divide Creek and Hunter Mesa (57 mi2 area) . increased the population to 3,580 elk in the intensive study area. In January 1995, a single nonrandom flight produced a Lincoln-Peterson mark-resight estimate of 3,411 .elk and adding 399 elk counted on private lands in lower Divide Creek and Hunter Mesa to this estimate increased the population to 3,810 elk in the intensive study area in 1995 (Freddy 1995). Population estimates for the larger "pool" of elk within which radiocollared elk were moving ranged from 3,969-4,004 elk as estimated by the IEJHE and BE mark-resight estimators (Table 12). This "pool" estimate would include elk on the Divide Creek and Hunter Mesa private lands and some additional elk on areas adjacent to the western boundary of the intensive study area. �95 Population estimates from quadrat flights 1 (2,165 elk) and 2 (3,175 elk) were lower than the IEJHE mark-resight estimate (3,415) (Table 12). Although both quadrat flights were subject to sampling error, the lower estimate from the first flight may reflect a less favorable distribution of elk during that flight. Confidence intervals overlapped between quadrat flight 2 and IEJHE suggesting estimates were not different. Quadrat estimates will be adjusted upward for Sighting bias and at that time more conclusive comparisons to markresight estimators will be made (Freddy 1995, Samuel et al. 1987). Elk densities ~ 50/mi2 occurred within the Garfield Creek High, west Divide Creek West High, and Grass Mesa Low strata which together encompass about 25 mi2 of winter range (Table 13). Elk Moyements In December 1994, 33 male and 36 female calves and 8 adult females were radiocollared. Of these elk, 27 male and 31 female calves and all 8 adults survived to 30 November 1995. None of the adult females dispersed during winter 1995-96 to a winter range outside of the intensive study area in GMU 42. However, 25 (9M, 16F) or 43~ of the surviving yearling elk dispersed to winter ranges outside the intensive study area (Table 14). Rates of dispersal were 33% for males and 52% for females. Calves trapped in zones E and F, which encompass Alkali, Dry Hollow, and the Mamm creeks, comprised 68% (17) of the dispersing yearlings but only 45% of the surviving yearlings. Calves trapped in zones E and F dispersed to winter ranges near the towns of Collbran, Parachute-Rulison, and Paonia. Calves trapped in zones C and D which encompass West Divide Creek comprised 28% of the dispersing juveniles and these elk primarily moved south to winter ranges near Paonia and Paonia Reservoir. We are currently creating a GIS land database for the area frequented by radiocollared elk. This is a cooperative effort through the National Biological Service and Rocky Mountain Elk Foundation. We believe plots of elk locations and movements using this database will be forthcoming in 1996. 12 10 _ CALVES _ ADULT HALES 1-:·:-:-:-1 ADULT fEl1ALES ~ 8 H co 6 4 ·······································.1·········11··: .... 2 ~~ -<l,.~ r:.:j ~~ <:?-Q;- ~ . . .. ...... .................. l·lT o . IH ~.;. ~~~ J I ~<y ~ ~(j ~ § 0 ::: ~ ~G . ~ ~\) ~G ~ MONTH Figure 1. Timing of deaths for calf and adult 12 months old and adults were >12 months old. elk, 1993-1996. Calves were 6- �96 CONCLUSIONS We obtained acceptably precise estimates of calf and adult survival rates and recommend continuing our current sampling effort of capturing and radiocollaring 70 calves in 1996-97 and continuing to monitor > 75 radioed adult females and> 25 radioed adult males in 1996-97. We recommend conducting replicate aerial surveys in 1996-97 to estimate elk density using sample quadrats, mark-resight estimators, and sighting bias correction models to complete our evaluation evaluating techniques to estimate elk density. LITERATURE CITED Freddy, D. J. 1993. Estimating survival rates of elk and developing techniques to estimate population size. Colo. Div. Wildl. Game Res. Rep. July: 83-117. Freddy, D. J. 1994. Estimating survival rates of elk and developing techniques to estimate population size. Colo. Div. Wildl. Game Res. Rep. July: 27-42. Freddy, D. J. 1995. Estimating survival rates of elk and developing techniques to estimate population size. Colo. Div. Wildl. Game Res. Rep. July. Halfpenny, J. C., and E. A. Biesiot. 1986. A field guide to mammal in North America. Johnson Books, Boulder, co. 161pp. tracking Samuel, M. D., E. O. Garton, M. W. Schlegel, and R. G. Carson. 1987. Visibility bias during aerial surveys of elk in northcentral Idaho. Wild1. Manage. 51:622-630. SAS Institute Inc. 1988. SAS/STAT Cary, NC. 1028pp. User's Guide, 1982. Procedures Texas Agric. Expt. 6.03. SAS Institute, D. A., and J. E. Browns. livestock and wildlife. White, G. C. 1996. NOREMARK: Population estimation surveys. Wildl. Soc. Bull. 24:50-52. White, G. C., and R. A. Garrott. 1990. Analysis of wildlife data. Academic Press, Inc., San Diego. 383pp. by ~--------------------------David J. Freddy Life/Science Researcher Inc., for evaluating predation on Sta. Publ. B-1429. 42pp. Wade, Prepared J. from mark-resighting radio-tracking �Table 1. Elk capture objectives and numbers of elk radiocollaredin 8 trapzones,December 1993,1994.and 1995 within Game Management Unit 42. Elk Capture Elk Collared' Calves Collared Trap Objectives Total/Year Helicopter Corral Years Males Females Years Adult Females Collared Zone Name 93 94 95 93 94 95 93 94 95 93 94 95 Total 93 94 95 93 94 95 Total 93 94 95 Total A B C D E F G H Garfield Gibson Uncle Bob \JestDivide Hightower Middle Mamm \JestMamm Dry Hollow All 8 20 24 26 10 10 8 44 5 8 13 13 10 8 8 21 6 12 11 11 10 9 10 6 150 86 75 8 25 29 21 17 0 6 35 6 17 13 11 17 13 0 6b 4 7 14 18 14 9 5 0 141 83 71 8 19 29 21 17 0 6 0 6 4 7 4 13 14 11 18 17 14 13 9 0 5 0 0 0 0 0 6 10 3 0 0 0 0 0 0 0 0 0 0 0 0 o 0 0 35 6 0 18 49 56 50 48 22 11 41 3 5 10 5 4 0 2 7 100 67 68 41 16 3 295 36 3 5 7 1 9 8 0 0 1 4 10 8 8 5 1 0 33 37 1 8 7 6 3 0 3 9 3 3 6 3 6 4 8 10 8 6 5 4 0 4 0 0 14 31 44 38 38 22 10 16 4 12 12 10 10 0 1 19 37 36 34 213 68 0 6 0 2 0 0 0 6 0 0 0 0 0 0 0 0 4 18 12 12 10 0 1 25 14 0 82 • Helicopter = Helicopternet-gunning,Corral = corral-trapping. bAll 6 adult females captured 2 March 1995 Table 3. Survival rates of calves with sexes pooled for 1 December - 14 June each year 1993-94, 1994-95, and 1995-96. Survival rates (S) calculated as a mean estimate of (alive)/(alive + dead) and variance S(lS)/n collars. Elk Aqe and_Time Period (dates) Calves Cal\res Calves 12/01/9312/01/9412/01/9506/14/94 06/14/95 06/14/96 Survival L 95% CI U 95% CI n collars Censored' Died Nonhunting Hunting 0.92 0.85 0.98 73 0 6 6 0 0.90 0.83 0.97 69b 0 7 7 0 0.88 0.81 0.96· 69 2c 8 8 0 • Censored denotes collar failure and/or animal life/death status not known. b Includes (173.949/94) male whose collar failed 12/94 but seen alive 1/95, 1/96. e Censored elk were (174.619/95) for slipped collar and (174.800/95) for trap-related mortality. !!3 �98 Table 2. Causes of deaths in radiocollared elk between 1 December 1993 and 14 June 1996. Calves (M=male, F=female) were 6-12 months old and collared at 6 months of age. Yearling males and females were 12-18 months old and collared as 6 month old calves. Juvenile males and females were 18-23 months old and collared as 6 month old calves. Adult males and females were ~ 24-months old at time of death. Elk Age/Sex Class at Death Calves Adult Yearling Juvenile Total Cause of Death M F M F M M F M F F All 2 2 o o o o o 1 o o o o o o o o o o o o o o 1 0 2 2 0 1 1 0 2 0 o o o o 1 o o o o Legal Hunting Archery Muzzleloading Archery/Muzzle Rifle-First Rifle-Second Rifle-Third Rifle-Late 0 0 0 0 0 0 0 0 0 0 0 0 0 0 o o o o o o o 1 o o o o o o o o o o o o 5 1 1 0 o 1 527 101 011 628 246 426 Wounding Loss Archery/Muzzle Rifle-Regular Rifle-Late 0 0 0 0 0 0 1 o o o 1 112 4 3 2 o o o o 2 o o o o Illegal Hunting Rifle-Regular Out-of-Season 0 0 0 0 sa o o o o o o o o o 0 505 1 011 Presumed Hunting Disappear-Rifle Seasons 0 0 o o Unknown Cause 2 0 o 1 o o o 2 12 9 7 3 1 o 22 32 Malnutrition Unknown-Suspect Malnutrition Predation-Lion Predation-Bear unknown Predator Unknown-Suspect Predation Accident-Birthing Totals 5 o o o o o o o o o .0 o o o o o o 0 0 1 2 3 1 0 5 3 8 2 2 4 3 6 3 0 101 011 1 235 2 o 6 2 2 4 4 2 5 o o 224 055 o 336 235 42 44 86 • These elk were illegally shot as spike-antlered yearling males: (173.190/93), (173.232/93), (173.320/93), (173.919194), (174.059/94). • These elk disappeared during rifle hunting seasons and are presumably dead and legally harvested: (172.207/93), (172.649193), (172.800/93), (173.309/93), (173.390193), (173.439/93). One exception may be spike-antlered yearling male (173.309193) that disappeared during the first rifle season in 1994 when spike-antlered bulls were not legal. �Table 4. Estimated causes of mortality and associated body condition for 21 radiocollared months old, 1 December 1993 - 14 June 1996. Age' Trap BodY" Marrow. Frequency ID/ Date Dead Zone Sex (months) Wt. (kg) % Fat Ca.useof Death Year CaI2tured 172.899/93 173.000/93 173.262/93 173.289/93 173.461/93 173.469/93 173.589/94 173.640/94 173.789/94 173.870/94 174.119/94 174.140/94 174.170/94 174.220/95 174.230/95 174.240/95 174.339/95 174.500/95 174.609/95 174.689/95 174.789/95 C G C C H H B C E D F F B F F E E C B D C F F M M M M F F F F M M M M M M F F F M M 9 10 9 9 8 10 12 9 9 8 9 9 10 9 10 10 9 10 10 7 11 • Approximate age at death assuming 15 June birthdate. • Whole body weight at capture in December of 1993, 1994, or e Fat content as percent dry matter of bone marrow from either d Fat content as percent dry matter of bone marrow from either e Fat content as percent dry matter of bone marrow from either 86.0 102.0 108.0 111.0 82.0 77.0 117.0 89.0 100.0 86.0 140.0 112.0 140.0 127.0 120.0 115.0 79.0 78.0 120.0 100.0 03/18/94 04/25/94 03/18/94 03/22/94 02/07/94 04/25/94 06/01/95 03/30/95 03/20/95 02/21/95 03/01/95 03/14/95 04/25/95 04/08/96 04/24/96 04/17/96 03/27/96 04/16/96 04/15/96 02/05/96 OS/22/96 28.50d 8.03c 28.88d 0.21c 0.27c 51.71c 14.71c 94.97c 64.44c 76.05c 41.08c 86.28c 62,97c 34.56c 10.28c 91.90c 75.32· calf elk 6-11 & Code No. Lion Predation-3 Malnutrition-6 Unknown-11 Unknown-11 Malnutriton-6 Malnutrition-6 Unknown; suspect predation-30 Unknown; suspect predation-30 Unknown; suspect malnutrition-31 Lion Predation-3 Bear Predation-35 LionPredation-3 Lion Predation-3 Unknown; suspect malnutrition-31 Lion Predation-3 Lion Predation-3 Unknown Predator-5 Unknown; suspect malnutrition-31 Malnutrition-6 Lion Predation-3 Unknown; suspect predation-30 1995. femur of carcass. mandible of carcass. humerus of carcass. s �...• 8 Table 5. Survival rates for the cohort of 6-month old elk calves radiocollared in December 1993 for 6 time periods from 1 December 1993 through 14 June 1996 when elk were 35 months old. Survival rates for male and female calves pooled when 6-11 months old were 0.92 (95% CI 0.86-0.98, n=73). Survival rates (S) calculated as a mean estimate of (alive)[lalive + dead) and variance_S(l-S)/n collars. Elk Age (months) and Time ~eriod (dates) 6 - 11 12 - 17 18 - 23 24 - 29 30 - 35 6 - 35 12/01/9306/15/9412/01/9406/15/9512/01/9512/01/9306/14/94 11/30/94 06/14/95 11/30/95 06/14/96 __ 06/14/96 MALES Survival L 95% CI U 95% CI n collars Censored" Died Nonhunting Hunting 0.89 0.78 0.99 36 0 4 4 0 0.88 0.76 0.99 32 0 4b 0 4 FEMALES Survival L 95% CI U 95% CI n collars Censored" Died Nonhunting Huriting 0.95 0.87 1. 00 37 0 2 2 0 0.97 0.92 i .00 35 0 1f 0 1 1. 00 28 0 0 0 0 1. 00 34 0 0 0 0 0.21 0.06 0.37 28c 0 22d 0 22 1. 00 0.00 0.00 . 3 3e 0 0 0 0.09 0.00 0.19 33 3 30 4 26 0.97 0.91 1. 00 34 0 1 0 1 0.97 0.91 1. 00 33 0 1 0 1 0.86 0.98 0.75 37 0 5 2 3 • Censored denotes collar failure and/or animal life/death status not known. b Collar (173.309/93) "disappeared" during Fall 1994 hunting seasons and assumed dead. e Includes collar (173.241/93) that failed in August 1995 but bull seen alive January 1996. d Three collars (173.390/93,173.402/93,173.439/93) "disappeared" during Fall 1995 hunting seasons and assumed dead . e Three collars censored; (173.241193) failed, (173.340/93) not heard spring 1996, (173.381193) slipped off 12/01/95. r Collar (172.800/93) "disappeared" during Fall 1994 hunting seasons and assumed dead. �Table 6. Survival rates for the cohort of 6-month old elk calves radiocollared in December 1994 for 4 time .periods from 1 December 1994 through 14 June 1996 and survival rates for the cohort of 6-month elk calves radiocollared in December 1995 for one time period.. Survival rates for male and female calves pooled when 6-11 months old were 0.90 in 1994 (95% CI 0.83-0.97, n=69) and 0.88 in 1995 (95% CI 0.81-0.96, n=69). Survival rates (S)_Gillculatedas a mean estimate of (alive) / (aliy~__±_d.eaclLand variance S (l-S) /n collars. Elk A91L_(mQntb_~_and T:i.mePeriod (dates) 1994 Calf Cohort 1995 Calf Cohort 6 - 11 12 - 17 18 - 23. 6 - 24 6 - 11 12/01/9406/15/9512/01/9512/01/9412/01/9506/14/95__ 11/30/95 06/14/96 06/1AL~L________ 06/14/96 MALES Survival L 95% CI U 95% CI n collars Censored" Died Nonhunting Hunting 0.91 0.81 1. 00 33b 0 3 3 0 .0.90 0.79 1. 00 30b 0 3 0 3 0.95 0.87 1. 00 22 5" 1 1 0 0.75 0.59 0.91 28 5 7 4 3 0.86 0.74 0.98 35 2d 5 5 0 FEMALES Survival L 95% CI U 95% CI n collars Censored" Died Nonhunting Hunting 0.89 0.78 0.99 36 0 4 4 0 0.97 0.91 1. 00 32 0 1 0 1 0.97 0.90 1. 00 30 l' 1 1 0 0.83 0.70 0.96 35 1 6 5 1 0.91 0.81 1. 00 34 0 3 3 0 • Censored Includes , Censored d Censored e Cenosred b denotes collar failure andlor animal lifeldeath status not known. collar (173.949/94) that failed in December 1994 but male seen alive in January 1996 elk were (173.949194) failure, (173.981/94), (174.019/94), (174.090/94), (174.160/95) elk were (174.619195) slipped collar. (174.800195) capture mortality . elk was (173.719/94) slipped collar. not heard spring 1996. ...• s �....•• 2 Table 7. Survival rates for all radiocollared adult female elk ~12-months old during 5 periods from 1 December 1993 through 14 June .1996. Survival rates (S) calculated as a mean estimate of (alive)/(alive + dead) and variance S(l-S)/n collars. . Elk Aqe and Time Period (dates) Adult Adult Adult Adult Adult 12/01/9306/15/9412/01/9406/15/9512/01/9506/14/94 11/30/94 06/14/95 11/30/95 06/14/96 0.96 0.87 0.96 0.94 0.95 Survival 0.91 0.8a 0.92 0.90 0.91 L 95% CI 1.00 0.94 1.00 0.98 0.99 U 95% CI 68b• 100bf 95bg 129bh 119; n collars o 0 0 0 2i Censored' 3 13c 4 8d 7 Died 1 110 4 Nonhunting 2 12 3 8 3 Hunting • Censored denotes collar failure andlor animal lifeldeath status unknown. Includes collar (172.011193) that failed 4/1994 but seen alive in 1/1996 e Collars (172.800/93,172 .649/93) "disappeared" Fall 1994 hunting seasons,assumed • Collar (172.207/93) "disappeared" Fall 1995 hunting seasons, assumed dead. e Composition is 6 females 18+ and 62 females 30+ months old. Composed Composed h Composed ; Composed j Censored t I b dead. of 35-12+,6-24+, and 59-36+ months old females. of 34-18+, and 61-30+ montho old females. of 32-12+,34-24+, and 63-36+ month old females. of 30-18+ and 80-30+ monthos old females. (172.Dl1/93) failure and (173.719/94) slipped collar. Table 8. Survival rates for the group of adult female elk ~12-months old radiocoll~red in December 1993 for 8 time periods from 1 December 1993 through 14 June 1996. Survival rates. (S) calculated as a mean estimate of (aliv~)/(alive + dead) and.variance S(l-S)/n collars. Elk Aqe and Time Period (dates) Adult Adult Adult Adult Adult Adult Adult Adult 12/01/9306/15/9412/01/9406/15/9512/01/9512/01/9312/01/9412/01/9306/i4/94 11/30/94 06/14/95 11/30/95 06/14/96 11/30/94 11/30/95 06/14/96 0.96 0.82 0.93 0.88 0.93 0.78 0.84 0.58 Survival 0.91 0.72 0.85 0.78 0.85 0.68 0.75. 0.46 L 95% CI 1.00 0.91 0.99 0.97 1.00 0.88 0.93 0.70 U 95% CI 68b• 65bf 53bf 49bf 42f 68bf 53bf 67f n collars o 0 0 0 19 0 0 19 Censored' 3 12c 4 6d 3 lSc 10d 28cd Died 1 110 3 2 1 6 Nonhunting 2 11 3 6 0 13 9 22 Hunting • Censored denotes collar failure andlor animal lifeldeath status unknown. b Includes collar (172.011/93) failed 4/1994 but seen alive 1/1996. c Collar (172.649/93) "disappeared" Fall 1994 hunting seasons, assumed dead. • Collar (172.207/93) "disappeared" Fall 1995 hunting seasons, assumed dead. Composed of 6-18+ and 62-30+ month old females. r Composed of 100% females 24+ months old. 'Censored (172.011/93) failure of unknown status Spring 1996. e �Table 9. Frequency distribution of who~e body weights for male and female elk calves trapped in Game Management Unit 42, December, 1993, 1994, and 1995. Percentage 2er weight class shown in 2arentheses. Bod~ Weight 100-109 Class (kg) 110-119 120-129 60-69 70-79 80-89 90-99 M 0(0.0) 0(0.0) 0(0.0) 1 (2.9) 1(3.0) 0(0.0) 1 (2.9) 1(3.0) 0(0.0) 3 (8.6) 2 (6.0) 4(11.4) 6(17.1) 6(18.2) 3 (8.6) 12(34.3) 13(39.4) 8(22.9) 10(28.6) 6(i8.2) 9 (25.7) F F F 0(0.0) 0(0.0) 1(3.1) 1 (2.9) 2 (5.7) 2 (6.3) 4 (11.4) 4 (11.4) 1 (3.1) 8 (22.9) 7(20.0) 3 (9.4) 12(34.3) 10(28.6) 14(43.8) 4(11.4) 10(28.6) 5(15.6) 6 (17.1) 2 ( 5.7) 6(18.8) Year Sex 1993 1994 1995 M 1993 1994 1995 M Bod~ measurements 130-139 140-149 Total 1 (2.9) 2 (6.0) 10(28.6) 1 (2.9) 2 (6.0) 1(2.9) 35(100) 33(100) 35(100) 0(0.0) 0(0.0) 0(0.0) 0(0.0) 0(0.0) 0(0.0) 35 (100) 35 (100) 32(100) for elk calves tra22ed in Game Management Unit 42, December, 1993, 1994, 1995. Male Calves Female Calves Min___ Max Mean SD n Mean SD Min Max .n Table 10. Year Measurement 1993 112.7 Body Weight (kg) 191.2 Body Length (em) Hindfoot Length (em) 56.3 0.59 Condition Index· 13.7 10.2 2.2 0.05 77.0 164.0 52.0 0.43 141. 0 210.0 61. 0 0.67 35 36 36 35 103.8 188.9 54.8 0.55 12.2 9.8 2.1 0.05 76.0 167.0 51.0 0.46 123.0 207.0 59.0 0.64 35 35 34 34 1994 113.5 Body Weight (kg) Body Length (em) 190.3 Hindfoot Length (em) 56.9 Condition Index 0.59 14.2 9.2 1.8 0.05 71.0 168.0 50.0 0.42 140.0 204.0 59.0 0.69 33 32 32 32 103.0 186.3 55.1 0.55 13.3 8.8 2.1 0.06 70.0 166.0 50.0 0.42 128.0 207.0 59.0 0.65 35 36 36 35 1995 119.1 Body Weight (kg) Body Length (em) 194.0 Hindfoot Length (em) 57.4 0.61 Condition Index 13.6 9.3 2.2 0.05 92.0 166.0 53.0 0.50 141. 0 206.0 61. 0 0.71 35 36 36 34 105.7 189.4 54.9 0.56 15.2 11.4 2.3 0.06 60.0 158.0 47.0 0.38 129.0 203.0 59.0 0.66 32 34 34 32 115.1 Body Weight (kg) All 191.9 Years Body Length (em) Hindfoot Length (em) 56.9 0.60 Condition Index 14.0 9.6 2.1 0.05 71.0 164.0 50.0 0.42 141.0 210.0 61.0 0.71 103 104 104 .101 104.1 188.2 54.9 0.55 13.5 10.0 2.1 0.06 60.0 158.0 47.0 0.38 129.0 207.0 59.0 0.67 102 105 104 101 ....•• • Condition index = (Weight/body length). 8 �104 Table 11. Summary of flights to estimate elk population size and density on 132 mi2 of winter range in GMU 42 south of Rifle and Newcastle, Colorado, January/March 1996. Flight Total Marked Unmarked Marked Date Elk Elk Flight Elk Elk Flight (M/D) Counted Counted Available Counted Time (hrs) Quadrat 1 NonRandom 1 NonRandom 2 Quadrat 2 4 Flights Pooled Flight Quadrat 1 NonRandom 1 NonRandom 2 Quadrat 2 1/15-18 1/19-20 2/29 3/1-3 128 128 137 137 32 54 80 49 853 1300 1918 1324 885 1354 1998 1373 19.6 8.5 8.0 19.1 154" Marked/ Unmarked 0.0375 0.0415 0.0417 0.0370 Marked/ Marked+Unmarked 0.0362 0.0399 0.0400 0.0357 Proportion of Available Marks Seen (+/- 95% CI ) 0.2500 (0.0749) 0.4219 (0.0857) 0.5839 (0.0825) 0.3577 (0.0803) Changes in Total Elk Observed: Quadrat 1 vs. Quadrat 2: +488 elk or 55% increase over Quadrat 1 NRandom 1 vs. NRandom 2: +644 elk or 48% increase over NRandom 1 Changes in Total Marked Elk Observed: Quadrat 1 vs. Quadrat 2: +17 marked elk or 53% increase over Quadrat 1 NRandom 1 vs. NRandom 2: +2.6 marked elk or 48% increase over NRandom 1 Marked Elk Deaths: During the population estimate interval (1/15/96 - 3/3/96), 2 radiocollared elk died: 174.689/95 male calf on 2/5/96 and 172.070/93 adult female on 2/12/96 . • Marked elk available with immigration/emmigrationmovements involving 41 elk; 25 elk moved onto and 16 moved off the intensive sampling area during the time interval between 1120/96 and 2/29/96. Table 12. Estimates of population size and density for elk on 132 mi2 of winter range in GMU 42 south of Rifle and Newcastle, Colorado, January/March 1996 . Pvt. Density population Model , . /mi2 Landa 95% Conf. Int. Flights Estimator Estimate 3,168-3,840 26.3 165 4 Flights Pooled JHEb 3,472 3,129-3,807 25.9 165 4 Flights Pooled IEJHEc 3,415 BEd 2,764-3,872 24.8 165 Quad I+NonRaridom 1 3,272 27.4 165 3,169-4,130 Quad 2+NonRandom· 2 BE 3,618 3,621-4,446 4 Flights Pooled IEJHE 3,969" 3,560-4,504 4 Flights Pooled 4,004f BE StRSg 165 1,356-2,974h 16.4 Quadrats Flight 1 2,165 2,205-4,145h 24.1 165 Quadrats Flight 2 StRS 3,175 • Elk on private land enumerated by direct counts must be added to population estimate for estimated total elk in the intensive study area. h Joint Hypergeometric Mark-Resight Estimator thatassumes a geographically closed population and homogeneous sighting probabilities of individual elk. c Immigration/Emigration Joint Hypergeometric Mark-Resight Estimator that allows for movement of elk on/off intensive study area but assumes homogeneous sighting probabilities of elk. d Bowden Mark-Resight estimator allows for some violation of a geographically closed population but allows for heterogeneous sighting probabilities of elk. e Estimate of total pool of elk within which radiocollared elk are distributed on or just off the intensive study area; equates to N' estimate. r Estimate of total pool of elk; equates to N'. • Stratified random sample of quadrats with counts of elk not adjusted for sighting bias. h 90% confidence intervals. �Table 13. Counts of elk on quadrats during 2 replicate flights in February and March, 1996 in GMU 42 south of Rifle and Newcastle, Colorado. Strata Quadrats Elk Elk/Quadrat Population Size Strata Size (mi2) . N n Counted Mean StdErr. Total + 90% CI Quadrat Flight 1 Garfield High 10 10 5 133 26.60 16.107 266 265 Garfield Low 16 16 4 83 20.75 17.970 332 473 E. Divide.Low 16 16 5 44 8.80 4.023 141 i06 W. Div. East High 6 6 4 95 23.75 11.800 143 116 W. Div. East Low 10 10 3 45 15.00 6.535 150 107 W. Div. West High 5 5 3 28 9.33 5.283 47 43 W. Div. West Low 9 9 3 13 4.33 2.762 39 41 Hightower High 8 8 4 44 11.00 4.103 88 54 Hightower Low 19 19 6 185 30.83 16.425 586 513 D. Hollow High 3 3 3 15 5.00 0.000 15 0 D. Hollow Low 9 9 3 0 0.00 0.000 0 0 W. Mamm High 5 5 3 161 53.67 27.282 268 224 W. Mamm Low 7 7 3 3 13.00 5.674 91 65 Grass Mesa Low 9 9 3 0 0.00 0.000 0 0 Totals 132 52 885 16.40 2165 809 90% Conf. Int. on population: 1356 to 2974 90% Conf. Int. as percent of estimte: 37.38% -----------------------.------------------------------------------------------------------------------Quadrat Flight 2 Garfield High Garfield Low E. Divide Low W. Div. East High W. Div. East Low W. Div. West High W. Div. West Low Hightower High Hightower Low D. Hollow High D. Hollow Low W. Mamm High W. Mamm Low Grass Mesa Low Totals 10 16 16 10 16 16 5 5 5 6 6 4 10 10 3 5 9 8 5 9 8 3 19 19 3 9 5 7 9 3 9 5 7 9 132 3 4 6 3 3 3 3 3 53 90% Conf. Int. on population: 2205 to 4145 386 121 34 147 53 149 .11 56 41 6 77 87 16 189 1373 77.20 24.20 6.80 36.75 17.67 49.67 3.67 14.00 6.83 2.00 25.67 29.00 5.33 63.00 24.05 38.032 7.976 2.627 9.937 8.030 6.786 1.905 7.921 3.608 0.000 20.957 10.973 1.764 39.466 772 387 109 221 177 248 33 112 130 6 231 145 37 567 3175 626 210 69 98 132 56 28 104 113 o 310 90 20 584 970 90% Conf. Int. as percent of estimate: 30.56% ...•. fil �106 Table 14. Radiocollared elk captured on winter range in GMU 42 during December 1994 that dispersed out of GMU 42 to a new winter range as of 27 March 1996. Locations (GMUs) determined by aerial telemetry. Trap New Frequency Location Description Zone Age" Sex GMU 173.621/94 LOWER WEST MUDDY CREEK C 1+ F 521 173.659/94 MCCLURE PASS HIGHWAY C 1+ F 521 173.66.9/94 DEADHORSE CREEK D 1+ F 521 173.679/94 TERROR CREEK D 1+ F 521 HIGHTOWER MT. BUZZARD CREEK 173.689/94 E 1+ F 421 173.699/94 SMALLEY GULCH E 1+ F 421 173.709/94 DEADHORSE CREEK D 1+ F 521 173.719/94 VEGA RESERVOIR D 1+ F 421 MCCLURE PASS HIGHWAY 173.739/94 E 1+ F 521 173.750/94 LEROUX CREEK E 1+ F. 52 173.760/94 COLLIER CREEK E 1+ F 421 MORRISANA MESA F 173.799/94 F 42 West 1+ P!~CHUTE, WALLACE CREEK 173.829/94 F 42 West F 1+ MCCLURE PASS HIGHWAY F 173.840/94LE 521 F 1+ TERROR CREEK D 173.859/94 521 F 1+ PAONIA RESERVOIR C 173.990/94 521 M 1+ BUCK MESA SOMMERSET 174.030/94LE E 521 M 1+ 174 ..039/94 HAWXHURST CREEK E 421 M 1+ VEGA RESERVOIR F 174.049/94 421 M 1+ HUBBARD CREEK E 174.069/94 521 M 1+ HAWXHURST CREEK E 174.080/94 421 1+ ..~ GRASSEY GULCH F 174.109/94LE 421 1+ M LEROUX CREEK F 174.129/94 1+ 52 M PORCUPINE CREEK 174.150/94 F F 42 West 1+ LOWER EAST MUDDY CREEK B 174.181/94 M 521 1+ a b Age of elk in years as of March 1996 LE = Location elk routinely located. �107 Appendix I. Mortalities of 86 radiocollared elk from 1 December 1.993through 14 June 1996. Age is approximate age in years of elk at death; C=calf 6-12 months old, Y=yearling 12-18 months old. Body weight measured in December when elk captured as calves. Frequency 10/ Trap Body Date Year Captured Site Zone Sex Age WT.(kg) Heard Dead Cause·of Death & Code Number 172.030/93 BR A F 5+ Legal harvest rifle season-28 10/27/95 172.039/93 GR B F 2+ Unknown; suspect predation-30 06/14/95 172.070/93 BR A F 11 Unknown-11 02112196 172.080/93 GM A F 5+ Legal harvest·rifle season-28 11/05/94 172.090/93 GR B F 11 Wounding loss late rifle season-26 01/16/94 Wounding loss rifle season-25 172.139/93 BC C F 17 10/19/95 Legal harvest rifle season-28 172.160/93 CC C F 2+ 10/23/94 172.181/93 MC C F 3 Wounding loss rifle season-25 11/03/94 172.201/93 GS C F 3 Wounding loss rifle season-25 11/15/94 172.207/93 SG 0 F 2+ Disappear rifle season-22 11/30/95 172.258/93 HY C F 2+ Legal harvest archery/muzzle season-27 10/04/94 172.277/93 GS C F 3 Wounding loss archery/muzzle season-24 10/04/94 172.290/93 SG 0 F 5+ Legal harvest rifle season-28 10/23/94 172.369/93 AC E F 2+ Legal harvest archery season-33 09/18/94 172.369/94 FM B F 2+ Legal harvest late rifle season-29 01/14/96 172.409/93 AC E F 8 Wounding loss late rifle season-26 12/29/94 172.509/93 FS H F 3 Legal harvest rifle season-28 10/21/95 172.542193 FS· H F 6 Lion predation-3 02/01/94 172.549/93 PG H F 5+ Legal harvest late rifle season-29 11/28/95 172.570/93 PG H F 16 Wounding loss rifle season-25 11/03/74 172.581/93 PG H F 2+ Legal harvest rifle season-28 10/17/94 172.581/94 FM B F 2+ Legal harvest late rifle season-29 12/08/95 172.590/93 PG H F 2+ Unknown-11 04/24/96 172.610/93 PG H F 3 Accident; birthing/calving-32 10/04/94 172.639/93 PG H F 16 Accident; birthing/calving-32 06/19/96 172.649/93 PG H F 2 Disappear rifle season-22 11/30/94 172.670/93 PG H F 4 Legal harvest late rifle season-29 01/16/95 172.678/93 PG H F 9 Wounding loss late rifle season-26 12/22/94 172.690/93 FM B F 8 Illegal kill-7 01/24/94 172.699/93 FM B F 5+ Legal harvest rifle season-28 11/05/95 172.800/93 SR B F Y Disappear rifle season-22 105.0 11/30/94 172.899/93 BC C F C Lion predation-3 86.0 03/18/94 172.950/93 GH 0 F 2 Legal harvest rifle season-28 107.0 10/18/95 173.000/93 YM G F C Malnutrition-6 102.0 04/25/94 173.041/93 GM A M 2 Legal harvest rifle season-28 107.0 10/19/95 173.060/93 MH E F 2 Legal harvest late rifle season-29 99.0 12/27/95 173.120/93 GM A M 2 Legal harvest rifle season-28 99.0 10/14/95 173.190/93 BC C M Y Illegal kill rifle season-9 111.0 10/29/94 173.201/93 GR B M 2 Wounding loss rifle season-25 106.0 11/30/95 173.210/93 GC B M 2 Legal harvest rifle season-28 108.0 . 10/19/95 173.219/93 BC C M 2 Legal harvest rifle season-28 111.0 11/06/95 173.232/93 BR AMY Illegal kill rifle season-9 101.0 11/15/94 173.262/93 BC C M C Unknown-11 108.0 03/18/94 173.269/93 MC C M 2 Legal harvest rifle season-28 121.0 11/06/95 173.289/93 MC C M C Unknown-11 111.0 03/22/94 173.300/93 GS C M 2 Legal harvest rifle season-28 119.0 10/24/95 173.309/93 HY C M Y Disappear rifle season-22 124.0 11/30/94 173.320/93 HY C M Y Illegal kill rifle season-9 118.0 10/20/94 173.332/93 GH 0 M 2 Legal harvest muzzleloading season-34 125.0 09/12/95 173.351/93 SG 0 M 2 Legal harvest rifle season-28 103.0 11/05/95 173.359/93 AC E M 2 Legal harvest archery season-33 111.0 09/06/95 173.370/93 MH E M 2 Legal harvest rifle season-28 141.0 11/08/95 173.390/93 MG 0 M 2 Disappear rifle season-22 113.0 11/30/95 173.402/93 MG 0 M 2 Legal harvest rifle season-28 118.0 10/18/95 173.410/93 WM G M 2 Wounding loss rifle season-25 90.0 11/02195 173.420/93 WM G M 2 Legal harvest rifle season-28 125.0 10/24/95 173.429/93 MH E M 2 Legal harvest archery season-33 126.0 09/14/95 173.439/93 PG H .M 2 Disappear rifle season·22 0.0 11/30/95 173.450/93 PG H M 2 Legal harvest rifle season-28 113.0 10/15/95 173.461/93 PG H M C Malnutrition-6 82.0 02107/94 Wounding loss archery/muzzle season-24 173.461/94 CM AMY 123.0 09/19/95 173.469/93 FS H M C Malnutrition-6 77.0 04/25/94 173.479/93 FS H M 2 Legal harvest archery season-33 122.0 09/10/95 Legal harvest archery season-33 173.492/93 FS H M 2 110.0 09/03/95 Legal harvest archery season-33 173.510/93 FM B M 2 91.0 08/28/95 Legal harvest rifle season-28 173.521/93 FM B M 2 124.0 10i17/95 Legal harvest archery season-33 173.549/94 HM B F Y 106.0 09/07/95 Unknown-11 173.580/94 PR A F Y 114.0 12121/95 �108 Appendix I. Continued. Frequency 10/ Trae Year Captured Site Zone 173.589/94 173.640/94 173.789/94 173.870/94 173.919/94 174.059/94 174.101/94 174.119/94 174.140/94 174.170/94 174.220/95 174.230/95 174.240/95 174.339/95 174.500/95 174.609/95 174.689/95 174.789/95 OG MC MR SM BC MR MM MM MM FM MS MS MD MD LB GB AP I./D B C E 0 C E F F F B F F E E C B 0 C Sex Age F F F F M M· M M M M M M M F F F M M C C C C Y Y Y+ C C C C C C C C C C C ~ \.IT. (kg) Date Heard Dead 117.0 06/01/95' 03/30/95 89.0 100.0 03/20/95 02/21/95 86.0 11/02/95 114.0 126.0 10/17/95 117.0 OS/24/96 .140.0 03/01195 112.0 03/14/95 140.0 04/25/95 127.0. 04/08/96 120.0 04/24/96 115.0 04/17/96 79.0 03/27/96 78.0 (J4/16/96 120.0 04/15/96 100.0 02/05/96 0.0 OS/22/96 Cause of Death Unknown; suspect predation-30 Unknown; suspect predation-3D Unknown; suspect malnutrition-31 Li.on predation-3 Illegal kill rifle season-9 Illegal kill rifle season-9 Unknown; suspect predation-30 Bear predation-35 Lion predation-3 Lion predation-3 Unknown; suspect malnutrition-31 Lion predation-3 Lion predation-3 Unknown predator-5 Unknown; suspect malnutrition-31 Malnutrition-6 Lion predation-3 Unknown; suspect predation-30 �109 Colorado Division of Wildlife Wildlife Research Report July, 1996 JOB PROGRESS REPORT State of Colorado project NO. ~WL-~1~5~3~-~R~-~9~ _ Work Plan No Moose Inyestigations Job No. Development of census methods and determination of movements, habitat selection, and degree of calf mortality of moose in North Central Colorado. Period Covered: Author: Mammals Research July 1, 1995 - June 30, 1996 R. C. Kufeld Personnel: D. Bowden, D. Younkin. ABSTRACT Instrumented moose captured during 1991, 1992, 1993, and 1994 were located at approximately 2-week intervals from January, 1992, through November, 1995, and at approximately monthly intervals from December, 1995 through June, 1996. Data analysis was begun to determine seasonal movements, home range size, and habitat selection. Results will be presented in a future report. ��111 DEVELOPMENT .OF CENSUS METHODS AND DETERMINATION OF MOVEMENTS, HABITAT SELECTION, AND DEGREE OF CALF MORTALITY OF MOOSE IN NORTH CENTRAL COLORADO Roland P. N. C. Kufeld OBJECTIVES 1. To determine the proportion of moose counting moose in North Park. 2. To determine 3. To determine the degree of dispersal of young animals, and seasonal movements, home range size, and habitat selection of North Park moose. the extent of moose actually observed calf mortality when aerially in late winter. SEGMENT OBJECTIVES 1. To determine the extent of moose 2. To determine the degree of dispersal of young animals, and seasonal movements, home range size, and habitat selection of North Park moose. STUDy The study area was described by Kufeld METHODS AND calf mortality in late winter. AREA (1992). MATERIALS Instrumented moose captured during 1991, 1992, 1993, and 1994 (Kufeld 1992, 93, 94) were located at approximately 2-week intervals from January 1992 through November 1995, and at approximately monthly intervals from December 1995 through June 1996. Most 2-week locations were made by aerial telemetry using a Cessna 185 aircraft with a 2 element, "H" configuration rec·eiving antenna mounted on each strut. A switchbox permitted the telemetry operator, a passenger in the aircraft, to operate antennas jointly or separately. Most monthly and some 2-week locations were made by tracking on the ground until the animal was observed when the airplane was not available. Moose locations were plotted on USGS 1:50,000 scale maps and recorded by UTM coordinates. Vegetation type was also recorded for each moose location. �112 RESULTS Hoose monitoring Analysis of data for movements, home range size, habitat use, mortality and survival for all tagged moose will be presented in a future report when periodic monitoring of moose and data analysis is completed. Results of the P. N. objective "To determine the proportion of moose actually observed when aerially counting moose in North Park" was published during this segment (Bowden and Kufeld 1995). Colorado State Forest Strategic Plan A report entitled "Status and management of moose in the Colorado State Forest and adjacent area of North Park was prepared based on findings of this study and included in the Colorado State Forest Ecosystem Planning Project Strategic Plan, prepared and approved by the Colorado State Board of Land Commissioners (Kufeld 1996). LITERATURE Bowden, D. C., and R. C. Kufeld. 1995. size estimation applied to Colorado 851. CITED Generalized mark-sight population moose. J. Wildl. Manage. 59:840- Kufeld, R. C. 1992. Development of census methods and determination of movements, habitat selection, and degree of calf mortality of moose in North Central Colorado. Colo. Div. Wildl. Wildl. Res. Rep. July:95108. Kufeld, R. C. 1993. Development of census methods and determination of movements, habitat selection, and degree of calf mortality of moose in North Central Colorado. Colo. Div. Wildl. Wildl. Res. Rep. July: (119124) • Kufeld, R. C. 1994. Development of census methods and determination of movements, habitat selection, and degree of calf. mortality of moose in North Central Colorado. Colo. Div. Wildl. Wildl. Res. Rep. July: (4347) • Kufeld, R. C. 1996. status and management of moose in the Colorado state Forest and adjacent area of North Park. 13 pages in Appendix of Colorado state Forest Ecosystem Planning Project strategic Plan. Colorado state Board of Land Commissioners, February, 1996. Prepared by _ Roland C. Kufeld ·LS Res/Sci III. �113 Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS State of Project Work Colorado No. W-153-R-9 Mammals Plan No. ~S~P~I~ _ Job No. Period REPORT Deer Research Inyestigations Regulation of Mule Deer Population Growth by Fertility Control: Laboratory, Field, and Simulation Experiments Covered: Authors: Personnel: July 1, 1995 - June 30, 1996 Dan L. Baker T. M. Nett, and N.Thompson P.E. Bleicher, Hobbs C. W. McCarty, J. Griess ABSTRACT The Rocky Mountain Arsenal has the potential to become one of the premier wildlife viewing areas in the Nation. However, realizing that potential depends on wise management. In particular, the mule deer population contained within the boundary fence must be regulated in balance with the resources the area offers. Contraception offers a potential alternative to recreational hunting and professional culling to achieve this objective. Of the techniques potentially available for reducing reproduction in wild deer, conjugates of gonadotropin rele~sing hormone and cytotoxins appear to the most promising noninvasive method for'providing lasting infertility after a single treatment. In 1995-1996, we attempted to treat female mule deer with GnRH~toxin conjugates but initial' in :vit~cj laboratory experiments were unsuccessful. Other efforts, however, were more successful. We developed an analytical model to evaluate the ability of contraception to regulate animal populations. This model compares the effects of contraception on population dynamics relative to the effects of culling. ��115 REGULATION OF MULE DEER POPULATION GROWTH BY FERTILITY LABORATORY, FIELD, AND SIMULATION EXPERIMENTS Dan L. Baker and N. Thompson CONTROL: Hobbs P •N • OBJECTIVES 1. To develop a practical and acceptable method populations using GnRH-toxin conjugates. 2. To demonstrate the feasibility the Rocky Mountain Arsenal. 3. To predict simulation population modeling. impacts for controlling of such control of alternative mule deer in a field application contraceptive regimes at using SEGMENT OBJECTIVES 1. Evaluate the effectiveness and duration of a single dose application GnRH-toxin conjugate to prevent normal production of reproductive hormones in captive mule deer. 2. Develop a mathematical model regulate animal populations. to evaluate the ability of contraception of to Introduction The Rocky Mountain Arsenal, located in close proximity to Denver, Colorado, offers an exceptional opportunity for an urban population to view and enjoy Colorado's wildlife. However, the Arsenal presents unique problems as well as unique opportunities. For reasons of security, the perimeter of the area was fenced in 1990. While this fence dpes not impede the usual movements of birds and small mammals, it does create an unnatural barrier to the movements of ungulates, particularly mule deer (Odocoileus hemionus). Before the Arsenal was fenced, mule deer numbers were held at a relatively constant equilibrium of about 300 animals. As a result of preventing those movements, deer numbers within the Arsenal will certainly rise exponentially during the next decade. Experience with enclosed deer populations elsewhere has shown that such increases will lead to degradation of habitat, widespread starvation, and eventually to catastrophic declines in animal numbers. The only way to prevent this outcome is to control the abundance of the enclosed deer population. This, is usually done by public hunting or by professional culling of animals. However, public hunting cannot be allowed on the Arsenal because of concerns for security. Moreover, although professional culling would eliminate excess animals, it would also reduce the value of the deer population as a watchable resource. In this situation, alternatives to hunting as a means of regulating ungulate numbers are needed. Fertility control offers a viable alternative'to hunting population control when hunting is infeasible. However, as a means of current fertility �116 control technology does not provide a means of controlling ungulate numbers practically and economically. Here, we propose to develop a practical and economical method of fertility control in mammalian wildlife that overcomes many of the shortcomings of current technology, particularly problems of treatment duration and environmental safety. We propose to use conjugates of gonadotropin-releasing hormone (GnRH) and cellular toxins to selectively destroy gonadotropin-producing cells in the anterior pituitary gland, thereby preventing gamete production by the ovaries and testes. This research project consists of laboratory, field, and modeling phases, each of which is designed to address questions that must be answered before widespread application of hormonal-toxin conjugates is possible. Details of the experiments to be conducted in each of these phases are described by Baker (1992). In this report, we discuss: 1) results of laboratory studies with captive mule deer to determine the most effective dose of GnRH-toxin conjugate,2) development of an analytical model that compares the use of non-lethal and lethal methods for regulating ungulate populations, and 3) reproductive and genetic characteristics of the Rocky Mountain Arsenal mule deer population. 1. EVALUATION OF SINGLE DOSE APPLICATION OF GNRH-TOXIN CONJUGATE One of the most promising ,new approaches to contraception involves linking synthetic analogs of gonadotropin-releasing hormone (GnRH) to cytotoxins. GnRH is a molecule produced in the hypothalamus of the brain. It directs specific cells in the pituitary gland to synthesize and secrete two important reproductive hormones; follicle stimulating hormone (FSH) and luteinizing hormone (LH). These latter two hormones, known as gonadotropin, control proper functioning of the ovaries in the female and testes in the male. By coupling a superactive analog of GnRH to a cytotoxin, it should be possible to specifically target that toxin to LH and FSH-secreting cells in the anterior pituitary gland. Of the techniques potentially available for reducing reproduction in wild deer, conjugates of gonadotropin releasing hormone (GnRH) analog and cytotoxins appear to be the most likely noninvasive method for providing lasting infertility after a single treatment (Baker et ale 1993, Nett et ale 1993). consequently, research, is underway to evaluate the efficacy of a GnRHcytotoxin conjugate in reducing fertility in mule deer (Baker 1994). During 1995-1996, in vitro laboratory experiments using conjugates of GnRH analogs and either a diphtheria or plant toxin were moderately successful in reducing LH secretion from the pituitary but neither conjugate maintained its effectiveness over time. Therefore, in vivo experiments using mice, domestic sheep and mule deer are pending until laboratory evaluation and testing of other toxins has been completed. 2. DEVELOPMENT See Appendix OF FERTILITY A. CONTROL MODEL �117 3. REPRODUCTIVE AND GENETIC MOUNTAIN ARSENAL CHARACTERISTICS OF FEMALE MULE DEER AT THE ROCKY Applying GnRH-toxin conjugates to control the growth of deer populations at the Rocky Mountain Arsenal will require that wildlife managers choose specific tactics for treating animals. Choices must be made on the number and age to treat, the frequency of treatment, timing of treatments and so on. We will provide support for these decisions by developing an interactive model of mule deer population dynamics. This model will combine knowledge of deer biology with an understanding of the constraints intrinsic in the GnRH-toxin conjugate technique. Fundamental to the insights offered by this model is knowledge of the reproductive and genetic characteristics of the Rocky Mountain Arsenal mule deer population. Here, we present results of reproductive studies conducted during 1995 - 1996. The objectives of this study were to 1) measure reproductive rates of female mule deer and compare these rates with those for 1993 and 1994 2) collect the anterior pituitary gland and hypothalamus from pregnant females in order to more precisely estimate dosage requirements for contraception 3) collect tissue samples to evaluate genetic diversity of male and female mule deer populations. Methods Collection of Deer A total of 103 mule deer(74 females, 27 males, 2 unknown) were collected during the period October 11, 1995 to March 20, 1996. Animals were shot through the spine in the cervical region; all died within 5 minutes; no wounding loss occurred. Estimation of Age The left incisor tooth was collected from each deer. Deer with deciduous dentition were assigned an age using the tooth replacement chronology of Robinette et al. (1957). Deer with permanent dentition were assigned an age from counts of dental cementum in the first incisor (Erickson and Seliger 1969) • Pregnancy Rates Reproductive tracts of 74 females were examined for pregnancy. Does lacking identifiable embryos or fetuses after December 29 were regarded as current breeding failures. If corpora lutea were present, we used only those does in which there was gross evidence, through presence of embryos or pigmented degenerating corpora lutea, that the doe either had been or was currently pregnant. Reproductive data was summarized by the following age classes: yearling, 2-year old, prime, old, and unclassified. Yearlings examined for pregnancy were 18-23 months of age; 2-year olds were 30-35 months of age; prime does were those estimated .to be 3-7 years old, inclusive, and old does were 8 years and older. Unclassified does were those known only to have been older than fawns. �118 Anterior Pituitary/Hypothalamus The brain was completely removed from the cranium by first making three-four cuts on the skinned skull with a bone saw or cleaver, two along the edge of the frontal and parietal bones, and one perpendicular to the longitudinal axis of the skull, and at or just above, the postorbital process of the frontal bone. Following removal of the brain, the anterior pituitary gland was removed from the sella turcica by cutting the diaphragm sellae at the rim of the sella turcica. The pituitary and hypothalamus were wrapped in aluminum foil to facilitate fast-freezing, labeled and placed on dry ice. Genetic Evaluation Muscle, liver, and blood samples were collected from all females. Tissue samples were placed in plastic bags and kept on dry ice during collections, then stored at -70C until analyzed. Samples will be subjected to horizontal starch gel electrophoresis and mtDNA restriction enzyme analyses. RESULTS AND DISCUSSION Age Characteristics of Females Approximately 62% of the females collected were of prime breeding age (3-7 yrs),22.6% were 2 years old, 9.5% 8 years or older, and 5.7% fawns and yearlings(Table 1). The oldest female collected was 10 years of age. Most(> 95%) females examined were judged to be in good to excellent condition. This estimate was based on visual assessment of body fat reserves and overall body condition. It should be noted that these collections were not a random sample of the entire breeding population since larger females were intentionally selected over smaller animals. Thus, the proportion of yearling females may be underepresented in these collections. Pregnancy Rates We were unable to determine pregnancy in females collected before January 11.Reproductive data from 25 does collected after January 11 are shown in Table 1. Examinations of these does were made sufficiently long after conception that fetal counts could be made and breeding success of failure determined. Before this date, pregnancy could not be determined by gross examination of the reproductive tract, thus 49 females were excluded from this data set. Pregnancy rate averaged 95.8% was 1.83 fetuses per pregnant Anterior across doe. all age classes. Fetal rate for all does Pituitary/Hypothalamus Analysis these samples is currently in progress at the Department Physiology, Colorado State University, Ft. Collins, Colorado. of �119 Genetic Evaluation Analysis of these samples is currently in progress Ecology Laboratory, Aiken, South Carolina. at the Savannah River LITERATURE CITED Abramowitz, M., and LA. Dover Publications Stegun. 1968. Inc., New York, Himdbook of mathematical NY, pp. 645-652. f'unoti.Lone , Baker, D. L., N. T. Hobbs, T. M. Nett, M. W. Miller, and R. B. Gill. 1993.Regulation of mule deer (Odocoileus hemionus) population growth by fertility control: laboratory, field, and simulation experiments. Contraception in Wildlife Management Symposium, Oct. 26-28, Denver, CO, (Abstract) • Baker, D. L. 1994. Regulation of mule deer population growth by fertility control: laboratory, field, and simulation experiments. Quarterly Report, July-Oct 1994, Colo. Div. Wildl., Ft. Collins, CO, 6pp. Freund, R.J., models. Hobbs, R.C. Littell, and P.C. spector. 1986. SAS Institute, Cary, NC, 187-201. SAS system for linear N.T. 1994. ~egulating populations by controlling fertility: general, stage structured models. Ecological Applications: in review. Miller, M. W., N. T. Hobbs, and M. C. Sousa. 1991. Detecting stress responses in Rocky Mountain bighorn sheep (Ovis canadensis): reliability of cortisol concentrations in urine and feces. Can. J. Zool. 69:15-24. Miller R.G. 1966. Simultaneous New York, NY, pp. 152-168. Morrison, D.F. New York, statistical 1976. Multivariate NY, pp. 145~194. inference. statistical methods. McGraw-Hill McGraw-Hill \ Book Co, Book Co, ��121 APPENDIX A Regulating Populations by Controlling Fertility: General, Stage structured Models N. Thompson Hobbs Research Center" Colorado Division of Wildlife 317 W. Prospect st. Fort Collins, CO 80526 and Natural Resource Ecology Laboratory Colorado State University Fort Collins Colorado 80523 Phone: 303-484-2836 ext 360 Fax: 303-490-2621 Email: nthobbs@lamar.colostate.edu �122 Abstract There is growing interest in using chemical fertility control to regulate the abundance of wildlife populations. Although current technology is clearly effective in its ability to regulate the reproduction of individuals, it remains unclear whether that technology is effective in regulating populations. I offer simple models of populations regulated by several fertility control regimes. I compare dynamics of those models to dynamics of models representing populations regulated by culling. If contraceptives are delivered before breeding, fewer animals would have to be treated with long-lived contraceptives than would have to be culled to maintain populations at levels significantly below carrying capacity. Achieving meaningful reductions in steady state density will require that a large proportion (P) of the female population must be infertile. However, if contraceptives are long lived, then the proportion of the population that must be treated annually at steady state (a) can be substantially less than the proportion that are currently infertile, or the proportion that would have to be culled. Because steady state population density is a non-linear function of P and a, small errors in delivery rates can have large consequences for population equilibria. If these parameters are not estimated and applied properly, populations will overshoot target densities or will go extinct. Models predict that regulating populations of fecund, short-species at steady states substantially below carrying capacity is not feasible using agents delivered after breeding has occurred. �123 INTRODUCTION Regulating the abundance of animals is a prevailing function of contemporary wildlife management. There are a variety of motivations for such regulation, but the most compelling one is that overabundance causes problems, problems that may be biological, economic, or political in scope (e.g., see papers in Jewell and Holt. 1981). Resolving these difficulties requires controlling population growth. Although the growth of populations can be controlled by manipulating births or deaths, populations of wildlife have traditionally been regulated by influencing the death rate using harvest or culling. However, these traditional methods are not always feasible--hunting may not be safe in urban areas; there are ethical concerns about culling of some species, and harvest may conflict with other management objectives, for example, in national parks and preserves (Leopold 1963; Wright 1993). As a result, there is growing interest in controlling the growth of animal populations by influencing fertility (Kirkpatrick and Turner 1985; Brush and Ehrenfeld 1991; Wright 1993) During the 1980's, interest in non-lethal approaches to population control stimulated efforts to develop chemical contraceptives that could be periodically delivered to wildlife. The techniques that emerged from these efforts are clearly effective in controlling reproduction of individuals (Garrott et ale 1992; Eagle et ale 1992; Plotka et ale 1992; Turner et ale 1992), but it remains unclear whether they are effective in thei.r larger purpose--to control the growth of populations (e.g, Hoffman and Wright 1990). Mathematical modeling offers a first step in evaluating the ability of contraception to regulate animal populations. Although several existing models represent effects of fertility control on populations of individual wildlife species (Sturtevant 1970; Knipling and McGuire 1972; Garrott 1991; Garrott and Siniff 1992; Boone and Wiegert 1994), the insights offered by these models tend to be narrow, and a general theory of population regulation via fertility control is only beginning to emerge (Barclay 1981; Caughley et ale 1992; Hone 1992). Here, I offer general models of the dynamics of populations regulated by manipulating fertility. I use these models to compare effects of contraception on population dynamics relative to the effects of culling. I examine how timing of delivery and duration of efficacy of fertility control agents influence their ability to regulate population growth. METHODS Model Development Developing general models of population dynamics requires simplifying assumptions. I begin by assuming that the number of female offspring recruited to the breeding stage can be adequately represented as linear function of the breeding density of adult females. Thus, recruitment is treated phenomenologically--it subsumes effects of density on age of first reproduction, litter size, and juvenile survival. I assume further that the probability of survival of adults is independent of density and remains constant with age. This assumption represents the life span of the animal as the outcome a series of independent Bernoulli trials--species with high survival rates live longer on average than those with low survival, but senescence per se does not occur. When contraceptives are delivered to a portion of the population, the number of fertile animals declines and the number of infertile ones increases. �124 Representing these dynamics in a discrete time model requires assumptions on the sequence of events in the animal's life cycle (Fig. 1). In the models that follow, I assume that census occurs immediately after breeding and that all fertile animals are bred at the time of census. I choose breeding rather than births as the point of reference for census because the state of the animal that influences model behavior is its fertility rather than its age. That is, it doesn't matter whether a female is fertile on her birthday, but it does matter at the time of breeding. I consider two factors that influence the efficacy of fertility control regimes. The first factor is the duration of efficacy of contraceptives. I treat this factor by assuming that effects of contraceptives that last for a fixed number of reproductive seasons or that last for the lifetime of the animal. The second factor is timing of delivery, which I assume can occur before or after breeding. These assumptions motivate models representing 4 cases: (1) fixed duration contrace.ption, prebreeding delivery (2) fixed duration contraception, postbreeding delivery (3) lifetime contraception, prebreeding delivery (4) lifetime contraception, postbreeding delivery I compare these with models of culling, where animals are assumed to be culled before breeding and after. Thus, I offer 6 models, 4 representing fertility control and 2 representing culling (Table 1, 2). Detailed derivation of these models is found in Appendix A. Model Analysis I analyzed model behavior at steady state (Appendix B). steady state occurs when the finite rate .of population growth, A, = 1 and N* = Nt+1 = Nt. To solve for the steady state density, I derived characteristic equations for all models (Table 2), set A = 1 and solved for N. I solved for the proportion of fertile and infertile animals in the steady state population by substituting the expression describing N* for N in each model , extracting the dominant eigenvector, and normalizing its elements to sum to 1. The number of animals treated or culled annually to maintain a given N* (i.e., the treatment rate at equilibrium, T', in females/yr) was computed by solving for c in terms of N* for all models and substituting the resulting expressions in equations describing the treatment rate (for an illustration, see Appendix B). I analyzed local stability of the culling and lifetime contraceptionmodels by linear approximation of the effect of small deviations from steady state (Appendix B). These approximations were obtained by expanding expressions for steady state in a Taylor series, and dropping higher terms. I then solved the characteristic equation of the expanded system and derived conditions for which the absolute value of both of its roots < 1. Numeric illustrations of analytic results were obtained by setting model parameters to values faithful to life history characteristics of whitetailed deer in a population with a carrying capacity of 100 animals (m = 1.87, S = .9, fi = .018, McCullough 1979:88). For fixed duration models, I set r = 1 because effects of current, widely used contraceptives persist for about 1 year. I used numerical simulation to examine trajectories of populations approaching steady state under alternative control regimes. Symbolic and numeric computations were performed using the Maple V computer algebra system (Char et ale 1991). (N°) �125 RESULTS Steady Effects = State Population Density of delivery rate; In the absence of external predict that populations will equilibrate 0), all models Ke S + m - 1 =------ regulation at (i.e., c (1) ~ As the culling rate or the rate of delivery of contraceptives increase, the steady state density is reduced by an amount that depends on the control regime (Table 3, Fig. 2). Models predict that pre~breeding culling and fertility control exert identical effects on steady state densities per unit change in delivery or culling rate (Table 3, Fig. 2). However, at low delivery rates, models predict that contraceptives delivered post-breeding can have a greater effect on the steady state population density than postbreeding culling does if the recruitment rate is sufficiently large (Table 3, Fig. 2). At high delivery rates, these effects are reversed; culling depresses density more than fertility control does. These patterns occur because models show that low-intensity culling of species will elevate the steady state above the maximum steady state that would occur in the absence of control efforts, given sufficiently large m. Such increases are attributable to the compensatory response of recruitment to reduced, post-culling density (Table 3, Fig. 2). Models of culling and pre-breeding fertility control predict that populations will go extinct if 100% of the fertile females are treated annually (Table 3, Fig. 2). In contrast, the post-breeding fertility control models show that treating 100% of the fertile females will simply reduce the population to a new, equilibrium point, N° > 0, whenever m > 1. The cause of this behavior is revealed by setting c = 1 in the equation for the steady state of the post-breeding fertility control models (Table 3) and solving for N°. For both fixed and lifetime duration models, the minimum steady state (N°m1n) that can be maintained using fertility control delivered after breeding is N*, ml.n = m - 1 ~ = K e (m - 1) (2) m + S - 1 At this density each fertile female produces exactly 1 female offspring surviving to reproductive age. Moreover, at this density, the proportion of fertile females in the population equals the adult mortality rate (See below, "Composition of The Steady State Population"). Consequently, recruitment exactly replaces mortality, and the population stabilizes at a steady state, ».; > o. Effects of contraceptive duration; For a given delivery rate, the steady state population density of predicted by fixed duration models (N°F) and the density predicted lifetime models (N°L)e differs by an amount that depends on adult survival and the duration of the contraceptive. In the before breeding case that difference is N* - N* F L and in the arter breeding = CST ~(l-c) case, (3) �126 N* - N* F L = ST+lc(l - C) (4) ~ (1 - CST) Thus, models predict that effects of fixed effect contraceptives approach those of lifetime contraceptives as adult survival declines or as the duration of the contraceptive increases. Composition of the Population at Steady State Effects of timing of delivery: All fertility-control models predict that the proportion of infertile animals in the population increases monotonically with increasing delivery rate (Table 3, Fig. 3). However, the specific form of these predictions depends on timing of delivery. When contraceptives are delivered before breeding and m > 1, the entire female population will be infertile (i.e., p' = 1) whenever 100% of the females are treated population (i.e., c = 1). In contrast, when contraceptives are delivered after breeding and before births, the steady state proportion of infertile females is simply equal to the adult mortality rate (i.e., p' = 1 - S) whenever all fertile . females are treated (c = 1) (Table 3, Fig. 3). Under these circumstances, the number of fertile animals recruited to the adult stage each year exactly replaces the adults that die, as described above (see Steady State Population Density) • Effects of Contraceptive Duration: If inhibition of fertility persists for one year then the proportion of the population that is infertile at the steady state (p') is linearly related to the delivery rate (c) (Table 3, Fig. 3). In contrast, whenever the effects of contraceptives exceed one year, the proportion of infertile animals in the steady state population will exceed the delivery rate needed to m.aintain the steady state, (Le., p' > c, Table 3, Fig. 3). The magnitude of the difference between p' and c increases with increasing duration of contraception (r) and increasing adult survival. When lifetime-duration contraceptives are delivered to long-lived species, the relationship between the proportion infertile animals in the population and the delivery rate is markedly non-linear, and p' can be several fold greater than c, particularly when c is small (Table 3, Fig. 3). In this case, small increases in delivery rate can produce large increases in the equilibrium proportion of infertile animals in the population. population Infertility and Steady State Density: Models predict that increases in the proportion of the population that is infertile (p') do not yield proportionate reductions in the equilibrium population density (Fig. 4). This is revealed by solving for c in terms of p' (Table 3) and substituting these solutions in the expressions for steady state density, N' (Table 3). In so doing, we find the following nonlinear relationship between p' and N' for all fertility control models: e: = 1- (1 - S) m - (5) ~N* The term (1 - S)/(m - ~.) in equation 5 is the proportion of the population that is fertile at steady state. This proportion is the ratio of the adult mortality rate divided by the rate of recruitment to the adult stage. When the proportion of fertile animals in the population equals this ratio, recruitment exactly balances mortality and the population stabilizes. �127 When p' is large, models predict that small changes in p' can produce large changes in the equilibrium density (Table 3, Fig. 4). This is important because by setting N' 0 in equation 5, we see that the population will go extinct whenever = 1 - e: s 1- S (6) m Thus, to prevent extinction, the proportion of fertile females in the population must exceed the ratio of the adult mortality rate divided by the maximum possible recruitment rate (i.e., recruitment at 0 density). If the proportion of fertile females is maintained at levels less than this ratio, then the number of adult deaths will exceed the number of recruits at all population densities and the population will eventually decline to O. (However, recall that such values of p' may not be obtainable in the post breeding case.) The steady state density is a nonlinear function of the proportion of infertile animals in the population (Fig. 4) because of the dependence of the recruitment rate on the steady state density, N'. That is, when N' is small, the per capita recruitment is high and a greater proportion of the population must be infertile to maintain steady state than when N' is large and recruitment is low. This dependence means that large reductions in steady state density require disproportionately large delivery rates (Fig. 2) and a disproportionately large fraction of infertile animals in the population (Fig. 4). Approach to steady State and Local stability Dynamics of the approach to equilibrium differ among control regimes (Fig. 5). Given values of m, S, and c providing local stability, approach to equilibrium predicted by the culling models is substantially more rapid than the approach predicted by lifetime contraceptive models (Fig. 5). Fertility control models predict that populations would approach steady state via damped oscillations around the equilibrium density. Models predict that populations will return to steady state following small perturbations if the maximum rate of recruitment is sufficiently small relative to adult survival and to the rate of contraception or culling (Table 4). Both contraception and culling exert a stabilizing effect on the steady state. However, given the same per capita rate of delivery of contraceptives .or culling, models predict that higher rates of recruitment are sustainable in locally stable populations treated by contraceptives than in populations treated by culling. Number of Animals Treated at Steady state In the subsequent section, it is important to remember that the treatment rate is an absolute rate (females treated) while the delivery rate is a per capita rate (females treated per female). Models show that the relationship between treatment rate and steady state density differs among control regimes (Table 5, Fig. 6). Pre-breeding models of culling and lifetime contraceptives predicted that more animals would need to be culled than would need to be treated to maintain any given steady state density less than carrying capacity (Ke) (Table 5, Fig. 6). This result follows logically from the observation that the proportion of the fertile females treated or culled annually (c) to achieve a given steady state (N') is the same for the culling and the �128 lifetime/before breeding model (Table 3). However, the number of fertile females in the population treated by long lived contraceptives will always be less than the number of fertile females in the culled population, and hence, the number treated at equilibrium will be less. This difference follows from the simple fact that all females are fertile when the population is regulated by culling (i.e., F = N°), while a small proportion is fertile when the population is regulated by fertility control (i.e., F = (1 - P')N°) for the population treated with lifetime contraceptives. Differences between treatment rqtes of the fixed duration model and the culling model in the before breeding case depend on life history characteristics of the animal (m, S, ~) and the equilibrium population size (Fig. 6). Treatment rates for contraception will be less than for culling as long as the steady state density falls within the range o m + S - mS' < N* < - 1 (7) ~ ( 1 - S' ) Thus, delivery of contraceptives before breeding will allow a lower treatment rate than culling when steady state density is low. In contrast, delivery of contraceptives after breeding will allow a lower treatment rate than culling when steady state density is high (Fig 6). Expressions defining the range over which treatment rates required by culling rates will exceed those required fertility control are quite cumbersome. However, based on these expressions, we can conclude that when contraceptives are delivered after breeding, the treatment rate for contraceptives will be less than that required by culling only when the steady state density is close to ecological carrying capacity. Interactions between Reproductive State and Survival It is plausible that survival of infertile animals might exceed survival of fertile ones because infertility eliminates energy demands of pregnancy and lactation. To examine this potential interaction, I define e as the proportional reduction in the mortality rate of infertile animals that results from contraception. Thus if e = 2, the mortality rate for infertile animals is half that of fertile ones. This allows calculation of the survival rate of infertile animals (Sr) as SI = S + e - 1 (8) e It follows from equation 8 that S < Sr < 1 for all e > 1. Substituting S1 for S where appropriate in the models above, deriving new characteristic polynomials in Sand Sr and solving for the population density at steady state shows that interactions between adult survival and reproductive condition will not affect the steady state density if effects of contraceptives persist for the lifetime of the animal. However, the proportion of the population that is infertile at steady state increases as e increases. For the lifetime duration, pre-breeding model we have e: = and for the ce 1 + Sc + ce - S - c lifetime, post-breeding model, (9) �129 See e: = (10) 1 + See - S In contrast, the fixed duration models predict that the steady state population will increase if infertility enhances survival of infertile animals. For example, when the duration of the contraceptive is 1 year T = 1) and delivery occurs before breeding we have N* = S + and in the after breeding = S + - S) (11) case, m - 1 ~ (1 ~e(l-e) ~ N* e m - 1 (i.e., eS(l ~ (e - S) (12) + e - eS - ce ) The first term of the right hand side of equations 11 and 12 is simply Ke, the carrying capacity. The second term is the effect of fertility control. As e increases, the magnitude of the reduction in the steady state density resulting from fertility control approaches O. DISCUSSION The models described here forego detail in favor of general insight (Levins 1966). Such insight, of course, has limitations (e.g., see the excellent discussion of Bomford 1990:16-21). In particular, I have ignored social structure, and many of the effects of fertility control are quite sensitive to changes in social interactions (Caughley et al 1993). I have ignored effects of immigration, which is clearly important in determining population behavior. However, bearing these caveats in mind, several general results emerge from the modeling exercise. Efficacy of Fertility Control By definition, agents that regulate population growth cause population density to return to a steady state whenever densities exceed or fall below ~hat steady state (Sinclair 1989). It follows that the efficacy of contraceptives as agents of population regulation must be judged by their ability to maintain a population at.an equilibrium that differs meaningfully from the equilibrium that would occur in the absence of control efforts. An agent is efficacious if can bring about such regulation, it not efficacious .Lf it cannot. Models suggest that timing of reproductive intervention has a strong effect on its efficacy. If fertility control agents are delivered while animals are pregnant, if those agents do not interfere with the success of pregnancy, and if offspring breed before agents are delivered again, then fertility control regimes may be unable to maintain populations at levels significantly below ecological carrying capacity regardless of treatment effort. This is the case because offspring produced by young of the year always escape treatment and reproduce at least once. When this is the case, Equation 5 shows that it is infeasible to use fertility control to regulate fecund, short lived species. Fertility control can not regulate such population because the minimum density that can maintained when all fertile females are treated approaches the ecological carrying capacity (Equation 5). �130 Given that the females of many species spend the greater part of their life pregnant and given that their offspring breed before their first birthday, there is a relatively brief interval during which delivery of contraceptives must occur to avoid these limitations. These results illustrate that the conclusion of Stenseth () that r-selected species are the most suited for regulation by fertility control depends on the understanding the breeding cycle and the timing of delivery. In the case of contraceptive delivered post-breeding, it will be true that r-selected species (i.e., large m, small S) will be the least likely to be controlled by reproductive intervention. Efficiency of Fertility Control vs Culling There are two ways to look at efficiency of population control techniques. The first view sees efficiency as the number of animals that must be treated annually per unit reduction in steady state density. The second view sees efficiency as the amount of time required to bring a population at carrying capacity to some new steady state < carrying capacity. In this section, I consider these two perspectives. Efficiency measured by treatment effort; Using the first view, we can compare efficiency of control regimes by comparing treatment rate curves (e.g., Table 5, Fig. 6). All models predict a humped relationship between the treatment rate (the number of animals treated or culled annually) and the equilibrium population size (Table 5, Fig. 6). These relationships mimic sustained yield curves (indeed, for the culling models, these relationship are such curves) and occur for the same reason. Whenever the steady state density is large (i.e. N·-->Ke), then the delivery or culling rate needed to maintain that density is small because per-capita recruitment is diminished by density. When the steady state population is small (i.e., N·-->O) then the culling or delivery rate (c) must be large to offset the greater recruitment rate, but nonetheless, there are not many animals to treat. Thus, because the number of animals treated is a function of both the delivery rate and the steady state density, there are usually two steady states that can be maintained with the same treatment rate, but there is a single steady state requiring the maximum rate of treatment (Table 5, Fig. 6). The exception to this occurs for postbreeding models for which treatment curves are truncated (Table 5, Fig. 6). Such truncations occur at densities where c = 1 and reflect the fact that there are values of steady state density that are not attainable using contraceptives delivered after breeding (see above, "Steady State Population Density") • Thus, if we define efficiency as the magnitude of the reduction in steady state density per animal treated or culled, then treatment with lifetime duration contraceptives can be substantially more efficient than culling as a means of regulating animal populations, particularly for longlived species. Moreover, treatment. with short duration agents delivered before breeding can be more efficient than culling if per capita rates of recruitment are high and the target density is low (i.e., equation 7, Fig. 6). Thus, the results offered here agree in part with those of Stenseth (1981) who concluded that the greatest opportunity for efficacy of fertility control was for r selected species who are short lived and fecund. However, these results add to those of Stenseth (1981) by illuminating the efficiency of long lived agents in regulating long-lived, K-selected species. Models illustrate that efficiency of fertility control (as measured by tretment effort) depends strongly on the duration of the effect of the contraceptives (i.e., r). Although the proportion of the population that must be infertile to achieve a given equilibrium is not affected by the duration of �131 the contraceptive, the number of animals that must be treated annually to maintain a given level of infertility in the population depends strongly on the duration of infertility. If effects of contraceptives are brief, then the preponderance of the population must be treated each year to maintain a significantly reduced equilibrium density. However, whenever the duration of the contraceptive exceeds 1 year, the proportion of the population that is infertile at equ~librium (i.e., pO) will be less than the proportion of the fertile females that must be treated annually (i.e., c). The magnitude of the difference between p' and c increases with increasing duration of efficacy and with increasing adult survival. For long lived species treated with lifetimeduration contraceptives, the proportion of the population that must be treated annually will be much smaller than the proportion of the population that is infertile at equilibrium. For example, presume that we wish to regulate a population of whitetailed deer ( ) at 50% of carrying capacity by treating them with contraceptives delivered prebreeding. Models predict tha.t maintaining such an .equilibrium requires that 94% of the females must be infertile. If the effects of contraceptives last one year, then 94% of the population would need In to be treated annually to maintain the population at the target density. marked contrast, if the effect of contraceptives last for the lifetime of the animal, only 4.7% of the population would have to be treated each year to maintain 94% infertility and an steady state at 50% of carrying capacity. The 20 fold difference in treatement effort requred to achieve the same objective illustrates fundamental importance of duration of the effect of contraceptives in determining efficiency of fertility con~rol as a population regulator, particularly for long lived species. Efficiency measured by time to steady state; If we define efficiency in terms of the time required to reduce a population from carrying c.apacity to a new equilibrium point, then it is clear than culling is more efficient than fertility control is. This is the case because the maximum rate of decline achievable by controlling fertility is simply the natural rate of adult mortality (i.e. A ~ 1 - S). In contrast, the rate of decline resulting from culling can be several fold greater than the natural mortality rate (i.e. A « 1 - S). Thus, when adult survival is high and when control efforts are initiated when density dependent recruitment is low, then effects of fertility control on total density may not be detectable for several years. In contrast, the effects of culling at the same per capita rate would be immediately expressed. Approach to steady State Differences in approach to steady state between contraceptive and culling models result because of the delayed effect of contraception on adult density, and hence, on recruitment. In contrast to animals that are culled, animals that are treated with contraceptives remain in the population and continue to depress recruitment via density dependent effects. Thus, while culling has instantaneous effect on adult density (and hence, on recruitment), the effect of contraception is delayed. The magnitude of this delay depends on the adult survival rate--the higher the rate of adult survival, the greater the delay. Thus, increasing adult survival rates amplifies the magnitude of oscillations in the approach to equilibrium predicted by the lifetime contraceptive models. �132 Implications for Population Management The ability to detect the response of a population to management is a prerequisite for adapting management actions to a changing environment. Because effects of culling are immediate while the effects of·fertility control can be delayed, adaptive feedback between the state of the population and the actions of population managers will often be stronger for culling than for fertility control. Thus, it is vital that managers not underestimate the consequences of failure to monitor the effect of fertility control on natural populations •.. These consequences loom large because the relationships between the decisions that a manager makes (i.e., delivery rate) and the responses of the population (i.e., steady state density, proportion infertile) are highly nonlinear. This non-linearity means that small errors in decisions can have large consequences. For example, assume that we deliver fixed effect contraceptives with a duration of 1 year to a population of white-tailed deer (m = 1.87, fi .018, S .9). Presume our target steady state density is 50% of carrying capacity (Ke)' If we treat 90% of the fertile females annually, we hit the target--the population settles at 50% of Ke' If we treat 85% of the fertile females, the population would substantially exceed the target, equilibrating at 68% of Ke' Treating 95% each year assures extinction. Given the imprecision inherent in most census techniques, it is unlikely that one could accurately estimate a delivery rate within ±5 percentage points. However, but this interval (90 ± 5%) produces outcomes ranging from overshoot to extinction. The potential for excessive treatment, and hence, the possibility of driving populations to extinction is particularly problematic for agents that have irreversible effects. Models illustrated that substantial reductions in equilibrium densities could be achieved only by maintaining a relatively small proportion of fertile animals in the population. As a result, the effective, breeding population may be a mere fraction of the total, steady state population. Because the breeding population will be substantially smaller than the total female population, it follows that populations regulated by controlling fertility are more susceptible to chance extinction than would be the case for populations regulated by culling, where the breeding population and the female population are the same. Thus, the same features that make lifetime agents efficient as .control agents also make them undesirable for use in regulating small populations. This is to say that while contraceptives can be viewed as non-lethal agents when their effects are evaluated at the level of the individual, they can be quite lethal at the population level, particularly when populations are small or when managers fail to closely track the effect of reproductive intervention on the population's level of fertility. = = Feasibility Using of Regulating Fertility populations Control Model results suggest that given the proper circumstances, fertility control can be a feasible and efficacious means of regulating population growth when compared with culling. This finding is based purely on biological considerations and ignores the economics of implementing fertility control regimes. For many species of vertebrates, culling is accomplished by hunters who volunteer their time and pay for the opportunity to do so. contraceptives, however, are usually delivered by paid professionals. Thus, �133 for many vertebrates, the cost fertility control is likely to exceed the cost of culling by large measure. Part of the cost of regulating populations using fertility control will be the expense of obtaining information. Regulating populations using fertility control will require substantial knowledge of the reproductive state of the population. Such knowledge is required to deliver fertility control agents in proper proportion to the size of the reproductive pool. If proportionate delivery is not properly achieved, the population will likely to increase to unacceptably high levels, or will be driven to extinction. There are many dimensions to the decision on how to best regulate wildlife populations, dimensions that include economics, ethics, politics, and culture. However, the models I offer suggest that the option to regulate populations by controlling fertility should not be rejected, out of hand, on the basis of biological efficacy alone. LITERATURE CITED Barclay, H. J. 1981. Population models on the release of chemosterilants for pest control. Journal of Applied Ecology. 18:679-695. Boone, J. L. and R. G. Wiegert. 1994. Modeling deer herd management: sterilization is a viable option Ecological Modelling 72:175-186 Bomford, M. 199. A role for fertility control in wildlife management? Bureau of Rural Resources Bulletin No.7, Department of Primary Industries and Energy, Canberra, Australia. Brush, C. C. and D. W. Ehrenfeld. 1991 •.Control of white-tailed deer in non-hunted reserves and urban fringe areas. Pages 59-66 in L. W. Adams and D. L. Leedy, eds. Wildlife Conservation in Metropolitan Environments. National Institute for Urban Wildlife, Columbia Maryland, USA. Caughley, G., R. Pech, and D. Grice. 1992. Effect of fertility control on a population's productivity. Wildlife Research 19:623-627. Char, B. W., K. o. Geddes, G. H. Gonnet, B. L. Leong, M. B. Monagan, S. M. Watt. 1991. Maple V Library Reference Manual. springer-Verlag, New York, New York, USA. 698 pp. Eagle, T. C., E. D. Plotka, R. A. Garrott, D. B. Siniff, and J. R. Tester. 1992. Efficacy of chemical contraception in feral mares. Wildlife Society Bulletin 20:211-216. Garrott, R. A. 1991. Feral horse fertility control potential and limitations. Wildlife Society Bulletin 19:52-58. Garrott, R. A., and D. B. Siniff. 1992. Limitations of male-oriented contraception for controlling feral horse populations. Journal Wildlife Management 56:456-464. Garrott, R. A., D. B. Siniff, J. R. Tester, T. C. Eagle, and E. D. Plotka. 1992. A comparison of contraceptive technologies for feral horse management. Wildlife Society Bulletin 20:318-326. Hoffman, R. A., and R. G. Wright. 1990. Fertility control in a non-native population of mountain goats. Northwest Science 64:1-6. Hone, J. 1992. Rate of increase and fertility control. Journal of Applied Ecology 29:695-698. Jewell, P. A. and S. Holt. 1981. Problems in management of locally abundant wild mammals. Academic Press. 360 pp Levins, R. 1966. The strategy of model building in population biology. American Scientist: 54:421-431 Kirkpatrick, J. F., J. W Turner, JR. 1985. Chemical fertility control and wildlife management. Bioscience. 35:485-491. �134 Knipling, E. F. and J. U. McGuire. 1972.· Potential role of sterilization for suppressing rat populations: a theoretical appraisal. United States Department of Agriculture, Agricultural Research Service Technical Bulletin No. 1455: 1-27. Leopold, A. S. 1963. A study of wildlife problems in national parks. Transactions of the North American Wildlife and Natural Resources Conference 28:28-45. McCullough, D. R. 1979. The George Reserve deer herd: population ecology of a K- selected species. University of Michigan Press, Ann Arbor. 271 pp. Plotka, E. D., D. N. Vevea, T. C. Eagle, J. R. Tester, and D. B. Siniff. 1992. Hormonal contraception of feral mares with silastic rods. Journal of Wildlife Diseases 28:255-262. Sinclair, A. R. E. 1989. Population regulation in animals. Pages 197-241 in J. M. Cherrett ed. Ecological Concepts: The Contribution of Ecology to an Understanding of the Natural World. Blackwell Scientific Publications, Oxford, U. K. stenseth, N. C. 1981. How to control pest species: application of models from the theory of island biogeography in formulating pest control strategies. Journal of Applied Ecology 18:773-794. Sturtevant, J. 1970. Pigeon control by chemosterilization: population model from laboratory results. Science 170:322-324. Turner, J. W, JR. I. K M. Liu, and J. F. Kirkpatrick. 1992. Remotely delivered immunocontraception in captive white-tailed deer. Journal Wildlife Management 56:154-157. wrigth, J. B. 1993. Rocky Mountain Divide: Selling and Saving the West. University of Texas Press, Austin Texas. 275 pp. �135 APPENDIX A Given the assumptions described in the text (see METHODS, Model Assumptions), the dynamics of a population of females unregulated by contraception or culling can be represented by a simple variation of the discrete logistic model, (13) where N is the density of females, S is the per capita adult survival rate, m is the maximum per capita rate of recruitment to the adult stage (females recruited/female/year), {3 is the slope of the relationship between recruitment and survival ({3= m/Km where Km is the population density at which recruitment = 0), and t is time (Table 1). The units of time must correspond to the interval between breeding pulses, which, for convenience, I will assume to be 1 year. Representing the dynamics of fertile animals in a population treated with contraceptives begins with the same model as above (14) except that Ft is the density of fertile females and N is the total female density (fertile + infertile). I now model 2 cases, the dynamics of which depend on the timing of delivery of the fertility control agent. In both cases, I aaeume that the effects of the contraceptive last for a fixed number of breeding seasons. Thus, I will refer to these as models of fixed duration contraceptives. Fixed duration. delivery before breeding; When contraceptives are delivered before breeding and after births (Fig. 1) then the number of fertile animals treated annually is (15) where c is the per capita delivery rate. The delivery rate is defined as the proportion of fertile females who are alive at the time of treatment who receive a fertility control agent. The number of fertile animals is reduced annually by the number of animals treated; thus, subtracting the right hand side of 15 from equation 14 and rearranging yields Ft+1 = Ft ( 1 -c ) (S + m - ~Nt) (16) However, when the effect of contraception is transient, animals that are treated become fertile again at a later time, thereby increasing the density of fertile females. To represent such transitions, I define T as the duration of efficacy of the contraceptive. I define ITt as the number of animals who are in their final year of infertility during the interval t-->t+l. I assume that a portion (i.e., c) of the infertile animals are retreated in year T and those that are not retreated become fertile again. Thus, the total treatment rate (Tt, in females/yr) becomes (17) and the dynamics t+l = Ft ( of the fertile 1 - c)' ( S + m - ~Nt) portion + I 1t of the population S ( 1 - c) • can be portrayed as (18) �136 Representing the infertile portion of the population requires additional stages in the model. These stages are needed to keep track of the animal's state of infertility; hence, the number of stages is set by the duration of efficacy of the contraceptive. For example, if inhibition of fertility persists for 2 years, there will be two infertile stages, one for the first year after delivery and another for the second year. The number of animals in their first year of infertility (L1t+1) is equivalent to the number of animals treated during the previous time step, Ii t.l = C ( 13 N) Ft S + m - + C S I Tt , while the number of animals in subsequent stages function of the adult survival rate, i.e, I I 2t+l 3t+1 I. ~ t+l I T t+l = Ii = I 2 t = Ii-1 = I T t t -1 t (19) of infertility is simply a S (20) S (21) S (22) S (23) where Lit is the density of infertile adults at time t, i years after treatment. Given these multiple stages, the total adult density at time t (Ne) is (24) Thus, equations 14,16-24 define the model delivery of fixed-duration contraceptives, for fertility control pre-breeding. assuming Fixed Duration. Deliyery Post-breeding; When contraceptives are delivered post-breeding (i.e., while animals are pregnant, Fig. 1) this model must be modified if two conditions hold. These conditions are (1) that the contraceptive does not affect the success of pregnancy and (2) that young of the year can breed before contraceptives are reapplied to the population. Under these circumstances, the current year's offspring will escape treatment because they are in utero when contraceptives are delivered. As a result, they will be able to breed at least once (i.e., at 6 months of age, Fig. 1) regardless of the intensity of delivery of contraceptives to adults. In the post-breeding case, the treatment rate is (25) subtracting the right hand side of equation 25 from equation 14 and following the same steps as above, I derive the following model for the post-breeding case; +1 = Ft S ( 1 -c ) + E; (m - 13Nt) +I T t S(l-c (26) (27) �137 (28) I r tTl = I S _ T 'tl Lifetime Duration Models; delivered before breeding, then (29) When lifetime duration contraceptives are (30) (31) and in the after breeding FtTl = r, S ( 1 -c case, ) + r, (m - ~Nt) (32) (33) and in both cases Note that these are special instances of the fixed effect models where r is effectively infinite (i.e., there is no return from the infertile to the fertile stage). Culling; I now construct 2 simple models of regulation by culling to compare with those representing fertility control. When animals are culled before breeding (Fig. 1), I assume that culling mortality adds to effect of natural mortality because culling is assumed to occur after all natural mortality has taken place (Fig. 1). When culling occurs before breeding the treatment rate is (35) where c is the number of females killed annually per fertile female alive at the time of culling. Note that in this case Nt = Ft because there are no infertile animals in the population. Thus, subtracting the right hand side of equation 37 from equation 14 gives the model for culling pre-breeding, (36) Unlike fertility control, culling affects adults and recruits regardless of when it is applied (i.e., before or after breeding). However, as with fertility control, it is necessary to develop a post-breeding model distinct from the pre-breeding case. This is necessary for 2 reasons. First, it is plausible that recruitment can respond in a compensatory fashion when culling occurs after breeding (Fig. 1). That is, the reduction in density that occurs after culling could potentially stimulate recruitment if juvenile mortality is compensatory rather than additive. Second, in contrast with equation 35, the number of animals culled after breeding (while animals are pregnant) is simply (37) �138 However, because breeding reduces next time step. e) to reduce the animals are pregnant when culling occurs, culling after the number of recruits as well as the number of adults at the Thus, subtracting eFt from equation 14 and adding a term (1effect of density on recruitment, I obtain (38) This Table is the culling 1. Symbols model, used post-breeding. in models Definition F Density of fertile, I Density adults of infertile, Total adult and fertility Variable Symbol N of culling female female adults (= Type State Variable State Variable female density control. F + State Variable I) steady state density females m s Variable Maximum per capita rate of recruitment to the adult stage Parameter Slope of relationship between per capita recruitment rate at time t + 1 and adult female density at time t Parameter Adult Parameter survival rate T Duration of efficacy contraceptives e Per capita rate of culling delivery of contraceptives Parameter of Derived Quantity is Derived quantity Number of females treated with contraceptives or culled annually to maintain steady state at N' Derived quantity Per capita rate of treatment with contraceptives or culling required to maintain steady state at N' Derived quantity or Proportion of population that infertile at steady state c Decision of adult �Table 2. Models of population regulation by fertility control and culling. derivation, see Appendix A. For definitions of symbols, see Table 1. For detailed Timing of Treatment Before Breeding Regime Fertility control, fixed effects F'+1 - F,(1 -c )(S PN,) + In - PI:') 11'+1 - F,c (S + In - After Breeding I-r,S(l - c) + + F'+I - F,S(1 I-r,Se -c) + II - F, Se '+1 F,(m + Itt+l - I-r-lt S 1-rt+l· - 1-r-l S t + - Eli Ft( 1 -e ) (S It+l - F,e (S + m - N, - FI ' '+ F'+l - Ft( 1 - c)(S m - PN,) P N,) + +.~ L..., I.I, i-I + F'+l- FtS(I-e I,S 1'+1 - m - )+Ft(m - PNt) F, S e +I,S N t - F , + I, N, - F, + I, " Culling - c) -r i-I Ft+ 1 I-r,S(1 , 12t+l - I It· S Nt - F, + It Se 12'+1 - I I, S -r Fertility control, lifetime effects PN,l PFt) Ft+l - F,O - e)[S + m - P(1 - e)F,l ..••. ~ �Table 3. Predicted steady state behavior of populations regulated by fertility control and by culling. Variables are adult survival rate (S), maximum per capita recruitment rate (m), change in recruitment rate per unit increase in density ({3), per capita rate of culling or contraception (c), and duration of efficacy of contraceptives (T). In the absence of external control (i.e., c = 0) the steady state density is K, = (S + m - 1)/(3. Population Density (N) Control Regime Proportion of Population Infertile (p.) o Culling before breeding K If! Culling after breeding Lifetime contraceptives delivered before breeding c Kif! Lifetime contraceptives delivered after breeding Fixed duration contraceptives before breeding o K _; c2 (m + S - 1) + c (2 - m - S) . If! P (1 _ 2c + c2) ~ If! Kif! _ - P c( 1 - S't) P( 1 - c) _ cS( 1 - S't) K If! P(l-cS't) -r cS + cS cS K _ cS delivered Fixed duration contraceptives delivered after breeding 1 - S 1- S c (1 - S't) 1 - S - cS't + cS cS(1 - S't) 1 - S -c S" + cS ....•. B �141 Table 4. Conditions for local stability of steady state of culling and lifetime fertility control models. If the maximum rate of recruitment to the adult stage (m) is within the bounds shown, then small deviations from the steady state will decay with time. Variables are adult survival rate (S), change in recruitment rate per unit increase in density «(3),and per capita rate of culling or contraception (c). Bounds for Local Stability Model No external regulation (i.e., c I-S<m<3-S :;:;;0) Culling before breeding Culling after breeding I-S(I-c) <m< 1 - c 3 - S( 1 - c) 1- c 1 - S( 1 - c) < m < 3 - S(1 - c) 1 C 1 - c _i Lifetime contraceptives delivered before breeding 1 - S( 1 - c) ----~--~ < 1- c Lifetime contraceptives delivered after breeding (cS2 - S2 - cS + 2S + 3)(cS - S + 1) 1 - S (1 - c) < m < ~:'___---:.___--~~--,----_;_ (1 - S)( 1 + S - cS) (cS2 - S2 - cS + 2S + 3HcS - S + 1) + S - cS) m < ~~~~--~------~~------~ (1 - S) (1 - c) (1 �..•. f:5 Table 5. Effort required to stabilize populations at steady state = N· using fertility control and culling. Variables are proportion of population infertile' (P), adult survival rate (S), maximum per capita recruitment rate (m), change in recruitment rate per unit increase in density «(3), per capita rate of culling or contraception (c), and duration of efficacy of contraceptives (r). Delivery Rate (c', female/female) Control Regime Culling before breeding Culling after breeding Treatment rate (T", females) m - pN· + S -' 1 m -pN· + S ! 2pN· + Vm2 + 2 N·c·(m - pN· m - pN· Lifetime contraceptives delivered after breeding m + S - pN· Nr c • (1 - p.)( m - S - 1 + m - pN· + S); s:« 2mS + S2, - 4PN* pN· Lifetime contraceptives delivered before breeding + P N· + S) S N·c·( - 1 1 - pC<) S Fixed duration contraceptives delivered before breeding m - pN· m - pN· Fixed duration contraceptives delivered after breeding m - pN· m S" -pN·S'" + + S- 1 N*C*[(l - P*)(m S - S'" + S - 1 - pN· + c·S'" S) + 1 N*C*[(1 - P*) + S - S'" + . 1 + c·S"'(l + C·S",-l + c" + '(I - S) c" S - c •.S'" - S - S) 1 1 - S - 2c·S'" I If c· is calculated based a specifie_d target density (N) and model parameters (m, f3:-S),Htfien- P·can be calculated by substituting e fore-in expressions for p* in Table 3. This provides all of the information needed for calculating 7!. �143 Breeding Births Breeding Pre-breeding treatment Post-breeding treatment t t+1 I' Mortality Recrui ment Figure 1. Assumptions on timing of events in annual cycle of a population regulated by fertility control and by culling. Culling or delivery of contraceptives is assumed to occur at time marked "treatment". �144 Regime = Post - Breeding - 120, iC ~. ;?;- 100 ;----- en c Q) 0 --- 80 c 0 ~ 60 '"5 o, &. 40 E ::::s .£:: ,Q ·s CT w 20 Oi 0.00 0.25 0.50 0.75 1.00 Delivery Rate (c, female/female/year) Method Culling - - - - Fixed - - - - - - Ufetime Regime = Pre - Breeding -- 120 iC Z 100 .en ~ c; Q) ,0 -- 80 <, <, c 0 ~ 60 <, <, -, \ \ "'5 c.. &. \ \ 40 \ \ \ E ::::s '£:: ,.Q 20 \ \ 'S \ CT W \ 0 0.00 0.25 0.50 0.75 1.00 Delivery Rate (c, female/female/year) Method Culling - ~ - - Fixed Ufetime Figure 2. Toe steady state population density declines as the delivery or culling rate increases. Delivery or culling rate is defined as the number of females treated per female alive at the time of treatment. Parameter values used in this example are m = 1.87, S = .9, ~ = .018. �145 .-.. .0.. 100 .•.... - - I- Q) c: c: a .•.... ro -----~-~ ____ - -- . Q) 80 ,..,. ./ ./ /' ./ ./ 60 ./ ./ ./ ./ 0. - 0.. .' 40 I .: - "..,..., 20 .y .' /' /' /' ./ .Y .~ './ V I- ~~ Q) 0.. .: . . ./ /' a .•.... c: Q) o ./ ./ :J a , O~--------~--------~--------~---------. 0.75 0.50 0.25 Delivery Rate (c, female/female/year) 0.00 Method - ~ - Fixed/After - - Lifetime/After Figure 3. The proportion of the population depends on the delivery rate (C). Parameter = 1.87, S = .9, fi = .018. 1.00 - - - - - - Fixed/Before Lifetime/Before that is infertile at equilibrium values used in this example are m �146 100 --Z ..__.. iC C c Q) ·00 75 0 c 0 15 :J Q_ 50 s: E :J ·c .0 25 ·s 0W o 25 50 75 100 Percentage of Population Infertile (P*) Figure 4. The steady state population density depends on the proportion of the population infertile in an identical fashion for all fertility control models. Parameter values used in this example are m = 1.87, S = .9, fi = .018. �Reglme- CulllnWBefore Regime 10011 i l 75 .~ ~ 50 c3 8 50 , ,, a. I I I ~ I I 25 ,, :f2 I I I 25 ,, I I , ,, o ,,' ,, ,, , , I I o.J "---",, " 40 20 0 60 20 40 60 40 60 Years Years Regime = Ufetime I After Regime = Ufetime I Before 100 100 75 z- ~ i l , _,------- i 0 ~ CulllrzyAfter 100 75 ~ ~ ~ 1:1 75 .~ 0 50 .5 50 ,/ L .12 ~ 25 I 25 I I I I I , I o ' 0 o ' 20 40 Years 60 0 20 Years Figure 5. Simulations of approach to steady state using lifetime fertility control Parameter values used in this example are m = 1.87,· S = .9, f3 = .018. models. and culling ..• ~ -...j �148 "Regime:;::Post - Breeding - 75- __ - -... !>.. /' ~ ! E /' " \ /' \ /' /' \ /' \ /' 50 . ...~........ . \ .JE \ \ \ \ \ t: 0) ts 0: 25- I \ C 0) E \ \ ro lE F \ . ~ 0" 0 20 40 60 80 100 120 Equilibnum Population Density (N*) Method • Culling ----Fixed Ufetime Regime = Pre - Breeding - - 75~ !>.. .......... CI) 0) "'ffi E /'/' 50~ 0: - .. 25-' I . 0) E ro lE .F I 'O! \ \ ..•.•..... . . . ... ....". \ • .// /' \ \ \ \ \ \ \ \ \ // /' , i I c:: " /'/' t: 0). .....•••... /' ~ -ro --- /" /' //' /' /' // /' /' 0 I • I . 20 40 60 80 \ 100 120 Equilibrium Population Density (N*) Method • • Culling - - - - Fixed Figure 6. Number of animals that must be treated maintain a given steady state density. Parameter are m = 1.87, S = .9, ~ = .018. Ufetime or culled annually to values used in this example �149 Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS Title: Period Author: Heart Rate as a Potential Indicator Sheep Exposed to Human Disturbance. Covered: July REPORT of Stress: Application to Bighorn 1, 1995 - June 30, 1996. M. A. Wild and D. L. Baker. Personnel: P. E. Bleicher, R. B. Heath, M. Painter, Ritchie, J. L. Schaefer, E. Wheeler. D. L. Piermattei, J. ABSTRACT To better understand the effects of disturbances on bighorn sheep, a reliable, longterm, noninvasive, quantitative indicator of stress is required. We proposed using remote monitoring of heart rate to meet this need and began performing experiments to investigate the correlation of heart rate and serum cortisol in captive bighorn sheep. We surgically implanted heart rate transmitters in 10 additional bighorn sheep. All domestic goats and bighorn sheep with transmitter implants remained healthy. Transmitters failed in the two goats about 12 mo after implantation. Transmitters failed in five bighorn sheep. Three failures occurred 3-4.5 mo after implantation and two occurred 13-14 mo after implantation. Normal battery demise was the likely cause of failure in transmitters failing ~12 mo. Earlier failures were likely associated with battery or transmitter failure, possibly due to fluid leakage into the transmitter. Transmitters in 10 bighorn sheep have functioned normally for 8-14 mo. A calibration study indicated our data collection system was a reliable indicator of true heart rate in bighorn sheep. We began investigations into the line of sight distance range of the signal from the transmitters. Initial results suggest that althQugh variation may occur with the individual, body position, or replication, overall, the" data collection system receives usable signals up to about 800 m line of sight from the transmit'Fer when the 'animal is sta.nding. Distances for collection of transmitter signals appeared to be slightly reduced when the animal was bedded. These are promising findings and 'suggest good applicability of this system to animals under free-ranging conditions. We performed graded stressor experiments to determine the correlation between heart rate and serum cortisol levels in 15 captive bighorn sheep.' Animals performed well and experimental techniques were successful. We are currently analyzing data from this experiment. Additionally, we performed the first two of four 24 hr collections to assess the influence of circadian and seasonal cycles on heart rate and serum cortisol levels. These experiments all contribute to the evaluation of the utility of heart rate transmitters as a devise to monitor disturbance imposed by human activity on bighorn sheep. ��151 HEART RATE AS A POTENTIAL INDICATOR OF STRESS: APPLICATION TO BIGHORN SHEEP EXPOSED TO HUMAN DISTURBANCE Margaret A. Wild P. Develop a technique to monitor wildlife viewing areas. N. and Dan L. Baker OBJECTIVES disturbance to bighorn sheep from humans in SEGMENT OBJECTIVES 1. Develop a safe, reliable, and unobtrusive system to remotely rate in bighorn sheep over an extended period (~ 1 year). 2. To identify applicability, including an automated data acquisition system monitor heart accuracy, range, and ease of use, of for collecting heart rate data. METHODS AND MATERIALS We generally followed methods reported in the Program Narrative for this study (Wild and Baker 1995); however, we slightly modified our approach to assessing signal strength of transmitters and applying graded stressors to bighorn sheep. We·determined line of sight range of the signal at nine distances (20, 100, 200, 300, 400, 500, 600, 700, and 800 m) at 2 mo intervals. For the initial sampling, data were collected with bighorn sheep standing facing toward, away from, and at right angles to (left and·right) the receiver and laying down at a right angle (right side) to the receiver. Subsequent collections were performed using two replication with bighorn sheep standing and laying down at a right angle (right side) to the receiver. Heart rate data and blood samples were collected from bighorn sheep exposed to graded stressor as described in the Program Narrative with the following modifications. Planning discussions and bighorn sheep training sessions suggested that persons collecting blood samples should be visible to the bighorn sheep rather than visually isolated. Bighorn sheep appeared more calm when they could see the individual adjacent to them rather than only hear and smell them. Technicians also found this arrangement to be important in order to avoid tangling of the blood collection tubing. Pilot work during bighorn sheep training also suggested that modifications be made to the type and duration of stressors used. All stressors were applied for 3 min to assure that bighorn sheep would see the stressor and respond with the desired change in heart rate. The 3 min collection period also assured that the datalogger compiled heart rate data during several collection periods from each bighorn sheep during the stress. Stressors used were also modified. For the mild stress a person dressed uniquely would appear and walk in front of the holding stalls with a minimal to moderate amount of noise and extra movements. The same general procedure was followed for application of the medium stress but activity and noise were increased (increased arm waving, waving a 0.5 x 1 m nylon sheet, jumping). For the high stressor we used a motorcycle driving in front of the holding stalls. �152 RESULTS ~urgical Implantation of Heart AND DISCUSSION Rate Transmitters in Bighorn Sheep Heart rate transmitters were implanted successfully in 10 additional bighorn ewes. Body weights of the ewes ranged from 62-82 kg. Although we initially questioned whether yearling ewes possessed adequate body size and muscle mass for successful implantation of the transmitters, we found that the two yearlings implanted performed well. The surgical procedure was as previously described. In addition, we added one far-far near-near suture in the muscle split at the cranial end of the transmitter. We hoped that this would decrease the likelihood of the transmitter migrating cranially under the scapular cartilage. This technique appears to have helped because transmitter migration is not visible in any of the ewes implanted in this manner. No post-surgical complications were observed. We attribute the success procedure to our developed technique and strict aseptic technique. Transmitter Function in Domestic Goats and Bighorn of the Sheep Heart rate transmitters implanted in two domestic goats in November 1994 failed in November 1995. Batteries apparently failed after· about 12 months of use. This was shorter than the initially projected 20 mo likely because the goats' heart rate was higher than expected. Battery life depends, in part, on the number of heart beats and subsequent signals emitted by the transmit.ter. The goats' heart rate was generally about 110 bpm instead of the 60 bpm used in calculating expected battery life. We anticipated that transmitters would continue to function longer in bighorn sheep because resting heart rate is generally 50-80 bpm. On necropsy, goats were in excellent body condition. Transmitters and leads were surrounded by, and anchored in place by, a thin layer of fibrous tissue. No other tissue reaction was apparent. Both transmitters had migrated cranially from the initial site of implantation, but were currently anchored in place by fibrous tissue. Unlike the transmitter recovered from the first goat euthanized (February 1995), the waxy covering over the transmitters was intact. These findings support clinical findings that transmitters produce minimal tissue reaction. Transmitters are simply walled off by the body without impairment to the animal. Heart rate transmitters in three bighorn sheep failed 3-4.5 months after implantation. Cause of failure was not apparent; however, because failure was characterized by inability to detect the signal, malfunction of the battery or transmitter itself was suspected. In contrast, if the mortality signal had occurred, malfunction would likely have been due to lead breakage. Heart rate transmitters failed in two additional bighorn sheep at 13 mo and 14 mo postimplantation. These failures were attributed to normal battery expiration; however, batteries in these two transmitters may have failed earlier than expected due to occurrence of some spurious signals associated with muscle movement that contributed to battery discharge. Transmitters continued to function normally in 10 bighorn sheep. These transmitters have functioned normally for·8-14 mo. All bighorn sheep remained clinically healthy during the year. �153 Calibration Experiment We assessed the accuracy system through comparison electrocardiograph (ECG) counting the number of R collection period. This beat intervals collected collection system. of heart rate determination by the data collection with the "standard" of a hard-wired output. The "true" heart rate was determined by waves on the ECG strip output in a one minute was compared to the harmonic mean of output from 8 during the same one minute period using the data Results suggest that the data collection system was remarkably accurate in this application (Fig. 1). Although the r2 value has not yet been determined, the correlation between the two methods appears very strong. In fact, the small amount of discrepancy between the methods may actually be due to an inaccurate count off the strip ECG (assumed "true") due to interference associated with animal movement. This suggests that our data collection system should be a reliable indicator of true heart rate in bighorn sheep. Signal Strength Initial collections indicated that body position of the animal affected signal strength received. Based on this finding, we standardized body position of the animals in subsequent collections to minimize variation. Data were collected in September and November 1995 and January, March, and May 1996. Data collections will continue at 2 mo intervals until transmitters fail. Analysis of these data are not complete; however, initial evaluations suggest that although variation may occur with the individual, body position, or replication, overall, the data collection system receives usable signals up to about 800 m line of sight from the transmitter when the animal is standing. Unfortunately, line of sight evaluation at greater distances was not possible due to topography. changes at our study site. Signal strength appeared to decline with distance, at least initially; however, evaluation of signal strength at distances >400 m was difficult. Figure 2 illustrates an example of changes in signal strength with distance. Although some signals were collected from bedded animals at 800 m, distances for collection of transmitter signals appeared to be reduced when the animal was bedded. These are promising findings and suggest good applicability of this system to animals under free-ranging conditions. We found that the frequency which corresponded to maximum signal strength changed (drifted) from the original in three of the five transmitters. Maximum change noted was 0.006 MHz. This finding underscored the importance of checking a wide range of frequencies to obtain the optimal signal and therefore receive maximum distance of the signal. Graded Stressor Experiment We collected continuous heart rate data for ~5 min prior to and during the 150 min sampling period. Heart rate data has not yet been analyzed; however, heart rate during the control period was generally about 40-80 bpm, while during mild, moderate, and high stress heart rates peaked at about 80-150, 110-220, and 160-260, respectively. Heart rate appeared to return to near baseline rate within minutes after the stressor was removed. In general, successful. bighorn sheep performed well and experimental techniques were Weather was the primary limiting factor for sample collection. �154 EKG 200 190 180 170 160 150 t::, 140 , o t::, 130 o 0 o *o • * o * 120 110 100 90 80 o 70 5iI. cl t::, 60 * cJ'~ 50 0 50 60 70 BO 100 90 110 120 130 140 150 160 170 lBO HBPM ANIO *** 309 o 0 0 600 t::, t::, t::, 630 0 0 0 660 • • • 730 Figure 1. Calibration line of "true" hea:rt rate as determined by ECG (EKG) compared to the harmonic mean of data collected using the automated data co11ection'system from heart rate transmitters in bighorn sheep (n=5). �155 SIGNAL 240 I 230 /::,. 220 ~ 210 ~ B 200 ~ ~ 190 : 180 ~ /::,. 0 /::,. 170 ~ * i 0 160 • 150 * 0 140 ~ • 130 /::,. /::,. 0 ~ 120 /::,. § 110 ~ 0 100 8 • 90 0 /::,. 80 ~ I 70 /::,. 60 ~ 50 * ~ ~ i !• 8 • ~ ~ •• /::,. 0 .~ /::,. /::,. I El e ~ • €l ~ /::,. 0 0 I • 0 (:j • • ~ * ~ • ~ ~ ~ 700 800 j 40 0 100 300 200 400 500 600 OIST ANIO *** 309 o 0 0 600 /::,. /::,. /::,. 630 000 660 ••• 730 Figure 2. Signal strength relative to distance of receiver-data logger from transmitter of 5 bighorn sheep implanted with heart rate transmitters. �156 We were unable to consistently collect blood samples at the desired intervals when temperatures fell below about -8 C. We collected complete data sets from 12 bighorn sheep. Transmitters failed on the three remaining bighorn sheep during the trial period (two transmitters failed after the mild stress trial and one after the moderate stress trial). We are currently analyzing data to determine rate and serum cortisol levels. Circadian Cycle the correlation between the heart Experiment To study the circadian cycle of heart rate and serum cortisol levels we collected continuous heart rate data and blood samples every 2 hr from six bighorn sheep for a 24 hr period on 22-23 March and 18-19 June 1996. Blood samples were collected from unrestrained bighorn sheep and all except one (L894 time 0) were collected within 6 min of the time the technician appeared. Given the rapid collection of blood samples, cortisol levels should be near baseline and not altered from handling the bighorn sheep. This experiment will be repeated at the winter solstice and vernal equinox. When collections are complete, we will analyzed data to study seasonal cycles of heart rate and serum cortisol. LITERATURE CITED Wild, M. A. and D. L. Baker. 1995. Heart rate as a potential indicator stress: application to bighorn sheep exposed to human disturbance. Colorado Div. Wildl. Res. Rep., Jul 1994 - Jun 1995, Fort Collins. of � https://cpw.cvlcollections.org/files/original/eff6656c8b102e2baf69b9aae43c9043.pdf 4904d80c29dbfb29db1ec5a018b70fb2 PDF Text Text 157 Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS State of Project Colorado No. ~W~-~1~5~3~-~R~-~9~ Work Plan _ Author: Mammals Research Multispecies No. Job No. Period REPORT Inyestigations Animal and Pen Support Facilities for Mammals Covered: July 1, 1995 - June 30, 1996. M. A. Wild Personnel: Research and J. L. Schaefer. P. E. Bleicher, A. L. Case, L. A. Wallick. ABSTRACT The Colorado Division of Wildlife's Foothills Wildlife Research Facility (FWRF) maintained captive animals (up to 139 wild ungulates of 5 species and two domestic ungulates) and facilities supporting six major research projects. In fall 1995, we recruited 19 individuals into our captive herd. During the year, reductions in animal numbers were due to natural mortalities (18 adults and 2 neonates) and death and euthanasia associated with approved study protocols (16 animals). In FY1996, relatively high mortality rates occurred in captive mule deer due primarily to chronic wasting disease (CWO) (n=5) and failure to thrive syndrome (n=3). Given the likely start of a CWO epizootic in captive mule deer at FWRF, we have resolved to use the mule deer herd as an opportunity to study the epizootiology and diagnosis of naturally occurring CWO. In a retrospective study, we examined body weight dynamics of captive ungulates by compiling body weight data from the last several years on cohorts of femal~ mule deer, pronghorn, elk, and bighorn sheep. Examination of these data demonstrated informative trends in body weight related to growth, pregnancy status, and seasonal effects. Over the past 5 years, FWRF operation has emphasized economy arid utility' in function in addition to improvements in animal welfare. The high quality of animal care is in part reflected in the results of animal welfare inspections. FWRF was inspected by USDA APHIS in March 1996 and found to be in compliance with federal animal welfare standards. Economy and utility of FWRF function have increased in at least three areas. First, the use of volunteer workers has increased three-fold since 1993. Well trained volunteers contributed 584.5 man hours in FY1996. Second, a conservation oriented approach to the use of utilities and services for FWRF has lead to a reduction in resource use relative to the size of the facility over the past 5 years. And finally, as older portions of the facility are repaired and replaced, the need for unscheduled daily repairs appears to be decreasing; however, numerous maintenance and improvement projects were performed again in FY1996 to increase usefulness, efficiency, quality, and/or safety of facilities at FWRF. ��159 ANIMAL Margaret AND SUPPORT FACILITIES MAMMALS RESEARCH A. Wild and Jenelle FOR L. Schaefer P. N. OBJECTIVES 1. To provide and maintain captive wildlife populations and facilities supporting CDOW's Terrestrial Wildlife Research Program, as well as programs of CDOW cooperators. 2. To develop improved animal and facility management practices provide maximum research opportunities at minimum cost. 3. To enhance needs. facilities to serve a growing SEGMENT and modify animal diversity that will of anticipated research OBJECTIVES 1. Maintain research 2. Coordinate all animal activities. 3. Provide or oversee maintenance for up to 20 elk, 40 mountain sheep, 25 pronghorn, 45 mule deer, and 11 white-tailed deer under applicable federal and institutional animal welfare regulations and in suitable health for use in research experiments. 4. Conduct management experiments to increase efficiency and efficacy feeding, husbandry, and maintenance activities related to research facility operations. 5. Follow a conservation-oriented approach for providing services to operate research facilities. 6. Follow Standard Operating facility records. rearing, and holding training, Procedures METHODS facilities. maintenance, in maintaining and research utilities detailed of and animal and AND MATERIALS Over the past 5 years, FWRF operation has been generally guided by the objectives outlined in the 1990 Program Narrative (Miller 1990). Emphasis has been placed on economy and utility in FWRF function. These considerations have been concurrent with increased emphasis on quality of animal care and animal welfare. Animal care and facility maintenance standard operating procedures (SOP'S) were developed during FY1993 and have been followed for routine procedures since that time. Given this general guidance, and the direction required to meet upcoming terrestrial research needs, we performed the following tasks: �160 Animal Maintenance General: Again this year, routine feeding and caretaking of research animals, including health observations, training, weighing, and clean-up, was performed primarily by well trained work-study and temporary employees, as well as volunteers. FWRF was inspected by USDA APHIS for compliance with federal animal welfare regulations on 14 March 1996. In 1995, eight pronghorn fawns were hand-raised at FWRF. Five additional pronghorn fawns were seized by CDOW, hand-raised, and received by FWRF after weaning. Eight viable lambs were born to captive bighorn ewes in 1995. In 1996, no new animals were added to the captive herd. Nutritional Maintenance Feeding protocols: Feeding protocols were as previously described (Miller 1990, Wild et al. 1992, Wild 1993, Wild and Graffam 1994); however, we attempted to improve the mule deer ration due to maintenance of suboptimal body condition in that species despite ad libitum feeding. As an alternative to the standard feeding protocol of alfalfa hay and high energy pelleted ration (Baker and Hobbs 1985), we began clinical evaluation of partial supplementation with a pelleted browser maintenance ration (PMI Feeds, St. Louis, MO 63144). In July, mule deer in two of four pastures began receiving 1 kg/head/day of the browser ration in addition to the standard diet. In April, mule deer in the remaining two pastures began receiving this diet as well. Body weight data collected over the last several years in cohorts of animals from each species were graphed to examine seasonal trends and performance of animals over time. To examine body weight dynamics of captive female ungulates, we graphed the body weights of the 1992 cohort of mule deer, the 1991 cohort of pronghorn, the 1986 cohort of elk, and bighorn sheep ~4 years of age as of 1993. Bottle-raising neonates: Eight pronghorn fawns were hand-raised in the summer of 1995. We began weaning the fawns from ad libitum evaporated milk feedings starting at about 8 weeks of age. Fawns were weaned by about 16 weeks of age (30 September 1995). Health Maintenance General: We continued to monitor animal health using FWRF SOP's. Animal health care was provided as required and as mandated by the preventive medicine program and chronic wasting disease protocols. Chronic Wasting Disease: We followed protocols for the preventive medicine program (Wild 1995) and management of CWO (Wild and Graffam 1994). All animals at FWRF were monitored closely for clinical signs of CWO. Tissues from all mortalities occurring at FWRF were examined for evidence of infection with CWO. Facility/Maintenance/Repairs/lmprovements A variety of scheduled and unscheduled maintenance and repair activities were necessary to support facility operation and ongoing research programs. We worked toward a conservation-oriented approach for facility care by �161 undertaking preventive maintenance projects, and performing high-quality new construction and repairs to existing facilities. Facility repair and construction projects were prioritized based on animal welfare concerns and anticipated research needs. Research projects Facility operations offered support for pilot studies and for research projects conducted by CDOW personnel and other collaborators that were initiated, conducted, or continued using FWRF animals and facilities throughout the year. Educational Contributions Facility tours and educational lectures were provided to school, university, and professional groups visiting FWRF. We emphasized the importance of maintaining captive wildlife for performing controlled experiments and the contributions made by research projects performed at FWRF. FWRF animals and facilities were also used occasionally for hands-on training for professional groups. RESULTS Animal AND DISCUSSION Maintenance General: Use of volunteer workers at FWRF has increased three-fold since 1993. When these workers were carefully selected and trained, their contribution was remarkable. In FY1996, a record high 584.5 hr were contributed by 14 volunteers. These volunteers performed primarily caretaker tas~s and also assisted in weighing and collecting samples from animals. Contributions by volunteers represented a savings to FWRF of about 0.3 TFTE and $5439 (vs. cost of temporary employees). One to two YCC employees per Bummer have also contributed greatly to animal care and facility maintenance at a limited cost. The animal welfare inspection by USDA APHIS revealed all items examined to be in compliance. This finding highlights the high standards of animal care provided by FWRF employees and volunteers. At the close of FY1996, FWRF maintained 19 elk, 22 bighorn sheep, 11 whitetailed deer, 31 mule deer, and 22 pronghorn. During the year we recruited 8 bighorn sheep and 11 pronghorn into our captive herd. Twenty natural moralities occurred (Table 1). An additional. 14 bighorn sheep and 2 domestic goats died or were euthanatized as part of approved study protocols. The US Fish and Wildlife Service provided financial support to maintain 25 mule deer at FWRF. The USDA Animal Damage Control provided financial support for 11 white-tailed deer at FWRF. Nutritional Maintenance Feeding protocols: Individuals in all species maintained reasonable body condition on available diets. Subjective evaluation of clinical. health and body condition suggested that supplementation of mule deer diet with browser �162 Table 1. Summary Species Bighorn Mule of mortalities Animal sheep deer L93 Laa Ea3 L395 Q95 La7 Ma95 Ma7 A94 L794 L95 E295 C295 C294 M92 E994 L92 ID in hoof stock at FWRF during Age (yrs) 2 7 12 0.5 0.5 a 0.5 a 1 1 0.5 0.5 0.5 1 3 1 3 Cause FY 1996. of Death Brain abscessa Epizootic hemorrhagic disease Lumpy jaw; old age changesa Pasteurella pilot study Pasteurella pilot study Pasteurella vaccine study Pasteurella vaccine study Pasteurella vaccine study Pasteurella vaccine study Pasteurella vaccine study Pasteurella vaccine studya Pasteurella vaccine studya Pasteurella vaccine studya Pasteurella vaccine studya Pasteurella vaccine studya Pasteurella vaccine studya Pasteurella vaccine studya E91 R693 Q94 4 2 1 Undetermined neurologic disease Trauma Euthanatized due to aggression (primarily intraspecific P93 Aa93 Eb93 Ab93 S90 U93 Zb93 A91 193 T93 2 2 2 2 5 2 2 4 2 2 Failure Bb95 NK91 Nb95 231 0.3 4 0.3 10 Bk93 W93 2 2 aggression) a Pronghorn Domestic goats aEuthanatized to thrivea Pneumon La" Chronic Chronic Chronic Failure Trauma Chronic Chronic Failure Wasting Disease Wasting Diseasea Wasting Diseasea to thrive-chronic bloata Wasting Diseasea Wasting Diseasea to thrivea Enterocolitisa Epizootic Hemorrhagic Disease Enteritisa Pneumonia secondary to lumpy jawa Euthanized-Transmitter Euthanized-Transmitter failurea failurea �163 maintenance pellets may be beneficial; however, confounding effects of age, sex, and occurrence of disease (chronic wasting disease and failure to thrive syndrome) preclude analysis. Of the mule deer in two pastures initially supplemented with browser diet, the intractable nature of male mule deer in one pasture prevented collection of body weight data. Body weight data from mule deer in the second pasture are presented in Fig. 1. Although body weights increased after the initiation of browser supplementation in July 1995, this trend likely reflects weight increases associated with age. Examination of body weight data from cohorts of animals at FWRF revealed some interesting information. A cohort of nonpregnant mule deer exhibited marked weight gain during the second year of life. Thereafter, weight trends suggest that the. deer exhibited seasonal fluctuations in weight, with a peak in late autumn and nadir in mid-spring (Fig. 2). These fluctuations occurred despite ad libitum feeding year round. Nonpregnant pronghorn showed similar trends; however, minimum body weights occurred later in the year (during early summer) than in mule deer (Fig. 3). Weight trends suggest that when does were pregnant (parturition in June 1993 and 1995) they gained weight late in gestation after an apparent weight loss during the second trimester. Because fawns were pulled from does at about 24 hr of age, body weight trends do not reflect the usual lactation period. Elk exhibited marked weight gain through the first 2 years of life. Weight fluctuations in 1989-1990 reflect elk response to involvement in experiments at the Maybell facility. Six of eight elk gave birth and lactated in 1993. Despite limit feeding of the elk since 1992, they continued to maintain or slightly improve in body condition over time except during the period of calving and lactation (Fig. 4). Body weights of elk that developed chronic wasting disease are isolated for review as well (Fig. 4). The group of bighorn sheep were not as homogeneous as other species' cohorts. Bighorn sheep varied more than other species in initial age and body weight and reproductive status through the sampling period. six of the nine ewes sampled gave birth and lactated in 1994, eight of nine in 1995, and none were bred to lamb in 1996. Although weight trends were more difficult to appreciate given this disparity, body weights appeared to reach a minimum in November each year. Trends suggest that pregnant and open ewes can gain weight during the winter (Fig. 5). Weight trends suggest that body weights of animals at FWRF have remained stable or increased with age over the last several years. This confirms clinical evaluations that suggest the feeding regimens calculated by Baker (Miller 1990) and modified by Wild et al. (1992) and Wild (1993) are adequate for maintaining these captive ungulates. Bottle-raising neonates: Mean weaning weight of the 2 male fawns was 23 kg (range 22.7-24 kg). Mean weaning weight of 4 female fawns was 17.5 kg (range 16.8-20.2). Rota and corona virus as well as Salmonella group El was isolated from fawns at various times during the summer. Two male fawns (not included in body mass means above) exhibited chronic debilitation, likely from these infections, and were subsequently euthanatized. Health Maintenance General: Overall, captive wildlife maintained at FWRF remained healthy throughout the year. Most of the major sources of mortality listed 5 years ago in the Program Narrative (Miller 1990) have been controlled. Lumpy jaw persists only in a few old pronghorn and bighorn that were initially infected >5 years ago, traumatic injuries have been markedly reduced (especially in pronghorn), and no marked morbidity or mortality was attributable to naturally �164 75 70 65 ..-.. 0> e60 en ~ 55 C> w 50 S 45 40 35 JUN AUG OCT DEC FEB APR JUL OCT DEC FEB APR JUN AUG OCT DEC FEB APR JUN MONTHS Fig. 1. Body weight data from female mule deer born in 1993 at FWRF during period June 1994 (n=10) to June 1996 (n=7). . the 75 70 ..-.. 65 0> e60 en ~ 55 C> UJ S 50 45 40 35 JUN JUL AUG SEP OCT DEC JAN APR MAY JUN JUL SEP OCT NOV JAN MAR APR JUN MONTHS Fig. 2. Body weight data from female mule deer born in 1992 at FWRF during period June 1993 to June 1996 (n=14). the �165 52 50 48 38 36 -----_._-----_ ... _--- __ ._---_ ..-- 34 ... _---------_._----_._---- ._--------------_._.-.:.._--------- JUL SEP DEC MAR MAY AUG SEP NOV DEC, FEB JUN AUG SEP DEC FEB APR MAY AUG SEP NOV MONTHS Fig. 3. Body weight data from female pronghorn period July 1992 to December 1995 (n=6). born in 1991 at n~RF during the 350 --. - 300 HEALTHY Cl ~ 250 r- :r: CJ \ \ , 200 W ;.;: 150 >- Cl 100 CWO-AFFECTED 0 m 50 0 S J M S J M S J M S J M S J M S J M S J M S J M S J M S J JANUARY, MAY, AND SEPTEMBER . 1987 - 1995 Fig-. 4. Body weight data from female elk born ::ln1986 and maintained ,at FWRF. Four individuals that developed chronic wasting disease (CWD) are isolated for comparison to normal elk (n=7). �166 90 - 85 0> ~80 U). lI S2 75 w S 70 65 JUN JUL SEP OCT DEC JAN MAR APR JUN AUG OCT DEC JAN MAR APR JUL SEP OCT DEC APR JUN MONTHS Fig. 5. Body weight data for a group of adult female bighorn sheep maintained at TIvRF. Period noted is .from June 1993 to June 1996 (n=9 through October 1995, then n=7). �167 occurring respiratory disease in bighorn sheep lambs or adults last year. However, chronic wasting disease and failure to thrive syndrome in mule deer have emerged as significant sources of mortality. During FY1996, relatively high mortality rates occurred in mule deer (Table 1) due primarily to CWO (n=5) and failure to thrive syndrome (n=3). Epizootic hemorrhagic diseases (EHD) was responsible for two mortalities, one pronghorn and one bighorn sheep. EHD has not been previously reported in bighorn sheep. Following this diagnosis, we collected paired serum samples from the bighorn sheep herd to survey for exposure to EHD. Serum analyses will be conducted in fall 1996. Chronic wasting Disease: Despite of the strict protocols, CWO was diagnosed in five mule deer (one buck, one castrated male, three does) during JanuaryApril 1996. One other suspect case occurred in late 1995. Previous to this "outbreak", CWO had been diagnosed in only one deer at FWRF since the depopulation in 1985 (R91 in October 1994). Four of the five deer affected were born at FWRF, the other arrived at FWRF as a fawn in 1990. None had left the facility. In the first case in 1996, CWO was diagnosed as a factor contributing to the death of a buck in January. Weight loss and a chronic moderate bloat had been observed for about 1 month prior to death, but CWO was not highly suspected. Following this diagnosis, we critically evaluated health status of all deer at the facility. Five deer were subsequently euthanatized as CWO suspects. Two castrates were euthanatized due to poor body condition and three does were euthanatized due to subtle behavioral changes (primarily slight loss of tractability) and slight-moderate weight loss. Body weights of the three does ranged from 55-63 kg at the time of euthanasia. One castrate was incorrectly identified as CWO affected using clinical evaluation (T93); the other four were positive for CWO on histologic examination of the brain. CWO affected deer resided in three of five pastures which house mule deer at FWRF (Fig. 6). Given the increased occurrence of CWO at FWRF, RFAC met to discuss the future of the facility. All in attendance agreed that FWRF should maintain the infected herd because 1) a need exists for more information on the disease and its diagnosis and 2) FWRF is an ideal site to perform the research due to its current infected status and also its location in a CWO endemic area. The group also agreed that given our current level of knowledge, existing biosecurity measures are adequate. Facility Maintenance/Repairs/Improvements A conservation oriented approach to the use of utilities and services for FWRF has likely lead to a reduction in resource use relative to the size of the facility over the past 5 years. Analysis of changes in resource use are not possible because of confounding effects of facility size, intensity of use, and because prior to 1993, CDOW did not receive bills for electricity service which is the most significant utility consumed at FWRF (CSU unknowingly paid all FWRF electric bills). However, insulation of buildings, installation of automatic waterers in all west side pastures, revegetation of east side pastures with drought resistant grasses, and employee awareness of conservation techniques has likely helped minimize use of electricity and water. Trash production has been minimized through more efficient feeding programs which produce less waste. �168 • WI \...12- • IN W,3' • 0/11 \-lTD • -..Ill - - • A 2/12 t-ID MD 3/7 MD '" " •• 11"' •••••••• •• ••• ···"11 •• 111111 ••••••• ,"',.,,· •• , •• ,.,." ••• o 111"'111111'1"""111"""" II II II II I II E.I E:+. £5 0/3 MD Fig. 6. Housing location of six mule deer that have been diagnosed ,with chronic wasting disease at F\.J'RF since 1994. Totals shown are number CH'D positive/animals in pasture for mule deer (MD) and white-tailed deer (\-lTD). �169 As older portions of the facility are repaired and replaced, the need for unscheduled daily repairs appears to be decreasing. Maintenance projects continue to be important for animal safety and facility function. In addition to numerous other repair and maintenance projects, we performed several major improvements. Significant maintenance/repair/improvement projects completed at FWRF this year included: construction of a drainage ditch to catch and divert runoff from west side isolation pens. Upgrade of the electrical service on the east side to increase safety and meet code. Replacement of the outer wall of the east-west section of the west side alleyway. Replacement of the west side alleyway wall adjacent to W3- W4. Clean-up of the waterfowl facility. Clean-up and repair of the mobile home. Construction of a storage shed adjacent to the shop. Repairs to the metabolic cages. Repairs and modifications to west side isolation pens. Facility trash clean-up. Addition of road base to roughest/muddiest portions of facility roads. Repairs to old roofs after wind damage. Research Projects In addition to ongoing facility management experiments and improvements described above, the following pilot studies and research experiments were initiated, conducted, or continued using FWRF animals and facilities this year: Feasibility of using liposomes as deer immunocontraceptive oral vaccine carriers--L. Miller and B. Johns (ADC). Surgical implantation of heart rate transmitters in domestic goats as a model for bighorn sheep--M. Wild, Piermattei, D. Baker, and B. Heath. D. Evaluation of medetomidine and ketamine immobilization reversal with atipamezol in elk--M. Miller, W. Lance. and Experimental evaluation of a multivalent Pasteurella haemolytica toxoid-bacterin (Al, A2, TIO) in captive bighorn sheep (Ovis canadensis)--M. Miller, B. Kraabel, Conlon, and A. Ward. Heart rate as a potential sheep--M. Wild, D. Baker, Pronghorn winter wheat Strohmeyer et. al indicator of stress in bighorn D. Piermattei, and B. Heath. damage study feeding trials--D. J. �170 Educational Contributions FWRF provided formal educational instruction for special interest grade through university groups. Numerous other informal tours were provided individually to visiting professionals. school Youth in Natural Resources Pueblo Youth Naturally CSU Veterinary students in Summer Research Program CSU Zoological Medicine/Small Ruminant Medicine Clubs CSU Pre-vet Club Rocky Mountain High School zoology class Loveland High School zoology class Rocky Mountain Bighorn Society (two tours) Front Range Community College forestry and wildlife class (two tours) Front Range Community College animal health class Blevins Junior High School zoology class-mentoring program Three special interest grade school groups LITERATURE Baker, CITED D. L. and N. T. Hobbs. 1985. Emergency feeding of mule deer during winter: tests of a supplemental ration. J. Wildl. Manage. 49:934-942. Miller, M. W. Colorado Collins. 1990. Animal and pen support facilities for mammals research. Div. Wildl. Res. Rep., WP1a, J1, Jul 1989 - Jun 1990, Fort Wild, M. A. 1993. Animal Colorado Div. Wildl. Collins. and pen support facilities for mammals research. Res. Rep., WP1a, J1, Jul 1992 - Jun 1993, Fort Wild, M. A. 1995. Animal Colorado Div. Wildl. Collins. and pen support facilities for mammals research. Res. Rep., WP1a, J1, Jul 1994 - Jun 1995, Fort Wild, M. A, and W. S. Graffam. 1994. Animal mammals research. Colorado Div. Wildl. Jun 1994, Fort Collins. Wild, M. A, M. W. Miller, B. J. Maynard, and D. R. Magnuson. 1992. Animal and pen support facilities for mammals research. Colorado Div. Wildl. Res. Rep., WP1a, J1, Jul 1991 - Jun 1992, Fort Collins. and pen support facilities for Res. Rep., WP1a, J1, Jul 1993 - �Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESSS state of Project Work Colorado No. W-153-R-9 Mammals Inyestigations Mammals 2 Research Administration Job No. Author: Research Multispecies Plan No. Period REPORT Covered: July 1, 1995 - June 30, 1996 R. Bruce Gill Personnel: R. Bruce Gill and Diane K. Haerter ABSTRACT Thirteen federally funded jobs were planned, budgeted, supervised, and administered during the Segment. Three jobs involved multiple species dimensions, 2 invoved bighorn sheep research projects, 3 involved pronghorn research, 1 involved mountain goat research, 1 involved black bear research, involved GIS research, 1 involved kit fox research, 1 involved swift fox research, and 1 involved habitat research. All research objectives were accomplished within the allocated time limits and budget constraints. Two manuscripts/books were prepared by Mammals 1 staff members in professional journals and/or symposia proceedings. and published 1 ��173 Mammals 2 Research Administration R. Bruce Gill P.M. Administer research studies within productivity and the lowest cost. OBJECTIVE the Mammals Segment 1. 2 Research Unit for the highest Objectives Lead and administer research on mammalian species Research Program other than deer, elk, and moose. in the Mammals RESULTS Thirteen projects were completed successfully active during the segment. Segment objectives for all 13 projects. Highlights include: were { Approximately 35% of the time of Mammals Program Leader allocated to Mammals 2 Research was consumed by reorganization, Management Review implementation tasks, and temporary supervision of the entire Terrestrial Wildlife section. { Eleven articles/books authored by Mammals 1 Research accepted for publication during the Segment. { Three professional/technical manuscripts were in final stages of preparation by members of the Mammals 1 Research staff or were in the peer review process of professional journals. All program projects and activities were achieved allotted time and allocated fiscal resources. Prepared by: R. Bruce Gill Mammals Program Leader staffe within the were ��175 Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS state of Project Work Colorado No. W-153-R-9 Plan No. 1A Job No. Period REPORT Mammals Research Multispecies Inyestigations Monitoring and Managing in Colorado Covered: July Wildlife Health 1, 1995 - June 30, 1996 Authors: M. W. Miller Personnel: W. J. Adrian, J. Bredehoft, G. Byrne, A. L. Case, D. Clarkson, M. Cousins, B. Davies, H. Dietrick, R. Forde, D. Freddy, T. Fulk, D.M. Getzy, K. Green, J. Jackson, K. Kinney, S. Kolus, M. Lamb, M. Leslie, C. Leonard, R. Mason, K. Madriaga, C.W. McCarty, C.A. Mehaffy, B. Olmstead, J. Ritchie, G. Schoonveld, H. Spear, M.L. Stevens, R. Spowart, T. R. Spraker, A. N. Torres, W. Travnicek, J. Wagner, E. S. Williams, and E. Zimmerman ABSTRACT Wildlife populations throughout Colorado were monitored for occurrence of disease using a combination of extensive and intensive approaches. We continued to develop and modify a statewide surveillance program for acquiring, examining, reporting on, and summarizing sporadic wildlife disease cases occurring throughout Colorado. At least 80 carcasses and/or tissue samples representing 64 wildlife cases were submitted for diagnostic examination during July 1995-June 1996. Trauma, brain abscesses, locoism, bronchopneumonia, viral encephalitis,. and spongiform encephalopathy (chronic wasting disease; CWO) were diagnosed in wild cervids submitted, although case submissions were undoubtedly biased by intensive monitoring for CWO); all bighorn sheep submitted showed gross and/or histologic lesions of either bronchopneumonia or paratuberculosis. Among carnivore caseB~ parvoviral enteritis and neoplasia were diagnosed. Sporadic salmonellosis cases in passerine birds continued throughout late 1995 and early 1996. Other mammalian and avian cases appeared to represent isolated incidents or unusual maladies. For 15 cases, cause of death could not be determined. Aside from pneumonia epizootics in elk near Estes Park, locoism in South Park elk, salmonellosis in songbirds, and enzootic occurrences of CWO and paratuberculosis described previously, all cases completed to date appeared to represent isolated cases of trauma, intoxication, or disease. We continued our last annual survey of deer and elk hunters to collect sera for brucellosis screening. Of 2,001 elk hunters surveyed, 136(7%) returned �176 blood samples for brucellosis screening from animals harvested in select GMUs in southwestern Colorado during October-December 1995. Of samples returned, 68 (50%) were usable; marked hemolysis and/or contamination precluded evaluation of the remaining samples. All elk sera tested were negative for antibodies to Brucella spp. on the standard card test. overall, about 3.5% of the survey kits distributed to elk hunters in 1995-1996 provided usable samples, as compared to 7% in 1994-1995, 7% in 1993-1994, 7% in 1992-1993 and 5% in 1991-1992. Because Colorado's domestic cattle herd has been declared brucellosis free and nearly 30 yrs of surveillance have revealed no evidence of wildlife reservoirs of Brucella abortus in Colorado's wild ungulate populations, the annual statewide brucellosis survey will be discontinued. Seven cases of chronic wasting disease (CWO), a spongiform encephalopathy, were confirmed in free-ranging deer and elk in Larimer County between June 1995 and May 1996. Fifty-six free-ranging CWO cases have been confirmed in Colorado since 1981; to date, all but 2 of these cases have been from Larimer County (GMUS 9, 191, 19, or 20). To obtain reliable estimates for distribution and prevalence of CWO in wild cervids, we continued to survey for CWO in select deer and elk populations throughout Colorado. Brains from about 69 mule and white-tailed deer and 78 elk harvested in GMUs 8, 9, 191, 19, 20, or 96 (DAUs D4/E4, D10/E9, 044), were collected for examination for CWO. Histologic evaluation of all samples has not been completed, but of 343 deer and 212 elk brains from hunter-killed animals collected during 1991-1994 and examined to date, 3 deer were spongiform encephalopathy suspects; ancillary tests for confirmation are in progress. Based on survey data collected to date (and assuming all 3 suspects are confirmed positive), estimated prevalence of CWO in mule deer in DAUs 04/010 combined is about 0.009 (95% CI 0.002-0.025); the 95% CI for prevalence in DAU E4/E9 elk is 0-0.01 based on survey data (0/212) collected to date. We continued developing a generalized, stochastic, individual-based simulation model of infectious disease in wild ungulate populations. We constructed models for 2 wildlife disease problems of current interest: chronic wasting disease in a free-ranging mule deer population and bovine tuberculosis in a Michigan white-tailed deer population. Preliminary results of chronic wasting disease modeling suggest lateral transmission is probably an important component of maintaining this disease in a free-ranging population and that predicted impacts on poulation performance are substantially reduced if observed differences in prevalence between sexes (males » females) prove correct. Preliminary results of bovine tuberculosis modeling suggest assumptions for scenarios where Mycobacterium bovis first infected t~e Michigan wild white-tailed deer population >30 yrs ago are more plauBible.than assumptions required for scenarios where tuberculosis was introduced <io yrs ago, and that management strategies directed at reducing deer densities alone are unlikely to eliminate tuberculosis from the affected population within the next 25 yrs. �MONITORING AND MANAGING WILDLIFE HEALTH IN COLORADO M.W. Miller P. N. OBJECTIVES Develop and implement a program for enhancing statewide efforts manage health of Colorado's terrestrial wildlife populations. AGREEMENT to monitor OBJECTIVES 1. Modify and improve systems for submitting, diagnosing and reporting sporadic disease cases in wild animals throughout Colorado. 2. Develop and use databases for assimilating and analyzing problems identified through surveillance and surveys. 3. Design, conduct, paratuberculosis, elk populations. 4. and on data on disease and report results of surveys for brucellosis, and chronic wasting disease in specific deer and/or Provide assistance in investigating outbreaks in Colorado. and managing wildlife disease 5. Design experiments to develop and/or improve techniques for investigating wildlife diseases; begin conducting approved and funded research. Maintaining healthy wildlife populations is a fundamental component of sound wildlife management practices. Habitat degradation, high animal density, extreme weather, and disease can act singly or in combination to compromise the overall health of a wildlife population. As Colorado's wildlife managers, we have developed a variety of tools for monitoring and assessing the effects of habitat loss, animal numbers, and weather on wildlife populations. We have also invested considerably in developing tools to manage these factors to optimize performance of the wildlife populations in our stewardship. In contrast, monitoring and managing the effects of disease on wildlife population performance have received relatively little attention (with a few notable exceptions). This lack of attention may be rooted to some extent in a widely-held belief that wildlife diseases are symptoms of larger underlying population problems that will be resolved if those larger problems are managed properly. Despite this belief, disease can be a significant obstacle to effective and efficient wildlife management in Colorado. Disease outbreaks account for substantial mortality in some wildlife populations. Introduced pathogens have potential to decimate local wildlife populations. Some diseases depress wildlife population performance to levels below resource-based carrying capacity. Many wildlife diseases are shared with domestic animals and/or humans, and in some cas~s wildlife populations serve as reservoirs for these agents. Disease also detracts from the aesthetic value of wild animals, and may convey a perception of mismanagement to uninformed publics. For these �178 reasons, diseases should be regarded as an integral population dynamics and wildlife management. part of wildlife Select wildlife health problems have been monitored in Colorado for more than 30 years. These longstanding efforts have provided useful information on the diseases studied. However, because these efforts have not always been coordinated on a statewide basis, and because some findings have not been widely available to managers and policy makers, applications to overall management programs have been limited. In order to improve our collective ability to manage wildlife health in Colorado, we need a more coordinated and systematic approach for monitoring, investigating, and reporting on health problems in free-ranging wildlife. A more complete understanding of wildlife diseases and their effects on population performance is fundamental to comprehensive wildlife management. Enhanced surveillance efforts will provide a mechanism for detecting health problems throughout the state before serious impacts to wildlife populations occur. Assimilating diagnostic data will aid in assessing trends suggestive of population-level disease problems. Programs for conducting extensive and intensive surveys for potential and realized wildlife diseases will provide reliable prevalence and distribution data for managers and administrators to use in decision making. Expertise in investigating and managing epizootics and epornitics will ameliorate efficacy and efficiency of efforts to control outbreaks. Improved techniques for diagnosing and studying wildlife diseases will provide a firm foundation for health management programs designed to enhance the quality of Colorado's wildlife populations. MATERIALS Disease AND METHODS Surveillance We monitored wildlife populations throughout Colorado for occurrence of disease using a combination of extensive and intensive approaches. These organized and conducted as follows: Statewide were Surveillance We continued to develop and modify a program for acquiring, examining, reporting on, and summarizing sporadic wildlife disease cases occurring throughout Colorado. All carcass submissions were subjected to necropsy. Ancillary diagnostics, including histopathology, bacteriology, virology, serology, parasitology, and toxicolqgy were performed at the discretion of CDOW personnel and/or the attending pathologist. preliminary examination and/or test results were telephoned or faxed to CDOW's Wildlife Research Center Laboratory, usually within 3-5 days of completion, and a final report were usually provided within 15 business days of submission. Copies of reports were filed and sent to appropriate field personnel. Pertinent data from preliminary and final reports, including species, age, sex, location, number affected, diagnosis, and other information (as available) were entered into a permanent computerized database. This database was used to generate quarterly and annual wildlife morbidity and mortality reports. In addition, data are available for analysis of long-term trends in select wildlife disease problems. �179 Surveys Brucellosis Survev: We continued the statewide survey of deer and elk hunters to collect sera for brucellosis screening. We mailed about 2,000 blood sampling kits to elk hunters in selected GMUs in southwestern Colorado to gather samples for CDOW's annual brucellosis surveillance program conducted in cooperation with the Colorado Department of Agriculture's State/Federal Brucellosis Laboratory in Denver. Kits went to sportsmen with antlerless elk permits for second or late seasons in select GMUs. Returned samples were identified by GMU of harvest. Usable samples were centrifuged, and sera were tested for antibodies to Brucella spp. using a standard card test. Unused sera were banked and stored at -20 C for future use. Chronic Wasting Disease Survey: To obtain reliable estimates for distribution and prevalence of CWO. in wild cervids, we continued to survey for CWO in select deer and elk populations throughout Colorado. Brains from mule deer and elk harvested in various seasons during October 1994-January 1995 in GMUs 19 and 20 (DAUs D4 ,D10/E4, E9) were collected for examination for CWO. Brains from hunter harvested mule deer and elk were collected, usually within 24-48 hrs of death, and fixed in 10% buffered formalin for at least 3 months. Sections of medulla at the obex and frontal portion of the brain including basal ganglia, olfactory cortex and tract, and some frontal cortex were processed routinely for paraffin embedment. Histologic sections were cut at 56 pm, stained with hematoxylin and eosin, and examined under a light microscope. In addition to formal surveys, we continued to encourage increased surveillance efforts by field personnel statewide and submission of carcasses from deer or elk showing clinical signs resembling CWO, and held a workshop to inform field personnel about cwo and train them to in recognizing and properly submitting field cases. Disease Investigations Assistance was provided and Granite during July Experimental in investigating 1994-June 1995. pneumonia epizootics near Loveland Approaches We began developing a generalized, stochastic, individual-based simulation model of infectious disease in wild ungulate populations (Fig. 1). We plan to use this model in predicting consequences of disease introductions, improving understanding of the epizootiology of select disease problems, and evaluating potential disease management strategies. In this model, populations display density-dependent sigmoid growth in the absence of disease or other limiting processes. We employed a novel mathematical approach for estimating pathogen transmission within simulated populations, a~d assumed transmission probabilities are a function of prevalence. Initially, we incorporated parameters to simulate introduction of bovine tuberculosis into a wild elk population and examined probable consequences of such introductions. As a preliminary step, we examined results of replicated 50-year simulations (n SaO/parameter set) where 2 infected elk were introduced into a population of 500 wild elk. Our model incorporated population parameters estimated from a lightly hunted elk population (Forbes Trinchera Ranch). We assumed a constant �180 cow-calf transmission rate (0.95) and a 2-year incubation period before newly infected animals became infectious. We then made replicated simulations, varying transmission coefficient (tc = 0.3 or 0.5 new infections/infected individual/year) to assess the influence of transmission on potential outcome of tuberculosis introductions. RESULTS Disease AND DISCUSSION Surveillance Statewide Surveillance At least 80 carcasses and/or tissue samples representing 64 wildlife cases were submitted for diagnostic examination during July 1995-June 1996. Trauma, brain abscesses, locoism, bronchopneumonia, viral encephalitis, and spongiform encephalopathy (chronic wasting disease; CWD) were diagnosed in wild cervids submitted, although case submissions were undoubtedly biased by intensive monitoring for CWD); all bighorn sheep submitted showed gross and/or histologic lesions of either bronchopneumonia or paratuberculosis. Among carnivore cases, parvoviral enteritis and neoplasia were diagnosed. Sporadic salmonellosis cases in passerine birds continued throughout late 1995 and early 1996. Other mammalian and avian cases appeared to represent isolated incidents or unusual maladies. For 15 cases, cause of death could not be determined. Aside from pneumonia epizootics in elk near Estes Park, locoism in South Park elk, salmonellosis in songbirds, and enzootic occurrences of CWD and paratuberculosis described previously, all cases completed to date appeared to represent isolated cases of trauma, intoxication, or disease. We will continue adding new accessions throughout the coming fiscal year to our computerized database for diagnostic case information, as well as data from archived reports as they become available. Surveys Brucellosis SurVey: Of 9,825 elk hunters surveyed, 1,337 (14%) returned blood samples for brucellosis screening from animals harvested throughout Colorado during October 1994-January 1995. Of samples returned, 662 (49%) were usable; marked hemolysis and/or contamination precluded evaluation of the remaining samples. All elk sera tested were negative for antibodies to Brucella spp. on the standard card test. OVerall, about 7% of the survey kits distributed to deer or elk hunters in 1993-1994 provided usable samples, as compared to 7% in 1993-1994, 7% in 1992-1993 and 5% in 1991-1992. Chronic Wasting Disease Survey: Seventeen cases of chronic wasting (CWD), a spongiform encephalopathy, were confirmed in free-ranging elk in Larimer County during FY 1994-1995. Forty-nine free-ranging have been confirmed in Colorado since 1981 (Fig. 1); to date, all these cases have been from Larimer County (GMUs 9, 191, 19, or 20) disease deer and CWD cases but 2 of (Fig. 2). Brains from about 150 mule deer and 65 elk harvested in GMUs 19 and 20 (DAUS D4, D10/E4, E9), were collected for examination for CWD. Histologic evaluation of all samples has not been completed, but of 343 deer and 212 elk brains from hunter-killed animals collected during 1991-1994 and examined to date, 3 deer were spongiform encephalopathy suspects; ancillary tests for confirmation are in progress. Based on survey data collected to date (and �181 assuming all 3 suspects are confirmed positive), estimated prevalence of CWO in mule deer in DAUs D4/D10 combined is about 0.009 (95% CI 0.002-0.025); 95% CI for prevalence in DAU E4/E9 elk is 0-0.01 based on survey data (0/212) collected to tate. To obtain reliable estimates for distribution and prevalence of CWO in wild cervids, we continued to survey for CWO in select deer and elk populations throughout Colorado. Brains from about 25 mule deer and 29 elk harvested in GMUs 19 and 20 (DAUs D4 ,D10/E4, E9), from about 67 mule deer and 32 elk harvested on the Forbes Trinchera Ranch near Ft. Garland (DAU D31/E33), and from about 80 mule deer and elk harvested in GMUs 66 and 67 (GMU D25/E25) were collected for examination for CWO. Histologic evaluation of samples has not been completed, but all brains from hunter-killed deer and elk examined to date have been negative for spongiform encephalopathy. Disease Investigations No significant Experimental disease outbreaks were investigated during July 1992-June 1993. Approaches In examining preliminary results of 500 50-year simulations where 2 infected elk were introduced into a population of 500 wild elk, transmission coefficient (tc) assumptions markedly influenced outcomes. Under conservative assumptions (tc = 0.3 new infections/infected individual/year), the probability that tuberculosis became established (i.e., infection still present 50 years after initial introduction) in simulated populations was about 0.2 (Fig. 2), and prevalence in infected populations averaged about 0.03 (Fig. 3). Using a slightly higher tc (0.5 new infections/infected individual/year), the probability that tuberculosis became established increased to about 0.6 (Fig. 2), and mean prevalence in infected populations reached about 0.7 (Fig. 3). Our preliminary results suggest introduction of bovine tuberculosis into wild elk populations could represent a significant obstacle to national eradication goals. We plan to further refine parameter estimates for elk-tuberculosis simulations, and to explore application of this modeling approach to other real and potential disease problems affecting wild ungulate populations. Acknowledgments The statewide wildlife health monitoring and surveillance program described above relies heavily on efforts of dedicated field personnel throughout the Colorado Division of Wildlife, and truly represents a division-wide effort to improve our understanding and management of wildlife disease problems. In addition to those specifically listed, we collectively thank all of those regional and area biologists, district and area wildlife managers, and others who assisted by submitting diagnostic cases throughout the year. In particular, we thank personnel from areas 2, 4, 10, and 16, and from the Forbes Trinchera Ranch for assistance and logistical support in tuberculosis and CWO surveys and surveillance activities, and personnel from the stateFederal Cooperative Brucellosis Laboratory for their continued cooperation, assistance and logistical support in conducting annual brucellosis surveys. Prepared by Michael W. Miller Wildlife Research Veterinarian �182 Table 1. At least 80 carcasses and/or tissue samples representing 64 wildlife cases were submitted for diagnostic examination during July 1995-June 1996. FY1995-1996 DATE SPECIES AGE SEX DIAGNOSTIC REGION 6-15-95 8-14-95 8-30-95 8-14-95 9-8-95 9-11-95 10-2-95 10-13-95 10-30-95 11-14-95 11-16-95 11-19-95 11-22-95 12-12-95 11-14-95 12-12-95 1-2-96 raccoon C NA bighorn sheep NA SW white-tailed deer A NW mule deer A elk liver and kidney mule deer A NE red-tailed hawks (2) NA SE mule deer F SE mule deer Old M NE hawk NA mule deer SW A ·2+ NE mule deer mule deer. SW 5mos NE golden eagle A NE hawk (unspecified) NA A M golden eagle SE NE elk (2) Calf F 1-2-96 1-5-96 bighorn sheep elk 1-8-96 1-17-96 1-29-96 2-96 4.5 MIF bighorn sheep mule deer (head only) mule deer NA M mule deer Yrlng 2-5-96 2-96 2-14-96 2-21-96 2-22-96 2-27-96 3-6-96 3-13-96 3-18-96 3-26-96 3-27-96 3-29~96 4-2-96 4-96 4-3-96 4-3-96 4-3-96 4-3-96 4-3-96 4-4-96 4-5-96 elk 5+ F mule deer Yng NA Pine Siskin elk NA M pigeons NA mule deer F mule deer 2yrs M mule deer F raccoon NA red fox NA F mule deer 2yr F mule deer lyr NA mule deer mule deer mule deer 4 F mule deer 4 F elk 8 F mule deer 4 M elk A F mule deer 2.5 M sharp-tailed grouse (10) M Yng M A NW SE SUBMISSIONS CAUSE OF DEATH unknown unknown acidosis cellulitis/pododermatitis mercury normal brain abscess trauma unknown old age (shot) unknown drowning keratoconjunctivitis spongiformencephalopathy(CWD) brain abscess renal tubular necrosis unknown renal tubular necrosis fibrinous pneumonia (pasteurellosis) bronchopneumonia poxivirus infection(2° Actinomyces pyogenes Pasteurella multocida) CASE # 945-29765 956-03981 956-R0391 956-W1047 945-90082 956-06562 956-08397 956-09490 956-10815 956-12098 956-12318 956-12690 956-12770 ·956-14276 956-12098 956-14276 956-15876 956-15848 956-16105 956-16316 956-17054 956-18017 C (Grant) paratuberculosis (Johne's disease) keratoconjunctivitis NE spongiformencephalopathy NE subacute lymphoplasmacytic encephalitis NE unknown (trama?) NE encephalitis salmonellosis .(S. typhimurium) C abcess 2° to gunshot NE likely avitrol toxicity chronic glomerulitis NE abscess in cheek viral infection C unknown C likely-isletcell carcinoma in the liver NE spongiformencephalopathy NE unknown (not CWD) unknown (lung VI negative) fibropapilloma SW bacterial infection SW unknown C unknown (not CWD) C mandibulartrauma-mot CWD) SW chronic interstitialnephritis 956-21650 956W1374 56-22694 956-23518 956-25338 956-23862 956-24273 956-25740 956-24512 956-24513 956-24511 956-24515 956-24514 NE 956-24630 Corynebacterium pseudotuberculosis serologynegative for mycoplasmosis 956-18533 956-18745 956-18533 956-19650 956-19973 956-20365 ____________________________________________________ ~_~~~~~ 2~~~~~~~~ __ �183 DATE SPECIES 4-9-96 4-9-96 4-10-96 4-18-96 4-18-96 4-22-96 4-22-96 4-29-96 5-10-96 elk pronghorn mule deer elk turkey mule deer elk mountain lion swifts (6) 5-17-96 5-17-96 6-3-96 6-12-96 6-20-96 redfox sharp-shinned hawk Canada goose Aberts squirrel elk AGE SEX REGION F M Fawn A M NE 1.6 NE NE C 2.8 F A 1 mo. A F 2.5 F C SE SE C C NE NW SE CAUSE OF DEATH spongiform encephalopathy blunt trauma blunt trauma locoweed and elaeophorosis car blunt trauma unknown gunshot parvoviral enteritis severe pulmonary edema, cause unknown unknown unknown unknown myocarditis locoism CASE # 956-25079 956-25063 956-25132 956-25905 956-26179 956-26204 956-27055 956-28288 956-29030 956-29031 956-30367 956-31516 956-32407 �184 15 en (]) en ctS o 10 •••• MlJ1e q~er .••• Elk .. 0 White:-taile.ci deer o 10- (]) ..c E 5 ::l Z o 1981 1983 1989 1987 1985 1991 1993 1995 Year Fig. 1. Fifty-six cases of spongiform encephalopathy were diagnosed in freeranging deer and elk from northcentral Colorado between March 1981 and June 1996. Fifty-three of these 56 cases were submitted since 1990; this pattern may be a product of detection efforts, increasing prevalence, or both. '0 CD •• !em • C rado e o Mule deer (n=46) Elk (n=8) White-tailed deer (n = 2) Fig 2. All but 2 of the 56 documented CWO cases in wild deer and elk have originated in Larimer County (inset); most mule deer cases were clustered around Estes Park (n 19) or in the foothills between Fort Collins and Loveland (n = 20). Black diamonds indicate locations of 2 wildlife research facilities where CWO was described previously. Bar = 10 km. = �185 Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS Colorado State of Project No. Work Plan No. W-153-R-9 Mammals Research 2A Mountain Sheep Inyestigations Strategies for Managing Infectious Disease in Mountain Sheep Populations Job No. Period REPORT Covered: July 1, 1995 - June Authors: M. W. Miller, J. M. Bulgin Personnel: P. E. Bleicher, 30, 1996 B. J. Kraabel, A. L. Case, J. A. Conlon, H. J. McNeil, and and A. C. S. Ward ABSTRACT We examined efficacy and safety of a multivalent Pasteurella haemolytica vaccine (A1, A2, T10) in captive bighorn sheep (Ovis canadensis). In one experiment, 30 captive bighorns were divided into trios on the basis of age, sex, and previous history of pneumonic pasteurellosis; one bighorn from each trio was randomly assigned to receive 0, 1, or 2 vaccine doses. Mild, transient lameness in most vaccinated bighorns 1 day after initial vaccination was the only adverse effect observed. We identified 36 distinguishable biogroup variants among 464 P. haemolytica isolates from bighorns, but oropharyngeal (~ 75%) and nasal (~ 50%) isolation rates for P. haemolytica did not differ among treatment groups (P ~ 0.43). Bighorns receiving 1 or 2 vaccine doses showed marked elevations (P = 0.0001) in leukotoxin neutralizing antibody titers beginning 1 wk after vaccination. Agglutinating antibody titers to serotype A1 and A2 surface antigens were also elevated (P ~ 0.031) in vaccinated bighorns within 2 wk after vaccination; agglutinating antibody titers to serotype T10 surface antigen were relatively high in all three groups but appeared unaffected by vaccination (P = 0.495). Vaccination 7 to 14 wk prior to parturition elevated leukotoxin neutralizing antibody titers in colostrum (P = 0.0103), but neither leukotoxin neutralizing nor serotype A1 surface antigen agglutinating antibody titers differed (P ~ 0.6475) through 16 wk of age among lambs born to dams from different vaccine dose groups. In a separate experiment, we evaluated vaccine-induced protection from challenge with pathogenic P. haemolytica (biotype T, serotype 10, ribotype Eco; "Alamosa Canyon" strain). Fifteen captive bighorns were divided equally into 3 treatment groups: control (no vaccination), 1 dose 10 days prior to challenge, or 1 or 2 doses 57 wk prior to challenge. At challenge, each bighorn received about 5 ml (6.2 X 101 CFUs) of P. haemolytica suspension sprayed into the proximal trachea. Vaccination reduced mortality rates (P �0.1) and lung pathology (P = 0.08) in bighorns vaccinated 10 days prior to challenge, as compared to controls; although mortality rates and lung pathology in bighorns vaccinated 57 weeks prior to challenge did not differ from controls (P ~ 0.2), a trend in reduced mortality and pathology was apparent. Leukotoxin neutralizing antibody titers to P. haemolytica were elevated at challenge in bighorns vaccinated 10 days previously (P = 0.0034), and titers in bighorns from both vaccinated groups were elevated at postmortem ~ 7 days after challenge (P ~ 0.0044). In contrast, titers of agglutinating antibody to P.haemolytica serotype A1 or TIO surface antigens did not differ between vaccinated and unvaccinated bighorns (P ~ 0.19). Based on these data, we believe that this experimental P. haemolytica vaccine is safe and can stimulate protective immunity from pneumonic pasteurellosis in bighorn sheep. Further evaluation of this vaccine as a tool in preventing and managing pasteurellosis in wild bighorn sheep appears warranted. �187 EXPERIMENTS TO IDENTIFY AND MANAGE STRESS IN MOUNTAIN SHEEP POPULATIONS M. W. Miller B. J. Kraabel J. A. Conlon B. J. McNeil and J. M. Bulgin P. N. OBJECTIVE To develop population strategies for managing performance. infectious SEGMENT diseases affecting bighorn sheep OBJECTIVES 1. Design and conduct an experiment evaluating select humoral immune responses of captive bighorn sheep to a multivalent haemolytica vaccine. 2. Design and conduct an experiment evaluating efficacy of a multivalent Pasteurella haemolytica vaccine in protecting captive bighorn sheep from challenge with pathogenic Pasteurella haemolytica. MANAGEMENT OF BACTERIAL AND VIRAL DISEASES IN MOUNTAIN SHEEP and cellular Pasteurella POPULATIONS Inability to control infectious disease outbreaks and subsequent mortality in mountain sheep populations represents a significant obstacle to long-term success in their management. Although the "bighorn pneumonia complex" has been studied intensively for over 3 decades, little is known about many aspects of its etiology and epizootiology. Moreover, management interventions recommended for preventing or controlling this problem remain untested. Although viral, bacterial, and parasitic agents have all been incriminated in these outbreaks, Pasteurella spp. are perhaps the most common pathogens associated with bronchopneumonia in bighorns. Two species, P. haemolytica and P. multocida, and several biotypes and/or serotypes within those species, have been isolated from bighorns during epizootics. Unfortunately, despite extensive diagnostic and experimental investigation, the epizootiology of pasteurellosis in wild bighorn populations is poorly understood. In the absence of knowledge about the epizootiologY of pasteurellosis, effective strategies for managing pneumonia in bighorn populations have not emerged. Difficulties in understanding and managing pasteurellosis also plague the domestic sheep industry world-wide. Reported differences in species susceptibility and strain-specific pathogenesis notwithstanding, the epizootiology of pasteurellosis in domestic sheep is strikingly similar to that observed in bighorn sheep. Because of its wide distribution and sporadic nature, recent attempts to manage pasteurellosis in domestic sheep have focused on prevention through vaccination. The efficacies of vaccines developed for domestic sheep have varied widely (Gilmour and Gilmour, 1989; Donachie, 1994), and many either exacerbated or failed to prevent disease. �188 However, experimental vaccines containing leukotoxin and soluble cell surface antigens from P. haemolytica offered ~37% protection against experimental challenge (Sutherland et al., 1989; Alexander et al., 1995). Humoral immune responses stimulated by one of these vaccines (Sutherland et al., 1989) approximated those observed in lambs allowed to recover from experimental P. haemolytica infections (Donachie et al., 1986). Because captive bighorn sheep that survived pneumonic pasteurellosis have shown resistance during subsequent pneumonia epizootics (Miller et al., 1991b), it follows that vaccines stimulating antibody to P. haemolytica leukotoxin and soluble cell surface antigens might afford protection against naturally occurring pasteurellosis in bighorns as well. Here, we examined effects of a multivalent P. haemolytica vaccine (serotypes A1, A2, TIO) on humoral immune responses and resistance to P. haemolytica challenge in captive bighorn sheep. METHODS Management of Bacterial and Viral AND MATERIALS Diseases in Mountain Sheep Populations In conjunction with numerous cooperators, we continued developing and improving tools available for use in studying etiology, epizootiology, prevention or control of disease outbreaks in bighorn populations: and Experimental evaluation of a multivalent Pasteurella haemolytica toxoid-bacterin (A1. A2. T10) in captive bighorn sheep (Miller, Conlon, McNeil, Bulgin, and Ward): We used 30 captive Rocky Mountain bighorn sheep (0. canadensis canadensis) in this experiment. All bighorns were housed at the Colorado Division of Wildlife·'s Foothills Wildlife Research Facility (Fort Collins, Colorado 80526, USA; 40035'N, 105°10'W) throughout the study. We subdivided bighorns into groups by age and sex «1 yr, >1 yr rams, >1 yr open ewes, >1 yr pregnant ewes), and individuals within these subgroups resided together in 3 to 7 ha pastures throughout the study. In addition to natural forage, grass/alfalfa hay mix and pelleted high-energy supplement were provided throughout the study as prescribed under established feeding protocols for bighorn sheep in respective age/sex classes (Miller, 1990); fresh water and mineralized salt blocks were also provided ad libitum. a The general health of all bighorns was evaluated prior to and immediately after vaccination, and daily thereafter. We recorded health observations throughout the experiment, giving particular attention to detecting signs of respiratory disease (depression, segregation, anorexia, nasal discharge, coughing, labored breathing). Injection sites were examined weekly for 4 wk after vaccine administration to assess local reactions. All bighorns were weighed in conjunction with sampling. Health problems were evaluated and treated by attending veterinarians as necessary. Bighorns that died during our experiment were submitted to the Colorado State University Diagnostic Laboratory (Fort Collins, Colorado 80523, USA) or Wyoming State Veterinary Laboratory (Laramie, Wyoming 82070, USA), where carcasses were necropsied and subjected to histopathologic examination and ancillary diagnostic tests to determine .cause of death. The experimental P. haemolytica vaccine (Langford Laboratories, Inc., Guelph, Ontario, Canada; lot 940902) used here was a bacterial cell-free extract of culture supernatants from three P. haemolytica serotypes (A1, A2, T10) that contained leukotoxin, serotype-specific surface antigens, and an adjuvant (MUNOKYNIN~, Langford, Inc., Kansas City, Missouri, USA). Methods for �189 preparation and serotype A1 components of this vaccine were used in a commercially-available bovine vaccine (PRESPONSE® Laboratories, Inc.). the same as those Langford We measured the effects of experimental P. haemolytica vaccine administration on humoral immune responses and P. haemolytica isolation rates in captive bighorn sheep, and on antibody levels and isolation rates in lambs born to pregnant ewes included in our study. Resistance to experimental challenge with pathogenic P. haemolytica was not tested in this study, but we did not attempt to prevent the natural occurrence of pneumonic pasteurellosis in study bighorns. Our study was designed as a randomized complete block experiment with a repeated measures structure. In order to distribute treatments equally across the study population, subject animals were stratified by age «1 yr, >1 yr), sex (>1 yr rams, >1 yr open ewes, >1 yr pregnant ewes), and previous history of pneumonic pasteurellosis (health history). Within strata, individual sheep were assigned to blocks (n = 3 animals/block). One bighorn within each block was then randomly assigned to each of 3 treatment groups: 0 (control, no vaccination), 1 (1 vaccine dose), or 2 (2 vaccine doses 14 days apart. On 20 February 1995 (= wk 0), we aseptically injected 2 ml of experimental vaccine intramuscularly (1M) into bighorns in treatment groups 1 and 2; controls (group 0) were injected 1M with 2 ml 0.9% saline. Fourteen days later, bighorns in treatment group 2 were injected 1M with a second 2 ml vaccine dose; bighorns in treatment groups 0 and 1 were injected 1M with 2 ml 0.9% saline. All bighorns received vaccine or saline in the right hind leg on day 0 and in the left hind leg on day 14. We collected about 10-12 ml blood for antibody measurements from each bighorn on wk 0 (prior to vaccination), 1, 2, 3, 4, 6, 8, 12, 16, 25, and 29; we also collected oropharyngeal and nasal swabs (Baxter Healthcare Corporation, McGaw Park, Illinois, USA) from each bighorn on wk 0, 2, 4, 8, 12, and 29. Blood and oropharyngeal swabs were collected from pregnant ewes and their lambs within 12 hr postpartum, and about 1, 2, 3, 4, 6, 8, 10, 12, 14, and 16 wk postpartum; we also collected colostrum. from ewes within 12 hr postpartum. Blood samples were held for 1-4 hr at about 22 C, centrifuged, and serum collected. Serum and colostrum were stored at -20 C until analyzed at Ayerst Veterinary Laboratories (Guelph, Ontario N1K 1A8, Canada). Swabs were placed in transport tubes containing modified Cary and Blair medium (Port-A-Cul®, Becton Dickinson Microbiology Systems, Becton Dickinson and Company, Cockeysville, Maryland~ USA) and shipped overnight on.ice packs to the University of Idaho's Caine Veterinary Teaching and Research Center (CVTRC; Caldwell, Idaho 83605-8098, USA) for culture and analysis. Levels of leukotoxin neutralizing antibodies in bighorn sera and colostrum were measured using a modified in vitro leukotoxin neutralization assay (Greer and Shewen, 1985; Shewen and Wilke, 1988). Serial two-fold dilutions of test sera or colostrum were pre incubated with leukotoxic P. haemolytica culture supernatant for 30 min at 22 C; supernatant concentration was adjusted beforehand to produce a standardized titer with positive control bovine serum. We then transferred 50 ~l of each serum (or colostrum)/toxin mixture to microtiter plate wells containing bovine leukemia-derived B cell line (BL-3) cells and 50 ~l Roswell Park Memorial Instruments (RPMI) medium (Life Technologies/Gibco BRL, Toronto, Ontario, Canada). Plates were incubated for 1 hr at 37 C, and cell viability was measured by uptake of neutral red dye as �190 previously described (Greer and Shewen, 1986). All sera were assayed in duplicate, along with positive and negative control bovine sera. We expressed neutralization titers as the highest reciprocal 10g2 dilution that yielded ~ 50% neutralization of toxicity. Levels of serum and colostral antibodies against serotype-specific surface antigens were measured using a direct microagg1utination assay (Reggiardo, 1981) that incorporated washed formalinized P. haemolytica serotype A1, A2, or T10 cells as antigen. We expressed agglutination titers as the reciprocal 10g2 of endpoint dilutions. At CVTRC, oropharyngeal and nasal swabs were plated on both Columbia blood agar (Difco Laboratories, Detroit, Michigan, USA) with 5 % citrated ovine blood (CBA) and a selective medium containing Columbia blood agar, 7 % bovine blood, and antibiotics selective for Pasteurellaceae (Ward et al., 1986). Suspected P. haemolytica colonies were selected after 48 hr incubation at 37 C in 5% CO2 and further propagation was carried out on CBA~ Plates were examined for hemolysis. Identities and biochemical profiles of colonies yielding gram negative rods or coccobacilli that fermented triple sugar iron, glucose, and sucrose were further evaluated to determine species and biogroup identifications (Kilian and Fredericksen, 1981; Bisgaard and Mutters, 1986) based on these reactions. The biogrouping schema of Bisgaard and Mutters (1986) was expanded by CVTRC modifications that allowed use of negative test results to separate isolates into additional biogroups. Select isolates identified as ~ haemolytica were further characterized by serotype using rapid plate agglutination (Frank and Wessman, 1978). We calculated rates for isolating P. haemo1ytica, both in general and for select biogroups, from nasal and oropharyngeal sites. Although resistance to experimental challenge with pathogenic P. haemolytica was not tested in this study, we observed bighorns daily for signs of respiratory disease (nasal discharge, depression, segregation, anorexia, coughing, dyspnea). We defined clinical pneumonia as a combination of signs including mucopurulent nasal discharge, depression (with or without segregation from penmates), anorexia or failure to gain weight, and coughing or dyspnea, accompanied by estimated resting body temperature ~ 39.5 C. The probable etiology of all pneumonia cases was to be determined by ancillary diagnostic tests. We compared levels of neutralizing antibody titers to P. haemolytica leukotoxin and levels of antibody to P. haemolytica surface antigens in serum and colostrum, rates of Pasteurella spp. isol~rom oropharyngeal and nasal sites, phenotypic traits of P. haemolytica isolates, and rates of naturally-occurring pneumonic pasteurellosis among treatment groups. We analyzed serology data using least squares analysis of variance for general linear models (SAS Institute, Inc., 1989). Responses to treatments were analyzed with analysis of variance for 'a randomized complete block design with a repeated measures structure. We used vaccine doses (.0, 1, 2), age/sex and health history as main effects. Factors in the analysis were vaccine treatment, age/sex, health history, and interaction of vaccination and health history. Time was treated as a within subject effect using a multivariate approach to repeated measures (Morrison 1976). For postpartum serum and colostrum titer data, we also used least squares analysis of variance for a randomized complete block design with vaccine dose as the sole main effect. We calculated prevalence rates for P. haemolytica isolation from oropharyngeal and nasal sites and compared these among treatments using Fisher's exact probability tests (Mielke and Berry, 1992). We also compared prevalences of �phenotypically distinct strains of P. haemolytica among treatments for the ten most common biogroups isolated using Fisher's exact probability tests. Efficacy of a multivalent Pasteurella haemolytica toxoid-bacterin in protecting captive bighorn sheep (Ovis canadensis) from challenge with pathogenic Pasteurella haemolytica (Miller, Conlon, McNeil, Bulgin, and Ward): We used captive Rocky Mountain bighorn sheep (0. canadensis canadensis) (n = 15) in this experiment. All bighorns were housed at the CDOW's Foothills Wildlife Research Facility (FWRF) throughout the study. We subdivided bighorns into groups by vaccination status. Three or four individuals resided together in about 100 m2 isolation pens throughout the study; pen assignments were random, but accommodated social dif.ferences in order to minimize stress on cohoused individuals. Grass/alfalfa hay mix and a pelleted high-energy supplement was provided as prescribed under FWRF feeding protocols for bighorn sheep; fresh water and mineralized salt blocks were provided ad libitum. Administering live cultures of cytotoxic P. haemolytica into the laryngopharynx of bighorn sheep was intended to cause severe respiratory infections, at least in controls; this experiment was designed to evaluate the efficacy of vaccination in preventing or reducing these infections. We anticipated some sheep would become clinically ill or die in response to challenge. Our approach attempted balance the need to evaluate clinical responses of challenged bighorn with the need to relieve pain and suffering in affected animals. The health of the bighorns was closely monitored by attending veterinarians at least 3 times daily (early morning, mid-day, and dusk) after challenge. Analgesic or antiinflammatory drugs were not provided to affected animals because they might have interfered with vaccine-mediated responses to challenge. Moreover, because the goal of this research was to determine whether the vaccine protected bighorns in the face of infection with P. haemolytica, we did not euthanize animals demonstrating respiratory infection unless they become recumbent, extremely depressed, or moribund. All animals that dies or were euthanized during the experiment were necropsied to determine the cause of death. Surviving bighorn were euthanized 7 or 8 days after challenge and necropsied to evaluate lung pathology for comparison with natural mortalities. All bighorn were euthanized with about 100 mg/kg pentobarbital sodium solution (Beuthanasia*) administered intravenously (IV). We examined whether prior administration of experimental P. haemolytica toxoid-bacterin (AI, A2, T10) to bighorn sheep increased their resistance to experimental challenge with a cytotoxic strain of P. haemolytica in a randomized, complete block experiment. Bighorns were assigned to 1 of 3 groups based on their vaccination status: 0 (control, no vaccination), 1 (vaccinated 10 days prior to challenge), or 2 (vaccinated 57 wk prior to challenge). Bighorns in group 2 received experimental toxoid-bacterin in February 1995 during a previous vaccine study (Miller et al. 1996); bighorns in groups 0 and 1 were not vaccinated previously, and were assigned randomly to respective groups. The experimental P. haemolytica toxoid-bacterin (Langford Laboratories, Inc.) evaluated here was the same vaccine used by Miller et al. (1996). A P. haemolytica biotype T, serotype 10, ribotype Eco field isolate ("Alamosa Canyon" strain) was used for challenge. This isolate originated from tissues collected during a bighorn pneumonia epizootic in southcentral Colorado in 1990; previous in vitro and in vivo evaluations of this isolate demonstrated marked cytotoxin production and pathogenicity (Miller et al. 1995, B.J. Kraabe1, unpubl.). All challenge doses were prepared from a 12-hr brain heart �192 infusion broth culture centrifuged at 4000 g for 15 minutes sterile PBS to an optical density of 1.0 at 525 nm. and resuspended in After a 12 day acclimation period in the isolation pens, we aseptically injected 2 ml of experimental toxoid-bacterin intramuscularly (1M) into bighorns in treatment group 1; treatment group 2 and controls (treatment group 0) received 2 ml 0.9% saline, aseptically injected 1M. Ten days after vaccination or saline injection, each bighorn received about 5 ml (6.2 X 107 CFUs) of P. haemolytica suspension sprayed into the proximal trachea. To administer this chailenge dose, we sedated each bighorn with xylazine HCL (about 5~20 mg IV), held open its mouth with a speculum, anesthetized the larynx with lidocaine, placed a spray nozzle into the glottis with aid of a laryngoscope, and sprayed culture suspension into the proximal trachea. Sedation was reversed with yohimbine HCL (10-40 mg IV) immediately after administration of challenge dose. Efficacy of the experimental toxoid-bacterin was evaluated through comparison of morbidity and mortality rates associated with P. haemolytica challenge, as well as humoral immune responses. Resistance to experimental challenge was measured by comparing mortality rates and both clinical and post mortem evaluations among treatment groups. Clinical signs were observed and scored 3 times daily (early morning, mid-day, and dusk; observers were blinded to treatment group assignments. Seven or 8 days after challenge, all surviving bighorns were immobilized with a combination of ketamine HCL and xylazine HCl (about 40-120 mg ketamine + 4-12 mg xylazine IV) and euthanized with intravenous pentobarbital. All study bighorn were necropsied immediately after death, gross lesions recorded and photographed, and respiratory tract lesions scored using a system adapted from Black and Duganzich (1995). For serology, blood (10-12 ml) was collected from each bighorn immediately prior to vaccination, challenge, and death. All blood samples collected for serology were held for 1-4 hr at about 22 C, centrifuged, and serum collected. Serum was stored at -20 C until analyzed. Levels of cytotoxin neutralizing and agglutinating antibodies in sera were measured as described above. oropharyngeal and nasal swabs were collected from each bighorn in conjunction with sample collections for serology. Swabs were placed in transport tubes containing modified Cary and Blair medium (Port-A-Cul~, Becton Dickinson and Company) and shipped overnight on ice packs to Caine Veterinary Teaching and Research Center (CVTRC), Caldwell, ID for culture and analysis. Representative tissue samples from all organ systems were collected at necropsy and placed in 10% buffered formalin for histologic evaluation. Fresh lung, spleen, liver, and kidney tissue, as well as joint swabs, were also collected for bacteriology and/or virus isolation. Oropharyngeal, nasal, and joint swabs, as well as lung, spleen, liver, and kidney tissue from sheep will be cultured for detection of Pasteurella spp., and P. haemolytica isolates will be further characterized by biochemical profile and serotype using CVTRC protocols (A.C.S. Ward, CVTRC, unpubl.). Phenotypic differentiation of P. haemolytica isolates were based on biogrouping (Bisgaard and Mutters 1986, A.C.S. Ward, CVTRC, unpublished) and serotyping by rapid plate agglutination (Frank and Wessman 1978). We compared haemolytica antigen, 4) Pasteurella 1) mortality rates, 2) serum neutralizing antibody titers to P. leukotoxin, 3) serum antibody titers to P. haemolytica capsular clinical scores, 5) necropsy scores, and 6) recovery of spp. from lung tissue among treatment groups. Mortality rates �193 were compared using Fisher's exact test. Serology data were analyzed using least squares ANOVA for General Linear Models (Freund et ale 1986). Ranked clinical response scores were analyzed using ANOVA with repeated measures structure. Post mortem scoring data were analyzed by the Kruskal-Wallis technique using non-parametric paired comparisons. We also used Cochran's Q test to compare distributions of phenotypically distinct strains of P. haemolytica among treatment groups. We used a = 0.1 for all statistical comparisons. RESULTS AND DISCUSSION Experimental evaluation of a multivalent Pasteurella haemolytica vaccine (A1, A2, T10) in captive bighorn sheep: Mild, transient lameness in most vaccinated bighorns 1 day after initial vaccination was the only adverse effect observed. We observed no clinical signs of pneumonic pasteurellosis in any of the study bighorns. One ewe (L93) died between 20 February and 11 September, but her death appeared unrelated to vaccine treatment (she succumbed to a pyogranulomatous brain abscess that was probably caused by a migrating grass awn about 4.5 mo after vaccination; Actinomyces pyogenes was isolated from this lesion). We observed no abortifacient effects associated with vaccinating pregnant ewes (n = 13); no definite etiology was ascribed to one abortion that occurred in early April, but the affected ewe (A82) had a preexisting (~2 yr) history of late_'term abortions. Of 12 lambs delivered between 10 April and 23 May, eight survived to weaning. All lamb mortality occurred 5 5 days postpartum. We attributed two of the four perinatal losses to hypothermia (possibly complicated by prematurity), one to starvation, and one to stillbirth possibly caused by prematurity and/or inadequate maternal care; evidence of infectious disease was not observed in any of these lambs. None of the losses appeared related to vaccine treatment: two of the ewes that lost lambs had been vaccinated and two were controls. We observed no signs of pneumonic pasteurellosis in any of the eight surviving lambs. We recovered P. haemolytica from all captive bighorns used in this study. Oropharyngeal (~ 75%) and nasal (5 50%) isolation rates for P. haemolytica did not differ among treatment groups (~~ 0.43) In all, we identified 36 distinguishable biogroup variants among 464 P. haemolytica isolates from adult bighorns. We observed differences in relative prevalence of specific ~ haemblytica biogroups: ten biogroup variants accounted for about 87% of all isolates. Prevalence within respective biogroups was not affected by vaccine .treatment (.E....2! 0.185), although some temporal trends in relative prevalence were evident. Vaccination of dams also had no apparent effect on colonization of lambs: we initially isolated P. haemolytica from all neonatal lambs < 1 to 16 days after birth and routinely thereafter. Mean pretreatment titers for P. haemolytica leukotoxin neutralizing antibody were relatively low (~ 0.70) across all three treatment groups (Fig. 1A), and remained low (5_2.1) throughout the 29-wk trial in unvaccinated bighorns. In contrast, vaccinated bighorns showed marked elevations (~ = 0.0001) in leukotoxin neutralizing antibody titers beginning 1 wk after initial vaccination (~ = 0.0001). Mean responses peaked at 2 wk for both vaccine groups, and titers remained elevated above control titers ~ 8 but < 12 wk in group 1 (~5 0.0014) and ~ 16 but < 25 wk in group 2 (~5 0.0382). Mean pretreatment titers for agglutinating antibody to P. haemolytica surface antigens ranged from 5 3.9 for serotype A1 to ~ 9.3 for serotype T1D (Fig. 18- �194 D), but these also remained relatively stable in unvaccinated controls. Agglutinating antibody titers to serotype Al surface antigen were elevated (£ = 0.0017) in vaccinated bighorns beginning 1 wk after initial vaccination (£ 0.0001) and showed response patterns similar to those of neutralizing antibodies: mean responses peaked at 2 wk and titers remained elevated ~ 6 but < 8 wk in both dose groups (£ ~ 0.0036) (Fig. 1B). Although elevation (£ = 0.0310) of agglutinating antibody titers to serotype A2 was less dramatic, titers rose within 2 wk after initial vaccination (£ ~ 0.0018) and remained elevated ~ 4 but < 8 wk in dose group 1 (£ ~ 0.0106) and ~ 8 but < 29 wk in group 2 (£ ~ 0.0303) (Fig. Ie); in addition, during wk 4 mean titers in group 2 were higher (£ = 0.0407) than those in group 1. Titers of agglutinating antibody to P. haemolytica serotype T10 surface antigen were relatively high in all three groups and appeared unaffected by vaccination (£ = 0.4950) (Fig. 1D). We also detected differences in agglutinating antibody titers to serotype Al (£ = 0.0366) and T10 (£ = 0.0001) among age/sex strata. These differences appeared age-related, particularly among responses to serotype T10: highest mean titers tended to occur in adult ewes and lowest in 9 mo old bighorns. Antibodies to P. haemolytica leukotoxin and serotype Al surface antigen were detected in colostrum and periparturient serum from bighorn dams and in serum from their lambs (Fig 2). Vaccination 7 to 14 wk prior to parturition elevated leukotoxin neutralizing antibody titers in colostrum (£ = 0.0103), but serum titers in vaccinated dams and their 1-wk-old lambs did not differ from controls (~0.1713)(Fig. 2A); small sample sizes may have precluded detection of additional differences, particularly among· lambs. Serotype Al surface antigen agglutinating antibody titers in colostrum, as well as in serum from dams and lambs, did not differ among vaccine dose groups (~ 0.4053) (Fig. 2B). Neither leukotoxin neutralizing nor serotype Al agglutinating antibody titers differed (~ 0.6475) through 16 wk of age among lambs born to dams from different vaccine dose groups (Fig. 3). Both leukotoxin neutralizing and serotype Al agglutinating antibody titers in bighorn lambs declined steadily in lambs over the first 4 to 8 wk after birth. Mean agglutinating antibody titers subsequently began increasing when lambs were about 6 to 8 wk old and eventually returned to neonatal levels, but we detected no such active resurgence in bighorn lambs' leukotoxin neutralizing antibody titers. Efficacy of a multivalent Pasteurella haemolytica toxoid-bacterin in protecting captive bighorn sheep (Oyis canadensis) from challenge with pathogenic Pasteurella haemolytica: Data analyses are incomplete, and results reported here are preliminary. Vaccination reduced mortality rates (P = 0.1) and lung pathology (P = 0.08) in bighorns vaccinated 10 days prior to challenge, as compared to controls; although mortality rates and lung pathology in bighorns vaccinated 57 weeks prior to challenge did not differ from controls (P ~ 0.2), a trend in reduced mortality and pathology was apparent. Leukotoxin neutralizing antibody titers were elevated at challenge in bighorns vaccinated 10 days previously (P = 0.0034), and titers in bighorns from both vaccinated groups were elevated at postmortem ~ 7 days after challenge (P ~ 0.0044). In contrast, titers of agglutinating antibody to P.haemolytica serotype Al or T10 surface antigens did not differ between vaccinated and unvaccinated bighorns (P ~ 0.19). �A. Cytotoxin neutralizing antibodies B. Agglutinating 12 ~ 8 g 6 :; • 0 ••••• ..•. ,;; 10 .£? .£? ~ ~ ~ .!:! 1U 4 ... .• 10 15 20 25 to serotype ....0 ••••• antibodies 6 4 en <I: 2 ~en 0 30 0 10 5 to serotype A2 D. Agglutinating 12 -0 ..... • •••. 1 •••• 10 8 enN 0'" ~ ••••• 6 f.~~~~~~~~:~;~~~~~~~~~~"8 .s .= 4 .= c 1U ~ c 1U ~ Cl en <I: ~en 2 o en <I: ~~~~~~~~~~~~~~~~-L~~~ o 5 10 15 Week 20 25 10 E .s ~ 20 15 25 30 Week N 0 , 8 .= 12 en ..... ................•.. 10 Week C. Agglutinating A1 N en c ~ I"~ .: - ~,~;::""'" o o 5 antibodies 12 ....... ..~•.·,I"~ .••• 30 Ii'.·1' t-....- =-.; =...= ~ J.-:-" antibodies T -f- ..... - - '-'~_- to serotype T1 0 r . ... · .. ~ .~.-:-.~.-,l~~~ •.. ! J. ..•• 8 011 •••• .•.. , 6 .•... 4 2 o I; :. o r····· .,.' ..... 5 '" 10 " 15 " 20 " , 25 30 Week Figure 1. (A) Bighorns receiving one or two vaccine doses showed marked elevations in P. haemolytica leukotoxin neutralizing antibody titers (~= 0.0001) as compared to unvaccinated controls. (B) Titers of agglutinating antibody to P. haemolytica serotype Al surface antigen were also elevated·in vaccinated bighorns (~= 0.0017). (e) Titers of agglutinating antibody to P. haemolytica serotype A2 surface antigen showed less marked elevations (~= 0.0310) in vaccinated bighorns. (D) Titers of agglutinating antibody to P. haemolytica serotype TIO surface antigen were relatively high in all three groups but unaffected by vaccination (P = 0.4950). Points are mean observations; vertical lines are + 1 standard error of mean observations.- ~ ~ �196 A. Cytotoxin neutralizing antibodies 12 N C> Do ~ 10 •.... -- ........ <D 8 c 6 o as 1 dose .m 2 o doses doses ; .. N as •.... 4 ::J <D Z 2 o Dam serum Lamb serum Colostrum (1 wk old) (postpartum) B. Agglutinating antibodies to serotype A1 12 ~ C> o 10 - 8 •.... <D c o as c Do ~ ·doses 1 dose m 2 doses 6 4 2 o Dam serum (postpartum) Colostrum Lamb serum (1 wk old) Figure 2. (A) Vaccination 7 to 14 wk prior to parturition elevated leukotoxin neutraliiing antibody titers in colostrum (P = 0.0103), but serum titers in vaccinated dams and their 1-wk-old lambs did not differ from controls (P > 0.1713). (B) Serotype A1 surface antigen agglutinating antibody titers in colostrum and in serum from dams and lamhs did not differ among vaccine dose groups (P > 0.4053). Bars are mean observations; vertical lines are + 1 standard error of mean observations. �197 A. Cytotoxin ...-.. neutralizing antibodies 12 -0- 0 doses C\I 0> o -A- 10 ..•........••.. •... Q) •.... •.... 8 o •.... c 6 N 4 ::::I Q) -.Y-: ,2. d.oses . ·1································· m m •... •.... 1 dose 2 .m:::,.,_················· Z o o 2 4 6 8 10 12 14 16 Age (weeks) B. Agglutinating antibodies to serotype A 1 12 ~ 0 doses :.••- 1 dose -y- t.... Q) •.... •.... c o m 8 . ":::'. . . . .:..:::'. . . . .:.'..:.::.::::.:::::::::.. -.-.d~I··········· . .. . . . . . . . .. .. .. ••.•.•.• f. .•.• :~1::?i: . .". .... ---... ---i 6 •.... c •.... 4 .,\1 \ ....~~......... . . -: I;.~....; ' 'j:..'~...'.....[. ;';'..;,';'. ,·~·····t·. . . . . .. . "':-.'_ -'' :' ,+..:. .;,-/.-- '_ :-. -::-c:. '" ::::I 0> 0> 2 <{ o 2 doses ./ o 2 4 6 8 10 12 14 16 Age (weeks) Figure 3. Neither (A) leukotoxin neutralizing nor (B) serotype A1 surface antigen agglutinating antibody titers differed (P > 0.6475) through 16 wk of age among lambs born to darns that had previously received 0, 1, or 2 vaccine doses. Although titers to both antigens declined steadily in lambs over the first 4 to 8 wk of life, only agglutinating antibody titers eventually returned to neonatal levels. Points are mean observations; vertical lines are + 1 standard error of mean observations. �198 The challenge strain of P. haemolytica was recovered not recovered from oropharyngeal or nasal swabs from any of the study bighorns at vaccination or at challenge, but was recovered from lung tissue or oropharyngeal swabs from all bighorns at post mortem. In addition, several other distinguishable P. haemolytica biogroups were isolated from lung tissues of some bighorns at post mortem. The multivalent P. haemolytica vaccine evaluated here stimulated humoral immune responses that protected bighorn sheep from experimental challenge with a field strain of pathogenic P. haemolytica. Previous attempts to vaccinate bighorn sheep against pasteurellosis have yielded widely varied results. Although early studies of autogenous bacterins showed some promise of protection (Rufi, 1961; Post, 1962; Nash, 1972), these either failed in application (Howe, 1964) or were never fully evaluated or incorporated into bighorn management programs. In more recent studies, neither an autogenous bacterin (Foreyt, 1992) nor prechallenge inoculation with cytotoxic ~ haemolytica (Foreyt and Silflow, 1996) prevented captive bighorns from succumbing to pneumonic pasteurellosis. Moreover, use of a modified-live ~ haemolytica A1 vaccine apparently caused pasteurellosis in previously healthy captive bighorns (Onderka et al., 1988); the vaccine tested here did not cause or exacerbate pasteurellosis in bighorns. Other direct comparisons of data among our and others' studies are confounded by differences in respective methodologies: previous studies either did not measure antibody responses to vaccination (Onderka et al., 1988; Foreyt, 1992; Foreyt et al., 1996) or used serology methods that preclude reliable comparisons (Rufi, 1961; Nash, 1972). Vaccination had no measurable effect on carriage of various P. haemolytica strains enzootic in our captive bighorn herd. Bighorn responses to vaccination observed in this study suggest several potential applications in managing pasteurellosis in wild bighorns. Antibody levels in vaccinated bighorns rose rapidly and remained elevated for 12 to 16 wk. Such responses could be beneficial to wild sheep vaccinated either annually or early in the course of a pneumonia epizootic. Perhaps of equal importance, trends in antibody levels detected in colostrum and neonatal .lambs may reflect potential for passive transfer of protective antibodies from vaccinated bighorn ewes; similar patterns of passive antibody transfer have been shown between domestic ewes and lambs (Gilmour et al., 1980), and between dairy cows and calves (Hodgins and Shewen, 1994). Because pasteurellosis in neonatal bighorn lambs may be the single most important factor hampering population. recovery from an epizootic, vaccine-mediated protection could vastly diminish the long-term impacts of pasteurellosis on bighorn population performance. Conferring protection to young bighorn lambs through ewe vaccination could be an effective means of reducing lamb mortality that is a common sequela of pasteurellosis epizootics in wild bighorn herds (Onderka and Wishart, 1984; Festa-Bianchet, 1988; Foreyt, 1990), although our data suggest that ewe vaccination would not prevent lambs' colonization with P. haemolytica and that any conferred protection would likely be ephemeral at best. Such protection could, however, be sufficient to allow neonatal bighorn lambs to recover from pneumonic pasteurellosis and subsequently develop natural immunity to future infections (Donachie et al., 1986; Hodgins and Shewen, 1994) • Our data suggest that this experimental P. haemolytica vaccine is safe and Stimulation of dramatic stimulates protective immunity in bighorn sheep. antibody responses by a single vaccine dose greatly enhances prospects for its Moreover, potential for delivery via oral carriers use in wild bighorns. �199 (Bowersock et a1., 1994) or biodegradable implants makes application to freeranging bighorn populations a tangible goal. Consequently, we believe further evaluation of this vaccine as a tool in preventing and managing pasteurellosis in bighorn sheep is warranted. LITERATURE CITED ALEXANDER, A., M. ALLEY, AND J. CONLON. 1995. 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Comparison of protection of experimentally challenged cattle vaccinated once or twice with a Pasteurella haemolytica bacterial extract vaccine. Canadian Journal of Veterinary Research 59:179-182. DONACHIE, W., C. BURRELLS, A. D. SUTHERLAND, J. S. GILMOUR, AND N. J. L. GILMOUR. 1986. Immunity of specific pathogen free lambs to challenge with an aerosol of Pasteurella haemolytica biotype A serotype 2. Pulmonary antibody and cell responses to primary and secondary infections. Veterinary Immunology and Immunopathology 11:265-279. FESTA-BlANCHET, M. 1988. A pneumonia epizootic in bighorn comments on preventive management. Biennial symposium Wild Sheep and Goat Council 6:66-76. sheep, with of the Northern 1990. Pneumonia in bighorn sheep: effects of Pasteurella haemolytica from domestic sheep and effects on survival and long-term reproduction. Biennial Symposium of the Northern Wild sheep and Goat Council 7:92-101. FOREYT, W. J. 1992. Failure of an experimental Pasteurella haemolytica vaccine to prevent respiratory disease and death in bighorn sheep after exposure to domestic sheep. Biennial Symposium of the Northern Wild sheep and Goat Council 8:155-163. , AND R. M. SILFLOW. 1996. Attempted protection of bighorn sheep (Qyia canadensis) from pneumonia using a nonlethal cytotoxic strain of Pasteurella haemolytica, biotype A, serotype 11. Journal of Wildlife Diseases 32:315-321. �200 FRANK, G. H., AND G. E. WESSMAN. for serotyping Pasteurella Microbiology 7: 142-145. FREUND, R. J., R. C. LITTELL, models. SAS Institute, 1978. Rapid haemolytica. plate agglutination Journal of Clinical procedure AND P. C. SPECTOR. 1986. SAS system Cary, North Carolina, 187-201. for linear GILMOUR, N. J. L., J. M. SHARP, W. DONACHIE, C. BURRELLS, AND J. FRASER. 1980. Serum antibodies of ewes and their lambs to Pasteurella haemolytica. The Veterinary Record 107:505-507. , AND J. S. GILMOUR. 1989. 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Pages 16-18 in Federal Aid in Wildlife Restoration, Game and Fish Laboratory Research, Research Project Segment, Job Completion Report, Project Number FW-3-R11, Work Plan 1, Job 2W. Wyoming Game and Fish Department, Cheyenne, Wyoming, USA. KILIAN, M., AND W. FREDERIKSEN. 1981. Identification tables for the Haemophilus-Pasteurella-Actinobacillus group. pp. 281-287 in Haemophilus, Pasteurella, and Actinobacillus. M. Kilian, W. Frederiksen, and E. L. Biberstein, eds., Academic Press Ltd., San Francisco, California, 294 pp. MIELKE, P. W., AND K. J. BERRY. 1992. Fisher's exact probability test for cross-classification tables. Educational and Psychological Measurement 52: 97-101. MILLER, M. W. 1990. Animal and pen support facilities for mammals research. Pages 45-63 in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-3, Work Plan la, Job 1. Colorado Division of Wildlife, Fort Collins, Colorado, USA. _____ , N. T. HOBBS, AND M. A. WILD. 1991a. Experiments to identify and manage stress in mountain sheep populations. Pages 101-122 in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-4, Work Plan 2a, Job 4. Colorado Division of Wildlife, Fort Collins, Colorado, USA. �201 , AND E. S. WILLIAMS. 1991b. Spontaneous pasteurellosis in captive Rocky Mountain bighorn sheep (Oyis canadensis canadensis): clinical, laboratory, and epizootiological observations. Journal of Wildlife Diseases 27:534-542. MORRISON, D. F. 1976. Multivariate statistical Co., New York, New York, pp. 145-194. NASH, methods. McGraw-Hill Book P. 1972. Development of a Pasteurella bacterin for bighorn sheep (Qv.ll canadensis canadensis). Ph.D. Thesis, Colorado State University, Fort Collins, Colorado, 102 pp. ONDERKA, D. K., AND W. D. WISHART. 1984. from pneumonia in southern Alberta. Northern Wild Sheep and Goat Council A major bighorn sheep dieoff Biennial symposium of the 4:356-363. , S. A. RAWLUK, AND W. D. WISHART. 1988. Susceptibility of Rocky Mountain bighorn sheep and domestic sheep to pneumonia induced by bighorn and domestic livestock strains of Pasteurella haemolytica. Canadian Journal of Veterinary Research 52: 439-444. POST, G. 1962. Pasteurellosis of Rocky Mountain canadensis). Wildlife Disease 23:1-13. REGG IARDO , C. 1981. Epidemiological in Experimental bighorn sheep (Qv.ll Serological tests in bovine bacterial pneumonias. application and potential as a research tool. Advances Medicine and Biology 137:818-822. RUFI, V. G. 1961. Study of pasteurellosis in bighorn sheep. Pages 21-27 in Federal Aid in Wildlife Restoration, Game & Fish Laboratory Research, Investigations Projects, Job Completion Report, Project Number FW-3-R-8, Work Plan 1, Job 2W. Wyoming Game and Fish Department, Cheyenne, Wyoming, USA. SAS INSTITUTE, INC. 1989. SAS/STA~ Volume 2. SAS Institute, Cary, User's Guide, Version 6, Fourth North Carolina, 846 pages. Edition, SHEWEN, P. E., AND B. N. WILKiE. 1988. Vaccination of calves with leukotoxic culture supernatant from Pasteurella haemolytica. Canadian Journal of Veterinary Research 52:30-36. SILFLOW, R. M., W. J. FOREYT, S. M. TAYLOR, W. W. LAEGRIED, H. D. LIGGITT, AND R. W. LEID. 1989. Comparison of pulmonary defense mechanisms in Rocky Mountain bighorn sheep (Oyis canadensis canadensis)and domestic sheep. Journal of Wildlife Diseases 25:514-520. ___ ___ ' , AND • 1991. Comparison of arachidonate metabolism by alveolar macrophages from bighorn and domestic sheep. Inflammation 15:43-54. ___ , AND R. W. LEID. 1993. Pasteurella haemolytica cytotoxindependent killing of neutrophils from bighorn and domestic sheep. Journal of Wildlife Diseases 29:30-35. �200 SUTHERLAND, A. D., W. DONACHIE, G. E. JONES, AND M. QUIRIE. 1989. A crude cytotoxin protects sheep against experimental Pasteurella haemolytica serotype A2 infection. Veterinary Microbiology 19:175-181. WARD, A. C. S., L. R. STEVENS, B. J. WINSLOW, R. P. GOGOLEWSKI, D. C. SCHAEFER, S. K. WATSON, AND B. L. WILLIAMS. 1986. Isolation of Haemophilus somnus: a comparative study of selective media. American Association of Veterinary Laboratory Diagnosticians 29:479-486. WILD, M. A., AND M. W. MILLER. 1991. Detecting nonhemolytic Pasteurella haemolytica infections in healthy Rocky Mountain bighorn sheep (~ canadensis canadensis): Influences of sample site and handling. Journal of Wildlife Diseases 27:53-60. AND 1994. Effects of modified Cary and Blair medium on recovery of nonhemolytic Pasteurella haemolytica from Rocky Mountain bighorn sheep (Oyis canadensis canadensis) pharyngeal swabs. Journal of Wildlife Diseases 30: 16-19. Prepared by _ Michael W. Miller Wildlife Research Veterinarian �Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS State of Colorado Project No. W-153-R-9 Work Plan No. Job No. 2A REPORT Mammals Research Mountain Sheep Inyestigations Experimental Evaluation of Mountain Sheep Transplanting and Disease Treatment Period Covered: July 1, 1995 - June 30, 1996 Authors: M. W. Miller, J. Vayhinger, Jurgens Personnel: R. Dobson, J. Duran, B. Elkins, J. George, D. Getzy, R. Green, R. Hancock, M. Lamb, R. Myers, S. Ogilvie, G. Roberts, B. Thornton, R. Zaccagnini. S. Roush, T. Verry, A. Torres, and V. ABSTRACT We conducted a 4-year management experiment to examine the effects of alternative lungworm treatment strategies on lamb survival and population performance in Rocky Mountain bighorn sheep (Ovis canadensis canadensis) herds in southcentral Colorado. Beginning in December 1991, 2 bighorn herds in the Tarryall Mountains and 2 herds in the Collegiate Peaks were managed under 1 of 4 alternative lungworm treatment regimes: baiting with alfalfa hay and apple pulp treated with fenbendazole (about 3 g/adult ewe) (B/T), baiting with alfalfa hay and apple pulp without fenbendazole (B), placing fenbendazoletreated salt blocks (1.65 g fenbendazole/kg) on winter ranges (T), and withholding bait and fenbendazole (control) (C). Treatments were rotated annually under a predetermined, randomly-selected schedule. We monitored lamb production and survival among radiocollared ewes in each herd from May through October 1991-1995. Both production and survival varied widely and affected recruitment (lambs/marked ewes) through October. Annual recruitment rates through October ranged from 0.13 to 0.88, but mean recruitment rates did not differ among herds (P = 0.51) or between years (P = 0.80). Among the 4 study herds, sick and coughing lambs were observed every summer in 1 herd and 1 summer (1995) in a second. Mean lamb production (estimated by observations of marked ewes with < 2 wk old lambs at heel) ranged from 0.83 (B/T) to 0.95 (B), but did not differ among treatments (P ~ 0.23). Mean lamb survival (lambs in October/lambs born to marked ewes) through October ranged from 0.59 (B/T) to 0.81 (T), and was also unaffected by management treatment (P ~ 0.15); moreover, baiting combined with fenbendazole administration failed to prevent catastrophic (86%) lamb mortality in 1 herd. Overall, neither baiting (P = 0.45) nor fenbendazole treatment (P = 0.62) enhanced lamb recruitment among �204 these 4 apparently healthy bighorn populations during the 4 years of our study. Our results demonstrate that annual parasite treatment is not prerequisite for lamb survival among southcentral Colorado's wild bighorn populations. Moreover, annual parasite treatment may not prevent catastrophic losses of lamb cohorts among treated herds. Based on these data, we question the need for annual baiting and parasite treatment in the herds studied here and believe such practices need to be reevaluated elsewhere as well. In making these reevaluations, we encourage managers to weigh the potential benefits of baiting and treatment against the "costs ••of such interventions those costs may include locally increased bighorn densities (that could lead to infectious disease outbreaks), alteration or loss of movement and distribution patterns, increased vulnerability to predators (including man), and dependence on artificial feed sources. In light of the potential detrimental effects of baiting and treating wild sheep herds annually, we recommend that anthelmintic therapy be reserved for specific situations where verminous pneumonia in lambs is not only documented, but is occurring at rates sufficient to depress long-term performance of individual bighorn populations. �205 EXPERIMENTAL EVALUATION TRANSPLANTING OF MOUNTAIN AND DISEASE SHEEP TREATMENT M. W. Miller, J. Vayhinger, S. Roush, T. Verry, A. Torres, and V. Jurgens P. N. OBJECTIVE Design, conduct, and report on management experiments to evaluate efficacy transplanting and disease treatment practices for managing mountain sheep populations. of AGREEMENT OBJECTIVE Complete parasite a management level experiment control program. evaluating Colorado's mountain sheep Rocky Mountain bighorn sheep (Ovis canadensis canadensis) populations throughout North America are plagued by pneumonia outbreaks caused by a complex of bacterial, parasitic, and/or viral agents. These epizootics and subsequent population declines represent a significant obstacle to long-term success in bighorn sheep management. Treating with anthelmintics to reduce parasitic lungworm (Protostrongylus spp.) burdens has become an integral part an aggressive management program to reduce disease outbreaks and enhance productivity of mountain sheep herds in Colorado. Although some combination of treating, trapping and other management activities has increased sheep numbers statewide over the last 2 decades, it is unclear which strategies were most influential in this achievement. In light of the growing number of significant mountain sheep herds throughout Colorado, the apparent intensity of management required to perpetuate individual populations, and the costs associated with applying intensive management, it has becoming increasingly important to explore alternatives for efficient but efficacious management practices. Here, we describe a management experiment conducted to examine effects of alternative lungworm treatment strategies on bighorn lamb survival and population performance. Our experiment tested the hypothesis that bighorn lamb production and survival rates for herds would not differ in re'sponse to various management treatments. MATERIALS AND METHODS Beginning in December 1991, we began managing each of 4 study herds [Tarryall Mountains: Twin Eagles (TE) and sugarloaf (SL); Collegiate Mountains: Chalk Creek (CH) and Cottonwood Creek (CW)] under 1 of 4 alternative lungworm treatment regimes: Control no treatment -- bait and fenbendazole withheld (C); �206 Treat OnlyBait Only Bait/Treat- fenbendazole-treated salt blocks placed on bait stations (T); baited with alfalfa hay and apple pulp but not treated with fenbendazole (B); baited with alfalfa hay and apple pulp and treated with fenbendazole (BIT). Treatments were assigned to study herds as prescribed rotating schedule (Table 1.; Year 1 = 1992). by a randomly selected, Table 1. Treatment assignments for 4 bighorn herds included in a 4-year management experiment to examine effects of alternative lungworm treatment strategies on bighorn lamb survival and population performance. HERD QQLLE~IAIE YEAR CHALK CREEK MQIlliIAIHS TARRYALL MQIlNIAINS COTTONWOOD CREEK SUGARLOAF MOUNTAIN TWIN EAGLES 1992 B1 C T BIT 1993 BIT T B. C 1994 C B BIT T 1995 T BIT C B Treatment assignments: BIT = bait with alfalfa hay and apple pulp treated with fenbendazole; B = bait with alfalfa hay and apple pulp without fenbendazole; T = fenbendazole-treated salt blocks on bait stations; and C = withhold all bait and fenbendazole (control). We attempted to apply experimental treatments uniformly across the 4 study herds. We baited with third cutting alfalfa hay (about 2 kg/head/day) and fermented apple pulp (about 1 kg/head/day) at both B and BIT sites for about 8-10 weeks beginning in mid-December. In addition, B/T sites were also treated with fenbendazole (about 3 g/adult ewe) added to apple pulp on 2 occasions 1 to 2 wks apart. Four or five treated salt blocks (1.65 g fenbendazole/kg; 15 kg/block) were available to sheep at T sites during January to May; 1 additional block held in a wire cage during that same period was used as an environmental control. Blocks were replaced if they disappeared before 1 May. For Band B/T herds, we recorded baitsite attendance and receipt of fenbendazole treatments for each radiocollared ewe. For T sites, we weighed medicated blocks to estimate consumption but were otherwise unable to document exposure of marked ewes to the treatment. Control herds were monitored periodically but received no other management intervention. Adult bighorn ewes from each herd were initially captured using drop nets in February and March 1991 and marked with individually identifiable radiocollars; collar symbols, color combinations, and radio frequencies were �207 identifiable to individuals and herds of origin. Additional adult ewes were captured opportunistically in some herds each of the following winters to achieve target sample sizes (15 to 20 ewes per herd) or replace mortalities. We used net gunning, aerial darting, or darting from the ground in supplemental captures. No anthelmintics or antibiotics were used during supplemental captures. Each year, we assessed effects of winter treatments on lamb production and survival by observing radiocollared ewes (n = 11 to 18) from all 4 herds about once every 2 weeks from May through October to determine whether they produced lambs, and whether their lambs were still alive and healthy. We defined annual lamb production as the proportion of marked ewes seen with a lamb nursing or at heel at least once between May and October. Annual lamb survival was the proportion of lambs seen with marked ewes that were still alive at the end of October. Annual lamb recruitment, the product of these two processes, was the proportion of marked ewes with lambs alive at the end of October. We recognize that the foregoing approach may have slightly underestimated production and overestimated survival by misclassifying periparturient mortality as failed production. However, because vermipous pneumonia primarily affects survival of lambs >6 wks of age, we believed our approach more sensitively measured the treatment responses of primary interest in this study. In. addition to lamb survival data, we recorded approximate UTM coordinates, habitat type, and group size and composition for each radiocollared ewe observed. All field data were transcribed into a computerized database to aid in mapping seasonal range movements and determining annual lamb production and survival rates. Radiocollared ewes were also monitored every 2-4 weeks to detect mortality and movements during November through April in conjunction with a USFS/CDOW cooperative project to identify critical winter and transitional ranges of these 4 herds. These data will be reported separately elsewhere. RESULTS AND DISCUSSION All marked ewes maintained fidelity to their respective herds of origin throughout the study. Consequently, we believe treatments were applied ewes within each herd only as prescribed by the foregoing experimental schedule. Treatment to Rates As controls, herds received no treatments or intervention (aside from supplemental captures) and were essentially undisturbed by management activities other than monitoring or inventory during December-May. Use of treated salt blocks varied considerably among herds. Adjusting for estimated environmental losses, block consumption equated to about 13 ewe treatments (2 X 3 g/ewe) at CH, 26.5 at CW, 6 at SL, and 6.5 at TE; under a moderate dosing regime (3 X 0.75 g/ewe) (Foreyt and Coggins 1990), block consumption equated to about 35 ewe treatments at CH, 70 at CW, 16 at SL, and 17 at TE. We observed marked and unmarked sheep using blocks on several occasions in late January-April, but mule deer or elk also may have used treated blocks at some sites. Apparent differences in salt block consumption between study herds in the Collegiate Peaks and Tarryall Mountains are �208 supported by field observations suggesting marked differences in affinity for natural mineral licks between ewes in these 2 discontinuous mountain ranges. Whether these observed differences reflect natural mineral deficiencies in some ranges and/or greater abundance of salt associated with domestic stock grazing remains undetermined. However, our data suggest such differences, combined with consumption by other species, could influence the potential use and efficacy of fenbendazole-treated salt blocks among diverse bighorn sheep ranges in Colorado and elsewhere. Responses to bait, with or without fenbendazole, were generally more consistent than to treated blocks in 3 of the 4 study herds. This may be in part because all of the herds we used had been baited and treated annually for a decade or more preceding initiation of this experiment. In years when bait alone was applied, marked ewes averaged (± sd) 42 (± 13.7) days on bait at CH, 26 (± 15.1) days at CW, 38 (± 1.9) days at SL (6 days missing data), and 44 (± 1.0) days at TE. In contrast to the other 3 herds, most marked ewes at CW visited the bait site infrequently during December-January. Many ewes in this herd stayed on alpine winter ranges until heavy snows apparently forced them to lower elevations in February, when CW baitsite attendance increased markedly. In years when baiting was combined with fenbendazole treatment, marked ewes averaged (± sd) 51 (± 10.8) days on bait at CH, 23 (± 6.3) days at CW, 45 (± 1.1) days at SL, and 47 (± 4.5) days at TE. In addition to bait, 14 (93%) of 15 marked ewes at CW, 8 (50%) of 16 marked ewes at CW, and all 17 marked ewes at both SL and TE also were present to presumably receive at least 1 fenbendazole treatment during their scheduled treatment periods. The relatively low treatment rate at CW was attributed to failure of marked ewes to remain on low elevation winter ranges in 1995 despite daily baiting. Lamb Production and Survival Both production and survival varied widely and affected recruitment (lambs/marked ewes) through October. Annual recruitment rates through October ranged from 0.13 to 0.88; however, mean recruitment rates did not differ among herds (P = 0.51) or between years (P = 0.80). Among the 4 study herds, sick and coughing lambs were observed every summer in the Chalk Creek herd and during the summer of 1995 in the Cottonwood Creek herd; no sick lambs were seen in either of the Tarryall Mountain herds during the 4-year study period. Mean lamb production ranged from 0.83 (B/T) to 0.95 (B), but did not differ among treatments (P ~ 0.23). Mean lamb survival (lambs in October/lambs born to marked ewes) through October ranged from 0.59 (B/T) to 0.81 (T), and was also unaffected by management treatment (P ~ 0.15); moreover, baiting combined with fenbendazole administration failed to prevent catastrophic (86%) lamb mortality in the Cottonwood Creek herd in 1995. Although recruitment varied widely, neither baiting (P = 0.45) nor fenbendazole treatment (P = 0.62) enhanced lamb recruitment among these 4 apparently healthy bighorn populations during the 4 years of our study. Range Use and Movement Patterns Relatively consistent and predictable range use and movement patterns for each of the 4 study herds have e~erged since monitoring began in 1991. Plots of May 1991-March 1994 location data (UTM coordinates) for radiocollared ewes revealed apparent differences in distribution and movement patterns among �209 herds (Fig. 2). Ewes from the CW herd have consistently shown widest distribution and greatest movements; CH ewes were the most limited in their range use and movement. Although ranges of the SL and TE herds appeared to overlap considerably, to date we have observed no exchange of radiocollared ewes between these 2 herds. Disturbances by hikers (CW) and hunters (CH, SL) appeared to influence movements of ewe/lamb groups on occasion. Location data gathered since March 1994 will be added to further define key ranges and migration corridors for these 4 herds. Population Parameters and Performance Overall, noncapture mortality rates of adult ewes in these 4 herds averaged about 0.08 (se = 0.01) annually over the 56 months covered by our study, but causes and annual rates (ranging from 0 to 0.28 at CW in 1994) of ewe mortality appeared to vary among herds. No consistent health problems were detected among marked ewes. In total, 20 radiocollared ewes (2 at CH, 5 at TE, 5 at SL, and 8 at CW) died of noncapture causes between February 1991 and October 1995. Of these, lion predation appeared to have caused 6 losses in the Tarryalls (3 at TE and 3 at SL), injuries from falls may have killed 2 ewes (at CW), pneumonic pasteurellosis killed 1 ewe (at CW), and lightning claimed 1 ewe (at CW); causes of death or disappearance for 10 other ewes (2 at CH, 2 at TE, 2 at SL, and 4 at CW) could not be determined, although we speculated lightning strikes also may have been involved in 2 deaths and 2 disappearances at CW and 1 death at CH during late June-early July. Six of 10 ewe mortalities in the Collegiate Peaks herds since 1991 occurred in apparently healthy ewes during mid June-mid August and may have been lightning-caused; whether telemetry collars are a predisposing factor in these losses was uncertain. Despite observed variation in recruitment and adult mortality rates, winter range counts during 199.1-1995 suggest all 4 bighorn herds under study remained stable or grew during the course of this experiment. CONCLUSIONS AND MANAGEMENT RECOMMENDATIONS OUr results demonstrate that annual parasite treatment is not prerequisite for lamb survival among southcentral Colorado's wild bighorn populations. Moreover, annual parasite treatment may not prevent catastrophic losses of lamb cohorts among treated herds. These findings should not be altogether surprising -- there are clearly a multitude of factors besides disease that can influence lamb production and survival in bighorn populations. And even in herds where signs of respiratory disease are observed among lambs, anthelmintic therapy in ineffective in treating bacterial pneumonia (e.g., pasteurellosis) that appears to be far more prevalent than verminous pneumonia among bighorn.populations in Colorado and elsewhere. Based on these data, we question the need for annual baiting and parasite treatment in the herds studied here and believe such practices need to be reevaluated elsewhere as well. In making these reevaluations, we encourage managers to weigh the potential benefits of baiting and treatment against the "costs" of such interventions -- those costs may include locally increased bighorn densities (that could lead to infectious disease outbreaks), alteration or loss of movement and distribution patterns, increased vulnerability to predators (including man), and dependence on artificial feed sources. �210 In light of the potential detrimental effects of baiting and treating wild sheep herds annually, we recommend that anthelmintic therapy be reserved for specific situations where verminous pneumonia in lambs is not only documented, but is occurring at rates eufficient to depress long-term performance of individual bighorn populations. Prepared by Michael W. Miller Wildlife Reeearch Veterinarian �211 Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS State of project Colorado No. W-153-R-9 Work Plan No. 3A Job No. Period Author: REPORT Mammals Research Pronghorn Inyestigations Habitat Selection and population Performance of a Pioneering Pronghorn Population Covered: July 1, 1995 - June 30, 1996 T.M. Pojar ABSTRACT The rate of increase (ROI) for the Middle Park pronghorn population continues the downward trend as the population size increases. The projected late summer 1996 population size is 584 (Table 3). Using the linear regression technique (ROI on population size) to estimate carrying capacity, the projected K-value for this population is 625 animals (Figure 1). During late winter 1995-96, there was a distinct shift in winter distribution to the Wolford Mountain area. Deep, late winter snow and/or snowmobile traffic on the traditional Red Mountain wintering area may be the cause. The third cohort of fawns (10 males and 10 females) were equipped with solar powered radio ear tags in December 1995 as part of the objective for estimating differential natural mortality between males and females. Sixty radios have been put on fawns of the year (1993, 1994, and 1995). Monitoring continues. ��213 HABITAT S.ELECTION AND POPULATION PERFORMANCE OF A PIONEERING PRONGHORN POPULATION Thomas P • N. Describe population dynamics pronghorn population. M. Pojar OBJECTIVE and habitat use of a pioneering, expanding SEGMENT OBJECTIVES 1. Describe seasonal population. and annual distribution 2. Monitor natural mortality males and females. 3. Map areas of habitation 4. Monitor population dynamics of Middle Park pronghorn with: a. Ground counts to describe changes in population size. b. Ground counts to quantify population sex and age composition. and movement using the GIS of the Middle patterns Park pronghorn of radioed yearling format. STUDy AREA The study area is described (1993) in Pojar (1988) and a map of the area is in Pojar METHODS AND MATERIALS SEASONAL AND ANNUAL DISTRIBUTION Tracking was done mostly from the ground; fixedwing aircraft was used if an animal could not be located after a reasonable effort from the ground. Legal descriptions of animal locations were recorded to the nearest quarter mile then converted to UTM (U.S. Army 1973) coordinates for computer processing. All radioed animals have been located biweekly (with very few exceptions) since January 1, 1987. POPULATION SIZE AND STRUCTURE Herd structure estimates were obtained by classifying all animals that accompanied the animals that are radioed. The herd structure estimate used in population projections is the one with the largest sample size obtained in August or September. Total counts are made during winter by counting all animals associated with radioed animals. With the increased population size, it is not always possible to get an accurate count of total mature bucks (1.5 yrs and older) in the population. However, it is still possible to get very accurate counts of bucks in 60-80% of the population. The proportion of bucks in this portion of the population is then extrapolated to the total population to estimate total mature bucks. Total population count during winter, �214 estimated number of mature bucks from the winter count, and recruitment based on fawn to doe ratios from late summer are used for the population projection. Population 1. 2. 3. 4. NATURAL projections are based on the following assumptions: Winter counts represent the total population and the estimated number of mature bucks in Middle Park. Late summer age ratio estimates represent "recruitment" into the population. Annual survival of mature bucks and does and female fawns is 92.5%. Annual survival of males in their first year (after weaning) is 50%. (This severe mortality on male fawns is arbitrary, however, it allows the number of mature males in subsequent years to match fairly well with winter counts.) MORTALITY OF MALES AND FEMALES The methods for estimating differential natural mortality between female pronghorn are outlined in Appendix I of Pojar (1994). male and RESULTS SEASONAL AND ANNUAL DISTRIBUTION The winter dis~ribution during 1994-95 was similar to the previous winter distribution through mid-January. Groups of 50+ spent the early part of the winter in Sulphur Gulch and Antelope Creek vicinities. These groups all coalesced into a much smaller area on the south slopes of Wolford Mountain as the snow got deeper. They occupied this area through spring thaw and coexisted there with several hundred deer and elk. POPULATION SIZE AND STRUCTURE Total population size estimates are obtained during winter and herd structure estimates are obtained in late summer (Table 1). The annual changes in population size are used to calculate the rate of increase which is regressed on population size to project the K-value for the population. The ROI is calculated as where P1 is the population size at time 1 and P2 is the population at time 2 (Table 2). The rate of increase for 1995-96 is 0.15 (Table 2). Based on the relationship of population size and ROI, the projected K-value is 625 (Figure 1). �215 Table 1. Herd structure of Middle Park pronghorn based on a sample obtained by locating radioed animals in late summer. The population size is from the subsequent winter counts with harvest added back into the population to get the pre-hunt population size, e.g. 1995 pre-hunt population was 535, 520 winter count plus 15 harvest. YEAR POP. SIZE NO. RADIO RADIO B:lOOD RATIO F: lOOD RATIO SAKPLE % OF POP. 1986' 1987 1988 1989 1990 1991 1992 1993 1994 1995 80 122 160 223 261 308 347 425 466 535 7 24 22 17 13 39 31 58 52 5.7 15.0 10.2 6.5 4.2 11.2 7.3 12.4 9.7 36 54 40 56 22 23 26 10 29 32 77 77 32 50 47 65 48 66 46 42 47 63 108 161 148 148 286 266 332 437 59 52 68 72 66 48 82 63 71 82 , This year'.s data based 1986. % on the sample of the population trapped 16 December Table 2. Population size of the Middle Park pronghorn herd during winter and the calculated rate of increase. Population size reflects the removal of 1315 animals per year by harvest beginning in 1990, i.e. the 1994-95 winter population was 466 before harvest and 453 after harvest. YEAR 1986-87 1987-88 1988-89 1989-90 1990-91 1991-92 1992-93 1993-94 1994-95 1995-9.6 1996-97 (Projected) POP. 80 122 160 223 246 292 332 410 453 520 584 SIZE RATE OF INCREASE .52 .31 .39. .10 .19 .14 .23 .10 .15 .12 �216 Table 3. Population projection text for the assumptions. I POPULATION I BUCKS for the Middle I DOES Park pronghorn population. I I FAWNS See I TOTAL WINTER '95-96 90 302 128 520 WINTER MORTALITY 90 X .075 = 7 MORT 302 X.075 = 23 MORT 64X.5=31B 64X.075=5D 66 PREFAWNING 1996 90 - 7 = 83 MATURES + 21 YRLS TOTAL =104 302 - 23= 279 MATURES + 59 YRLS TOTAL = 338 LATE SUMMER 1996 MATURE 83 YRLS 21 TOTAL 104 MATURE 279 YRLS 59 TOTAL 338 442 @ 42F:I00D 338 X .42 = 142 FAWNS 584 The winter of 1995-96 was relatively mild through January with low snow fall and accumulation, and no extended periods of sub-zero (Fo) temperatures. About mid-January, heavy snows and deep accumulation in higher elevations forced several hundred deer and elk into the general vicinity of the pronghorn wintering area. The accumulation on the wintering area constricted pronghorn movement and limited foraging areas. There was no mortality of pronghorn observed on the wintering area and they did not appear to be severely stressed nutritionally. Both deer and elk did show signs of nutritional stress but there was no stress related mortality observed. NATURAL MORTALITY OF YEARLING MALES AND FEMALES For the second year, the net-gun technique (Helicopter Wildlife Management, Salt Lake City, Utah) was used to radio 10 male and 10 female fawns in December, 1995. Thus far, 48 pronghorn have been captured using this technique with 1 known capture related mortality. To avoid the problem of accommodating neck growth and neck swelling during the rut for males, solar power transmitters mounted on ear tags were used. As of this writing, the dependability of these radios is disappointing. Signal transmission is sporadic depending on the angle of the solar panels to direct sunlight. Six of the 20 radios have not been located for> 60 days. All of the radios (40) deployed in 1993 and 1994 be accounted for (Tables 4 and 5). One animal (female) with radio (149.510) moved to North Park and was seen on the south side of Independence Mountain (TIIN,R81W,S36,NW) on March 16,1996; the last location was also in North Park on June 21, 1994.· This is the farthest known movement of a pronghorn radioed in Middle Park and the longest time span between locations. Of the 20 solar powered ear tag transmitters deployed in December males and 3 females) have not been located for> 60 days. 1995, 6 (3 �217 Table 4. Record of sex, ear tag numbers, radio frequency, and status of fawns radio collared on December 14, 1993 in Middle Park, Colorado (T2N,R80W,S36). Ear tag designation Y=yellow and B=blue. Sex F F F F F F F F F F M M M M M M M M M M Ear Tag Y3 Y5 Y8 Y9 Y10 Yll Y12 B53 B54 B55 Y1 Y2 Y4 Y6 Y7 B56 B57 B58 B59 B60 Radio Frequency 148.500 149.230 149.150 149.272 149.502 149.512 149.490 148.760 148.730 149.430 149.650 149.172 149.450 149.410 149.470 149.390 149.190 149.550 149.530 149.132 Status Alive Alive Alive Alive Alive N. Park, Last Radio slipped Alive N. Park, Last Alive N. Park, Last Collar broke Collar broke Collar broke Alive Alive Alive Alive Radio quit Alive 3/19/96 . off, recovered 7/10/96 1/12/96 and recovered and recovered and recovered Table 5. Record of sex, ear tag numbers, radio frequency, and status of fawns radio collared on December 12-13, 1994 in Middle Park, Colorado. Ear tag designation Y=yellow and B=blue. Sex F F F F F F F F F F M M M M M M M M M M Ear Tag B61 Y20 Y13 B62 Y15 Y22 Y23 Y24 B64 Y19 B65 Y21 B67 B66 Y18 Y25 Y16 B63 Y14 Y17 Radio Frequency 148.030 148.050 148.060 148.100 148.150 148.160 148.190 148.280 148.300 148.320 148.010 148.020 148.040 148.070 148.080 148.090 148.120 148.140 148.170 148.180 Status N. Park, last 7/10/96 Alive Alive Alive Auto kill, 12/23/94 Alive Alive Alive Alive Alive Alive Alive Auto kill, 10/16/95 Alive Alive Alive Alive Alive Alive Trapping mortality, 12/17/94 �216 REFERENCES CITED McCullough, D. R. 1994. What do herd composition Soc. Bull. 22:295-300. Pojar, counts tell us? Wildl. T.M. 1988. Habitat selection and population performance of a pioneering pronghorn population. Colo. Div. Wildl. Res. Rep. July, 181-192. 1993. Habitat selection and population pronghorn population. Colo. Div. Wildl. performance of a pioneering Res. Rep. July, pp 199-207. 1994. Habitat selection and population pronghorn population. Colo. Div. Wildl. performance of a pioneering Res. Rep. July, pp 125-136. u.S. Army. 1973. Technical Manual: Headquarters, Dep~ of the Army, pp Universal Transverse Mercator Grid. Washington D.C. TM No. 5-241-8, 64 pp. 1 0.9 0.8 CD CI) 1995 0.7 CO Q) "-0 C 'f- 0.6 '87 0.5 0 CD CO • '89 0.4 ~ • 0:::0.3 '93 '91 0.2 0.1 '90 • '92 • '95 • • 0 0 100 200 300 Population Figure 1. population 400 500 600 Size Projected carrying capacity (K-Value) of middle based on regression of ROI on population size. Park pronghorn 700 �219 Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS State of Project REPORT Colorado No. W-153-R-9 Mammals Research Work Plan No. Pronghorn Job No. Detecting Density Dependence Natural Populations Period Author: Covered: July Inyestigations in 1, 1995 - June 30, 1996 T. M. Shenk ABSTRACT The final results of this study are reported in T. M. Shenk's Ph.D. Dissertation. The Abstract of that Dissertation is included in this report. A copy of the entire document can be obtained by contacting R. B. Gill, Colorado Division of Wildlife, 317 W. Prospect, Fort Collins, CO 80526. ��221 ABSTRACT DETECTING DENSITY OF DISSERTATIO~ DEPENDENCE IN NATURAL POPULATIONS A review of methods developed to detect density dependence from temporal trends in natural populations highlighted similarities, strengths, weaknesses and applications of the various techniques. Past evaluations and criticisms. of the methods were included in the review. Attempts to detect density dependence from regression techniques relating either population abundance or segments of popUlation abundance (key factors) at time t + 1 to population abundance at time t were criticized for spurious correlations resulting from the non-independence of.observations. Techniques developed to address this time-series dependence in population abundances (e.g., autoregression, and tests for randomness, limitation and attraction) either are not robust to sampling error or lack power. Tests for direction and significance of curvature in population growth curves show promise as a management tool to determine i~ harvested populations are above or below a maximum net productivity level. Techniques developed to relate survival or reproduction to population abundances through correlation or logistic regression are valid if (1) conducted on independent parameter estimates to avoid spurious results and (2) analyses are adjusted for overdispersion of the response variable. Detecting· density effects on survival may also be done within the extensive theory developed for parameter estimation using mark-recapture data. Testing for compensatory mortality as a specific form of density dependence has served as a management tool for harvesting of waterfowl. Of the methodologies reviewed, if test assumptions are met, or tests are robust to assumption violations, power increases with increasing sample size, strength of density dependent response, and intrinsic growth rate. Thus, if these tests are to be used for valid inference to support or refute the presence of density dependence in a natural population, the tests must be robust to the presence of sampling and process variation in the data. Therefore robustness of five methods was evaluated when sampling and process variation occurred in the data. First, Monte Carlo simulations were conducted to evaluate robustness of four tests developed to detect density dependence from series of population abundances, to the addition of sampling variance. Population abundances were generated from random walk, stochastic exponential growth, and densitydependent population models. Population abundance estimates were generated with sampling variances distributed as lognormal and constant coefficients of variation (CV) from 0.00 to 1.00. In general, when data were generated under a random walk, Type I error rates increased rapidly for Bulmer's R, Pollard et al.'s, and Dennis and Taper's tests with increasing magnitude of sampling variance for n > 5 years and all values of process variation. Bulmer's R* test maintained a constant 5% Type I error rate for n > 5 years and all magnitudes of sampling variance in the population abundance estimates. When abundances were generated from two stochastic exponential growth models (R = 0.05 and R = 0.10) Type I errors again increased with increasing sampling variance; magnitude of Type I error rates being higher for the slower growing population. Therefore, sampling error inflated Type I error rates, invalidating the tests, for all except Bulmer's R* test. Comparable simulations for abundance estimates generated from a density-dependent growth rate model were conducted to estimate power of the tests. Type II error rates were influenced by the relationship of initial population size to carrying capacity (K), length of time series, as well as sampling error. Given the inflated Type I error rates for all but Bulmer's R*, power was overestimated for the remaining tests resulting in density dependence being detected more �222 often than it existed. Population abundances of natural populations are almost exclusively estimated rather than censused, assuring sampling error. Therefore, because these tests have been shown to be either invalid when only sampling variance occurs in the population abundances (Bulmer's R, Pollard et al.'s, and Dennis and Taper's tests) or lack power (Bulmer's R* test), little justification exists for use of such tests to support or refute the hypothesis of density dependence. Alternatively, logistic regression can be used to detect density dependence by coupling abundances as the explanatory variable, with the binary demographic parameter hypothesized to be density-dependent as the response variable. Therefore, logistic regression was evaluated as a feasible method to detect density dependence from temporal trends in demographic parameters. Data were generated from a 3-age class population growth model designed to mimic mule deer (Odocoileus hemionus) population growth. Logistic regression was then performed with overwinter fawn survival as the response variable and density as the explanatory variable. To estimate Type I error rates all demographic parameters were held constant, resulting in density-independent growth. To estimate Type II error rates, overwinter fawn survival was modeled as density-dependent. Monte Carlo simulation was used to evaluate the effect of sampling variance in both the response variable, fawn survival rate, and the explanatory variable, population density. Fawn survival rate estimates were determined from a binomial distribution for a sample of fawns from the population. Density estimates were generated by adding a log-normally distributed sampling variance to the population densities. Increasing process variation, creating extra-binomial variation, was also included in the response variable by adding a random variable to the overwinter fawn survival rates. Data were analyzed using logistic regression assuming (1) only binomial variation in the fawn survival rates and (2) assuming survival rates were overdispersed. Process variation (overdispersion) increased Type I error rates beyond the expected 5% when extra-binomial variation was not accounted for in the analysis, artificially inflating power (1 - Type II error). Type I error rates remained at .5% for increasing process variation if an overdispersion parameter was estimated and used in the analysis. When overdispersion of the fawn survival rate was considered 'in the analysis, power increased as (1) estimates of fawn survival and density became more precise, (2) time series length increased, and (3) time series included wider ranges of densities. Manipulations were simulated to best achieve (3). The stability of the Type I error rate when sampling variance was. added to the response and explanatory variables as well as overdispersing the response variable, suggests logistic regression, adjusted for an overdispersed response variable, is a valid method for detecting density dependence in survival rates. Given the validity of the test, guidelines were suggested to maximize power and broaden inference. In summary, of the five methods evaluated in this study, only logistic regression is a feasible approach for detecting density dependence in natural populations. Department Tanya Marie Shenk of Fishery and Wildlife Biology Colorado State University �Colorado Division Wildlife Research July 1996 of Wildlife Report JOB FINAL REPORT state of Project Colorado No. W-153-R-9 Mammals Research Work Plan No. Pronghorn Research Job No. Pronghorn Winter Period Covered: Authors: July 1, 1992 - December D. C. strohmeyer, G. C. White, Wheat Damage study 31, 1995 and R. B. Gill ABSTRACT Wildlife managers have responded to winter wheat damage complaints by reducing pronghorn (Antilocapra americana) numbers. These reductions may not be necessary because our telemetry research supported the idea that pronghorn may not damage winter wheat. Winter wheat becomes vulnerable to grazing damage only after it enters the jointing developmental (phenological) stage. Marked, free-ranging pronghorn naturally stopped foraging on winter wheat before wheat began to joint in our study. Torbit et al. (1993) speculated that this behavior was stimulated by the concurrence of native forage regrowth and declining forage quality of wheat. Our feeding-trial research focused on pronghorn response to forage quality changes in wheat as it developed and became vulnerable to grazing damage. OUr treatments were 2 phenological stages of wheat: tillering (younger) and jointing. Our preliminary trials suggested that pronghorn selected the highest biomass when presented small quantities of wheat. We tried 2 more feeding-trial designs to eliminate this hypothesized biomass selection. The first equalized plant height, the second incorporated more biomass. The new designs altered pronghorn choice. Pronghorn response to the treatments appeared random (~ ~ 0.96) for both experimental designs. Our observations of pronghorn behavior suggested that the novel opportunity to consume wheat, vastly exceeded.interest in differentiating between the tillering and jointing stages. This implied that to quantify pronghorn preference for different stages of wheat, we need to increase the ecological scale of the test. That is, the amount of biomass presented and the length of the feeding trial needs to approach (at least) that encountered during normal pronghorn feeding bouts. A possible alternative design in which the ecological scale does not need to be increased is an operative-learning design. However, such a design could only determine if pronghorn can distinguish between tillering and jointing wheat, and would not quantify preference. ��~5 PRONGHORN WINTER D. C. Strohmeyer, WHEAT DAMAGE G. C. White, STUDY and R. B. Gill P.N. OBJECTIVES Elucidate pronghorn movement patterns concerning SEGMENT OBJECTIVES winter wheat use. 1. Monitor changes in pronghorn antelope use of winter wheat native grassland vegetation in northeastern Colorado. 2. Evaluate pronghorn antelope native grassland vegetation food preferences for winter using tame pronghorn. fields wheat and forage and INTRODUCTION This research had the potential to offset increasing winter wheat (Triticum aestiyum L.) damage complaints concerning pronghorn (Antilocapra americana). The Colorado Divisiori of Wildlife (CDOW) reimburses landowners for wildlifecaused damages to their property. In the 1983-84 fiscal year, game damage claims peaked at nearly $1 million (Colo. Div. Wildl. 1991). In 1990-91, $192,000 in game damage claims were approved, including $1,533 for pronghorn winter wheat use. These figures underestimate game damage costs because they do not include game damage prevention costs. The CDOW spends more than $300,000 annually to prevent game damage. One example of a game damage prevention cost is the time spent by CDOW employees responding to game damage complaints. In 1992-93, John Wagner, the District Wildlife Manager in the Weld county area, roughly spent 20% of his time in November handling pronghorn winter wheat damage complaints (J. Wagner, Colo. Div. Wildl., pers. commun.). These complaints peaked January-March and took 50-60% of his time. Complaints began decreasing in mid-March. set against other depredation costs pronghorn winter wheat damage complaints may seem trivial, but the political situation's powder-keg potential is noteworthy. In Colorado, >200,000 ha of native grasslands have been converted to winter wheat and approximately 75% of the pronghorn population resides in wheat growing areas (Torbit et al. 1993). Colorado's pronghorn population is currently at political carrying capacity (K. Kinney, Colo. Div. Wildl., pers. commun.). This implies any CDOW management objective that would increase pronghorn numbers would also increase CDOW depredation costs. Before 1983-84, farmer complaints concerning pronghorn winter wheat damage were intense (K. Kinney, Colo. Div. Wildl., pers. commun.). The CDOW responded by reducing pronghorn numbers through hunting and trapping removals. For example, in the Hugo area pronghorn numbers were reduced by about 50-55% from 1981 through 1983 (Torbit et al. 1993). Farmer complaints concerning pronghorn winter wheat damage are rising again; however, the solution of intensive hunting is not desirable for several reasons. One, animal right's organizations will probably oppose it. Colorado's constitutional amendment banning bear hunting is an example of how far a game management issue can be pushed. TWO, the CDOW will (again) not be able to meet hunter demands for �226 pronghorn if pronghorn numbers are severely reduced. Three, region-wide, near-eradication of game animals is not philosophically compatible with the CDOW's mission to perpetuate Colorado's wildlife resources and to provide opportunity for people to enjoy wildlife. Trapping removals are also not desirable, largely because of high costs and problems associated with finding places to put the animals. Torbit et al. (1993) suggested pronghorn may not damage winter wheat, implying the CDOW should not be incurring depredation costs for pronghorn winter wheat use. Assessing this statement requires integrating winter wheat physiology, winter wheat herbivore interactions, native vegetation physiology, ungulate foraging selection theory, and pronghorn movement patterns. This is not the first attempt to address this issue and gaps in past research are noted. Winter Wheat Farming About 95% of Colorado's wheat is winter wheat, which is planted in the fall from late August through mid-October (Jenkins 1983:213-4, Hay and Walker 1989:159-163). Wheat is basically a dryland crop, partly because more profitable crop species are grown if irrigation is available. Wheat is "strip cropped". That is, an area is divided into long strips, which are set perpendicular to the prevailing winds, and alternate strips are planted annually. Strips are 90-185 m wide, depending on soil type. Strip cropping minimizes wind erosion and conserves soil moisture. A wheat crop needs at ,least 10 cm of moisture. It usually takes 2 years to accumulate this much moisture in the Weld County area (M. Petersen, Soil Conserv. Serv., pers. commun. )• Winter Wheat Physiology After planting, wheat seed imbibes water and germinates. The first shoot appears in October and tillering (multiple shoot production) begins in November. Tillering duration is largely temperature dependent and is interrupted by a winter dormancy period. Tillering is important because (1) main shoots exceed ear-bearing tillers in yield, (2), 75% of the tillers do not produce ears, (3) not all tillers survive past the jointing stage, and (4) tillers inhibit reproductive shoot growth (Bremner 1969, Darwinkel 1980). These points suggest winter wheat produces biomass exceeding that which is optimal for maximum grain production. In February, after vernalization, spikelet and floret development begins. At this time, the growing point in the crown stops vegetative production and starts making reproductive parts. Terminal spikelet formation and stem elongation occur simultaneously. Nutrients begin shifting from the stems and leaves to the developing grain heads (Spiertz 1977). This nutrient shift is important because it reduces winter wheat forage quality (Torbit et al , 1993). During the jointing phenological stage, the growing point, via stem elongation, moves from belowground to above the soil surface. Anthesis (flowering) occurs in early May followed by grain-ripening in late May. Harvest begins in late June. Winter Wheat Vulnerability To Grazing Wheat should be invulnerable to grazing during tillering because excess biomass is produced (Torbit et al. 1993). From November until April, wheat is commonly grazed by livestock. Forage removal effects on wheat yields were examined with field experiments (Dunphy et al. 1982). Final forage harvests corresponded to 3 wheat phenological stages: early, mid, and late-jointing. �~7 Maximum grain yields were obtained when forage removal was terminated by the early-jointing stage. These results cannot be generalized to a specific, annual calendar date because jointing dates vary among years and among wheat cultivars. Torbit et al. (1993) specifically addressed winter grazing of pronghorn on wheat. Pronghorn densities used during the experiment were 166 pronghorn/km2, which greatly exceeded that found in .the wild, roughly 2 pronghorn/km2• Four winter grazing treatments were compared: early, late, continuous, and control (no grazing). All grazing was terminated by the mid-jointing stage. Grain yields among treatments were not significantly reduced (~ = 0.17). Tests for grazing treatment effects were somewhat insensitive in their ability to detect differences because variability was high among replicates within all treatments. A 21~ decrease in grain yields on grazed plots would have been required before differences (a = 0.05, ~ was not indicated) were detectable. Winter Wheat and Shortgrass Prairie Forage Quality Livestock winter wheat grazing has been initiated partly because wintertime native plant species forage quality is poor. During the winter, shortgrass prairie plant species may be deficient in protein, phosphorus and vitamin A according to cattle maintenance standards (Cook et al. 1977). Wheat crude protein concentration should exceed that of native plant species during the winter (Torbit et al. 1993). Wheat crude protein concentration starts decreasing in April when nutrients are relocated from stems and leaves to the developing grain head (Spiertz 1977). Crude protein concentrations of native plant species peak in April and May (Schwartz 1977). This sequential peaking in nutritional levels between winter wheat and native plant species can be expected annually. The sequence arises from differences in metabolic pathways. Wheat is a C3 species, while most native plant species are C4 species. Differences in carbon fixation between C3 and C4 pathways cause C3 species to have lower optimum temperatures for photosynthesis and to germinate sooner in the spring when temperatures are cool (Caswell et al. 1973, Salisbury and Ross 1992). Despite metabolic pathway differences, this peaking will remain synchronous because the metabolic rates of both pathways are affected by the same major environmental variables: irradiance, temperature, nitrogen supply, and water stress (Tenhunen et al. 1976a,bi Van Soest 1982; Woledge and Parsons 1986). Ungulate Foraging Theory The first decision an ungulate must make, after deciding to graze, is where. The plant-animal interface can be described in a hierarchical context: landscape, plant community, patch, feeding station, and plant (Senft et al. 1987, Coleman et al. 1989). Factors influencing grazing decisions at the landscape level include distribution of plant communities and accessibility of foci such as water and shade. Plant-animal interface models are currently lacking data concerning shifting and triggering needs of animals at the landscape level. The feeding station is defined as an area that is grazed without taking a step. Within a feeding station, animals generally select diets that are higher in crude protein and digestibility than the average of that among the available forage. Foraging decisions are also influenced by animal experience. Current diet selection theory is based on learning and memory �228 (Launchbaugh and Provenza 1991). It includes social elements, cautious sampling of novel foods, and the formation of food preferences and aversions based on gastrointestinal consequences. Mechanistic factors affecting diet selection include (1) ease of eating; (2) sensory factors like taste, texture, and odor; (3) water content; and (4) sward architecture (Colebrook et al. 1990). Hurd and Blaser (1962) lumped the above mechanistic factors into one category, palatability. They viewed grazing decisions to be based on palatability and concentration of essential nutrients. Ruminants must continuously sample a broad spectrum of available foods to monitor spatial and temporal changes in essential nutrients, fiber content, availability, etc. (Provenza et al. 1992). Food selection on the basis of nutrient content requires a perception of one's requirements, an internal recognition of a food's value in meeting rather specific requirements, but not necessarily a taste or smell for each nutrient (Robbins 1983:324). Animals given free access in a cafeteria-style experiment to 30-40 required nutrients selected a diet enabling growth about equal to that of animals consuming balanced laboratory diets. . Body size largely determines rumination efficiency (Welsh and Hooper 1988:114). Larger animals are more efficient in their digestion of plant parts, both within and between species, even when metabolic body size is the basis for comparison. Smaller ruminants, like pronghorn, are unable to ruminate large amounts of plant cell wall and must select higher quality material to survive. Pronghorn select forages which are relatively high in crude protein and low in cell wall content (Schwartz and Ellis 1981, Krueger 1986) • Timing Of Pronghorn Winter Wheat Use For the statement, "pronghorn do not damage winter wheat", to be true, pronghorn must be shown experimentally to stop foraging on wheat as wheat enters the jointing stage. Combining forage quality differences in wheat versus native plant species with pronghorn preference for high crude protein and low cell wall content implies pronghorn will use winter wheat until the forage quality of native plant species exceeds that of the winter wheat. This hypothesis requires that (1) native plant species' forage quality must exceed that of wheat as wheat begins to joint; and (2) pronghorn must respond to nutritional changes in forage quality. Alldredge et al. (1987) qualitatively examined monthly nutritional changes in pronghorn diets and compared a winter wheat diet to a shortgrass prairie diet. Shortgrass diet compositions were taken from 3 tame pronghorn using prairie adjacent to the CDOW's Foothills Research Facility. These diets were augmented by data from Schwartz (1977). Winter wheat quality falling below that of native diets depended on largely on 2 data points, May and June. Statistical tests of changes in relative nutritional levels are absent. Within month variation is not estimable because there is only one replication per month. Bi-monthly phenology measurements were recorded for native species, but winter wheat phenology was not reported. Thus, the relationship between winter wheat phenology and changes in winter wheat forage quality still needs to be quantified. Observational studies involving pronghorn and wheat have noted seasonal use patterns (Cole and Wilkins 1958, Hoover et al. 1959, Torbit et al. 1993). Pronghorn use of wheat is heavy in the fall and winter, while light in spring �and summer. Monthly aerial surveys in northeastern Colorado found unmarked, free-ranging pronghorn used wheat fields from November through April, then abandoned them by early May. Data from 5 radiocollared pronghorn also support this pattern. Marked pronghorn abandoned wheat fields by 19 April 1985 and by 12 May 1986, but the exact timing of the shift in vegetation type was not well documented. Explicit ties between the shift in vegetation used and winter wheat phenology are also lacking because winter wheat phenology was not reported. Quantifying Vegetation Nutrient Dynamics Concerning native vegetation forage quality, it is tempting to select a few plant species, prominent in pronghorn diets, and track their respective changes in percent crude protein and cell wall contents. Unfortunately this approach is inadequate. Once a food item is ingested, its. nutritional value becomes relative to the other ingested food items (Robbins 1983:324). Thus, assessing nutritional constraints on foraging behavior requires weighing plant species chemical composition results by a botanical composition of an appropriate diet. Vegetation phenology measurements involve chronologically assigning numbers to phenological stages of interest. Phenology of shortgrass prairie species are often divided into 14 stages (Dickenson and Dodd 1976). Winter wheat phenology is typically documented by the Feekes System or the Decimal Code System (Hay and Walker 1989:159). The Decimal Code System is the more recently developed system, arising from the need for more resolution in grain development stages. This resolution is not important for our hypothesis and choice of either system is arbitrary. Quantifying Pronghorn Response To Forage Quality Changes As previously mentioned, we need to establish that individual, wild, freeranging pronghorn actually follow Torbit et al.·s (1993) nutritional hypothesis. Techniques for measuring pronghorn vegetation use could range from focal-animal sampling (Altmann 1974) to binomial presence/absence data. For our hypothesis concerning pronghorn winter wheat use timing, binomial presence/absence data are adequate. Animal activity is less important than properly recording the vegetation types of animal locations. Binomial data requires less data collection time than behavioral data. Plus, binomial data can be statistically evaluated via logistic regression procedures. Design Choices For Quantifying Pronghorn Preference There is a continuum of potential experimental designs available to test pronghorn preference of different developmental stages of wheat. (Note that this approach assumes that the stimulus causing pronghorn to stop using wheat is in. the winter wheat.) At the continuum extremes are high external validity (using wild animals in a natural setting) and high internal validity (running feeding trials with tame animals). External validity is inversely related to internal validity (Kirk 1982:21). External validity describes the generalizability of research findings to and across populations of subjects and settings. Internal validity is concerned with correctly concluding an independent (predictor) variable is in fact, responsible for variation in the dependent (response) variable. In this case, we favor designs with high internal validity, because past research has had high external validity. �2W Wild versus Tame Pronghorn Pronghorn are excitable animals and are difficult to handle without incurring animal mortality. Thus, use of wild pronghorn is only realistic for experimental designs with almost no animal-human contact, for example, freeranging animals. still, tame animals will not be the perfect homolog of wild animals. Tame pronghorn will not have had experience with plant species in native rangelands. Wild pronghorn will be under greater nutritional constraints than tame animals eating balanced diets. Plus, wild pronghorn will be in the third trimester of pregnancy during the spring. Tame and wild pronghorn may use different diet selection criteria. Schwartz (1977) found diet composition of tame and wild pronghorn to differ, yet considered tame pronghorn to provide reliable data on wild pronghorn forage preferences. Olson-Rutz and Urness (1987) found behavioral differences between tame and wild deer were explained by the effect of experience with new environments rather than foraging per see Exposing tame animals to experimental conditions before starting data collection will reduce differences in diet selection criteria. Free-ranging Versus Captive Animals We decided to use captive pronghorn instead of free-ranging pronghorn because free-ranging pronghorn designs had lower internal validity. Working with wild, free-ranging pronghorn would require large sample sizes to adequately replicate all treatments. We would need many manipulated fields and many marked, wild pronghorn, because pronghorn use intensity changes across wheat fields. Low statistical power would result from small sample sizes, high variances, and a lack of balance (equal replication among cells). Working with tame, free-ranging pronghorn would.also be difficult because of the amount of transportation involved. Even tame pronghorn are skittish and susceptible to handling stress. Plus, transportation logistics would heavily constrain sample size. There are 2 types of captive animal experimental designs: pasture and feeding trials. Burns and Mochrie (1981) compared performance differences between continuous grazing (pasture trial) and feeding-trial preferences. Heifer performances between the designs were similar, except one treatment group. In this treatment group, the feeding trials showed avoidance, while the pasture trial resulted in one of the highest weight gains. Ideally we would do both designs, but logistics prevent this. Pasture Design season-long grazing trials offer several advantages over other designs. Both wild and tame pronghorn would be under similar nutritional selection pressures. Tame pronghorn would habituate to field conditions. Transportation and handling stress would be minimal. Data collection would be continuous, uninterrupted by transporting animals. Season-long grazing trials are easier to interpret than feeding trials, because their experimental setting has natural components at the landscape scale. Most criticisms of penned-animal experiments arise from the concern that the pen itself may affect the animal's behavior. Fencing costs will constrain sample size and would necessitate designing the pens with common fences. Common fences introduce the possibility that animals �231 in adjacent pens will influence each other's behavior. At least one experiment involving penned cattle has failed because the need for social contact concentrated animal use along fence lines, violating the assumption of uniform pen-use (L. Rittenhouse, Range Sci. Colo. state Univ., pers. commun.). At least one grazing study involving single pronghorn in adjacent pens was successfully completed (J. Liewer, Colo. Div. Wildl., pers. commun.). Liewer noted pronghorn in adjacent pens moved towards each other when frightened, but grazing occurred throughout the pens. Without question, social behavior of pronghorn in adjacent pens will not be independent, but we are not presenting a hypothesis involving social behavior. We suggest the primary mechanism influencing foraging decisions is nutritional quality. Shifts in vegetation types used for foraging should occur despite social behaviors and should parallel choices made by wild pronghorn. Concurrent monitoring of wild pronghorn will help determine if the pens themselves are influencing foraging decisions. A second concern is that grazing intensity inside the pens may change nutritional patterns and alter foraging decisions (L. Rittenhouse, Range Sci. Colo. State Univ:, pers. commun.; D. Swift, Nat. Resour. Ecol. Lab., pers. commun.). This is serious because our hypothesis involves foraging quality and preference as opposed to foraging quantity. Selection of the pen sites is also problematic. The selection process would not be random, but rather the result of logistical considerations. Also, diet composition of a given animal is limited by the plant species in the pen. Pasture Design: Potential Manipulations Potential ways to manipulate winter wheat phenology include fertilization, irrigation, and seeding. Fertilization and irrigation will not produce the desired phenological changes (D. smith, Agron. Dep. Colo. State Univ., pers. commun.; W. K. Lauenroth, Range Sci. Colo. State Univ., pers. commun.). Crop fertilization and irrigation may change phenology by 2 days. Spring fertilization of native vegetation is unlikely to affect phenology, but it may change species composition by increasing the proportion of CJ species (Petersen and Moser 1985). Dickenson and Dodd (1976) manipulated native range vegetation on the Pawnee Site (the proposed study area) and found no observable difference in phenological progression resulting from variations in grazing intensity or nitrogen fertilization. However, their water treatment did affect phenological progression of some species by delaying peak flowering by 5 days and shortening the flowering period by 10 days. Seeding is the most promising manipulation. There are 2 potential crop choices: spring wheat or a northern winter wheat variety (D. Smith, Agron. Dep. Colo. State Univ., pers. commun.). The respective phenologies of spring wheat and northern winter wheat will be roughly 4 and 2 weeks behind that of the typical winter wheat variety grown in northern Colorado. Choosing spring wheat has drawbacks. Spring wheat planting dates are largely weather dependent and range from mid-March to late April. Spring wheat does not tiller like winter wheat and would require higher seeding rates. A disadvantage of seeding different wheat varieties is wheat varieties may affect pronghorn response. However, if nutritional quality is the driving mechanism in pronghorn foraging decisions, then pronghorn will select the more nutritional forage. Note this implication assumes that pronghorn respond equally to both varieties of winter wheat (at the same phenological stage). �The validity of this assumption is questionable, since winter wheat cultivars vary in their susceptibility to grazing. Testing this assumption would require feeding trials. Ideally, manipulations would include changing the native range vegetation in addition to changing the winter wheat. Seeding C3 species in native rangelands is possible, but this is unsatisfactory for 2 reasons. One, this manipulation will change the habitat structure and could change foraging behavior (Roese et ale 1991). Two, this manipulation could damage the ecosystem (W. K. Lauenroth, Range Sci. Colo. State Univ., pers. commun.). C3 species are more likely to die during droughts and increasing the proportion will increase problems with erosion during droughts. Feeding Trials Feeding trials provide a valuable measure of forage quality, given proper feed preparation and use of standardized experimental procedures. These trials would provide the highest internal validity; however, the artificialness of the situation sacrifices external validity. Translation of the experimental results from a feeding station level back to a landscape (management) level will be difficult. Yet, our nutritional hypothesis is probably most effectively examined at the feeding station level. Animal preference for a resource is the likelihood that a resource will be selected when all resources are equally available (Johnson 1980). One method for estimating preference rankings of forages is to present several forages simultaneously to each animal. Each object/treatment accumulates a preference score during the test interval and the response variable is resource use rather that object selection. This method does not truly measure preference because it does not estimate the selection probability of a given forage. An alternative to multiple-choice (cafeteria) trials for estimating preference rankings is to use paired comparison trials where each subject chooses among forages offered in all possible pair-wise combinations (David 1988) and responses are either chosen or not chosen. The binomial responses from paired comparison designs can be used to estimate resource preference probabilities. (Contrast this to cafeteria trials which assume some unspecified relationship between resource use and resource preference.) Analysis of these responses is analogous to the normal-theory balanced incomplete block design, where treatments can have e~ther a one-way structure for estimating preference rankings or a factorial structure for modeling the effects of object attributes on preference. Paired comparison trials are favored when the selection factors of interest may be difficult to perceive. Feeding trials offer the following advantages (Minson 1981). One, accurate direct measurements of individual animal feed consumption and digestibility are obtainable. Two, the characteristics of forage, as consumed, are determinable. Three, the environment of experimental animals is controllable. Four, the forage growing season does not restrict the timing of the experiment. Disadvantages of feeding trials include the following when measuring digestibility and voluntary intake (Minson 1981). One, the chemical composition and nutrient availability of forage may change during the study or when conserved and stored. Two, feeding trials will not duplicate forage �233 selection of grazing animals. cuttings or preservation. Feeding Trials: vegetation Three, forage handling involves either daily Collection Two choices for collecting vegetation are (1) growing the plants in a greenhouse (or outdoors) and presenting potted plants which allows an animal to select plant parts, or (2) clipping naturally-growing plants and storing them until the experiment. The disadvantages of the former principally center around the greenhouse itself. Obtaining space in existing greenhouses can be difficult and building a basic greenhouse would be costly, >$15,000. Plus replicating the vernalization period of winter wheat would require the ability to control a wide temperature range. Disadvantages concerning clipping naturally-growing plants involve the clipping itself and storage without altering the nutrients. Hand-clipping inhibits the ability of animals to select plant parts, while storage destroys palatability factors. storage may also affect plant nutritional content. Note, clipping naturally-growing plants is not feasible without storage because we want to manipulate the time order of plant maturation. There are 3 methods used to limit nutritional changes in clipped plants: oven-drying, freezing, and freeze-drying (Minson 1981). Oven-drying is typically the most practical. Long drying times prolong the period when enzymatic changes and metabolic losses occur. Freezing and freeze-drying are more effective in retaining nutrients than oven-drying. Freeze-drying leads to the minimum changes in composition and doesn't require freezer storage. Freeze-drying offers the additional advantage that dried forage does not decompose in the feed trough. Feeding Trials: Vegetation Choice We are interested in exploring possible nutritional mechanisms that stimulate pronghorn to shift from winter wheat to native-range species. One possibility is to examine differences in pronghorn response to winter wheat in different phenological stages, tillering and jointing. If pronghorn make the anticipated choice, tillering wheat is selected over jointing wheat, then we may conclude pronghorn can detect and respond to relatively subtle differences in nutrition. Note palatability factors will confound this conclusion. If pronghorn do not select tillering wheat over jointing wheat then we may conclude: (1) pronghorn do not response to the differences in the wheat, and (2) the stimuli for the behavioral shift may be something in the native range vegetation. This approach implicitly assumes the stimulus for the behavioral shift is a negative stimuli in the wheat. Potential positive stimuli from native-range species are ignored. One drawback to this approach is wild pronghorn do not face the choices presented in the feeding trial. A second drawback is even if pronghorn (correctly) select tillering wheat, we cannot determine if they are responding to a positive stimulus or a negative stimulus. Ideally a feeding trial would contain both types of vegetation (wheat and native range) because we are concerned with nutritional stimulation across plant species. The issue becomes what is the best representation of the native-range complex? Use of multiple species to represent the native-range complex would be interesting, but is more appropriate for future research. A better starting point is to select one species, but which one? We want a �234 species having (1) high preference by pronghorn, and (2) a nutritional status that is lower than that of tillering wheat then higher than that of jointing wheat. None of the native species most heavily consumed by pronghorn clearly meet the second criteria for crude protein (Tables 1-2). Agropyron smithii (Agsp) may meet the second criteria for cell wall contents. Table 1. Forage species most heavily consumed by pronghorn from March through May. Data are taken from Schwartz (1977:30) and were chosen because this particular diet composition most closely matches native species observed (1993-4) on the Pawnee National Grasslands during these months. Percentage Taxa of pronghorn diet March April-May Agropyron smithii 29 Artemisia frigida 10 18 <0.5 Bouteloua gracilis 13 <0.5 22 28 ~ spp. Sphaeralcea 1 coccinea Total 17 75 63 Table 2. Comparison of winter wheat and native-range plant nutritional characteristics. Wheat data are from Alldredge Native species data are from Schwartz (1977:111). Chemical Crude Taxa Winter wheat ~gt:QPyt:Qn !i!mithii ~t:t~mi!i!ia!t:igiga gt:aQilia BQ!.lt~lQ!.la spp. ~ Sphget:alQ~g QQQQin~g Protein (%) species et al. (1987:44). constituents Cell Wall contents (%) March April May June March April May June 31-35 9.3 10.7 8.4 14.9 27-30 23.5 13-18 11-9 47-55 65.3 43.9 72.0 59.6 40-55 49.2 52-53 60-61 7.3 21.0 15.0 73.2 47.7 41.4 Given a feeding trial with wheat and a native plant. The treatment pairs would be tillering wheat-early Agsp, tillering wheat-late Agsp, jointing wheat-early Agsp, jointing wheat-late Agsp. If pronghorn make expected choices, they select the plant with higher nutritional quality, then we may conclude that pronghorn can respond to (nutritional) differences between 2 palatable species. Using AgrQPyrQn smithii for the native species implies �235 pronghorn can respond to differences in cell wall contents. If pronghorn always select one species, then we may conclude that pronghorn prefer the selected species over the other. If pronghorn selection appears random, then we may conclude that pronghorn are not stimulated by differences between the species. Presenting 2 species in a feeding trial has problems. One, biomass during the feeding trials would not be limiting, which may not be the case for wild pronghorn, especially in March. Two, existing data are sparse and the choice of a single, native species is not clear. Three, greenhouse logistics become more complex. Four, the risk of not getting the expected results is greater than trials examining differences within a species. Objective Our objective was to examine the hypothesis that pronghorn shift from winter wheat to shortgrass prairie species .when wheat begins jointing. We specifically examined nutritional mechanisms. Our study was correlational· in nature and causation may not be inferred. TELEMETRY STUDY AREA All fieldwork was conducted in Weld County, Colorado, and included the Pawnee National Grassland. The study area was at the northern edge of the shortgrass steppe region at an elevation of 1650 m (Lauenroth and Milchunas 1991). The climate was continental. Mean annual air temperature was 8.2 C, with a mean daily minimum -12.0 C, in January and a mean daily maximum of 26.5 C in July. The frost-free period was approximately 150 days. Average proportion of sunshine hours was 60-70%. Annual average wind speeds were >5 m/s. In this semi-arid region, precipitation fluctuated widely about 21 cm, and 70% of the annual precipitation occurred.as rain from April through September. Thornthwaite's potential evapo-transpiration rate was 60 cm/year. Topography consisted of gently rolling hills, broad ephemeral stream courses, and low flat-topped terraces (Yonker et al. 1988, Lauenroth and Milchunas 1991, Torbit et al. 1993). The region was underlain by late Cretaceous shales and interbedded sandstones of the Laramie Formation. Parent materials were derived with rare exception from surficial deposits of alluvium composed of material derived from Front Range (Colorado Rockies) sources. Soils were in four predominant subgroups: Aridic Argiustolls, Ustollic Haplargids, Ustic Torriorthents, and Ustic Torrifluvents. Soils, geologic substrates, and landforms showed a high degree of spatial heterogeneity, arising from a complex geomorphic history. Loamy soils were predominant, with soil texture varying from sandy loams to clay loams. Most soils in the area had a moderate wind erodibility. Soil pH was generally alkaline, but changed with water table depth. The average growing-season above-ground plant biomass was 48% warm-season grasses, 8% cool-season grasses, 9% forbs, 11% shrubs, and 24% cactus (Milchunas et al. 1989, Lauenrdth and Milchunas 1991, Torbit et al. 1993). The dominant grass was blue grama (Bouteloua gracilis). Other common grasses included red threeawn (Aristida longiseta), buffalograss (Buchloe dactyloides), and western wheatgrass (Agropyron smithii). Needleleaf sedge (~ eleocharis) was also present. Prickly pear cactus (Opuntia polyacantha) was prominent. Native forbs included scarlet globemallow �236 (Sphaeralcea coccinea), scarlet gaura (~ coccinea), and scurfpea (Psoralea tenuiflora). Suffrutescent shrubs included fringed sage (Artemisia frigida), and broom snakeweed (Gutierrezia sarothrae). Rubber rabbitbrush (Chrysothamnus nauseosus) and fourwing saltbush (Atriplex canescens) were common shrubs. Shortgrass prairie comprised about 65% of the study area, with cropland comprising the remaining 35% (Liewer 1988, Lauenroth and Milchunas 1991, Torbit et al. 1993). Dominant land uses were cattle ranching and dryland winter wheat farming. Winter wheat was grown in a one-year fallow system and a combination of crop residue management, minimum tillage, and wind stripcropping was necessary to offset serious wind erosion. Nitrogen and phosphorous fertilizers were commonly applied to winter wheat crops. TELEMETRY STUDY METHODS Intensive ground telemetry monitoring and vegetation measurements began in February continuing through June. Sampling periods were based on weeks (7 days) because this provided enough time for phenology to advance (and forage quality to change) but not enough time to miss phenological stages for most native plants. Two relocations per week per marked pronghorn were attempted. Vegetation sampling of native and winter wheat vegetation types was paired with 6 replicates per week. When marked animals were relocated the vegetation type was recorded as native range; winter wheat, or other. We omitted nocturnal sampling because it would have greatly increased the time required to collect a sample and pronghorn were thought to be less active at night (W. Alldredge, Wild1. BioI. Colo. State Univ., pers. communv) , Fourteen wild female pronghorn were captured by CDOW personnel in February 1993. A radiocollar and one red, numbered ear-tag were attached to each animal. Only female pronghorn were marked for 2 reasons. One, only a few animals could be used and dividing effort among sex classes would have lowered statistical power. Two, spring coincided with the final stages of gestation for pronghorn. Female pronghorn were under more nutritional stress than males (Robbins 1983:172), and should be more sensitive to changes in forage quality. Vegetation Collection Location of the vegetation samples was paired (native range, winter wheat) and was determined by marked pronghorn. Sample location for the second half of the pair was in the closest section of the appropriate vegetation type. Marked pronghorn were not intentionally flushed from a location, hence vegetation work began as soon as possible after the pronghorn left the site. Native plants collected were those species found in pronghorn diets (Schwartz 1977). One wheat phenology sample was the mean of 25 individual plants (Dickenson and Dodd 1976). Measurement units were documented with the Decimal Code System (Hay and Walker 1989:159). Phenological measurements preceded clipping. When clipping vegetation, green forage was preferentially sampled relative to dead forage, so sampling was more akin to pronghorn foraging behavior. Size of the wheat samples were roughly 50 g (dry mass) of material collected from a minimum of 25 wheat tillers. One sample of a native plant species was at least 5 grams (dry mass) .from a minimum of 25 plants. Five grams was the minimum amount needed to perform the desired chemical analyses (L. Stevens, �237 Colo. Div. Wildl., pers. commun.). Samples were chemically analyzed by the CDOW's nutrition lab. Standard methodology for crude protein (Horowitz 1980), cell wall constituents (Goering and Van Soest 1971), and in vitro dry-matter digestibility (Tilley and Terry 1963, Pearson 1970) were followed. The laboratory analyses were used to calculate diet qualities for 2 diet scenarios. The first diet was 100% winter wheat. The second diet was a compositional input of native species following Schwartz (1977). Statistical analyses Patterns in pronghorn diet nutritional quality, wheat phenology, pronghorn vegetative type use, and time were empirically described by fitting polynomial curves to observed data using general linear model procedures (McCullagh and Nelder 1989). . Logistic regression modelled pronghorn behavior. Pronghorn use of vegetation types (shortgrass prairie or wiriter wheat) was the dependent variable. This discrete, nominally-scaled variable was recorded as use versus non-use and was binomially distributed. Time (t), a continuous variable, and collar (c), a discrete, nominally-scaled variable, were the predictor variables: 1 (1) where fiw fiN = = probability probability that pronghorn that pronghorn use wheat, and use shortgrass prairie. The null hypothesis was that there were no changes in the vegetation type used by pronghorn over time, Ho: ~l = O. The alternative hypothesis was that the 10git of the fiw increased over time, Ha: ~l > o. We also used this model to quantitatively relate the period when there were no differences in pronghorn use of vegetation types, fiR = fiv = 0.5, to diet quality values for each marked pronghorn. Our second analysis examined diet quality changes. This analysis was repeated separately for dietary NDF and dietary crude protein. First, diet quality was modelled over t.Lme, Wheat and prairie diets were modelled separately: (2) p = a! 1-'0 + a! t 1-'1 + a! t 2 1-'2 ' (3) �238 where t W P time in days, winter wheat diet quality, and shortgrass prairie diet quality. We compared linear and quadratic models. The null hypotheses were that the linear models were better, Ho: ~2 = 0 and Ho: ~; = O. The alternative hypotheses were the quadratic models were better, Ha: ~2 1 0 and Ha: ~; 1 O. If wheat and prairie diet quality did not change over time (linear models with zero slopes were selected), then the analysis was truncated here. Otherwise, the best wheat and prairie models were used to generate diet quality values for the second hypothesis. Next, we related pronghorn habitat use patterns to pronghorn diet quality logistic regression. This analysis was repeated separately for dietary neutral detergent fiber (NDF) and dietary crude protein. The model was: via (4) where Ilv = probability IlR = W P = that pronghorn use wheat (Ilv = 1. - IlR), probability that pronghorn use shortgrass prairie, winter wheat diet quality from equation (2), and shortgrass prairie diet quality from equation (3). We used this model to quantitatively relate the period when there were no differences in pronghorn use of vegetation types, IlR = Ilv = 0.5, to diet quality values for each marked pronghorn. Lastly, we used analysis of variance to test quality differences between wheat and prairie diets within sampling periods. FEEDING TRIALS STUDy AREA AND METHODS We compared preferences of female, hand-reared pronghorn for phenological stages of wheat via two-way feeding trials. Our treatments were the tillering and jointing stages (Hay and Walker 1989). We did 144 experimental trials. We used 2 types of feeding-trial designs which differed with respect to wheat arrangement. The first involved adjusting plant height via an elevated box and will be referred to as the box experiment. Pronghorn saw only the tops of the wheat plants, the tips of which were at the same height. Twelve replications per animal were .3.ttempted. The second design had n6 adjustment for plant height. Four pots of wheat were placed in pairs (each pair had the same phenological stage) on the ground. This experiment was named the 4-pot experiment. Fifteen replications per animal were attempted. This experiment always followed the box experiment for logistic reasons. For both experiments, the protocol was the same. Wheat placement (position), wheat pots, pen, and time order were randomized to pronghorn. Animals were given 3 minutes to begin feeding. (If feeding occurred it most commonly began in <30 seconds.) The initial selection was scored as 1 (jointing wheat �.239 selected) and 0 (tillering wheat selected). Feeding was monitored minute. Pronghorn preference data were binomially distributed: for 1 ni = probability of pronghorn i choosing tillering wheat 1 - ni = probability of pronghorn i choosing jointing wheat, where i = an individual pronghorn (1-12). Animal Handling The animals were housed in 2 pastures (each about 4 ha) at the CDOW's Foothills Wildlife Research Facility. Alfalfa hay, pelleted supplement (Baker and Hobbs 1985), trace mineral block, and water were provided following Foothills Wildlife Research Facility protocols (Wild et a1. 1992). Some green vegetat10n was also present in each pasture. Animal training to the experimental procedures occurred before data collection and animals were exposed to all treatments (wheat stages) during this time. Conditioning increased the chance that preference rankings were an attribute of the experimental food types and not animal inexperience.· At this point, the 3 oldest animals were dropped from the experiments because they did not respond to training. Nine animals were placed into isolation pens (the location of the trials) before trials began and all were held in these pens until one replication (1 pair of both wheat stages was presented to each animal) was completed. One replication of feeding trials (one trial per animal) were attempted on a given day. Animals were put into isolation pens (11 m2) with water but no food prior to beginning the feeding trials. Food was presented to one animal at a time. If an animal did not feed within 3 minutes, the food was removed then presented again after all other animals have been tested. A maximum of 2 retests were attempted. If an animal began feeding, the feeding behavior was monitored for 1 minute. All animals were held in the isolation pens until the feeding trials were finished. . Vegetation We planted rows of spring wheat (Oslo variety) weekly at the Colorado Division of Wildlife's (CDOW) Foothills Wildlife Research Facility in Fort Collins. We used spring wheat because it did not require 6 weeks of vernalization but is otherwise similar to winter wheat. The spring wheat was transplanted into pots prior to the feeding trials. We varied wheat planting dates so several phenologies were available simultaneously. During the experimental trials, all feeds (plants at different phenological stages) had roughly the same mass and were in identical containers. Position of treatments (left or right) were randomized • .statistical Analyses First, contingency-table exact tests examined pronghorn behavior collectively. Did all animals share a common value? Second, a one-sample sign test assessed whether pronghorn behaved randomly or had a preference, ~: n = 0.5. An animal was considered to have a preference if she chose a treatment 75% of the time. (A more common definition of preference was to use an 80% response criterion. ) Third, a likelihood ratio t.est compared pronghorn choice to biomass. The null hypothesis was that animal choice was independent (random) of biomass. Fourth, a paired t-test and Pearson's correlation coefficients �240 assessed the consistency the 4-pot experiment. of pronghorn responses between the box experiment and RESULTS Telemetry, 1993 Pronghorn on winter wheat were moved to native range using CDOW aircraft several times during the 1992~93 winter. These hazing operations conditioned pronghorn to run from aircraft and when marked pronghorn were relocated aerially they were often running when seen. It was often unclear whether or not the marked animals were on wheat before they responded to the aircraft. In these instances no vegetation type was recorded, otherwise the data would have erroneously indicated higher use levels of native range then actually occurred. Consequently, only 4 of 13 marked animals had relocations on both winter wheat and native range, with the rest only relocated on native range. All animals that were relocated on both vegetation types used wheat then shifted to native range. The last marked animal relocation on wheat was April 10th (Table 3). Unmarked pronghorn were observed on wheat on May 4th. (Marked animals were again relocated on wheat on October 20th.) No vegetation data were collected. Table 3. Date of last relocations County, Colorado, 1993-1995. Year Collar 1993 292 570 598 660 108 171 292 340 ·550 570 598 660 108 292 550 598 660 1994 1995 on winter wheat of marked Date of Most Recent Wheat pronghorn, Weld Relocation April 10 April 10 March 21 March 20 March 27 April 12 March 27 March 13 April 16 April 14 April 8 March 3 March 18 March 18 March 21 March 23 February 9 Goodness-of-fit tests for the models did not appear to be violated; the values of deviance/df were 1.6-1.7, suggesting a tendency for although, �241 overdispersion. Differences among individuals was not detectable (£ A change in the vegetation type used was moderately seen (£ 0.09). 0.7). = The pronghorn shift in vegetation use from wheat to detectable in this sparse data set. This shift for occurred between April 10th and May 5th. Mean date difference in pronghorn use of vegetation types (nN models was April 4th. Telemetry, native range was the marked animals of the period of no = nw) in the regression 1994 Problems with the quality of aerial relocations were not bad because pronghorn were not intensively hazed from wheat fields during the 1993-94 winter. Relocations from 10 marked animals were collectable during the 1994 field season. The last marked-animal relocation on wheat was April 16th (Table 3). Wheat entered the jointing phenological stage during the week of April 17th. As of May 25th, a few unmarked animals were still seen in wheat fields. Most of these observations occurred in the same wheat field. Of the unmarked groups seen in wheat fields from April 16th through May 27th, males dominated the observations. Only 4 groups of females (15 females total) were observed in wheat fields. Data from 8 marked pronghorn were analyzed. Dates of last winter wheat field relocations ranged from February 15th to April 16th. The mean date of when marked pronghorn were last relocated on winter wheat fields was March 24th. Winter wheat began jointing the week of April 17th. Only one marked animal was seen in a winter whe'at field after jointing occurred. She was located in a half-harvested field on the last day of sampling which implied that she was not eating wheat. We used the logistic models of pronghorn behavior versus time to estimate a mean date for the period of no difference in pronghorn vegetation use, nR = nv = 0.5. The mean of these predicted values was February 21st. This was almost one month earlier than the mean date of the last wheat field relocations. One factor contributing to the models' predicting early dates was that when pronghorn were using wheat fields they were also relocated on 'shortgrass prairie. Use of wheat fields gradually decreased over time. All of the 8 logistic regressions of pronghorn habitat use versus time had positive slopes, ~l > O. This indicated the pronghorn were more likely to be relocated on shortgrass prairie as time passed (£ = 0.0039). Differences among individuals were not detectable (P = 0.2). A change in habitat use over time was detectable (P < 0.0001). Neutral detergent fiber data showed slight negative trends with respect to time for both prairie and wheat diets. Crude protein data showed a negative trend for wheat diets while increasing then decreasing for prairie diets. The period of no difference in pronghorn vegetation use corresponded to mean crude protein contents of 17% and 15% for wheat and prairie diets~ respectively. The relation between NDF and time was better described in a linear model than in a quadratic model, Ho: ~2 = a and Ho: ~i = 0, for both wheat and prairie diets (£ = 0.31 and £ = 0.22, respectively). Slope parameter estimates for the linear models were essentially zero, ~l = -0.0007 and ~i = -0.001 respectively. Pre-jointing dietary NDF means were 65 and 56 for prairie and wheat, respectively. Post-jointing dietary NDF means were 55 and 49, respectively. This lack of trend over time suggested NDF was not a potential �242 stimulus for the change in pronghorn habitat found by Alldredge and Torbit (1987:39). use. This contrasts with results Pre-jointing dietary crude protein means were 12 and 30 for prairie and wheat, respectively. Post-jointing dietary crude protein means were 15 and 18, respectively. Prairie dietary crude protein values did not differ (£ = 0.3) between pre-jointing and post-jointing of wheat. In contrast, pre-jointing wheat had significantly higher (£ < 0.001) crude protein than post-jointing wheat. Prior to wheat entering the jointing stage, all crude protein values for wheat diets were higher than that of prairie diets (£ < 0.01). Post jointing, crude protein differences were detectable among the diets only in the first 2 sampling periods (£ = 0.05 and £ < 0.01, respectively). These periods corresponded to the last week in April and the first week in May. These trends were similar to those found in Alldredge and Torbit (1987:41), whose data suggested shortgrass prairie dietary crude protein exceeded wheat dietary crude protein in early May. The relation between crude protein and time was better described in a quadratic model than in a linear model, Ho: ~2 = 0 and Ho: ~2 = 0, for both wheat and prairie diets (£ = 0.02 and £ < 0.01, respectively). These quadratic models were used to generate the independent variables for the logistic regressions of pronghorn habitat use versus dietary crude protein. Each logistic regression model was solved for the time when marked pronghorn use of the vegetation types was equal, ilv = ilR = 0.5. These times corresponded to mean crude protein contents of 26.9% and 16.1% for wheat and prairie diets, respectively. Backcalculating these values through the dietary crude protein versus time regression gave the corresponding dates April 12th and April 10th. Telemetry, 1995 Five pronghorn were relocateable during the 1995 field season. Five deaths were confirmed during the study. Two transmitter failures were visually verified. The fates of two marked pronghorn were not determined. Pronghorn shifted from winter wheat fields to shortgrass prairie as we expected (£ < 0.0001). .Differences among individuals were not detectable (£ = 0.12) The last marked-animal relocation on wheat was March 23rd, which was about one month earlier than previous years (Table 3). More importantly, this behavioral shift occurred prior to when winter wheat became vulnerable to grazing damage. All wheat had entered the jointing phenological stage and became vulnerable to grazing·damage on April 15th. Feeding Trials Pronghorn had symmetric (balanced) exposure to the relative position of treatments (wheat stages) in both feeding-trial designs.. However, the designs became unbalanced when individuals chose not to participate in certain trials. These no-response data were omitted from this analysis. ~even of 9 pronghorn showed a lateral tendency in the box experiment; meaning, they consistently (~67%) chose one side of the box independent of wheat stage. Five of these animals chose the north side of the box. The 2 animals, P and T, which chose the south side of the box were the youngest animals and some of the tamest. Only one animal, T, showed a strong preference for tillering wheat (Table 4). Consequently, the null hyothesis of random selection of �243 wheat by pronghorn was not rejected (E = 0.98). Biomass did not influence pronghorn selection (E = 0.54). Pronghorn consumed both wheat stages 65 ± 7 (SE) % of the time. Table 4. Pronghorn experiment. selection of wheat developmental stages via first bite by Number of Trials with a response Number of times tillering wheat was selected B 7 3 0.43 c 6 2 0.33 D 7 4 0.57 E 9 4 0.44 N 12 4 0.33 P 12 5 0.42 S 8 4 0.50 T 10 9 0.90 y 3 2 0.67 B 10 9 0.90 c 14 10 0.71 D 5 3 0.60 E 12 6 0.50 N 15 8 0.53 P 14 7 0.50 S 11 6 0.55 T 12 8 0 •. 67 y 0 0 Animal Mean response (n) Box experiment 4-pot experiment Only 8 pronghorn ate wheat in the 4~pot experiment; Y never approached the wheat. Three animals showed a lateral tendency in the 4-pot experiment. In each case, they favored the closer (east) group of pots. Distance between the 2 groups was <0.3 m. Only B showed a strong preference for tillering wheat (Table 4). Again, there was no evidence to suggest that pronghorn preferred tillering wheat (~= 0.96). C chose tillering wheat 71% of the time, while T chose it 67% of the time. Biomass did not influence pronghorn selection (E = �244 0.90). During the trials and ~3 pots were consumed animals ate from ~2 pots 89 ± 4 (SE) % of the time 59 ± 10 (SE) %. Individual pronghorn lacked consistency in the strengths of their lateral tendencies between the box and 4-pot experiments (r = -0.09, £ = 0.83). Individuals also showed little consistency in their selection of tillering wheat between the experiments (t = 1.65, df = 7, £ = 0.14). DISCUSSION Telemetry Our data supported the premise that female pronghorn may not damage wheat, because marked pronghorn use of wheat terminated prior to jointing. However, our data do not as clearly link changes in crude protein to when pronghorn stop using wheat. Whatever the mechanism is, males appear less sensitive to it than females which were in the la~t trimester of pregnancy at this time. We used our 1994 models to estimate a mean date for the period of no differences in pronghorn vegetation use. The mean of these predicted values was February 25th. This is almost one month earlier than the mean date from the telemetry data, March 24th. One factor contributing to the models' predicting early dates was that when pronghorn were using wheat fields they were also relocated on short-grass prairie. Use of wheat fields gradually decreased over time. This also caused poor fit of the regression lines. Recall our premise for pronghorn forage selection was simply that we expected pronghorn to select the most nutritious forage. The crude protein data suggested that we may want to modify this hypothesis to include some type of threshold effect. Below the threshold value foraging decisions with respect to vegetation type were based on nutrition. Above. the threshold these foraging decisions may be based on palatability factors, for instance. As "rule of thumb" adult ruminants require a diet with 10% crude protein to meet maintenance requirements. Our data suggested that this may be an appropriate threshold. Recall our last regression analysis found no difference in pronghorn use of vegetation types until the crude protein content of prairie diets reached 16.1%. Feeding Trials We -found that getting pronghorn to place their head in the box to eat anything was difficult. In fact, after months of training only 3 animals were consistently eating from the box. Serendipitously, we found that opening the isolation pen door dramatically improved the chance that the more timid animals would eat out of the box. Their dislike for being closed-in suggested that the general preference for the north position in the box experiment may stem from the presence of the south wall of the isolation pen. This implied the feed box was unsuitable for trials involving the simultaneous presentation of feeds. At this point, the conclusion that pronghorn cannot distinguish between tillering and jointing wheat was unwarranted. The general lack of selectivity coupled with the tendency for the animals to eat all of the pots suggested �245 that these tame animals were not highly motivated to discriminate between the two types of wheat. The pronghorn appeared to react to the wheat as a novel food source and the excitement of getting the wheat exceeded interest in selecting types of wheat. This rationale was further supported by watching the pronghorn respond to pots of wheat in their pastures. We believe that are some critical points concerning the planning of more feeding trials. One, increasing replications is a doubtful means of getting better (stronger selection) results. Two, increasing the amount of biomass presented and the length of the trials is probably necessary for making preferences detectable.. Both need to approximate (at least) that observed during normal feeding bouts. 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Bioscience. 37:789-799. �248 Spiertz, J. H. 1977. The influence of temperature and light intensity on grain growth in relation to carbohydrate and nitrogen economy of the wheat plant. Neth. J. Agr. Sci. 25:182-197. . Tenhunen, J. D., C. S. Yocum, and D. M. Gates. 1976a. Development of a photosynthesis model with an emphasis on ecological applications. I. Theory. Oecologia. 26:89-100. _____ , J. A. Weber, C. S. Yocum, and D. M. Gates. 1976b. Development of a photosynthesis model with an emphasis on ecological application. II. Analysis of a data set describing the pH surface. oecologica. 26:101119. Tilley, J. M. A., and T. A. Terry. 1963. A two-stage vitro digestion of forage crops. J. Br. Grassl. technique for the inSoc. 18:104-111. Torbit, S. C., R. B. Gill, A. W. Alldredge., and J. F. Liewer. 1993. Impacts of pronghorn grazing on winter wheat in Colorado. J. Wildl. Manage. 57: 173-181. Van Soest, P. J. 1982. Nutritional Corvalis, Oreg. 374pp. ecology of the ruminant. Brooks, Welsh, J. G., and A. P. Hooper. 1988. Ingestion and feed and water. Pages 108-116. in D. C. Church, ed. The ruminant animal: digestive physiology and nutrition. Preston Hall, Englewood Cliffs, N.J. White, G. C., D. R. Anderson, K. P. Burnham, and D. L. otis. 1982. recapture and removal methods for sampling closed populations. Alamos Nat. Lab. Report LA-8787-NERP, Los Alamos, N.M. 235pp. Wild, M. A., M. W. Miller, B. J. Maynard, D. R. Magnuson. 1992. Animal and pen support facilities for terrestrial wildlife research. Colo. Div. Wildl. Res. Rep. Fed. Aid Proj. W-153-R5, WP1A J1, Job Progr. Rep., July 1991-Ju~e 1992, Fort Collins. CaptureLos Woledge, J., and A. J. Parsons. 1986. Temperate grasslands. Pp. 173-197 N. R. Baker, and S. P. Long, eds. Photosynthesis in contrasting environments. Elsevier Sci., New York., N. Y. in Yonker, C. M., and D. S. Schimel, .E. Paroussis, R. D. Heil. 1988. Patterns of organic carbon accumulation in a semiarid shortgrass steppe, Colorado. Soil Science Society of America Journal. 52:478-483. �249 Colorado Division Wildlife Research July 1996 of Wildlife Report JOB State of Project Work No. W-153-R-9 3A Job No. Author: REPORT Colorado Plan No. Period PROGRESS Mammals Research Pronghorn Inyestigations Experimental Pronghorn Surveys Using Fixedwing Line Transects and Helicopter Quadrats Covered: July 1, 1995 - June 30, 1996 T.M. Pojar ABSTRACT The third year of a 3 year study was completed to compare pronghorn population density estimates obtained by fixedwing line transect and helicopter quadrat surveys. As was done in 1995, line transects were run on 1.609 km (1 mile) intervals in 1996 resulting in a sample of 16 transects for the surveyed area. Since attempts to stratify the quadrat sample based on the 1995 data resulted in no improvement in the variance, the sample was not stratified in the 1996 analysis. The 1996 estimated population was 5,937 ± 26% from the helicopter quadrat survey and continues the downward trend of the past 2 years. The 1996 line transect estimate ranged from 4,776 to 5,537 (depending on the analysis used). The:z:::e is a consistent relationship between the quadrat survey estimate and the line survey estimate in that both show a downward trend during the 3 years of data collection reflecting the management action to reduce the herd. Although the trend is similar, the line estimates are always less than the quadrat estimates. Using the group size as estimated by the regression in DISTANCE for all 4 distance irttervals seems to provide the population trend that most closely follows that of the quadrat estimates. A manuscript authored by Pojar, Bowden, Madison, and Gill will be prepared and submitted to the Journal of Wildlife Management. �250 MANAGEMENT RECOMMENDATIONS The relationship between quadrat and line transect surveys should, be established for each area. that line transect sampling is contemplated. The relationship could vary with terrain, vegetative type, and pronghorn density. The minimum data set to estimate this relationship is 3 years of data. It is then preferable to make this same comparison every third or forth year thereafter. Line transect methodology must be followed meticulously and observers must be trained in the principles and techniques of line transect methods to get the best results. Sampling intensity should be 10% for quadrats and line samples should be at 1 mile intervals. For the Craig experimental area, it should'be assumed that the line transect population estimate is 80% of the "true" number of animals (pending further analysis of this data set). The line transect population estimate should be adjusted upward by dividing it by 0.80 to obtain an estimate that, on average, would closely correspond with the quadrat estimate. For analysis, the halfnormal function for estimating f(O) should be used with the cosine adjustment feature of DISTANCE. Group size is a sensitive parameter in the calculation of population size because it is multiplied by group density and total area to get total population size (i.e. GS*GD*Area=pop size). In general, it appears the use of the regression in DISTANCE on all 4 distance bands to estimate group size will give satisfactory results. �251 EXPERIMENTAL PRONGHORN SURVEYS USING FIXEDWING HELICOPTER QUADRATS Thomas LINE TRANSECTS AND M. Pojar P •N. OBJECTIVE Compare fixedwing .line transect pronghorn density. and helicopter SEGMENT quadrat in estimating OBJECTIVES 1. Compare pronghorn density estimate and precision transect survey with helicopter quadrat survey. 2. Evaluate 3. Test accuracy of a Global geographic points. consistency surveys of line transect Positioning of fixed-wing line data analysis. System (GPS) in locating known STUDy AREA The study area is 1,171 km2 (452 mi2) of sagebrush steppe pronghorn north and west of craig. It is described in Pojar et al. (1995). METHODS habitat AND MATERIALS The methods are outlined in the Program Narrative, see pojar I. Modification to the methods are described in Pojar 1995. (1994), Appendix Because of the unavailability of the contractor that flew the fixedwing line transects in 1994 and 1995, a different·contractor was used in 1996. The major difference was that the 1996 contractor used a Cesna 185; the previous contractor used a Maule. The procedure for flying the lines and recording data was identical to the past 2 years. There was nothing in the appearance of the 1996 data set that would indicate a difference from the past 2 years. The quadrats were searched using a Bell-Soloy helicopter in 1996; Brad Petch (CDOW) served as navigator/secondary observer. The GPS external antenna was mounted on the rear boom behind the engine of the helicopter. The GPS was used exclusively to locate quadrat corners and navigate the quadrat perimeter; if old quadrat markers were observed they were ignored. The navigation was done via GPS by the second observer/navigator communicating directions to the pilot while the primary observer counted and recorded pronghorn groups on a tape recorder. �252 RESULTS The line transect survey was done on May 29 and 30, 1996 by Sky Aviation of Worland, Wyoming with Jeff Madison and Mike Bauman (CDOW) as observers. Methodology followed that described by Johnson et ale 1991. Analysis procedures for line transect data will follow those described by Buckland et ale (1993) and Laake et ale (1993). In addition, the data will be subjected to analysis by the program, TRANSAN (Routledge and Fyfe 1992), which implements the shape-restricted estimator of Johnson and Routledge (1985). The helicopter survey was done on May 28 and 29, 1996. The population estimate was 5,937 with a 90% confidence interval of ± 26% (Table 1). The lower population estimates since 1994 (8,465) was expected because management efforts to reduce the population (Jeff Madison, Pers. corom.). Table 1. Results of helicopter quadrat surveys to estimate on a 452 square mile experimental area NW of Craig, co. pronghorn density % Change Year Pop. Est. 90% C. L 1994 8,465 ± 30% 1995 7,708 ± 33% - 8.9% 1996 5,937 ± 26% - 23.0% In the following analysis I used the half-normal function for estimating f(O) and used the cosine adjustment feature of DISTANCE. Group size is a sensitive parameter in the calculation of population size because it'is multiplied by group density and total area to get total population size (i.e. GS*GD*Area=pop size). In table 2, I use some different group size estimates to demonstrate the differences in population size that can result. Hopefully, further scrutiny of the data sets,will lead to a "best" analysis procedure. There is a generally consistent pattern in the line transect data sets, i.e., they all have a downward trend (Figs. 1 and 2). An important product of this experiment is to establish if there is a relationship between quadrat and line transect survey results. Helicopter quadrats cost approximately 10 times more than line ,transects to execute. Therefore, for management purposes; a close relationship between the 2 results would permit using the cheaper method for most years' data. In general, the line estimates follow the 3-year downward trend of the population as estimated by the quadrat survey (Figs. 1 and 2). Using the regression estimate in DISTANCE (Laake et ale 1993) for all 4 distance bands ,appears to most closely track the slope of the quadrat regression (Fig. 2). �253 Table 2. Results of fixedwing line transect surveys to estimate density on a 452 square mile experimental area NW of Craig, co. pronghorn Year Group Density 1994 4.2510 3.16691 3.76472 3.55903 4.02804 6,085 7,234 6,838 7,740 4.7125 3.02661 2.60652 2.61873 2.37884 6,447 5,552 5,578 5,067 5.1174 2.39381 2.26142 2.06463 2.17894 5,537 5,231 4,776 5,040 Group Size Pop. Est. 1995 1996 4 Mean group size for all distance bands. Mean group size f.or the first 2 distance bands. DISTANCE generated regression estimate of group DISTANCE generated regression estimate of group distance bands. REFERENCES size from all groups. size from the first 2 CITED Buckland, S. T., D. R. Anderson, K. P. Burnham, and J. L. Laake. 1993. Distance sampling: estimating abundance of biological populations. Chapman and Hall, New York, N.Y. 471pp. Johnson, B. K., F. G. Lindzey, and R. J. Guenzel~ 1991. Use of aerial line transect surveys to estimate pronghorn populations in Wyoming. Wildl. Soc. Bull. 19:315-321. Johnson, E. G., and R. D. Routledge. 1985. The line transect method: a nonparametric estimator based on shape restrictions. Biometrics 41:669679. Laake, J. L., S. T. Buckland, D.' R. Anderson, and K. P. Burnham. 1993. DISTANCE user's guide. Version 2.0. Colo. Coop. Fish and Wildl. Res. Unit, Colorado State Univ., Fort Collins. 72pp. Pojar, T. M. 1994. Experimental pronghorn surveys using fixedwing line transects and helicopter quadrats. Colo. Div. Wildl. Res. Rep. July, pp163-172. ______ • 1995. Experimental pronghorn surveys using fixedwing line transects and helicopter quadrats. Colo. Div. Wildl. Res. Rep. July, PP???-??? ______ , D. C. Bowden, and R. B. Gill. 1995. Aerial counting experiments to estimate pronghorn density and herd structure. J. Wildl. Manage. 59:117-128. 1992. TRANSAN: Line transect estimates Routledge, R. D., and D. A. Fyfe. based on shape restrictions. Wildl. Soc. Bull. 20:455-456. �254 - 0 0 0•.. 'I"""" - 9 Em quads All :::::::: 2 bnds II Reg all :I:tt Reg 2 8 X Q) .•... co 7 E .p (I) w 6 r:::: 0 .p co :; 5 a. 0 a, 4 1995 Figure 1. Comparison of population estimates from helicopter quadrat and fixedwing line transect surveys. Four different estimates were generated for lines using 4 different estimators of group size. All = group size mean in 4 distance bands, 2 bnds = mean from the first 2 distance bands, Reg all = group size based on a regression for all groups, and reg 2 = group size based on a regression for groups in the first 2 distance bands. 0 0 0•.. 10 9 'I"""" X 8 Q) .•... co E .£i 7 r:::: 0 .p 6 :J a. 5 w co 0 a, 1995 1996 4._--------------------------------------------~ 1994 Figure 2. Regressions for helicopter quadrat and 4 estimates for line transect population size estimates.· legend description. See Figure 1 for �255 Colorado Division of Wildlife Wildlife Research Report July 1996 JOB PROGRESS REPORT state of Colorado Project No. ~W~-~1~5~3~-~R~-~9~ _ Mammals Research Work Plan No. __~4~AL- _ Mountain Goat Inyestigations Job No. Period Covered: Mountain goat numbers, distribution, and dispersal in the northern collegiate range. July 1, i995 - June 30, 1996 Author: D. F. Reed ABSTRACT This study of mark-recapture population estimates, distribution and habitat use, and the pioneering of mountain goats in the northern collegiate range has relied on additional monies from the bighorn sheep and mountain goat auction and raffle funds. During 1994-95, $13,500 was awarded to compliment P-R Project monies for the cost of radio~collaring the 50 mountain. goats (completed 10-15 August 1994) needed in the study. Subsequently, $12,500 and $9,800 were awarded for 1995-96 and 1996-97, respectively, for distribution and habitat-use work. It was found that fixed-wing flights provided limited useful distribution data and that additional helicopter flights were needed to collect this data. During two winters and two hunting seasons, 5 collared animals have died and 7 have been harvested. Seven additional telemetry collars were attached in 1996 to replace most of these 12. However, one of the 7 additional collars was put on an animal out of 'the mark-resight study area northwest of Mount Elbert. This mark that was put on an individual in a group of 20 goats was important for addressing the dispersal part of the study, but does not replace a mark in the "mark-resight" study sample (sample·size 43 vs 44). Helicopter counts were conducted 25 August 1994, 30 August and 1 september 1995, and 18 July and 2 August 1996. The results of these counts ranged from a low number of the marks (radio-collars) being observed and poor identification of distinguishing marks (numbered or color coded collars) (9 of 39 marks; sightability = 0.23) to a high number of marks being observed and good identification of marks (29 of 36; sightability = 0.81). The best estimate for the study area population (north of Clear Creek) appears to be from the count on 30 August 1995 (N = 173 [CI 147-203]). Movements of up to 17 km have been detected for both males and females. The group of 20 goats found northwest of Mount Elbert (in which Black 74 radio-collar was placed) was 13 km north of Lake Creek and Highway 82. This appears. at this time to be a significant pioneering group. ��257 MOUNTAIN GOAT NUMBERS, DISTRIBUTION, NORTHERN COLLEGIATE AND DISPERSAL RANGE IN THE Dale F. Reed P .N. OBJECTIVE To improve estimates of mountain goat populations by mark-resight methodology, to determine distribution, and to estimate dispersal rates in an increasing mountain goat population. SEGMENT OBJECTIVES 1. Replace marks (radio-collars) in the original the North Collegiates (Quail-La Plata). 2. Conduct helicopter counts to test mark-resight methodology (resight marks [obtain resighting estimates], count unmarked animals, and estimate population) and to determine distribution and habitat use. 3. Conduct habitat fixed-wing flights to assist· in use of telemetered animals. STUDy The study area estimating mountain goats distribution in and AREA is des·cribed in the Program METHODS· 50 marked Narrative (Reed 1995) .• AND MATERIALS The methods are outlined in the Program Narrative (Appendix A in Reed 1995). For net-gun capturing a Hughes. 500 D (Helicopter Wildlife Management) was used. Once the pilot and gunner located and netted an animal, the gunner was quickly landed to handle the animal while the pilot often left to bring in another person waiting nearby on the mountain to assist and finish collaring, taking samples, and releasing the animal. This allowed the gunner to be off netting another .animal. For counting a B-47 So loy (High country) and the Hughes 500 D were used. Count techniques included either one or two observers. When two observers were used in the B-47, the middle person counted the left side and forward, and the person on the right side, counted right. The middle observer also did orienteering and .recording of observations on layed-out quad maps. During the most recent flights, only one observer. was used to lessen the weight and therefore increase the capability of the helicopter for the close-in work of identifying the distinguishing collar numbers and/or colors. This is important because the elevations involved, the B~47 is often operating at its upper-limits depending on air conditions (temperature/air currents). To assist in identifying distinguishing marks (numbers and/or colors) use of binoculars would seem appropriate but vibration of the helicopter seriously diminishes clarity. This problem was largely overcOme by using a Gyro Stabilizer (Ken-Lab, Inc., 29 Plains Road, Essex, CT 06426, (860) 767-3235) attached to the binoculars. at �258 For estimating general locations by teiemetry a Ce.ssna 185 was used. The technique involved flying relatively high (15,000-16,000 ft) and listening to the strength of signals from two external antennas. Forty-five and five of the original transmitters had frequencies of 165.20-165.72 MHz and 173.0372-173.2127 MHz, respectively. A LOTEK (Suretrack STR1000) receiver was used for the 165 frequencies and a Telonics (TR-1) was used for the 173 frequencies (Channels 7-8, 16-18) • The 7 replacement collars were all in the 165 MHz band. Collars supporting the transmitters were either color coded or color coded and numbered for individual identification. RESULTS Consistent with the plans in the Program Narrative, 50 mountain goats were captured and radio-collared 10-15 August 1994. Most of the animals collared (n = 30) were adult females (Table 1). Subsequently, 5 more females and two 3year old males were collared. The efficiency of capturing and radio-collaring mountain goats via helicopter net-gunning was calculated as 2.4 animals/hour but variables likely influence such efforts (e.g. pilot and net-gunner performance, weather, animal distribution and group size, etc.). Predictably, the efficiency declined during capture of the 7 additional animals (1.2 animals/hour) since a different pilot was used and only selected groups of animals were being sought. Table 1. Number of mountain goats captured and radio-collared 10-15 August and 17-18 July 1996 by sex/age group and approximate hours of flight. Date Aug 94 10 11 14 15 Male ~!:lJ.ll:t Female 2-:i!iil~u:: Ql!:l Ma:le Female 4 6 2 12 0 3 (waited two days 8 0 0 6 4 0 ::r::!iili:!.t:ling Male Female 0 1 for animals 0 0 0 0 Total Approximate hours of flight 1 1 to adjust) 0 0 0 2 13 17 6 7 8 12 4 4 0 50 21 10 30 Jul 96 17 18 0 2 4 1 4 3 2 4 Total 2 5 7 6 Total 5 1 4 1994 It was found that fixed-wing flights provided limited useful distribution data and that additional helicopter flights were needed to collect this information. More definitive data on distribution and habitat use has yet to be fully analyzed and will be reported in the next Progress Report. During two winters and two hunting seasons, 5 collared animals have died and 7 have been harvested (Table 2). Seven additional telemetry collars were attached in 1996 to replace most of these 12. However, one of the 7 additional collars was put on art animal out of the mark-resight study area northwest of Mount �259 Elbert. This mark that was put on an individual in a group of 20 goats was important for addressing the dispersal part of the study, but does not replace a mark in the "mark-resight" study sample (sample size 43 vs 44). Table 2. Number of marks (telemetry collars) present (before fall hunting season), harvest, and mortality of marked mountain goats in the Quail Mountain-La Plata Peak study area. Year Marks Present Cause of mortality Hax::Yeateg Harvested Mortalities 1994 50 4 1 Unknown 1995 45 3 4 Avalanches implicated for 2; 2 unknown 1996 39 45 Total After 6 replacement marks put on in study area 17 and 18 Jul 96' . , a 7th mark was put on outaige the study area .NW of Elbert for a total of 7 replacement marks 7 5 Five helicopter counts have been conducted since 25 August 1994 (Table 3). The results of these counts ranged from a low number of the marks (radio-collars) being observed and poor identification of distinguishing marks (numbered or color coded collars) (9 of 39 marks; sightability = 0.23 [Table 4]) on 18 July 1996 to a high number of marks being observed and good identification of marks (29 of 36; sightability = 0.81) on 30 August 1995. Each of these occasions need to be examined before conclusions should be drawn. The first count (25 Aug 94) yielded 107 mountain goats, 74 north of Clear Creek and 33 south of Clear Creek. Of the 50 collared animals only 13 were observed. Gusty winds prevented classification of 21 animals and searching the head of Willis Gulch and the west side of Hope Mountain. Furthermore, we were unable to identify collar numbers from the helicopter on that occasion. Conversely, the next count (30 Aug 9S) exceeded expectations when 29 of. 36 marks were observed and igentifieg, a total of 124 animals counted, and a aightability of 0.81 calculated. After waiting 2 days, the next count (1 Sep 95) yielded only 16 of 37 marks and a total of 93 animals. The animals appeared to be scattered and the pilot was more cautious on this occasion. The 4th count was conducted using the Hughes 500 and.this was done after some of the animals in Galena were disturbed during helicopter capture efforts earlier in the morning and clouds and some rain may have influenced the results. The results on this occasion should perhaps be discarded. The last count (2 Aug 96) seemed to go well but only 16 of 43 marks �260 were observed and identified with 89 total animals being counted. On this occasion the gyro mounted binoculars was judged to help in identification of the collars. The best estimate for the study area population (north of Clear Creek) appears to be from the count on 30 August 1995 (N = 173 CCI 147-203]). (Table 4). Some other values, calculated from the occasions with low sightability, probably can be judged to be "outliers." Table 3. Year, date, helicopter type, and general results of total in study area; total goats counted) of helicopter La Plata Peak study area. Year Date of Flight (no. of marks sighted flights in the Quail- General Type results 1994 25 August B-47 (Soloy) 13 of 50 marks; n = 74 1995 30 August 01 September B-47 B-47 29 of 36 marks; 16 of 37 marks; n n = 124 93 18 July Hughes 9 of 39 marks; n = 51 02 August B-47 16 of 43 marks; n = 89 1996 500 D Table 4. Counts for the study area (SA;Quail-La Plata) and for the extended study area (ESA;Quail-La Plata plus Cross Mtn-Waverly), number of marks sighted and identified, total number of marks in area, total number animals counted, sightability, and mark-recap~ure population estimates with 95% confidence intervals (CI). NymJ:!sU: of marks Count date/ area Sighted & identified 1994 25 Aug 1995 30 Aug 01 Sep In area Total no. counted Sightability M-R Population Estimates 13 50 74 0.26 SA 29 36 124 0.81 173 (CI 147-203) ESA 31 40 153 0.76 229 (CI 191-279) SA 16 35 93 0.46 (Combined Occasions 1 and 2, NOREMARK) 107 16 37 ESA 164 (CI 146-195) 0.43 1996 18 Jul 9 39 51 0.23 02 Aug 16 43 89 0.37 �261 Movements of up to 17 km have been detected for both males and females (Table 5). Dispersal and/or special status of each telemetered mountain goat are noted and subject to change (Table 5). The group of 20 goats found northwest of Mount Elbert (in which Black 74 radio-collar was placed) was 13 km north of Lake Creek and Highway 82. This appears at this time to be a significant pioneering group. Table 5. Collar frequency or channel (color/number designation), location when collared (10-15 Aug 94), estimated location during helicopter flights 30 August 1995 and 2 August 1996, and noted dispersal and/or special status. Frequency/Channel (color/number) 10-15 Aug 94 165.200 (Blu 20) Galena 210 220 (Grn 21) (Grn 22) W " " Locations 30 Aug 95 (- same) (Grn (Grn (Grn (Y/B (Grn 290 300 (Yel 29) (Grn 30) Galena Willis 320 330 340 350 (Blu 32) (Or 33) (Yel 34) (Blu 35) NE Ellingwood W Galena (? ) 360 371 380 390 (Yel (Yel (Yel (Yel 410 420 (Blu 41) (Blu 42) 430 450 (Blu 43) (Or 45) Middle 460 (Or 46) Galena E Crystal L 5E Hope Middle Mtn (?) Cirque (? ) (Grn 23) (none - harvested 9/25/94 Lake 7 km 5E Middle Mtn) (- same) (- 2-3 km W) (- same as 300) NW Twin Pks 5W Willis Cirque Galena (?) (? ) W Galena " (- same) " " 240 250 260 270 280 36) 37) 38) 39) Dispersal/status Cirque 230 24) 25) 26) 27) 28) 2 Aug 96 " Mtn (?) Pear " " (mortality 7/19/95; replaced collar 7/17/96 on adult female; 7/17 to 8/22 96 moved 8 km from Continental Divide to La Plata Gulch) (- same) (harvested 9/7/95) (- same) (W Twin Pks) "(left side missing no.) (none - harvested 9/11/94; replaced as Blk 39 Jul 17, 96 on adult female in Continental Divide group) (- same) (5 Clear Ck, NE Waverly Mtn)(N Clear Ck) ("" (? ) near " ")(" ") (5 & low on Quail, 5-6 km NE) (mortality 11/30/95, recovered 7/20/96 N Middle Mtn) (5 Clear Ck, NE Waverly Mtn) (La Plata Gulch) (Distances between these point locations = 12 and 17 km) �262 Table 5 (continued) Frequency/Channel (color/number) 470 (BIU 47) 480 (Blu 48) 490 (Or 49) (? ) W Galena (- same) (La Plata Galena 3 km) (? ) 510 (Blu 51) (? ) 520 (Blu 52) (? ) 530 540 (Blu 53) (Blu 54) (? ) Upper Galena 550 (Or 55) Galena 590 (Blu 59) Middle Mtn 600 (Blu 60) 610 (Blu 61) 620 (Blu 62) 630 (Blu 63) (? ) (? ) SE La Plata (?) 640 Middle (?) Pk Mtn 650 (B/Y 65) 660 (Orange) 670 (Yellow) 680 (B & W) SE La Plata Pk S Willis Lake W Galena Cirque Sayres Gulch 710 720 740 SE Twin Pks SW " " (Black) (Blu/Or) (Black) 7 (Grn 7) 2 Aug 96 Dispersal/status (- same) (mortality 7/19/95, recovered 9/31/95; replaced collar 7/17/96 on adult female near Continental Divide) (S Clear Ck, NE Waverly Mtn) (Upper E Fork Sayres Gulch; moved 17 km from Waverly to Sayres) (? ) 500 (Blu 50) (Blu 64) Locations 30 Aug 95 10-15 Aug 94 N Middle Mtn Pk; moved from W (est W Quail) (La Plata Gulch; collar appeared Blue & White) (S Clear Ck, NE Waverly Mtn) (Willis Gulch; move 10 km) (S Rinker Pk)(Harvested 9/15/95 Twin Pks; replaced collar 7/18/96 on 3-yr male in Galena) (Twin Pks) (none - harvested 9/6/94 E Galena; replaced collar 7/18/96 on 3-yr male in Galena) (S Clear Ck, NE Waverly Mtn)(SE La Plata?) (Low & SW Quail)(E Quail & 13,130 by tele) (move 5 and 3 km) (S Independence Pass?) (La Plata Gulch) (Galena Gulch) (Willis) (Crystal Lake Creek)(S La Plata) none) (Mortality 9/1/95, recovered 8/13/96 about 2 km S La Plata Pk) (none - harvested 9/15/94 near Ann Lake 7 km S Middle Mtn) (N La Plata Pk)(La Plata Pk) (- same) (La Plata Pk by tele) ( )(?) (- same) (Mortality < 6/27/95, recovered 9/ /95 NE Winfield; moved 9 km. from Sayres to Winfield) (Replaced collar on adult female 7/17/96 near continental Divide) (- same) ( ? ) ( )(?) (none) (none) (Collared 7/18/96 NW Mt Elbert) (none - not found)(S Independence Pass; moved about 5 km to NW from N of Middle Mtn, i.e. the "Middle Mtn" that is near the Continental Divide, ~ the one near Cross Mtn) �263 Table 5 (continued) Frequency/Channel (color/number) 8 (Grn 8) 17 (Grn 17) 18 (Grn 18) Locations 30 Aug 95 10-15 Aug 94 SE La Plata Pk NE Ellingwood Middle Mtn (?) LITERATURE Reed, 2 Aug 96 Dispersal/status (S Silver King Lake; moved 16 km to SE)(?)16(Blue) W Willis Lake (Garfield Pk)(E Sayres; moved 7 km Willis to sayres) ( ? )(NW Galena) (S Silver King Lake; moved 9 km SE from Middle to Silver King) (Harvested 9/16/95 NW Browns Pk; moved 8 km from Silver King to Browns) CITED D. F. 1995. Mountain goat. numbers, distribution, and dispersal northern Collegiate range. Colo. Div. Wildl. Res. Rep. July, in the pp. �264 �2~ Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS state of Project Colorado No. Work Plan No. W-153-R-9 SA Job No. Period Author: REPORT Covered: Thomas Mammals Research Black Bear Research Development of black Inyentory Technigyes bear July 1, 1995 - June 30, 1996 D. I. Beck Personnel: T. Beck, S. Birch, R. Firth, M. Guy, A. Vitt, York, CDOW; D. Harper, D. Marlow, R. stevens, Colo. state USFS; D. Bowden, G. White, Colo. State Univ. L. Willmarth, D. Patrol; D. Schmidt, Abstract A total of 78 captures of black bear (~ americanus) were made over a 100day period with a total of 2,525 trap days. The capture sample was comprised of 7 adult males, 10 adult females, 18 subadult males, 13 subadult females, and 1 unknown. Five minor injuries were observed, the most serious a broken canine tooth. Sex and age composition of the sample was strikingly similar to the Uncompahgre study. Similarities to annual kill data proportions suggest a revision may be appropriate in simulation models; the net effect being to lower proportion of adult females and thus the reproductive potential. Body size of Middle Park bears is about 80% of that observed in southwest Colorado. ��267 DEVELOPMENT OF BLACK BEAR INVENTORY Thomas TECHNIQUES D. I. Beck P.B. OBJECTIVE 1. Evaluate a capture-sight program utilizing for estimating black bear density. 2. Document age and gender autumn hunting seasons. 3. Obtain density estimates of black bears in 3 heavily markedly different vegetation communities. bias in vulnerability SEGMENT 1. capture area. 2. Evaluate bears. and radio-collar METHODS set on bait of black bears hunted stations during areas of OBJECTIVES up to 75 black the use of infra-red cameras triggered bears in the Middle cameras to resight Park marked study black AND MATERIALS Study Area Description The Middle Park study area was located in GMU 18 and was chosen because this area was consistently the top area for hunter killed bears among the high elevation areas of the state.· Mountain areas vegetated with spruce-fir (~ - ~) and lodgepole pine (limui contorta) forests account for 30% of the statewide black bear habitat (Beck 1991). The 435 km2 area was divided into 2 42 quadrants, each 10.4 km • The area is roughly 22 km north-south and 19 km east-west. The west boundary is the E. Fork Troublesome Creek, the east is a north-south line over Little Gravel Mtn., the north boundary is just south of the Continental Divide, while the south boundary parallels the Colorado River. The south boundary is the only one with a distinctive vegetation change .with the lower boundary being the upper limits of extensive sagebrush (Artemisia spp.) hills. Elevations range from 2380 m to 3540 m. The average frost-free period is approximately 45 days. The southern row and western 2 columns of quadrats exhibit a mosaic of vegetation with substantial stands of aspen (Populus tremuloides). The remainder of the quadrats are characterized by relatively homogenous conifer stands interspersed with meadows. Capture and Marking Black Bears Capture procedures basically follow those outlined for the earlier study. on the Uncompahgre Plateau (Beck 1994). The goal of equal trapping effort for each quadrat was unattainable because of access restrictions and unusually high water. �268 Because of the poor ear-tag retention observed on the Uncompahgre Plateau (Beck 1995),. a new marking system was developed. All black bears were tagged in one ear with a DuFlex Round Hog Tags, numbered on the front and hand lettered on the rear with MP CDOW BB RES. Ear tags were yellow or white. Female bears were tagged in the right ear, male bears in the left. In addition, a unique marking system was affixed to the collar. Two colored dowels were attached to the collar opposite the transmitter so that they would normally ride on the top of the neck, extending 10 cm above the collar. The dowels were cut from 2.5 cm diameter birch dowel rod, in lengths of 10 cm. Colored 22 oz PVC (scraps provided by Jack's Plastic Welding, Aztec, NM) was glued over the entire surface of the dowel. Eight colors were used, allowing for unique marking of 64 black bears. Colors were red, white, yellow, blue, purple, orange, teal, and red-and-white stripes. Each dowel was pre-drilled at one end to accept a 4 cm woodscrew. To reduce dowel splitting, a hose clamp was tightened around the pre-drilled end. In addition to gluing, the plastic material was also stapled to the dowel. The relative position of the dowels (left, right) along with the colors provide the keys to unique identification. The dowels would be affixed to the collar after the collar had been fitted to an individual bear. After fitting, the collar would be marked opposite the transmitter, then removed. Two screw holes would be made with an awl, approximately 7 cm apart, through the collar material. A flat washer waS placed on each wood screw and glue placed in each dowel hole prior to tightening. After affixing the dowels, the collar would be placed on the bear. All bears over 27 kg were to be collared. Another change from the Uncompahgre study was that the canvas spacers placed in the collars were not treated with a preservative and they were lightly scored with razor to facilitate decomposition. This was in response to poor disintegration of the canvas observed with the treated collars. a The following characteristics were recorded for each bear: sex, age group (c~b, subadult, adult), color; weight, total length, chest girth, number of incisors, broken teeth, nipple diameter and color on females, pelage condition, breathing rate. Date, quadrant, and UTM coordinates were recorded. All crew members received detailed written and oral instructions on the Safe handling of trapped bears, emphasizing safety to the bears foremost. Addition~lly, all new crew members worked at least 10 bears with an experienced handler (Beck or Willmarth) before handling a trapped bear alone. Monitoring seasonal movements Aerial radio-tracking June, 1995. Location was conducted at biweekly intervals beginning was recorded as UTM coordinates. in late �269 RESULTS AND DISCUSSION capture and Marking Black Bears Over a 100-day trapping period with 2,525 trap days, we made 78 captures of black bears (Tables 1,2). Forty-eight individual black bears were ear-tagged, of which 47 were radio-collared. Also, a single yearling bear was released without tagging. No cubs were captured. Forty-five (92%) of the initial captures were made in the first 60 days of trapping effort. This is similar to the results from the uncompahgre Plateau, where 91% of captures were made in the first 59 days of trapping. In addition, 17 individuals were recaptured a total of 29 times. Only one recaptured.bear was immobilized in order to replace a missing ear tag~ All others were released without further handling. One subadult male was captured 6 times, while another subadu1t male was captured 4 times. These are the first bears ever captured more than 3 times during any of our Colorado bear studies. Table 1. Composition AGE/SEX of captured INITIAL CAPTURES black bears, Middle INDIVIDUALS RECAPTURED Park, 1995. TOTAL RECAPTURES· AD FEMALES 10 4 6 SA FEMALES 13 3 4 AD MALES 7 3 4 SA MALES 18 7 15 17 29 UNKNOWN TOTAL 1 49 Again this year, trap-related injuries to bears were low. Of 49 bears examined, 1 had a broken canine tooth, 1 had pulled off a claw sheath, cracked claw sheaths, and 1 had a small cut on a toe. Of the 29 bears released without handling, none appeared to have any visible injury. closely 2 had Capture rates were roughly half those as experienced on Uncompahgre Plateau, although the proportion of total capture through time was quite similar. Data from both areas suggest that at a trapping density of 1 trap/10.4 km2 the return on effort declines dramatically after 60 days; with most captures then being recaptures. Radio-tracking in Middle Park indicated that nearly all the collared bears were in the study area during the latter trapping periods. However, observations by hunters and preliminary photo resighting suggest a very high proportion of the bears on the site were marked. �270 2. Table Trapping PERIOD success NEW CAPTURES by time for black RECAPTURES bears, Middle TRAP DAYS Park, DAYS/ NEW CATCH 1995. CUM. CAPTURE 6/18-6/27 16 0 200 12.5 16 6/28-7/7 4 1 229 57.3 20 7/8-7/17 5 4 226 45.2 25 7/17-7/26 7 1 267 38.1 32 7/27-8/5 5 3 244 48.8 37 8/6-8/14 8 5 232 29.0 45 8/15-8/24 2 6 247 123.5 47 8/25-9/3 0 5 281 9I4-9/13 2 3 289 9/14-9/23 0 1 310 49 29 TOTAL Table Mesa, BLK MESA AVG WT RANGE AVG n = 14 SUBAD FEMALE n = 28 ADULT MALE n = 12 SUBAD MALE n = 35 WI (kg) between 51.5 3 Colorado RANGE AVG WT n = 11 46 20 23-84 n =13 117 16 96-152 n = 57 27 28-90 n = 24-77 n = 91-159 n = 34-114 n = 44 123 67 areas; Black RANGE 64 52-111 n = 15 49 MP 74 59-107 49 49 UNC 79 ADULT FEMALE 144.5 2,525 3. Comparison of black bear weight Uncompahgre Plateau, Middle Park. AGE/SEX 47 44-83 41 103 6 51 18 22-68 68-138 19-91 Both color phase proportions and body weight were different for Middle Park bears when compared to bears from the southwestern study areas (Table 3). Whereas black color phase comprised 10% of both Black Mesa and Uncompahgre study bears, they comprised 29% of Middle Park bears. Body weights of Middle Park bears of comparable age were roughly 80% of those from the southwest areas. �271 Two collared bears were killed during 1995. A subadult female, which frequented several guest ranches and summer homes, was killed after entering the kitchen at one ranch. A subadult.male was killed by a sheepherder near Meeker, CO on 8/30/95. This bear was located in the study area on 8/15/95, 103 km east of the kill location. Age and sex composition of capture in Middle Park was nearly identical to the Uncompahgre study (Table 4). When examining only females it is interesting .that 44% captured were judged adult in both studies. Since adulthood in females is based on having had a litter, rather than a set age, this is an unambiguous measure. Percent of adults among females has also been measured during the 1993, 1994, and 1995 hunting seasons. The percent adult among females was 40, 41, and 42 percent for those hunting seasons. This is significant when one analyzes the reproductive potential of a bear population. Preliminary simulation work based on black bear survival rates from the Black Mesa study would have.the proportion of females as adults over 50%. The Black Mesa area was nominally closed to legal hunting and as such the survival rates estimated are probably higher than exist statewide. The proportion adults from the 2 capture studies and 3 years of hunting suggest this to be the case. Either that, or all methods and times suffer the same consistent bias. If the samples truly represent reality, -then estimates of population growth will need to be adjusted in the models; primarily by reducing female survival rates. This will have a net effect of lowering the reproductive potential of Colorado black bears. Table 4. Age and sex composition of 2 trap samples of black bears, Colorado. % of Sample AREA MIDDLE ADULT MALE PARK UNCOMPAHGRE Monitoring seasonal ADULT FEMALE SUBADULT MALE SUBADULT FEMALE 15 21 38 27 20 20 34 26 movements The berry crop in 1995 was quite poor in the study area. Collared bears moved extensively during the september period but no concentrations near any berries were detected. Examination of bear scats. and actual observation strongly indicated that nearly all feeding during September was on mushrooms. There .was one area of concentrated use near Little Gravel Mtn. and again, there were no berries found but mushrooms were abundant. Increased efforts in 1996 will focus on September movements. Prior to this study, it had been suggested by the principal investigator that a proportion of the bears would migrate northwest to more productive berry production areas approximately 45 km away. However, no such movements were detected from the collared bears. By June 1996 all bears collared as sub-adult males were "missing" from the study area. This provides some support for the accuracy of age designation. �272 Another subadult female was translocated on June 6 over 60 km to the southwest because of chronic nuLaance activity. She had returned to the study area and her habitual range by June 13. LITERATURE CITED Beck, T.D.I. 1991. Black bears of west-central Colorado. wildlife Tech. Publ. 39. 86 p. Colo. 1994. Development of black bear inventory techniques. Wildlife Fed. Aid Prog. Rep. W-1s3-R-7, WP SA, J2. 8 p. Div. Colo. Div. 1995. Development of black bear inventory techniques. Colo. Div. Wildlife Fed. Aid Prog. Rep. W-1s3-R-8, WP SA, J2. 11 p. �273 APPENDIX . PROTEIN USE, MUSCLE FIBER TRANSFORMATION, AND CITRATE SYNTHASE ACTIVITY IN FREE-RANGING BLACK BEARS DURING HffiERNA TION by Daniel B. Tinker A thesis submitted to the Department of ZOOlogy and Physiology and The Graduate School of The University of Wyoming in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE 10 ZOOLOGY AND PHYSIOLOGY Laramie, Wyoming December, 1995 . �274 ACKNOWLEDGEMENTS This research was made possible by funding from the Colorado Division of Wildlife, as a part of their project No. W-153-12, and grant monies from the National Aeronautic and Space Administration and University of Wyoming Planetary and Space Science Center, Nasa Grant # NGT -40050. The CDOW also provided invaluable aid by providing our crews with trucks, snowmobiles, ATV vehicles, and other bearhandling gear. lowe my thanks to the United States Forest Service for access to the Uncompahgre National Forest, and for the use of the Cold Springs Work Station for the duration of our research. In addition, the University of Wyoming, Department of Zoology and Physiology provided both fmancial assistance and . logistical assistance in the form of graduate teaching assistantships and the use of departmental vehicles used in the field work. I would also like to thank the L. Floyd Clark Scholarship Fund for summer support used forthe completion of lab and data analyses. For their guidance and support during the entire process 1would like to thank my graduate committee, Drs. Fred Lindzey, Jason Lillegraven, William Romme, and particularly I would like to thank my major advisor and committee chairman, Dr. Hank Harlow, for his daily instruction, encouragement, and support, both in the field, in the classroom, and in the laboratory. I also want to thank Mr. Tom Beck, c0principal investigator on this research, for logistical support and instruction in bear tracking and handling, and more importantly, for allowing me access to bears previously collared by him and his crews. For their help in the field, I want to thank my primary crew, Lyle Willmarth and Shawn Lechman, without whom the field work would have been impossible. In addition, I would like to thank the many volunteers for their. time and effort, often during cold, adverse conditions: Ron Hayes, Zach Harlow, Ron Grogan, Mark Conrad, George Montopoli, Nick Visser, Bill Romme, and Sally Tinker. TABLE OF CONTENTS Section Introduction Materials and Methods Results '......................................................... Discussion ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Literature Cited '. . . . . . . . . . . . . . . . . ... . . . . . . . . . . . .. Appendix A - Protein Assay by Dye Binding . . . . . . . .. .. . . . .. .. .. .. .. .. . . . .. . . . . . . . . . . . . . . . .. Appendix B - Citrate Synthase Assay , . . . . .. . . . . .. . . . .. . . . Appendix C - Muscle Tissue Staining. . . . . . . . . . . . . . . . . . .. . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Appendix D - Tables: Individual Bear Data , 275 279 285 291 296 299 301 303 305 �275 INTRODUCTION Contrasting opinions regarding winter dormancy in the American black bear (Ursus americanus) have abounded throughout the years. The use of the word "hibernator" to describe bears in general has been a topic of scientific debate for decades. When compared to small mammal hibernators, bears have a uniquely different strategy for winter survival. Specific differences will be explored in this research project, along with some ideas about the selective pressures which may have been instrumental informing bears' extraordinary adaptations. However, before investigating the physiological and behavioral specifics of bear hibernation, some general background information on winter sleep in beats is essential. NATURAL HISTORY OF WINTERING BLACK BEARS For the purposes of this report, time frames relevant to denning behavior and physiology will be that of bears found in the Rocky Mountain region, unless otherwise noted. Unlike hibernators that experience rhythmic alterations of deep torpor and arousal, bears remain in a relatively shallow torpor for five to seven months. Remarkably, the bears do not eat, drink, urinate, or defecate during the entire dormancy period (Folk 1974; Nelson 1973). During this time, bears experience no regular periods of arousal, and maintain nearnormal body temperatures of31-35 °C "(Nelson et aZ. 1983). In late summer and early fall, black bears enter hyperphagia, a period of prodigious food intake, often increasing their daily caloric intake from an average of approximately 33,000 Kl/day to as high as 84,000 Kl/day (Nelson et al. 1983). Nelson (1980) observed bears in the wild feeding 20 hours per day during late fall in anticipation of hibernation, and our personal observations indicate that bears will continue to forage long past the time when they appear to be canying a surplus off at for hibernation. During this period of hyperphagia, bears generally begin to locate and prepare winter dens for occupancy. . Apparently, black bears utilize many different types of natural shelters, often denning in rock caverns, large depressions beneath uprooted trees, or on the surface of the ground under dense shrubs (Beck 1991). In Great Smoky Mountains National Park, Pelton et al. (1977) noticed that all the black bear dens they observed were in some way associated with large mature hardwood trees and many of the den entrances were located as high as 17 meters above the ground. Unlike grizzly bears (Ursus arctos horrtbtlisy; black bears usually do not perform a great deal of modification to their chosen den sites, rarely excavating a den completely (McNamee 1990). Multiple dens are often prepared and, if disturbed during occupancy of one den, black bears will often relocate to another previously prepared den, even in mid-winter (Beck 1991). Contrary to anecdotal beliefs, protection from possible predation, rather than shelter from adverse climatic conditions, seems to constitute the primary consideration for den selection, most bears being well able to supply their own insulative needs (Beck 1991). Black bears often enter their dens for very short periods of time prior to fmal den entry; however, the specific mechanisms that trigger final entry remain somewhat unclear. Beck (1991) suggests that, because of the consistency of den entry and exit between years, the bears probably respond to various indogenous physiological changes rather than to external climatic conditions. The actual time of den entry varies considerably within the United States. Black bears in the Southeast often delay entry until mid-December, while the optimum period in the U.S. Rocky Mountain region seems to span from mid-October to midNovember (Lindzey and Meslow 1976, Tietje and Ruff 1980, Beecham et al. 1983, Schwartz et al. 1987). Den occupancy ranges from three to seven months, based upon regional variation in weather conditions and food availability (Nelson et al. 1983). Duffy (1971) illustrated the variation in bear denning periods and the apparent genetic propensity for specific lengths of denning. He established that native bears of Louisiana remained active throughout the entire winter, while Minnesota bears transplanted to Louisiana regularly denned for extended periods of time. If left Undisturbed, bears will normally remain in a state of shallow torpor for the duration of the denning period and will rarely leave their dens for any reason (Hamilton and Marchinton 1977). Female black bears give birth to cubs while occupying their dens, usually in January, and will nurse their young both during hibernation and following emergence in the spring (Nelson et al. 1983, Nelson 1973). �276 In most of the Rocky Mountain region, bears emerge from their dens any time from mid-March to mid-April, although some bears in the interior of Alaska do not emerge until late-April to mid-May (Beck 1991, Schwartz et al. 1987). Post-denning lethargy is evident in black bears upon emergence from their winter dormancy, and most are anorectic for several days following emergence (Nelson et al. 1983, Nelson 1980, Hock 1958). Bears often exhibit hypophagia for a period of 1-2 weeks after leaving the den, even if ample food is available (Nelson et al. 1983). PHYSIOLOGICAL AND METABOLIC ADAPTATIONS TO HIBERNATION Basic similarities exist between bears and deep hibernators (e.g., distinct decreases in metabolic rate, heart rate and body temperature), but bears are truly unique in their physiological adaptations to winter dormancy. While deep hibernators (rodents, insectivores, bats) lower their body temperatures to near ambient conditions, often near 0° C, and arouse periodically to eat and/or drink, bears only lower their body temperature a few degrees C, and, if undisturbed, do not arouse for the duration of the denning period (Nelson et al. 1983, Folk 1974). The metabolic and physiological changes that bears undergo have been the subject of study for decades, so the general features of these mechanisms are, for the most part, fairly well understood. However, certain aspects of physiological adaptations to hibernation remain unclear, and two are the focus of this study. Protein and Fat Utilization During Hibernation On the surface, bear hibernation appears to be a simple biological process but, in actuality, it is a marvelously complex event involving literally every organ system of the body. Bears are relatively large mammals, and while hibernating utilize some 17,000 KJ per day, yet do not eat, drink, urinate, or defecate for the duration of winter dormancy (Nelson 1973; Nelson et al. 1983). This required energy is believed to be provided almost exclusively by utilization of fat reserves, with little or no net muscle protein loss (Nelson et al. 1973, 1975; Lundberg et al. 1976). A problem commonly experienced by fasting animals that rely primarily on fat as a metabolic substrate is ketosis (the accumulatin of by-products of fat catabolism). Krilowicz (1985) found that hibernation in Belding's ground squirrels was accompanied by ketosis,which assists the animal in conserving glucose, leading to conservation of muscle protein. She further suggested that unlike hibernators.ketosis in fed or fasting animals does not prevent the uptake of glucose, therefore protein conservation is not affected. Hibernating bears seem to circumvent the problem of ketosis altogether. Palumbo et al. (1983) suggested that an increase in triglyceride turnover in hibernating black bears presumably prevents the development of ketosis. Nelson (1980) also found no evidence of ketosis in hibernating bears. Small mammal hibernators, such as ground squirrels and other rodents, typically experience some degree of muscle atrophy during hibernation (Wickler and Hoyt 1990; Loughna et al. 1986), which is characterized by loss of lean muscle tissue. This loss is the result of protein breakdown in the muscles. While it is claimed that black bears do not utilize protein as a source of metabolic energy during hibernation (Nelson et al. 1973, 1983; LUndberg et al. 1976; Alquist et al. .1976; Nelson 1980), conflicting ideas regarding protein breakdown and synthesis in bears as well as other hibernators have been offered. It has been suggested that the common end products of protein catabolism are not accumulated during black bear hibernation, indicating an absence of protein breakdown. For example, total serum protein, urea, uric acid, total amino acids and ammonia do not increase during hibernation (Nelson et al. 1983, Nelson et al. 1973). Ahlquist et al. (1976), and Lundberg et al. (1976) obtained similar results using captive black bears and believe that this absence of catabolic end products is due in part to an increase in the effectiveness of protein anabolism, wherein rather than entering the urea cycle, amino acids from protein breakdown reenter protein anabolic pathways. Bears possess the ability to passively move urea, a nitrogenous end product of protein catabolism, into the lower intestinal tract where it is hydrolyzed into ammonia. This ammonia is then moved into the blood and eventually the liver, where reamination to nonessential amino acids occurs. Later, Nelson (1980) also found no measurable loss of lean body mass, perhaps due to the entry of free amino acids into protein anabolic pathways. �The low ratio of serum urea-creatinine, and the reduction in gluconeogenesis (glucose production from protein catabolism) as reported by Nelson et al. (1984), also indicates a vel)' low rate of protein catabolism by hibernating bears. Harlow et al. (1990) suggested that blood serum cortisol, which can stimulate gluconeogenesis, but can also enhance fat mobilization, was significantly higher in bears during hibernation than in the summer months. However, a concomitant reduction in plasma urea (Nelson et al. 1984) indicated metabolic utilization of fat reserves rather than protein breakdown. Even though these studies suggest no net protein loss in overwintering bears, protein turnover, i.e., the cyclic degradation and synthesis of protein, may still be high. For example, Lundberg et al. (1976) found that winter protein turnover increased three- to fivefold over summer levels without a measurable net protein loss. Even though it is generally thought that there is a marked reduction in protein catabolism during hibernation (Bintz et al. 1979; Yacoe 1983; Krilowicz 1985), it is becoming more evident that a minimum amount of protein breakdown is required for all fasting or overwintering animals (Cahill 1976; Yacoe 1983; Le Maro et al. 1981). A basal level of protein catabolism may be needed to: 1) sustain a continuous utilization of fat by providing short-chained carbon intermediates such as pyruvate to the krebs cycle (Bintz et al. 1979); and 2) be a source of water during periods oflimited food and water intake (Bintz et al. 1979). Some believe that the resulting metabolic water produced by fat catabolism replaces most of the respiratory water loss (Nelson et 01. 1983). Bintz et al. (1979) challenged this paradigm and suggested that because fat is such a high energy fuel and contains such a small amount of bound water, it actually can result in a negative water balance if it were the only substrate metabolized. For a fixed metabolic demand, protein provides over four times more net gain of water (metabolic + bound water) than fat, which is released when proteins are catabolized (Riedesel and Steffen 1980). Under normal circumstances, the release of this water is potentially negated by its loss in urine to detoxify urea .. However, bears recycle urea during winter hibernation (Nelson et a11973; 1975). Urea produced by the urea cycle in the liver is first excreted into the bladder. It is then moved across the bladder wall into the blood and reabsorbed by the lower intestinal tract where it is hydrolyzed into ammonia. This ammonia is eventually reaminated to nonessential amino acids once it has been transported to the liver (Nelson 1980; Guppy 1986). A decrease in fat mass for energy and protein mass for water would, therefore, be anticipated during the winter, especially during periods of lactation. The problem of maintaining a metabolic water balance is exacerbated during gestation and lactation due to the extra requirements of both energy and water needed for fetus and inilk production. Nursing cubs place an additional demand for muscle protein breakdown by the lactating female. These cubs undergo a rapid growth process requiring a large amount of milk containing both water and protein. Black bear milk contains approximately 55% water and 11% protein (Jenness et al. 1972; Cook et al. 1970; Baker et al. 1963). The need for this water and nitrogen for cub growth further taxes protein reserves (skeletal muscle and non-muscle) of the lactating female. These physiological challenges would seem to suggest that protein use during hibernation, especially by lactating females, is inevitable. HYPOTHESIS 1: Overwintering black bears, particularly lactating females, utilize protein tissue in addition to fat reserves as a source of metabolic energy and water, and therefore exhibit a net protein loss over winter. Muscle Disuse Atrophy and Fiber Type Transformation Another phenomenon typically associated with protein degradation in hibernators involves the atrophy of skeletal muscles. One of the most consistent observations made by bear biologists in the field is the lack of impaired locomotor ability and muscle tone in bears aroused during hibernation. It remains unclear how bears remain inactive, within a confmed space, for 4-6 months without any apparent impairment of muscle tone. Even after several months of inactivity, hibernating bears in captivity do not appear to experience any significant muscle protein loss (Koebel et al. 1991). Many muscle disuse models have been . described primarily using either rodent hibernators or some type of limb suspensionldenervation, but none yet address the specific mechanisms by which muscle fiber integrity could be maintained by black bears that remain inactive for extended periods of time. �278 Peter, et al. (1972) described a classification system for skeletal muscle fibers based on i) contraction time of the fiber; ii) oxidative capacity; and iii) glycolytic capacity. Since that time, numerous studies have been done to determine the relative effects of muscle disuse atrophy on each fiber type. Type I, slow-twitch muscle fibers are typically small-diameter cells and are characterized by slow-acting myosin ATPases, abundant myoglobin which stores oxygen and increases the aerobic capacity of the fibers, and high mitochondrial density. Glycogen concentration is low and they are extremely fatigue-resistant. Type II, fasttwitch muscle fibers, on the other hand, are usually larger in diameter than slow-twitch cells and contain fastacting myosin ATPases, little myoglobin and few mitochondria. They do, however, maintain large glycogen reserves and depend on the anaerobic lactic acid pathway to generate ATP, hence they fatigue quicldy (Marieb 1992). Musacchia, et al. (1983) suggested that it was logical that slow-twitch (Type I) oxidative fibers should be affected more by disuse than fast-twitch (Type II) glycolytic fibers due to the fact that slowtwitch fibers receive continual neural input in order to maintain tonic weight-bearing functions, whereas fasttwitch fibers are recruited only during more prolonged or intense activity. However, as described below, other studies have yet to reveal any discernable pattern or targeting of a particular fiber type with regard to muscle atrophy. . There have been two traditional models for studying muscle disuse atrophy in mammals: hind-limb suspension and muscle denervation. These conditions of disuse cause muscle atrophy due to a decrease in muscle fiber protein, cell size and number, and altered ratio of muscle fiber types, resulting in an impaired muscle function. Denervation and hind-limb suspension typically result in fewer aerobic, Type I muscle fibers and a greater proportion of Type II, anaerobic muscle fibers (Booth and Kelso 1973). . These models appear to be inconsistent, however, as evidenced by the broad range of fmdings. Nicks, et al.(l989) immobilized the hindlimbs of rats and found that relative percentages of fiber type did not change, but that cross-sectional area of the fibers decreased by 42%. Other similar studies reported a decrease in Type I slow-twitch fibers of from 20 - 40% following some form of immobilization (Templeton et al. 1984; Thomason and Booth 1990). Specifically, these models have limitations in that I) they are "unnatural" conditions, and 2) they do not allow for repeated sampling of the same individuals over time. Recent studies on hibernating small mammals have partially circumvented the first concern and have identified some unique properties in this "natural" muscle disuse model. For example, hibernation immobilization does not cause the same magnitude of protein loss or fiber type alteration as artificially restrained animals (Wiclder et al. 1991). Additionally, citrate synthase activity appears to be elevated in the muscle of hibernating ground squirrels, which may .partially compensate for the loss in percentage of Type I fibers and thereby help to retain a relatively high aerobic muscle capacity throughout the winter. However, there are two major limitations to this hibernation muscle disuse model as well. First, because of their small size, the same individual cannot be repeatedly sampled and different groups of animals must be killed and compared during stages of hibernation. Second, small hibernators arouse with violent muscle shivering approximately every 15-20 days, which may provide sufficient exercise to maintain muscle fiber type ratios and normal citrate-synthase activity levels. The black bear allows us to circumvent these problems: It is large enough to resample during the winter, and it does not undetgo periodic arousal with violent shivering. Despite the obvious advantages of using hibernating bears as natural disuse models, relatively little work has been done in this area. Koebel et al (1991) found that concentrations of glycogen, triglycerides, and other principal energy-supplying substrates of muscles were unchanged in black bears during hibernation, and suggested that this might contribute to the bear's ability to exhibit good muscle tone upon spring emergence. However, this study was conducted on 3 bears held in captivity, and does not necessarily reflect the dietary, thermal, and activity profiles of naturally foraging and denning bears. The same study revealed that citrate-synthase (CS) activity, an indicator of muscle oxidative capacity, was also unchanged during the denning period. This is in contrast to nuinerous reports of a significant increase in CS during hibernation in smaller rodent hibernators (Wickler 1981; Wiclder et al. 1987; Wiclder and Hoyt 1990; Yacoe 1983; Thomason and Booth 1990; Van Breukelen et al. 1990; Steffen et al. 1991). One possible explanation for this increase is that an elevation in CS could facilitate shivering thermogenesis during episodic arousal (Wickler and Hoyt 1990). If this is true, and since bears do not .undergo these periods of arousal, this increase in CS activity would not be expected in hibernating bears. �279 HYPOTHESIS 2: Overwintering black bears in a natural denning situation do not exhibit a dramatic loss of Type I aerobic muscle fibers, nor do they experience a significant decrease in cross-sectional area of these fibers. HYPOTHESIS 3: Citrate-synthase activity, a mitochondrial enzyme that is an indicator of muscle oxidative capacity, does not increase during hibernation, although it does increase in smaller hibernators. The specific questions that will be addressed in this study are: i) What is the amount of fatty tissue used by lactating and non-lactating females during the hibernation period"; ii) How much (if any) protein is catabolized from specific muscle areas by lactating and non-lactating females"; iii) Are muscle fiber type and cross-sectional area preserved during hibernation, resulting in maintenance of muscle tone for spring emergence"; and (iv) Does citrate-synthase activity remain relatively consistent throughout the denning period? By comparing results obtained from early-denning samples to those of late-denning samples, I was able to describe for each individual bear sampled: 1) the total amount of body fat utilized during the denning period; 2) the amount of protein catabolized and which of the muscles sampled were the major source of the protein; 3) the relative numbers of Slow and fast twitch muscle fiber types to see if they remain constant during the denning period, and if there is a decrease in cross-sectional area.of these fibers; and 4) seasonal trends in citrate-synthase activity from the different muscle types. METHODS Study AreaThe study area is in west-central Colorado and is part of an ongoing study by the Colorado Division ofWildllfe (No. W-153-12) wherein principal investigator Tom Beck is studying new inventory techniques for black bears. The area is in Mesa County, and encompasses much of the northwestern end of the Uncompahgre Plateau, south of Grand Junction and northwest of Delta, Colorado (Figure 1). The plateau is cut by numerous steep canyons and drainages. The elevation generally ranges from 2400 - 2800 meters, though one of the study bears used a den that was located at slightly over 1800 meters, in pinyon-juniper forest (Pinus edulis - Juniperus osteosperma). The primary canopy cover in most of the study area consists of ponderosa pine (Pinus ponderosa), quaking aspen (Populus tremuloides), and gambel oak (Quercus gambelii), while other canopy and understory species such as blue spruce (Picea pungens), serviceberry (Amelanchier alnifolia), and sagebrush (Artemesia sp.) are somewhat less abundant. The majority of the study area is in game management units 61 and 62, and is under the jurisdiction of the U.S. Forest Service. Study Population - . 106 bears were captured within the study area by Colorado �280 STUDY AREA to _(I" '" O( lII .••U Figure 1 - Map of the Study Area - Uncompahgre Plateau in west-central Colorado. The study area was located at the northwestern-most end of the Uncompahgre Plateau, near the Utah border. Access.to the study area was west of Whitewater, Colorado, on State Highway 141, to U.S.F.S. Divide Road. �281 Division of Wildlife researcher Tom Beck's research crews during the summer of 1993. Of these, 89 bears were ear-tagged, and all bears over 23 Kg were fitted with radio tracking collars transmitting in the 150-152 mHz band. These collars are designed to disintegrate and falloff within approximately 24 months if not removed prior to that time by the bear or by researchers. Field Methods Preliminary field work - A trip to Washington State University was made on December 3-5, 1993, to work with Dr. Charlie Robbins at his captive bear facility. The purpose of the trip was to become familiar with the operation and use of a biological impedance analysis (BIA) instrument, to be used to determine total body fat content of bears in our study. An initial 10 days were spent in our study area, from December 28, 1993 through January 7, 1994 working with Tom Beck. This trip was undertaken to field test equipment and techniques, including radio-tracking and den location, field surgery, as well as field logistics in general. Field Strategy and Radio-tracking - . All field work was based out of the Cold Springs Ranger Station, in the Uncompahgre National Forest. The field work was divided into two main efforts: the late fall, or early denning period (September-December 1994) and late winter/early spring, or late denning period (March-April 1995). It was our initial intention to obtain data from a minimum of 14 bears, 7 female, nonlactating bears and 7 female, lactating bears. The decision as to which females.were most likely to be bred, and therefore lactating, was based on visual observations from the summer and early fall of 1994, as well as capture records from the summer of 1993. If a female bear was observed with cubs in the fall, she was automatically assumed to be non-lactating for the upcoming denning season. If a female was observed travelling with a boar during breeding season, and was (based on capture records) reproductively mature, she was considered a likely candidate to be lactating during the upcoming denning season. Both aerial and ground radio-tracking began in mid-September (aerial reconnaissance was provided by the Colorado Division of Wildlife and the Colorado State Patrol). In both instances, dens and bears were located using the Telonics/Y agi radio tracking receiver and antenna. Prior to heavy snowfall, 4-wheel drive trucks and four-wheel ATVs were used to get as close as possible to each bear and bear den. Once snow depth prohibited the use of trucks, snowmobiles were used exclusively to transport both personnel and supplies. Once a den was identified and it was determined that a target bear was indeed inside, the bear was anesthetized using ketamine hydrochloridelxylaxine hydrochloride (200 mg ketamine and 50 mg xylaxinel ml) in a jab-stick at a dosage of 4.0 mg ketaminelKg bear weight (the maximum capacity of anesthetic for the jab-sticks was 5.0 cc of the cocktail, and this was. the dosage that was normally administered). If the bear did not respond to the initial dose of anesthetic, a second, full dose was given since the effects of ketaminelxylaxine are not additive until the bear has succumbed to the drug. After the bear was immobilized, which usually took about 15 minutes from the time of injection, she was removed from the den and placed on an inflated Therma-rest pad and heavy plastic tarp which provided insulation from the snow and cold. It was discovered early in the study that it was imperative that animals be kept completely dry during all phases of the handling, but specifically during the BIA measurements. All bears were extracted from the den by placing them on a heavy plastic tarp while still in the den, and dragging the tarp out. If it was snowing or there was blowing snow, a shelter was constructed over the bear, thereby effectively keeping it dry. Measurements/Surgical Procedures - . All physical measurements and surgical procedures were performed on nine female bears during the fall field season and on seven of the nine bears during the spring field season (Bears #6 and #7, which were sampled during the fall season, had already left the den sometime during mid- to late-March and could not be . resampled in the spring). Physical measurements and surgical procedures were identical during each season, i.e., the surgical biopsies were all removed from the left hind limb during the fall sampling and from the right hind limb during the spring sampling. Six of the seven bears sampled in the spring had cubs and were . therefore considered lactating. Bioelectrical Impedance Analysis (BIA) - Total body fat of the animal was estimated using a BIA 101-Q (RJL Systems) impedance plethysomograph. This instrument was used to determine snout-tail resistance (in ohms) for the bear, and these data used to calculate total body water and total body fat of the �282 animal (Farley and Robbins, 1993). The instrument consists of a small hand-held digital impedance meter and two pairs of dual electrical leads, each lead equipped with small alligator-type clips at the end. While various electrical lead configurations are possible, we used the snout/tail resistance path, as recommended during our visit to Dr. Robbins' facility. This method is the best suited to field conditions due to its ease of lead placement with respect to the body position of the bear, as well as ease of repeatability. One pair of leads was attached to the snout, i.e., one leadto either side of the snout on the inner surface of the upper lip, just opposite the upper canine tooth. The other pair ofleads was attached on either side of the animal's tail by first inserting I", 20-gauge vacutainer needles into the fat deposits on either side of the tail and attaching the alligator clips to the protruding needle. Resistance and reactance were both measured, and then immediately remeasured to insure precision. However, while these procedures are seemingly straightforward, many variables affect results of this measurement: the bear must be completely dry and clean; the bear must not have any open wounds or sores along the electrical pathway between the electrodes; the position of the torso and limbs of the bear must be absolutely consistent between seasonal measurements; and very accurate estimates of body mass (to within 100 g) must be obtained for each animal. These variables were, in fact, well controlled in the field. However, open wounds around the neck were often encountered as a result of rubs from radio collars and may have resulted in suspect calculated values for total body fat (see Discussion). Physical measurements - Each bear was weighed on a digital load scale (Dyna Link, Model MSI7200), accurate to within ± 0.1 kg. This accuracy is necessary for dependability of body fat calculations based upon BlA measurements. If a tree was located very near the den, the bear was suspended from either the trunk of the tree or a large limb and hoisted off the ground using a pulley system. If no trees were available, a metal tripod was carried into the field from which the bear could be suspended for weighing. Snout/vent length was measured to the nearest centimeter. A metal retractable carpenter's tape was used; the measurement was made from the tip of the nose along the contours of the head andtorso to the base of the tail. Surgical Procedures - Two small (::::500mg) muscle tissue samples were surgically removed from the hind-limb of the bear (left-hind in the fall and right-hind in the spring), one from the lateral head of the gastrocnemius and one from the biceps femoris. Preliminary studies on the location, contour and accessibility of each muscle were conducted by dissecting legs from bear carcasses collected by the Colorado Division of . Wildlife. In order to be confident that each biopsy was removed from the same region of each muscle.careful measurements were made on each bear prior to making the surgical incision, as follows: a) gastrocnemius the distance from the calcaneus bone to the bend behind the knee was measured and 80% of that distance was calculated in centimeters. This distance was then measured up from the calcaneus to a point between. the two heads of the gastrocnemius. The distance from this point laterally to the medial edge of the fibula was measured and the incision was made at the midpoint of that line, obliquely pointing towards the patella, along the "grain" of the fibers; b) biceps femoris - using a piece of elastic rubber tubing, a contour line was laid out from the anal vent to the patella, along the contour of the upper leg and hip; this distance was measured and the incision was made at the midpoint of this line, obliquely towards the patella, again along the "grain" of the muscle fibers .. Once the exact point of incision was correctly identified, an area of approximately 25 cnf surrounding the planned incision was shaved, first by removing the long guard hairs with scissors, and then shaved closely with a disposable, double-edged razor, using shaving cream to soften the dense hairs. This surgical area was cleaned with soap and a Betadine iodine solution. The long hair surrounding the shaved area was taped down with duct tape and covered with a fenestrated, sterile drape to prevent contamination of the incision. At this point in the surgical procedure, one of the researchers donned surgical gloves, and maintained a sterile environment throughout the biopsy removal. All surgical instruments, including suture, were maintained in a liquid sterilant, and a sterile surgical field was preserved during the procedure. A 3 cm-Iong incision was made, and a rectangular tissue sample approximately l.5 em long, 0.5 cm wide and 0.5 em deep was removed and immediately given to the surgical assistant. The muscle tissue was then pulled together snugly with dissolvable OO-gutsuture and the skin was closed using non-dissolvable 0- �283 silk suture. A small drop of superglue was applied to each knot of the external, interrupted mattress stitches to ensure the integrity of the closure. A topical antibiotic (Furazine) was applied to the suture line. Each biopsy sample was placed onto a glass surface and carefully cut into three equal parts by the surgical assistant. One of the three pieces of tissue was immediately placed into a small, labelled ziplock bag and quickly put into liquid nitrogen, which was carried into the field in vented thermos bottles. This first piece of tissue was later used for assays of enzymatic activity. The second portion of tissue was mounted onto a small round piece of cork using Cryoform tissue freezing matrix, with the muscle fibers oriented perpendicular to the surface of the cork. This tissue/cork was placed into liquid isopentane cooled by liquid nitrogen and, after being frozen, was put into a labelled ziplock bag and into the liquid nitrogen thermos. This mounted tissue was later used for fiber type and histological analysis. The fmal piece of muscle tissue was then put into a labelled ziplock bag and placed into the liquid nitrogen for later analysis of protein content. All samples were transported frozen to the field station where they were transferred to a large 35liter liquid nitrogen dewar for transport to the laboratory at the University of Wyoming. There was always an ample volume ofliquid nitrogen for freezing and transport of tissue to cover each field test season. Following the removal of the two tissue biopsies, 15-20 cc of blood were removed from the femoral vein with a 20gauge needle, and stored in styrofoam carriers on ice for transport to the field station. There, the blood was spun down in a IEC, Model CL clinical centrifuge and the plasma frozen at -4°C until it was taken to the laboratory at the University of Wyoming. The bear was then replaced in the den by sliding it on the plastic tarp to as close to its original position as possible. The den entrance was then covered with vegetation (and snow, if snow was present upon our arrival at the den). This entire procedure was repeated on seven of the nine bears during the late denning season (MarchApril). At that time, we reweighed each bear, repeated the BIA measurements and surgically removed two additional muscle tissue samples from the opposite hind-limb (right hind-limb) that was sampled in the fall (left-hind limb). As during the fall sampling, all bears were replaced in their dens as close to the original position as possible before leaving the den site. Particular effort was made with regard to females with newborn cubs, to try to replace both mother and cubs back in the den exactly as we found them in order to reduce the chance of injury to the cubs. Laboratory Methods A period of laboratory analysis followed each of the field seasons in the Biological Sciences Building at the University of Wyoming. Upon arrival at the laboratory from the field sessions, the frozen tissue samples were taken from the nitrogen dewar and immediately stored in a Forma" ultra-cold Bio-freezer at 70°C. Each of the three tissue sections from each muscle was previously labelled either "fiber", "protein", or "enzyme", and was analyzed in those groups as follows: Fiber type and number analysis - Frozen sections were previously mounted on cork discs in Cryoform tissue freezing matrix as described above. Thin sections (5-6 microns) were serially cut using an American Optical Histostat freezing microtome knife at -20°C and mounted on glass slides. Cutting continued until 5 high-quality slides for each bear muscle were obtained. These sections were air-dried overnight and then stained for myofibrillar ATPase activity and NADH activity (see Appendix for complete protocol). Since the slides were cut serially, and were only 6 microns thick, each of the 5 slides was virtually identical to the other in each group. The "best" slide, based on the quality and evenness of the staining, was selected from each group of five for analysis. Using a Zeiss Universal compound microscope, a mti® videoscope, a Caliber 486 dx-66 desktop computer, and morphometry software known as Flexible Image Processing System (FIPS), the entire image of each of the selected slides was stored on a 3.5" floppy disc for analyses by FIPS. In addition, each complete image was visually divided into 4 sub-images, and enlarged at 6.3x magnification; these sub-images were also stored on a 3.5" floppy disc for analysis. Each of the four enlarged sub-images was projected onto the computer monitor individually, and all of the individual fibers on each screen were measured and analyzed, for an average total of 350-400 fibers per muscle sample. The diameter and cross-sectional area of each fiber was measured. All fibers from each sub-image were classified either as slow oxidative (SO) or fast glycolytic (FG) according to the classification method of Peter et al. (1972), i.e., SO fibers stained a light gray color and �284 FG fibers stained a very dark gray. When all fibers were analyzed, fiber-type ratios (percentages of each .fiber type) were calculated for each muscle and mean cross-sectional area and fiber diameter were calculated for each muscle using the FIPS morphometry software. Additionally, total nwnber of fibers for each muscle sample was determined by adding the nwnber of fibers analyzed from each of the four sub-images. Each subimage consisted of approximately 975 J.lm2,and the total area of all four sub-images was approximately 3900 J.lm2,or 3.9 mm". Protein concentration and water content - Protein concentration of each muscle was determined by two separate assays: i) modified Bradford (1976; Kley and Hale, 1977; Pierce and Suelter, 1977) colorimetric assay (see Appendix for complete protocol); and ii) carbon-hydrogen-nitrogen (CHN) ignition method. The CHN analysis was performed by Dr. Steve Boese, Department of Geology, University of Wyoming, using a Carlo Erba Model 4500 CIN analyzer. Tissue preparation - Protein concentrations for both assays were determined for dry weight samples which were prepared by freeze-drying the samples in a VirTis 10-145MR-BA freeze-dryer for 24 hours. Prior to freeze-drying, each tissue sample was weighed to the nearest 0.001 mg using a Cahn C-31 microbalance. Samples were weighed again following freeze-drying, and percent water was calculated. For the Bradford assay, the dried samples were again weighed to the nearest 0.00 I mg and homogenized in an Omni 5000 electric tissue grinder in a 1:40 weightlvolwne (milligram:microliter) of O.IM Phosphate buffer, pH 7.4. The homogenate was diluted 1:19 with protein dilution buffer, e.g., 20 J.lIof homogenate: 380 J.lIof dilution buffer, arid 100 J.daliquots of this mixture were combined with 1 ml of assay buffer for spectrophotometric analysis at 595 nm. For the CHN analysis, a portion of each dried muscle was weighed and ignited for analysis, and the results were reported in weight-percent of nitrogen for each sample. Total nitrogen (mg) was calculated based on the dry weight of the sample and this value was multiplied by a factor of 6.25 to calculate total protein (mg) in the sample (Runde and Hilditch 1974), and expressed as mg protein per dry weight of muscle tissue. Citrate-synthase activity -The third section of the tissue sample was used to measure maximal enzyme activity of citrate synthase, which is representative of the oxidative potential of the muscle. Tissue preparation - Citrate synthase activity was determined from wet-weight samples weighing approximately 20-40 mg. The wet tissue samples were homogenized in the Omni tissue homogenizer in a l g: 19 nil weight.volume ratio of tissue homogenizing mediwn as described by Shepherd and Garland (1969) (see Appendix for complete assay protocol). Homogenates were subjected to four freeze-thaw cycles by immersing the culture tubes containing the homogenates into liquid nitrogen until frozen. The culture tubes were then placed into a warm water bath (30°C) until thawed. Following the fourth freeze-thaw, the homogenates were centrifuged at 2700 rpm for 10 minutes at 4°C by placing the centrifuge inside a refrigerator. 50 J.lIof the supernatant was pipetted into a new culture tube to which 1 nil ofTris buffer (PH 8.0)was added; this mixture was vortexed briefly and placed on ice until analysis. The cuvette chamber in the spectrophotometer was maintained at 30°C during the procedure by a YSI Model 5214 water bath and pwnp system. Duplicatesamples and a blank were read at 412 nm. Statistical Analyses - A paired t-test with repeated measures design was used to detect significant differences in mean values of the measured parameter. A standard t-test was used for non-repeatable measurements, eg., muscle fiber cross-sectional areas. One-tailed tests were performed when changes were expected in a specific direction, otherwise two-tailed tests were used. Percentage data were normalized before analysis using the arcsin - square root transformation. All alpha-levels are at 0.05 unless otherwise noted. �285 RESULTS Values for each parameter measured are listed as "fall" and "spring" and "change". Fall measurements were made very soon after the bears entered their dens; spring measurements were made shortly before the bears left their dens. Change refers to the difference between these two measurements during hibernation. Tables containing measurements for individual animals for each parameter tested are found in Appendix D. Body mass/fat content (BIA) The mean percent body mass loss for each bear during hibernation was 24.3% ± 6.1%, and ranged . from 15% to 33% (Figure 2). These figures are consistent with expected body mass loss for bears during hibernation. Although body fat measurements suggested an increase in total body fat during hibernation (Figure 3), the difference between fall and spring values was not significant (p. > 0.05). Additionally, I believe that these data are erroneous due to inconsistencies with BIA instrument performance and the condition of some bears during both fall and spring measurements (see Discussion). CHANGES IN BQDY WEIGHT 180 • FALL WEIGHT 160 ......-.. SPRING WEIGHT 140 (9 ~. .•.....•.•. lI (9 120 I 100 W S 80 I 60 o 1 2 3 4 5 6 7 8 BEAR # Figure 2 - Changes in body mass for each bear during hibernation. Fall weights were obtained during the fall .sampling in November and December (open circles) and spring weights during the spring sampling in March and April (solid squares), shortly before the bears emerged from their dens. Mean body mass loss for all bears was 27.8 kg (± 10.4kg) and ranged from 16 kg (Bear #9) to 51.2 kg (Bear #2). �286 CHANGES IN BODY FAT DURING HIBERNATION FALL % BODY FAT SPRING % BODY FAT 60 .,. 50 ~ 40 LL >o o (Q 30 'Cf2. 20 10 o 1 2 3 4 5 6 7 8 BEAR # Figure 3 - Comparisons of Fall (open bars) and Spring (hatched bars) Body Fat as measured with the Bioelectrical Impedance Analysis instrument. Resistance (in ohms) was measured on each animal during early and late hibernation by snout/tail electrode placement of the instrument electrodes. Total body water was calculated from the measured resistance by the regression equation; Total body water (TBW) = -0.224 + 0.197 (svI2*/stailr**) + 0.137 (BM***) * - snout/vent length (em) squared ** - snout/tail resistance (ohms) *** - Body mass (kg) Percent body fat was calculated from total body water by the equation: Percent body fat = 98.01 - 1.28 (TBW) Protein utilization/muscle tissue water content Bradford Assay - Results from the Bradford Assay of peptide bonds indicated that there was a . significant decrease in skeletal muscle protein concentration during hibernation in both the gastrocnemius (GAST) muscle, Mean 6 = -17.2 (± 6.83) mg/g dry wt, P = 0.045, and in the biceps femoris (BIFEM) �287 muscle, Mean t:. = - 44.6 (± 13.59) mg/g dry wt, P = 0.017. Six of the seven bears sampled experienced a decrease in protein concentration in the GAST and all seven bears showed a decrease inprotein concentration in the BIFEM (Figure 4). CHN Assay - Protein concentration, as calculated from percent nitrogen in each muscle sample, did not significantly change in either the GAST or the BIFEM during hibernation, p = 0.28 and p = 0.15, respectively (Figure 5). Percent water content ofBIFEM increased significantly during hibernation, Mean Il= 2.68% (± 0.87), p = 0.02, while water content in the GAST did not differ significantly, p = 0.42 (Figure 6). Muscle fiber type composition, number, and cross-sectional areaThere was a significant increase in the percentage of fast-twitch versus slow -twitch muscle fibers in the BIFEM, Mean t:. = 9.9l'1o(± 4.5), p = 0.03, during hibernation. There was also an apparent increase in percentage of fast-twitch fibers in the GAST, however, this change was not statistically significant, p = 0.23 (Figure 7). Cross-sectional area of fast-twitch muscle fibers in both the GAST and the BIFEM was not significantly altered (p = 0.58 and p = 0.48, respectively) .. Similarly, there was no significant change in crosssectional area of slow-twitch muscle fibers in either muscle, p = 0.18 for GAST, and p = 0.17 for BIFEM (Figure 8). There was no significant change in the mean number of total fibers counted (per 3.9 mrrr'), for either the GAST p = 0.58 or the BIFEM p = 0.63 (Figure 9). . PROTEIN CONCENTRATION - BRADFORD ASSAY 450 FALL SPRING I- 425 :c o w S >cr:: * 400 * 0 375 o -.... o ~ 350 GAST BIFEM Figure 4 - Changes in protein concentration(mglgdry wt.) during Fall (open bars) and Spring (hatchedbars) in the GAST and BIFEM muscles of seven bears using the BradfordProtein Assay. Asterisk depicts a significantreduction (p < 0.05) in protein content during hibernation. GAST mean change in concentration = -17.2 mglg dry wt.·(± 6.83), p = 0.045~BIFEM mean change in concentration = -44.6 mglgdry wt. (± 13.59); p = 0.017. Verticallines represent + SEM. �288 PROTEIN CONCENTRATION - CHN ASSAY 1000 900 lI FALL 800 C) 700 ~ 600 o 500 >0:: SPRING ~ C) ~ 300 200 GAST BIFEM Figure 5 - Fall (open bars) and Spring (hatched bars) protein concentration (mg/g dry wt.) in GAST and BIFEM muscles of seven bears using the eHN tissue-ignition assay, which measures total % nitrogen in each sample. Neither the GAST nor the BIFEM showed a significant change in protein concentration using this assay (p = 0.28 and p = 0.15, respectively). Vertical lines represent + SEM. . MUSCLE FIBER WATER CONTENT 80 * 70 SPRING 60 0::: c=J FALL 50 ill I- 40 ~ cf? 30 20 10 0 GAST BIFEM Figure 6 - Mean % water ofGAST and BIFEM during Fall (open bars) and Spring (hatched bars) for seven bears following freeze-drying for protein concentration analysis. Weights were obtained before drying, and again after 24 hours in the freeze-dryer for both Fall and Spring samples. Asterisk depicts a significant (p < 0.05) increase in water content in BIFEM during hibernation. Vertical lines represent + SEM. �289 FIBER TYPE TRANSFORMATION DURING HIBERNATION 80 (f) 70 * 0::: W .c=J FALL % FAST-TWITCH rn 60 LL ~ 50 I FIBERS SPRING % FAST-TWITCH FIBERS 0 J- 40 ~I J- 30 (f) « LL 20 "CF. 10 0 BIFEM GAST Figure 7 - Percentages offast-twitch fibers for Fall (open bars) and Spring (hatched bars) for GAST and BIFEM muscles of six bears. Asterisk depicts a significant increase in the ratio offast-twitch to slow-twitch fibers as indicated by the increase in percentage offast-twitch fibers during hibernation and the reciprocal decrease in percentages of slowtwitch fibers for the same period (only fast-twitch fibers are represented in the figure). Mean increase in % fast-twitch fibers in the GAST was 3.56 (± 5.0), p = 0.23; in BIFEM, 9.9 (± 4.5), p = 0.03. Vertical lines represent + SEM. MUSCLE FIBER X-S AREA 7000 (f) 0::: w 6000.,. W ~ 5000 - J- o FALLX-S AREA ~ SPRING X-S AREA 0::: o 4000 - ~. a 3000 - « w 2000 - (f) 0:: « (f) 1000 - I >< o _J..__----'--L. 'FAST 'SLOW GAST '-'--FAST SLOW BIFEM Figure 8 - Mean cross-sectional areas Jlffi2 ofboth fast- and slow-twitch fibers for Fall (open bars) and Spring (hatched bars) , in GAST and BIFEM for seven bears. Neither muscle exhibited a significant change in fiber area in either fast- or slowtwitch during hibernation, p = 0.18 and p = 0.17 for GAST and BIFEM respectively. Vertical lines represent + SEM. �290 CHANGES IN MUSCLE FIBER NUMBER « ill 400 0::: « I- 300 ~ SPRING c:::=J FALL Z :::> =H:. 0::: ill en LL 200 100 GAST BIFEM Figure 9 - Number of muscle fibers per 3900 JllD2 (in four video sub-images) of combined fast and slow twitch muscle fibers during Fall (open bars) and Spring (hatched bars) for seven bears. There was no significant change in number of fibers per 3900 JllD2 in either muscle during hibernation. Vertical lines represent +SEM. Citrate-synthase activity Citrate-synthase (CS) enzymatic activity decreased significantly in the BIFEM muscles during hibernation, Mean '" = -8.41 umol/g/min (± 2.99), p = 0.015. However, CS activity in the GAST muscle exhibited only a vel)' slight and non-significant alteration during the hibernation period Mean '" = -1.11 umcl/g/min (± 2.95), p = 0.72 (Figure 10). 30 CS ACTIVITY .•........ z -::J ~ c=:J 25 FWIJ· SPRING (9 0 FALL 20 ~ __.. :::> 15 ~ > I- 10 0 « (J) 5 0 0 GAST BIFEM Figure 10 - Mean CS activity for Fall (open bars) and Spring (hatched bars) in GAST and BIFEMmuscles of seven bears sampled. Vertical axis is CS activity in umol/gm tissue/min. Asterisk depicts a significant decrease in CS activity during hibernation; mean decrease = -8.41 umol/g/min (± 2.99), p = O.QlS. Vertical lines represent + SEM. �291 DISCUSSION Den preparation and entry times . The entire denning chronology of our sample bears was quite a bit later than other bears in different locations in Colorado (Beck, 1991), and could have been a result of several factors, including a very warm, dry fall, and an excellent hard mast crop. We observed what appeared to be obese bears feeding on acorns and pinon nuts well into December until heavy snowfall made foraging for the mast difficult for the bears. This suggests that some bears, at least in this particular habitat, will continue to forage as long as it remains energetically beneficial to do so, and they may not be stimulated to enter their dens by photoperiod or any physiological trigger until then. In fact, radio tracking signals from the second week of December indicated that several bears (mostly subadult and adult males) were still active and moving throughout the study area. Sampling intervals - Table 1 lists the number of days between samples for each bear. The mean interval was 123 days (± l3.7); the shortest interval between samples was 108 days and the longest interval was 152 days. Bears were resampled in the spring in approximately the same order as they were handled in the fall in an effort to maintain fairly equal intervals between samples. TABLE 1 - Sampling interval (days between the Fall and Spring sampling periods) for each bear lkm:.! Fall sample date 1 2 3 4 5 6 7 8 9 11-10-94 11-12-94 11-14-94 11-15-94 12-1-94 12-2-94 12-5-94 12-9-94 12-9-94 Spring sample date 3-17-95 3-18-95 4-16-95 3-20-95 3-19-95 No sample No sample 4-1-95 ·3-31-95 Interval 127 days 127 days 152 days 125 days 108 days N/A N/A 113 days 112 days As previously mentioned, bears Nos. 6 & 7 left their dens before we were able to res ample them in the spring. Bear No.6 was tracked to a location near her den in March, but was already out of the den and . moving, making resampling impossible. No cubs were visible during this time (Bear No.6 was subsequently observed during May, 1995 breeding with two different males, only minutes apart). Bear No. 7 was never relocated either by aerial radio-tracking or ground tracking, and it is assumed that she left the den early to begin to forage; she was the smallest of all the bears and was visually thinner than the other bears when she was handled in the fall. In fact, she was the only one of the nine bears handled that did not appear to be in excellent condition early in the denning season. It seems highly unlikely that she would have produced any cubs, given her somewhat poor physical condition in the fall and the fact that she left the den when cubs, if present, would have been too small to travel. Bears entered their dens later than expected (Beck, personal communication). Lindzey and Meslow (1976) found that female black bears in southwestern Washington entered their dens earlier than males and this chronology seemed to hold true for bears in our study area, although all bears were still foraging into .November. The abundant hard mast crop and a relatively mild fall could help explain the apparent delay by the bears in entering their dens. In addition to the two bears that left their dens in mid-March, we observed evidence that a few of the other bears may have already been out of their dens briefly during the spring. �292 Body MasslFat Measurements Bears lost an average of 24.3% of their total body mass, as expected, during hibernation. There was no significant correlation between individual body mass loss and litter size. However, it is interesting to note that Bear #7, the only bear sampled that was not lactating during the study period, lost a smaller percentage of body mass (15%) than any of the bears with cubs, and considerably less than the average of the seven bears. Body fat, as calculated from resistance readings generated by the Bioelectric Impedance Analysis instrument, appeared to increase. However, I believe this pattern was erroneous. It is well-documented that bears utilize adipose tissue almost exclusively as a metabolic substrate during hibernation (Nelson 1973; Nelson et al. 1973; Lundberg et al. 1976), therefore one can only conclude that these measurements were in error since total body mass decreased by an average of 24.3%. Field calculations performed after the fall BIA measurement on Bear #2 indicated that only 10% of her total body mass was fatty tissue; this bear weighed 345 Ibs. and appeared obese. We assumed at the time that the instrument reading was in error due to the fact that we pulled this bear out of her den in the midst of a snowstorm andshe became wet very quickly. Therefore, subsequent to the handling of this bear, we took even greater pains in the field to ensure that all bears were completely dry and in a consistent body position when using the BIA instrument. Nevertheless, BIA readings continued to seem erroneous during the fall field season, as some of the smallest, and visually leanest bears produced calculated body fat values of upwards of 50%. BIA measurements during spring sampling again provided no clear pattern or correlation with body mass, and all of the calculated body fat percentages appeared to be too high, given that the bears were about to emerge from their dens after approximately four months of fasting. Conversations with Dr. Mike Vaughn, a biologist at Virginia Polytechnic Institute, revealed that he too was experiencing inconsistencies in measuring total body fat on his captive bears using the BIA instrument. He went on to say that he felt that the range of variability of each measurement prevented him from placing much trust in either the accuracy or the precision of these measurements. This protocol, as reported by Farley and Robbins (1993) was, at the time of my study, the very latest in "cutting edge" technology for field measurements of body condition. As previously mentioned, time spent in Dr. Robbins' laboratory at Washington State University ensured our understanding of the equipment and its operation; , however, the poor performance in the field prevents endorsement of this procedure for field applications. The fact that even in a completely controlled laboratory setting researchers were unable to use this BIA instrument with confidence (Vaughn, personal communication 1994) places extreme doubt as to the utility of this method, especially under field conditions. Obviously, better methods for estimating body fat on free-ranging bears, such as portable ultra-sound instruments, would make these measurements much more useful. Protein Utilization During Hibernation Thirteen of fourteen muscle biopsies taken during the spring from the gastrocnemius (GAST) and biceps femoris (BIFEM) indicated a net loss of protein concentration when compared to corresponding biopsies taken during fall sampling. While these results support our hypothesis that lactating female black bears utilize some amount of protein during hibernation, they are contradictory to previous studies that did not identify net loss in bear skeletal muscles during early and late hibernation (Nelson 1973; Lundberg et al. 1976; Nelson 1980; Koebel et al. 1991). For example, Lundberg et al. (1976) found that protein turnover, a measure of both protein catabolism and protein synthesis, increased three- to five-fold in captive bears during hibernation, but that skeletal muscle protein concentration was unchanged. This statement seems to imply that net protein loss does not occur during hibernation in bears. Since it has been shown that bears do not eat or drink during hibernation, the onlysource of free amino acids that could be used for protein synthesis would have to come from protein breakdown: Therefore, if protein degradation increased during hibernation, it follows that an equivalent amount of protein synthesis would also have to increase in order to provide an exact protein replacement. Urea recycling could potentially assist in achieving this nitrogen balance. As previously mentioned, Nelson et al. (1975) determined, through the use ofisotope-labelled urea, �293 that bears have the ability to passively move urea (the nitrogenous end product of protein catabolism) into the lower intestinal tract where it is hydrolyzed by urealytic bacteria to ammonia. This ammonia is then actively moved from the intestinal tract into the blood and subsequently reaminated to nonessential amino acids in the liver (Lundberg et al. 1976). The recycling of this nitrogen, in the form of amino acids back into structural proteins has been offered as an explanation for the apparent lack of protein loss during hibernation (Lundberg et al. 1976). This scenario suggests, however, an extremely efficient (almost 100%) process of nitrogen recycling in order to achieve high protein turnover with no protein loss, a highly unlikely condition. Data from this study only partially support the conclusions drawn from the previously discussed studies on protein loss in bears. The results from the CHN assay suggested that the mean percent nitrogen from both GAST and BIFEM was not significantly reduced during hibernation. However, unlike other studies, the peptide bonds of skeletal muscle protein content, as revealed by the Bradford assay, did not remain unaltered but were actually reduced. Protein catabolism is characterized not just by a reduction in muscle protein, but also by an increase in free amino acids, ammonia, and urea as metabolic end products. A reduction of peptide bonds (results of Bradford assay) without a concomitant reduction in total nitrogen (results ofCHN assay) suggest that some of the nitrogen from catabolized protein is being identified as other nitrogenous end products or intermediates. Therefore, data from this study not only suggests an increase in protein mobilization but a significant muscle loss. Also, it is possible that if protein were being continually degraded and synthesized, some amount of repartitioning of the proteins may be occurring, i.e., proteins from the particular skeletal muscles we biopsied may be catabolized, and some of the free amino acids synthesized and synthesized into protein in another muscle that was not sampled. These results support our hypothesis that bears in general, but particularly lactating bears, must utilize protein during hibernation in order to provide such things as: 1) replacement water for insensible water loss (Bintz et al 1979); 2) short chain carbon compounds for Krebs cycle intermediates (Bintz et al 1979); and, 3) a source of water and energy for fetal development and milk production (DelGiudice et al. 1991). Further, it is apparent that hibernating bears provide an excellent model for the study of muscle disuse. This is important considering the conflicting reports of protein use during periods of inactivity by both hibernators and non-hibernators. Some studies suggest that no change in protein concentration occurs in . hibernating ground squirrels (Musacchia, et aI., 1989) or that there are no seasonal changes in protein concentration in mouse skeletal muscle (Wickler 1981). Other investigators have reported that protein concentration decreased in certain muscles during hindlimb suspension (Steffen and Musacchia 1984) and hibernation (Wickler and Hoyt 1990), while remaining unchanged during hibernation in others (Steffen et al. 1991). Finally, while Lundberg et al. (1976) reported a 3- to 5-fold increase in protein turnover in bears during hibernation, Loughna et al. (1986) measured a decrease in protein turnover in rat soleus muscle following hindlimb suspension. It is clear from the variability of results in these studies that considerable work needs to be conducted in order to clarify the various biochemical mechanisms used by immobile mammals. In addition to the apparent variability in the literature regarding protein usage, and citrate synthase activity as well, there continues to be a wide range of reported values of these parameters. Reported protein concentration ranges from 36 mg/g wet wt. (Ji et a/1991) to 302 mg/g wet wt. (Yacoe 1982), and is also reported as a percentage of dry wt. (Koebel et aI1991). Similarly, the reported range of citrate synthase activity is from 4.7 umol/g/min wet wt. (Ji et a/1991) to 352 umol/g/min wet wt. (Yacoe 1983). While variability is to be expected, the magnitude of these discrepancies indicate that standardization is necessary, with tissue preparation, type of assay, and methods of expressing units. Muscle fiber transformation, cross-sectional area, and citrate synthase activity There was a significant decrease in percentage of Type I, slow-twitch aerobic muscle fibers during hibernation in the BIFEM, suggesting that some transformation from slow- to fast-twitch fibers had occurred. Interestingly, unlike other hibernators, there was no compensatory increase in citrate synthase (CS) activity in this muscle. These results support my hypothesis regarding muscle disuse. In point of fact, CS activity actually decreased in the BIFEM from fall measurements, which seems logical for the following reasons. First, as mentioned earlier, bears do not lower their body temperature to the same extent as more 'classic' hibernators, and therefore do not undergo the violent periods of shivering thermogenesis during periodic �294 arousals. This seems to be a driving force behind the increase in CS activity in rodent and insectivore hibernators (Wielder and Hoyt, 1990; Wielder, et al., 1991).· Second, there is no supportive evidence that denning bears demonstrate periodic muscular activity such as isometric contractions (Brian Barnes, personal communication). Third, a decrease in overall CS activity would be expected, based upon the observed . decrease in SO muscle fibers which utilize CS in aerobic metabolic processes. Finally, it seems logical that bears would not increase CS activity during hibernation to maintain aerobic muscle capacity since any demand for muscle activity would typically be following some type of denning disturbance. Should this disturbance occur, the period of muscle recruitment would be brief and intense, either during a relocation or defense of the den. Type II glycolytic fibers would typically be recruited during such a scenario. In the GAST, slow/fast fiber type ratios and CS activity were essentially unchanged. These results agree with Koebel, et al. (1991), who also found no significant alteration in CS activity in post-denning biopsies from GAST of captive black bears. When viewed collectively, there is a striking difference in CS activity from hibernating animals compared to animals subjected to some form of suspension/immobilization. Most studies using rodent hibernators have reported a significant increase in CS activity during hibernation (Wiclder, 1981; Wiclder,etai., 1987; Thomason and Booth, 1990; WiclderandHoyt, 1990; Steffen,etal., 1991). Onthe other hand, hindlimb suspension/immobilization models largely depict a decrease in CS activity during periods of muscle disuse (Booth, 1977; Desplanches, et al., 1987; Fell, et aI., 1985). This suggests that these suspension models may not be effective in duplicating physiological conditions experienced during hibernation. One study in particular, however, made an interesting observation regarding muscle position and CS activity. Booth (1977) found that rat GAST and soleus muscles exhibited an even greater decrease in CS activity when immobilized in a position that was flexed to less than resting length. This may be similar to the curled, compact position assumed by most bears during hibernation, which could cause a flexed state less than the resting length, thereby reducing any potential increase in CS as reported for other hibernators. Interestingly, the alteration of CS for bears in this study is more like that reported on standard disuse animal models than it is for hibernating small mammals. In this study, the mean number of fibers per unit area for either the GAST or the BIFEM did not change during hibernation. Mean cross-sectional area of the two muscles sampled also did not change over the four month period, indicating no measurable atrophy of these skeletal muscles. In addition, muscle tissue weights taken before and after freeze-drying for protein concentration analysis showed there was no significant change in water content of the GAST sampled during hibernation; however, the percent water in . the BIFEM actually increased significantly, indicating that hydration was not a critical factor affecting muscle cell shape or size and that no dehydration occurred during winter. This increase in water content in the BIFEM could be a result of water influx down an osmotic gradient created by protein catabolites. Under normal metabolic conditions, a relatively isosmotic state is maintained between the muscle cells and surrounding extracellular space. Since the BIFEM exhibited a significant decrease in protein concentration, indicating an elevation of protein catabolism, the resulting osmotically-active free amino acids, ammonia, and nitrogen may have created a hyperosmotic condition within the muscle cells, which resulted in net water movement into the cell to equilibrate the cellular environment. Steffen and Musacchia (1984) suggested that muscle atrophy results from a decrease in muscle cell size, rather than a reduction in the number of fibers. This idea is supported by a number of studies on both hibernating small mammals and animals subjected to various types of immobilization/suspension. For example, Musacchia, et al. (1989) found that hibernating ground squirrels experienced a decrease in crosssectional area of muscle fibers during hibernation, but no change in fiber number .. Similarly, using immobilized rats, Nicks, et al. (1989) found no change in the number of fibers, but discovered that crosssectional area decreased by over 42%. It appears that bears in this study did not experience significant muscle atrophy in terms of muscle fiber size or number during hibernation, but did show a loss in protein and alteration of fiber type ratios. In spite of this, bears manage to maintain good muscle tone for emergence in the spring. This unique maintenance of muscle tone could be due either to periods of mild shivering (although never observed) or to some type of isometric activity during denning not previously identified in �295 captive bears. It seems logical since, with the exception of a decrease in protein concentration and altered fiber type, bears apparently do not undergo the physiological changes common to other muscle disuse atrophy models, and must therefore be doing something different in order to exhibit such normal muscle function following several months of inactivity. Monitoring muscle activity of hibernating bears using EMG telemetry and data recorders could clarify this phenomenon. CONCLUSIONS - Based upon the muscle biopsies obtained during early and late stages of denning and hibernation, black bears, and particularly lactating bears, seem to require some protein degradation and net protein loss during hibernation. This protein is presumably used as a source of metabolic energy in the form of Krebs cycle intermediates, as a source of replacement water for insensible water loss, as well as a source of nitrogen and water for cub production and growth. The observed reduction in protein concentration was most likely influenced by lactation, needed to support the production and maintenance of cubs in the den. The bears' unique ability to recycle urea, typically a toxic by-product of protein catabolism, allows them to utilize protein as a metabolic substrate, without accumulating urea and necessitating nitrogen and water excretion through urination. With regard to muscle atrophy during hibernation, bears do not appear to experience significant muscle loss during several months of inactivity, in contrast to other hibernators. There was no reduction in cross-sectional area or number of muscle fibers (indicators of muscle atrophy)· and no increase in citrate synthase enzymatic activity, commonly reported for small mammal hibernators. The transformation of slowtwitch aerobic fibers to fast-twitch anaerobic fibers exhibited by these bears is typical of extended periods of inactivity reported for other disuse atrophy models (including hibernators), but the concomitant decrease in CS activity is more characteristic of non-hibernating immobile mammals. Hibernating bears do not completely fit characterizations of any single muscle disuse model, but share similarities with each of these models. For example, bears in this study experience little muscle atrophy during hibernation, but, unlike small hibernators, bears do not increase their muscle aerobic metabolic. capabilities by increasing citrate synthase activity. Additionally, when compared to human disuse models and hindlimb suspension models that exhibit significant atrophy and decreased citrate synthase activity, bears do not show measurable muscle atrophy, but do decrease citrate synthase activity in conjunction with slow- to fast-twitch fiber transformation. Clearly, bears employ a unique physiological mechanism to maintain muscle tone during extended periods of inactivity such as hibernation. Physiologists working on bears are just beginning to understand the complex adaptations to hibernation that these large mammals have acquired. New research that would help to understand these phenomena include field testing of more accurate ways to obtain measurements of body fat, possibly using portable ultra-sound devices. This would allow closer estimates of relative amounts of fat and fat-free tissues utilized during hibernation. Further, to better calculate protein utilization, simultaneous assays of protein, urea, free amino acids, and free nitrogen, along with connective. tissues such as collagen, could be used to determine the sources of protein catabolism and synthesis. Finally, the use of ultra-sensitive EMG recorders would allow scientists to determine if bears maintain some type of shivering/isometric activity during hibernation, thus allowing them to avoid the debilitating atrophy typical of other mammals, including humans, that remain inactive for extended periods. It is clear that there are important broader scientific implications associated with our understanding of the phenomenon of hibernation. This is true not only for increased appreciation of bears as a component of species diversity, but also for management of bears as a game animal. The stress of winter anorexia, compounded by lactation, result in the loss of fat and protein, making the period of spring emergence critical to survival both of the female and her cubs. Consequently, spring is not an appropriate time to subject bears to additional stresses by hunting. The understanding of bears as a muscle disuse model is important for human applications as well, such as extended periods of convalescence and spaceflight. .Ursid hibernation is, without a doubt, a unique physiological and behavioral strategy for winter survival. �296 Literature Cited Ahlquist, D.A, Nelson, RA, Jones, J.D., Ellefson, RD. 1976. Glycerol and alanine metabolism in the . hibernating black bear. Physiologist. 19: 107. Baker, B.E., Harington, C.R, and Symes, AL. 1963. Polar Bear Milk. I. Gross composition and fat constitution. Can. J. Zool. 41: 1035-1039 .. Beck, T.D.1. September 1991. pp.86. Black bears of west-central Colorado. Technical Publication No. 39. Colorado Division of Wildlife. 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Energy provision, tissue utilization, and weight loss in prolonged starvation. British Med. Journal. 2: 352-356. Schwartz, C.C., Miller, S.D., and Franzmann, A.W. 1987. Denning ecology of three black bear populations in Alaska. Int. Conf. Bear Res. and Manage. 7: 281-29l. Shepherd, D., and Garland, P.B. 1969. Citrate synthase from rat liver. Meth. Enzym. 8: 98-108. Steffen, 1M., and Musacchia, X.J. 1984. Effect of hypokinesia and hypodynamia on protein, RNA, and DNA in rat hindlimb muscles. Am. 1 Physiol. 247: R728-R732. . Steffen, 1M., Koebel, D.A., Musacchia, X.l, and Milsom, W.K 1991. Morphometric and metabolic indices of disuse in muscles of hibernating ground squirrels. Compo Biochem. Physiol. 99B(4): 815-819. �298 Templeton, G.H., Padalino, M., Manton, 1, Glasberg, M., Silver, CJ., Silver, P., DeMartino, G., Leconey, T., Klug, G., Hagler, H., and Sutko, lL. 1984. Influence of suspension hypokinesia on rat soleus muscle. 1AppI. PhysioI. 56(2):278-286. Thomason, D.B., and Booth, F.W. 1990. Atrophy of the soleus muscle by hindlimb unweighting. 1AppI. PhysioI. 68(1): 1-12. Tietje, W.D., and RL. Ruff. 1980. Denning behavior of black bears in boreal forests of Alberta. 1. Wildl. Manage. 44: 858-870. Wickler, SJ. 1981. Seasonal changes in enzymes of aerobic heat production in the white footed-mouse. Am. 1PhysioI. 240: R-289-R294. . Wickler, S.l, Horwitz, B.A., and Klott, K.S. 1987. Muscle function in hibernating hamsters: a natural analog to bedrest? 1Therm. BioI. 12(2): 163-166. Wickler, SJ., and Hoyt, D.F. 1990. Disuse atrophy in rodents: a hibernator responds atypically. Physiologist. 33: A12!. (Abs) Wickler, SJ., Hoyt, Donald F., and van Breukelen, Frank. 1991. Disuse atrophy in the hibernating goldenmantled ground squirrel, Spermophilus lateralis. Am. 1PhysioI. 261: RI214-RI217. Yacoe, M.E. 1983. Protein metabolism in the pectoralis muscle and liver of hibernating bats, Eptesicus fuscus.L Compo PhysioI. 152: 137-144. �299 APPENDIX A - Protein Assay by Dye Binding (modified Bradford method) Theory - Protein concentration may be determined quantitatively by the binding of Coomassie blue dye G250 to protein in an acid-denaturing solvent. The concentration of protein-bound dye is determined by the spectrophotometric absorbance at 595nm. Reagents Coomassie Brilliant Blue G250 (Sigma # B 1131) 95% Ethanol 85% Phosphoric Acid Defatted Bovine Serum Albumin (Sigma # A6003) Sodium Chloride Triton X-I00 HPLC-grade H20 Solutions A. Protein Dilution Buffer - 0.15M NaCI + 0.1% Triton X-I00 B. BSA stock solution - lmg/ml fmal volume in protein dilution buffer C. Assay Buffer (makes 1 liter final volume) 1) Dissolve 100mg Coomassie Blue in 50ml95% Ethanol 2) Add 100ml85% Phosphoric Acid to the CoomassielEthanol solution 3) Dilute to 1 liter with HPLC-grade H20 4) Store refrigerated Protein Standards - Standards are mixed in 12 x 75 mm disposable culture tubes using the BSA stock solution and the protein dilution buffer as follows: . 1OJ.1g1100,d= 200J.1lBSA + 1800J.1lprotein dilution buffer = 2ml final volume 20J.1g1100J.1l = 400J.1lBSA + 1600J.1lprotein dilution buffer = 2ml fmal volume 30J.1g1100J.1l = 600J.1lBSA + 1400J.1lprotein dilution buffer = 2ml fmal volume 40J.1g1100J.1l = 800J.1lBSA + 1200J.1lprotein dilution buffer = 2~ fmal volume 50J.1g1100J.11= 1000J.1lBSA + 1000J.1lprotein dilution buffer = 2ml fmal volume - vortex all fmal volumes Unknown Preparation - Tissue samples are freeze-dried overnight in a freeze-dryer and accurate wet/dry weights are obtained (to O.OOlmg). - Dry tissue sample is homogenized in a culture tube in 1:40 weight volume (mg/ul) O.IM phosphate buffer, pH 7.4, eg., 11.399mg dry tissue in 456 J.1lphosphate buffer. - Unknown homogenates are diluted 1:20 with protein dilution buffer, i.e., 20J.1lunknown homogenate + 380J.1lprotein dilution buffer - vortex Procedure - Each unknown sample should be prepared in triplicate. 1) Add 100J.1lof each protein standard to labelled culture tubes. 2) Add 100J.1lof dilute unknowns to labelled culture tubes (remember to prepare in triplicate) .3) Add 100J.1lof protein dilution buffer to labelled culture tube to be used as blank .. 4) Set timer for 15 minutes. 5) Add lml Assay Buffer to the "blank" tube which contains only dilution buffer - vortex 6) Start timer - add Iml Assay Buffer to each remaining tube, both standards and unknowns, vortexing each tube briefly after the addition of the Assay Buffer. 7) After the first tube (blank) has incubated for 15 minutes (timer reaches 0), put the blank in the �300 spectrophotometer and "zero" the absorbance of the instrument at 595nm. 8) Read each tube, in order, at 595 om, at approximately 30 second intervals so that each tube has incubated for at least 15 minutes before reading. 9) Generate a regression equation for concentration using the 5 standards as a standard curve and calculate the concentration for each unknown using this regression equation. NOTE: A new standard curve should be generated for each group of unknowns. References: Bradford, M.M. 1976. A rapid and sensitive method for the quantification of microgram quantities of protein utilizing the principal of protein -dye binding. Anal. Biochem. 72: 248-254. Kley, H.V. and S.M. Hale. 1977. Assay for protein by dye-binding. Anal. Biochem. 81: 485-487. Pierce,; 1. and C.H. Suelter. 1977. An evaluation of the coomassie brilliant blue 0250 dye-binding method for quantitative protein determination. Anal. Biochem. 81: 478-480. �301 APPENDIX B - Citrate Synthase Assay REAGENTSI. Homogenizing medium - Potassiwn phosphate (KPi) buffer (50mM, pH 7.4) A. Dissolve 0.68 g monobasic KPi (anhydrous K2P04) in 100 ml distilled H20. B. Dissolve 1.141 g dibasic KPi (K2PO4) in 100 ml distilled H20. C. Combine 19 ml of the monobasic stock with 81 ml of the dibasic stock - this gives a 50mM KPi buffer with a pH of 7.4. D. Add the following reagents to this buffer: 1) EDTA disodiwn salt(l mM) - 37.2 mg 2) MgCI2 (2 mMj - 40.7 mg 3) ADP (2 mM) - 85.4 mg (Sodiwn salt - Sigma # A-6521) 4) Dithiothreitol (0.5 mM) - 7.71 mg (Sigma # D-0632) The homogenizing mediwn should be stored at 4°C. ll. Tris buffer - (100 mM, pH 8.0) A. Dissolve 7.9 g Tris HCI in 500 ml distilled H20. B. Dissolve 6.05 g Trizma base in 500 ml distilled H20. C. Combine appropriate volwnes of stock Tris HCI and Tris base to achieve pH 8.0. Store at 4°C. (Note: This Tris buffer - not distilled water - will be used to make the OAA and the DTNB describied below as Reagents III and IV.) Ill, QM - (10 mM) (Oxalacetic acid, Sigma # 0-4126) - Dissolve 13.2 mg OAA in 10 mll00 mM Tris buffer, pH 8.0. Store at 4°C. IV. 5,5'-dithiobis-(2-nitrobenzoate) - (DTNB, 1 mM, Sigma # D-8130) - Dissolve 3.9 mg DTNB in 10 mll00 mM Tris buffer, pH 8.0. Store at 4°C. V. Acetyl CoA - (1.5 mM) (Lithiwn salt - Sigma # A-2181) - Dissolve 12.14 mg in 10 ml distilled H20. Can be stored frozen at -10°C for a few days. HOMOGENIZATION PROCEDURE- 1. Frozen muscle samples should be transported to the lab in a small dewar ofliquid nitrogen. A small piece of tissue (::: 4Omg) is quickly removed from the liquid nitrogen and placed on powder paper and weighed, and then immediately placed into a culture tube on ice. 2. Homogenizing mediwn is added in a lg: 19m1concentration to the culture tube and the tissue is homogenized using an Omni tissue homogenizer. The homogenization is performed with the culture tube immersed in a beaker of ice. 3. The homogenate is immediately placed into the freezer for the first of four freeze-thaw cycles. (Liquid nitrogen may be used to freeze the homogenate if care is used to gradually freeze the culture . tube to avoid breakage.) �302 (Note: Based on the amount of time required to run each assay, two muscle samples seems to be the optimum number to attempt to assay at one time.) ASSAY PROCEDURE - (Note: Spectrophotometer should be set up with circulating pump to maintain chamber temperature at 30°C) 1. ,Following the fourth freeze-thaw cycle, the homogenates are centrifuged at 700 x g (2700 rpm) at 4°C (refrigerator) for 10 minutes.' , 2. While the homogenates are spinning, prepare six culture tubes (label duplicate samples and a blank for each muscle sample) by adding the following to all six tubes: - 100 mM Tris (PH 8.0) 650 JlI - l.0 mM DTNB 100 JlI - 1.5 mM Acetyl CoA 100 JlI Place the six culture tubes containing the above reagents into a 30°C water bath to equilibrate before assay. 3. Remove the homogenates from the centrifuge after 10 minutes; dilute 50 JlI of each supem:atant with 1 ml of 100 mM Tris buffer (PH 8.0). Vortex each mixture and place on ice until analysis. The activity is stable for at least 1.5 hours. 4. When ready, add 50 JlI of one of the diluted samples on ice to the first of two duplicate culture tubes from the water bath; vortex and immediately transfer to a disposable cuvette. 5. Place into the spectrophotometer and monitor the change in absorbance at 412 nm every 30 seconds for 3 minutes. Remove the cuvette and add 100 JlI of OAA to the cuvette - cover with a small square of parafilm and shake vigorously for 5 - 10 seconds. Replace the cuvette into the spectrophotometer and monitor the change in absorbance at 412 nm every 30 seconds for an additional 10 minutes. The highest linear activity should occur between 6 10 minutes. 6. Repeat steps 4 and 5 for the second duplicate tube in the water bath. 7. Repeat steps 4 and 5 for the blank, however, do not add OAA to the blank. In addition, the blank should be monitored every 30 seconds for 10 minutes, omitting the initial 3-minute pre-OAA time as monitored for the duplicate samples. ' CALCULATIONS - 1. A mM extinction coefficientof 13.6 is used for calculation. 2. Dilution of muscle is as follows: 1 g musclel20 ml homogenate x 0.05 ml homogenatell.05 ml diluted homogenate x 0.05 ml diluted sample/cuvette = 0.000119 g muscle/cuvette. 3. Citrate synthase activity (umol/min x g) is calculated according to the formula: (Mean II absorbance/min of sample after OAA addition - Mean II absorbance/min of blank) + (13.6 per Jlmol)(0.000119g) = umol/min x g. �303 APPENDIX C - Muscle tissue staining and fiber typing procedure ATPase Activity Guth, L., and Samaga, F.J. 1970. Research Note: Procedure for the Histochemical demonstration of Actomyosin ATPase. Experimental Neurology. 28:365-367. I. Cross-sectional "pie" of muscle sample should be mounted on cork so that fiber orientation is perpendicular to the cork. Surround tissue with cryoform, leaving top portion of tissue exposed to promote uniform freezing. Freeze tissue mounted on cork using isopentane (3-methyl butane) cooled in liquid nitrogen. 2. Store frozen tissue in plastic ziplock bags at -70°C. 3. Warm tissue up to -20°C in a cryostat and cut tissue into 5 -7 Jim thick slices. Pull cut tissue off of the cryostat platform using a room temperature glass slide. Dry tissue at room temperature for 1524 hours, or overnight. Label slides with pencil, NOT an ink pen, otherwise the ink will come off in the stain. 4. Prepare solutions when ready to stain. (important to prepare solutions 3, 4, and 7 immediately before use because these tend to absorb CO2 and thus pH changes.) STAINING PROCEUDRE: 1) 5 min Solution 1 Fixative (5% formalin buffered at pH 7.6) 20 ml formaldehyde + 30 ml H20 = 50 ml formaldehyde solution 31 g Na cacodylate (MW 160, Sigma # C-0250) 10 g CaCI2 (MW 147) 115 g sucrose (MW 342) Bring to fmal volume of 1 liter with H20 2) 1 min Solution 2 - agitate every 20 seconds, then blot on Kim-wipes Rinse Solution (18 roM CaClz in 100 mM Tris. pH 7.8) 12.1 g Trizma base (tris-hydroxymethyl aminomethane, MW 121) 2.65 g CaCI2 in 100 ml = 100 ml of 0.18 M CaCI2 900'mlH20 Adjust pH to 7.8 with HCI (l-6N) and bring to 1 liter wI H20 3) 15 min Solution 3 Alkaline Preincubation (l8mM CaCI2 in 100 mM buffer, pH 10.4) 1.0 ml Sigma No. 221 buffer (2-amino, 2-methyl, l-propanol. 95% - FW 0.2646 g CaCl2 80mlH20 Adjust pH to 10.4 with HCI and bring to 100 ml wI H20 4) 1 min Solution 2 5) 1 min Solution 2 (repeat step 4, blot before step 6) = 89.14) �304 6) 30 min Solution 4 - @ 37°C (use warm water bath) Incubation Solution (2.7 mM ATP. 50 mM KCl, 18 mMCaCI2 in 100 mM buffer. Jili.ti 1 ml Sigma No. 221 buffer 0.2646 g CaCl2 0.370 g KCl (MW 75) 0.152 g ATP disodium (MW 551.2, Sigma # A-7699) 80 ml distilled H20 Adjust pH to 9.4 with 6N HCl and bring to final volume of 100ml with H20 7) 30 sec Solution 5 Wash solution (1% CaCl2, wtiyol) 10 g CaCl2 (MW 147) 1000 ml H20 . 8) 30 sec Solution 5 (repeat) 9) 30 sec Solution 5 (repeat, blot) 10) 3 min Solution 6 Cobalt chloride solution (2% wt/vol) - light sensitive 2 g CoCI2 (MW 238) 100 ml H20 11) 30 sec Solution 7 Alkaline washing solution (100 mM buffer, pH 9 4) 1.5 ml Sigma No. 221 buffer 450ml H20 Bring pH to 9.4 with HCI and adjust to 500 ml with H20 12) 30 sec Solution 7 (repeat) 13) 30 sec Solution 7 (repeat) 14) 30 sec Solution 7 (repeat, blot) 15) 3 min Solution 8 . Ammonium sulfide solution (1 % wtiyol - stinks, use 1.0 ml Ammonium Sulfide (light) 100ml H20 16) 4 min Running tap water 17) 1 min 50% Ethanol 18) 1 min 80% Ethanol 19) 1 min 100% Ethanol 20) 1 min Xylene 21) Mount in Permount and allow to air dry Black staining fibers reflect fast twitch fibers. hood) �305 APPENDIXD Table D-I - Fall and Spring estimates of body mass and mass loss for each bear during hibernation. Nwnbers in parentheses indicate nwnber of cubs present during Spring sampling (0,1,2,3). Mean values are mean± SE. . Ikm:.1l 1 (I) Fall Mass Spring Mass Mass Lost % Lost 24 kg 22% 110.6 kg (243Ibs) 86.6 kg (190Ibs) 2 (3) 157.0 kg (345Ibs) 105.8 kg (232Ibs) 3 (1) 86.6 kg 58.8 (190Ibs) (129Ibs) 115.6 kg (254Ibs) 84.6 kg (186Ibs) 31kg 5 (1) 108.2 kg (238Ibs) 85.6 kg (188Ibs) 22.6 kg 21% (50Ibs) 6 (N/A) 79.0 kg No weight (173Ibs) (left den early) 7 (N/A) 63.8 kg No weight (140Ibs) (left den early) 8 (2) 106.0 kg (233Ibs) 84Akg (186Ibs) 9 (0) 111.8 kg (245Ibs) 95 kg (209Ibs) Mean 104.3 kg 85.8 ±24.9 ± 13.2 27.8 ± lOA 2291bs± 54 61lbs ± 22 4 (2) 188lbs ± 29 (53Ibs) 51.2 kg 33% (113lbs) 27.8 kg 32% (61Ibs) 27% (68Ibs) 21.6 kg 20%. (47Ibs) 16.8 kg 15% (36Ibs) 24.3 ± 6.1 �306 Table D-2 - Fall and Spring percent body fat and percent change in body fat during hibernation as calculated from total body water measured by Bioelectrical Impedance Analysis. Values are mean ± SE. Ikm:.it Fall Body Fat % 1 2 3 4 5 6 7 8 9 Mean 30.48 10.41* 46.77 32.08 . 31.62 52.40** 53.73** J9.85 zazi 31.4±4.2 Spring Body Fat % 43.30 40.62 54.59 40.80 37.11 No measurement No measurement 35.48 uzi 40.4±2.8 % Gain/Loss +12.82% +30.21% +7.82% +8.72% +5.49 NIA NIA -4.37% ±2..lQ 9.0±4.0 *The BIA measurement for this bear was suspect for the fall sampling due to the fact that the bear got very wet during a heavy snowstorm. ** Since a paired t-testwas used for analysis, these two measurements were not included in calculation of Mean Fall body fat%. TABLE D-3 - Protein concentration (mg/g dry wt.) using the Bradford Protein Assay in the GAST and .BIFEM muscles. Mean values are Mean ± SE. * Indicates significant change. �307 TABLE D-4 - Protein Concentration (mg/g dry wt.) using the CHN Assay for total nitrogen in the GAST and BIFEM muscles. Total protein concentrations are calculated from % nitrogen contained in each sample. Mean values are mean ± SE. GASTROCNEMnJS Fall cone. ~ 1 812.5 2 900.0 3 887.5 4 893.7 5 875.0 6 N/A 7 N/A 8 906.2 9 868,7 Mean 877.6 ± 11.95 BICEPS FEMORIS 1 712.5 2 862.5 3 843.7 4 862.5 5 587.5 N/A 6 N/A 7 8 893.7 9 868.7 Mean 804.4 ± 42.51 Spring {<Qru,:, Cbange 899.9 862.5 912.5 937.5 931.2 +87.4 -37.5 +25.0 +43.8 +56.2 N/A N/A N/A N/A 875.0 868,7 -31.2 Q 898.2 ± 11.4120.5 ± 17.44 812.5 887.5 881.2 899.9 912.5 +100.0 +25.0 +37.5 +37.4 +325.0 N/A N/A N/A N/A 912.5 831.2 +18.8 -37,5 876.7 ± 14.9173.3 ± 44.78 �308 TABLE D-5 - Changes in the percent offast-twitch fibers during hibernation. Values are in percent fasttwitch fibers; Arcsin-square root transformation was used for all analyses of percent data. Mean values are Mean ± SE. * This value was not used in the calculation of the mean percent of fall fat-twitch fibers due to uncertainty as to the staining results. ** Indicates significant change. GASTROCNEMIUS ~ Fall % fast-twitch 1 3.8* 2 60.1 3 70.4 4 78.3 5 67.0 6 7 8 9 Mean Spring % fast-twitch 73.1* 79.3 75.8 69.5 65.2 Change +69.3* +19.2 +5.4 -8.8 -1.8 N/A N/A N/A N/A N/A N/A 73.0 65:1 89.4 56.1 +16.4 -9.0 68.9±2.6 72.6±4.0 +3.56± 5.0 = 0.23) (p BICEPS FEMORIS 1 2 3 4 5 6 7 8 9 Mean 46.3 57.4 59.0 52.4 61.1 73.7 53.2 63.2 67.6 56.9 +27.4 -4.2 +4.2 +15.2 -4.2 N/A N/A N/A N/A N/A N/A 52.1 57.2 69.5 71.2 +17.4 +14.0 55.1±1.9 65.0±2.9 ** +9.9±4.5 (p = 0.03) �309 TABLE D-6 - Muscle fiber mean cross-sectional areas (J.l2) for all seven bears sampled for both fast- and slow-twitch fibers in the GAST and BIFEM muscles. Mean values are Mean ± SE. FAST TWITCH FIBERS GASTROCNEMIUS Fall x-s area Spring x-s area Cban~ 4702.0 ± 553.3 5705.2 ± 717.6 403.2 ±704.5 BICEPS FEMORIS 4193.6 ± 330.2 4576.7 ± 654.5 383.1± 512.5 SLOW-TWITCH FIBERS GASTROCNEMIUS 5835.3 ± 587.5 5222.3 ± 557.5 -613.1±410.9 BICEPS FEMORIS 4095.8 ± 314.4 4648.6 ± 527.6 552.8± 361.3 TABLE D-7 - Citrate synthase activity in the GAST and BIFEM muscles during hibernation. are Mean ± SE. * Indicates significant change. GASTROCNEMIUS cs activity Spring CS activity ~ 1 16.4853 2 13.1339 3 27.3863 4 13.0604 5 21.3859 6 N/A 7 N/A 8 21.7400 9 5.6215 rsn Mean BICEPS FEMORIS 1 2 3 4 5 6 7 8 9 Mean Chan~ 21.2648 9.8483 15.4801 15.1569 19.3107 +4.7795 -3.2856 -11.9062 +2.0965 -2.0752 N/A N/A N/A N/A 13.3310 16.6114 -8.4090 +10.9899 16.9733 ± 2.7158 -1.1157 15.8576 ± 1.4214 ± 2.9555 26.2049 11.9136 26.8889 26.6096 33.6097 29.3644 14.1386 12.5976 15.1504 24.9446 3.1595 2.2250 -14.2913 -11.4592 -8.6651 N/A N/A N/A N/A N/A N/A 28.8037 21.9970 12.9662 8.0097 -15.8375 -13.9873 25.1468 ±2.5649 16.7388 ±2.8602 * -8.4080 ±2.9967 (P = 0.015) Mean values ��311 Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS state of Project Colorado No. W-153-R-9 Work Plan No. 9A Job No. Period Author: REPORT Mammals Elk Investigations Spatial Covered: July K. R. Wiison, Research Analysis of Elk Survival 1, 1995 - June 30, 1996 D. A. Werle, and N. T. Hobbs ABSTRACT We developed a landscape simulator to evaluate a proportional hazard model and logistic regression to detect annual differences in animal in relation to habitat use. No additional progress was accomplished this period. rate survival during ��313 Colorado Division Wildlife Research July 1996 of Wildlife Report Wildlife JOB PROGRESS State of Project Colorado No. W-153-R-7 Work Plan No. lOA Author: Mammals Research Kit Fox Studies Kit fox (Vulpes macrotis) in Colorado Job No. Period REPORT Covered: July status 1, 1995 to June 30, 1996 J. P. Fitzgerald Personnel: J. P. Fitzgerald, J. Eussen, C. Hatch, J. Prather, L. Dent, D. Finley, J. Olterman, T. Beck. S. Lechman, ABSTRACT In 1368 trap nights of effort 8 new kit foxes were captured. Five of them were from a family group (1 adult female, 2 female pups, 2 male pups) captured in July 1995 in Peach Valley. An adult male was captured at Cheney Reservoir and a juvenile male pup and adult female were captured at Corcoran Point. The total number of individual kit foxes trapped since 1992 is 47'with 33 of them from the Peach Valley-Montrose East complex. A total of 17 individual kit foxes were radio-tracked during 1995-96. Nine (53%) of the 17 animals died during the study period. Four deaths were attributed to coyotes, 2 to drowning, 1 to an automobile, and 2 from undetermined causes. Two litters of pups were born in the Montrose East area in 1996 with female F23 being killed by a coyote after her pups came above ground. The fate of those p~ps is unknown. One other female living between Peach Valley and. Montrose East may have young. J. Eussen will be working on kit fox food habits. as a'thesis project in 1996.' The draft. final report of project ac.tivities was completed and reviewed by T. Beck. Revisions are being made and the final report will be delivered by August 1. . ��315 KIT FOX (VULPES MACROTIS) James P. Document the geographic Western Colorado. STATUS IN COLORADO P. Fitzgerald N. OBJECTIVE distribution and relative abundance of kit fox in SEGMENT OBJECTIVES 1. Continue Garfield to monitor counties. 2. Continue search 3. Complete draft radio-collared foxes in Montrose, Delta, Mesa and for new kit fox populations. of final project report • . METHODS Field methods for live-trapping were similar to those reported previously (Fitzgerald and Link 1993, Fitzgerald and Verbeck 1993). Limited trapping was also conducted using 1.5, soft-catch, leg-hold traps at baited dirt-hole sets. Trapping efforts were concentrated in the Gunnison and Colorado River Drainages in Mesa, Garfield, Delta, and Montrose counties and in northwestern Moffat county. Field personnel continued to monitor and document locations of radio-collared foxes using searches with a fixed wing aircraft piloted by J. Olterman, or by ground tracking. RESULTS Live Trapping AND DISCUSSION Effort From July 1, 1995 to June 30, .1996 we conducted 1368 trap nights of effort resulting in capture of 8 new foxes (Table 1.). All trapping was .done in Moffat County or in the Gunnison River-Lower Colorado River drainages in Mesa, Garfield, Delta and Montrose counties. Two foxes were captured at Corcoran point, a juvenile male and an adult female. An adult male was captured at Cheney Reservoir. In Peach Valley, 2 juvenile females, 2 juvenile males, and an adult female were captured at a whelping den (Table 2). Field crews again failed to capture any foxes in Moffat County, although scat believed to be from kit or swift foxes was discovered and a CDOW biologist (C. Braun) reported seeing a swift or kit fox crossing the road near Vermilll.on Creek. The total number of individual kit foxes trapped since the study began in 1992 is 47. Thirty-three of the captures have been made in the Peach Valley~Montrose East complex. �316 Table 1. by county Trapping effort and numbers of individual and area, July 1 1995-June 30, 1996. County/Area Trap Nights kit foxes captured New Captures Mesa-Garfield Counties Prairie Canyon to Corcoran Point Areas 347 2 Mesa-Delta Counties Cheney Reservoir to East of Delta Airport 493 1 Montrose-Delta Counties Peach Valley-Montrose East complex 368 5 Moffat County Browns Park-Vermillion Creek 160 o Table 2. frequency Location Location, sex, age class, for previously uncaptured Sex/Ear Tag ear-tag number and radio-collar kit foxes, 1995-96 fiscal year. Radio-collar Date Captured Status Corcoran Point M(J)310 F(A) 311 150.189 151.186 8/12/95 11/3/95 Unknown Dead M(A)312 151.083 12/4/95 Dead F(A)309 F(J)306 F(J)308 M(J)305 M(J)307 150.029 150.728 150.004 150.920 151.009 7/28/95 7/26/95 7/27/95 7/26/95 7/27/95 Alive Dead Dead Dead Alive Cheney Reservoir Peach Valley �317 Monitoring of Radio-collared Foxes During the fiscal year, field crews periodically located radioed animals during ground searches. Jim Olterman also made a number of flights to locate animals. A total of 17 foxes (9 males, 8 females) were located and monitored during 1995-96, including 2 male and 2 female pups marked in July 1995, three of which have died (Table 3). Two litters of pups came above ground in May 1996 in Montrose East, one litter (F23's) had 4 pups, the other litter (F21's) has an unknown number of young. F309 living between Peach Valley and Montrose East is also believed to have pups but the whelping den has not been located. Nine animals died during the year with deaths of 4 attributed to coyotes (F306, F308, F23, M312). Two died from probable drowning (M30, F311), 1 was road killed (M305) and two (M26, F309) died from undetermined causes. other Activity The first draft of the final report wae finished and reviewed by T. Beck. It is being revised and will be submitted by the end·of August pending receipt of some additional information from the Montrose Office of the BLM. The state of Utah has been contacted to see whether UNC can do some trapping in late summer along the Utah-Colorado border to assess kit fox populations in Utah that might provide migrants to Colorado stocks. J~ Eussen is collecting kit fox scat and will be doing a food habits analysis for a M.A. thesis using his materials and material collected by other field workers. By: James P. Fitzgerald, Contractor, University of Northern Colorado �318 Table 3. Kit foxes alive during status as of June 30 1996. Site & Date of Capture/ Tracking Corcoran 8/12/95 11/2/95 3/17/96 5/16/96 11/3/95 3/17/96 5/16/96 6/18/96 Sex/Age Ear Tag # JM 310 150.189 AF 311 collar at private AM 33* Peach Valley AF 5 9/28/92 4/6/94 4/14/94 8/15/95 10/11/95 moved to Alkali 12/4/95 7/26/95 2/14/96 4/9/96 year and their Location Status as of 30 June AM 30 T9S,R100W,S25 T9S,R100W,S34 T10S,R100W,S10 too weak to locate n.c. signal Delta Airport 11/19/95 8/24/94 1/10/95 4/13/95 10/11/95 2/14/96 4/9/96 Radio-collar Frequency fiscal Point Cheney Reservoir AM 312 12/4/95 moved to Wells 5/16/96 6/18/96 8/26/94 12/22/94 1/10/95 1/27/95 10/11/95 2/14/96 6/18/96 the 1995-96 T1N,R1W,S10 T10S,R100W,S10 T1N,R1E,S29 T1N,R1W,S36 Dead T3S,R2E,S24 T4S,R3E,S13 T4S,R3E,S13 Dead 151.160 T14S,R96W,S36 Unknown 150.338 150.956 150.873 T50N,R9W,S9 T50N,R9W,S9 T51N,R9W,S32 T51N,R9W,S7 T51N,R9W,S29-32 TI5S,R97W,SI Unknown T49N,R9W,S7 T49N,R9W,SI2 T49N,R9W,SI2 T51N,R9W,S29 T51N,R9W,S29-32 T51N,R9W,S30 T51N,R9W,S29 Dead 151.186 residence 151.083 Gulch Gulch 151.107 moved to PV from ME 151.209 30/151 collar in Selig Canal 151.131 JM 26 moved to PV from ME 151.056 JF 306 Unknown 150.728 T49N,R8W,S7 T49N,R8W,S7 T50N,R9W,S9 T51N,R9W,S29 T50N,R9W,SI6 T50N,R9W,SI6 Dead T50N,R9W,SI6 T50N,R9W,SI6 T50N,R9W,S16 Dead �319 Table 3 (continued) Site & Date of Capture/ Tracking Sex/Age Ear T~g # JM 305 7/26/95 12/25/95 7/27/95 2/14/96 4/9/96 Montrose 5/29/94 S/26/94 4/17 /95 2/14/96 6/1S/96 150.920 JF 30S 150.004 .moved to ME moved back to PV JM.307 7/27/95 5/16/96 6/1S/96 7/2S/95 2/14/96 . 6/1S/96 Radio-collar Frequency n. c. 151.009 AF 309 moved to ME moved between 150.029 ME and PV Location Status as of 30 June T50N,R9W,S16 T15S,R95W,S33 Dead T50N,R9W,S16 T49N,RSW,S24 T51N,R9W,S30 Dead T50N,R9W,S16 T49N,R9W,S6 T49N,R9W,S6 Alive T50N,R9W,S16 T49N,RSW,S9 T49N, RSW, S3 Dead T49N,RSW,S7 T49N,RSW,S7 T49N, RSW, S7 T49N,RSW,S7 T49N,R9W,S6 Alive East AF 21 150.947 52/52 151.2SS 150.09S 5/29/94 S/25/94 1/10/95 AF 22 5/29/94 9/15/94 7/7/95 2/14/96 6/1S/96 AF 23 6/3/94 S/25/94 4/1S/95 10/11/95 2/14/96 5/16/96 6/1S/96 AM lS4 S/27/94 12/29/94 1/4/95 10/11/95 11/19/95 12/4/95 JM 33 150.403 151.146 150.33S 151.042 n. c. 150.70S 151.237 moved 151.160 north of Delta moved to Alkali Airport Gulch T49N,RSW,S7 T49N,RSW,S7 T49N,RSW,S7 Unknown T49N,RSW,S7 T49N,RSW,S7 T49N,RSW,S5 T49N,RSW,S7 T49N,RSW,S9 Dead T49N,RSW,S7 T49N,RSW,S7 T49N,R8W,S7 T50N,R9W,S18 T50N,R9W,S16 T50N,R9W,S26 T49N,R9W,Sll-12 Alive T49N,RSW,S7 T49N,RSW,S6-7 T49N,RSW,S12 T14S,R69W,S36 T14S,R96W,S36 T15S,R97W,Sl Unknown �320 �321 Colorado Division Wildlife Research July 1996 of Wildlife Report JOB PROGRESS State of Colorado Project No. W-135-R-9 Work Plan No. lOA Mammals Research Swift Fox Studies Swift Fox Investigations Colorado Job No. Period Covered: Author: REPORT in July 1, 1995 to June 30, 1996 James P. Fitzgerald Personnel: Colorado Division of Wildlife: R. Kahn, T. Beck, B. Gill. University of Northern Colorado: J. Fitzgerald, D. Finley, B. Roell, J. Eussen, L. Dent, L. Irby, M. Schafer, B. Cross, C. Wagner, S. Lechman Abstract Sampling of distribution of swift foxes on randomly selected 3x4 mi trapping plots o~ the eastern plains of Colorado has resulted in capture of 134 foxes (68 males, 65 females and 1 of unknown sex) since March 1995. During the 199596 reporting period 30 plots were inventoried with 79 foxes (41 males, 38 females, and 1 of undetermined sex) captured on 17 of the plots. Numbers captured per plot varied from 1-14 with a mean of 4.6. Success averaged 3.4 foxes per 100 trap nights of effort. Studies continued on the biology of foxes on 2 intensive study sites in Weld County in northeastern Colorado. Since mi4 October 1994 85 foxes have been captured and radio-collared (42 males, ,43 females). TWenty-nine are dead with coyotes accounting for 72% of the mortality. In 1995 and 1996 16 litters and 41 pups were counted. Reproductive success in 1996 was higher than in 1995. Den and site characteri~tics were measured at 63 dens including 15 whelping dens. All dens were on sites dominated by blue grama communities. There was a significant difference in numbers of dens on lightly grazed vs heavily grazed areas with more dens present on the heavily grazed pastures. ' ��323 swift Fox Investigations Dr. James in Colorado P. Fitzgerald P.N. Objective Determine recommend habitats. the status and trend of swift fox populations in Colorado and conservation strategies to maintain existing populations and Segment in eastern Objectives 1. Survey sites swift fox. Colorado which are believed 2. Conduct intensive research on populations of swift to the Central Plains Experimental Range to: to be inhabited by fox on and adjacent a. Measure recruitment in swift fox populations and identify potential factors which contribute to the regulation of recruitment; b. Develop estimates of swift fox home range range size to environmental variables; c. Test the potential of automated photography systems to estimate swift fox numbers using mark/resight population estimation models; d. Collect, analyze, and report on data describing characteristics and use patterns of swift fox natal dens and den occupancy; e. Document incidents of swift fox mortality from coyote predation and evaluate the importance of coyote predation to the population biology of swift fox. Methods size and relate home and Materials The eastern plains survey began in March 1995 with selection of 72 livetrapping plots. Plots were selected using 1:100,000 scale vegetation maps of the eastern plains and N-S oriented latilong blocks dividing the state into 4 tiers. Each latilong block was gridded into 3x4 mile plots with the amount of native prairie estimated for each plot and for the entire block. Plots with 75% > prairie and 50-74% prairie were grouped by tier and plots were randomly drawn by tiers. We estimated there were 676 plots encompass,ing 8,112 mi2 with > 75% prairie and 449 plots of 5,388 mi2 with 50-74% prairie. We tried to select 44 of the 72 plots from the greater than 74% plots and 28 from the 74% or less plots. After selection of plots local Colorado Division of Wildlife (CDOW) field officers contacted local landowners, explained the project and asked for permission to trap. In most cases we were allowed access to the selected plots, however, in some instances the CDOW officer changed plot locations. As a result of such changes we sampled 10 plots (14%) with < 50% native prairie. Live-trapping consisted of placement of one Tomahawk, 12x10x32" wire-mesh live trap at each section corner of the 3x4 mi grid (20 traps per plot) providing an effective trapping area of 20 mi2• Traps were �324 usually baited with culled turkey chicks and a commercial chicken and fish lure (Erickson's On Target, ADC). Traps were run for an average of 4 days (range 3-8). Traps were checked daily starting before sunrise. Captured foxes were ear tagged (Style 893 Jiffy National Wing Bands, Newport, Ky), sexed, and condition noted before release. Crews also recorded vegetative type by section, distance to and size of any prairie dog towns, and other wildlife taken in the traps. Intensive site investigations: Swift foxes are being studied on 2 sites in northeastern Colorado in Weld County. One site the Central Plains Experimental Range, managed by the U.S.D.A. Agricultural Research Service, is approximately. 96 km2 in size. The area also serves as a Long Term Ecological Research site managed by Colorado State University under funds from the Biotic Systems and Resources Section of the National Science Foundation. NSF provided initial money for swift fox research. The second area located 13 km east of the CPER is a 160 km2 unit centered around a 20 mi2 live-trapping grid on the Pawnee National Grassland. This site was used in earlier swift fox investigations in the late 1970's and early 1980's (Loy 1981, Cameron 1984, Fitzgerald et al. 1981). Live-trapping methods used on the Gl and CPER sites follow those described for the eastern plains survey although in some cases traps were set at specific locations (Le. dens) to trap particular individuals. Generally, we grid-trapped each area in 3-8 day sessions depending on weather and trap success. Most captured fox were fitted with radio-collars (150-151 HZ, Model HLPM 2180 LD, Wildlife Materials) averaging 52g. Tracking is with hand held or truck mounted antennaes using WMI or Lotech receivers. Radio-collared animals. are monitored on a regular basis to determine: locations of dens, movements, home range, mortality, and habitat use. Two CDOW flights were made in 1995 to try to locate foxes and see if air counts of fox or coyotes were possible. Field testing of infrared sensitive camera units was started using systems and methods described by Beck (1995) for black bear population estimation. Results Eastern plains swift fox inventory: Since March 1995 we have completed 42 of the 72 plots (58%). Trapping has resulted in capture of 134 foxes (68 males, 65 female, 1 of undetermined sex). During the 1995-96 reporting period we captured a total of 79 foxes (41 males, 38 females, 1 of unknown sex) from 17 plots (range 1-14 animals/plot). In 2303 trap nights of effort this yielded an average of 3.4 foxes/100 trap nights with highest success in october and November (Table 1). The second and third nights of trapping were slightly more productive than the first, fourth, or fifth nights (Table 2). Foxes were not ca~tured on 13 plots (Table 3). Most captures were on short grass prairie. Table 1. Numbers of swift fox captured by year and month in eastern Colorado for the 1995-1996 field season. Trapping was not conducted December 1995April 1996. July 1995-June 1996 (30 plots) Aug Sep Oct Nov May Jun Total Captures 1 14 472 340 Trap Nights Catch/lOO 0.2 Traps 4.1 10 260 27 360 13 79 9 332 5 79 2303 3.8 7.5 16 2.7 1.1 Jul 460 3.4 avg. �325 Central Plains Experimental Range studies: From mid October until July 1, 1996, we have captured and radio collared 85 swift fox. Twenty nine (15 adult males and 14 adult females) were captured on site G1, 55 (27 males and 29 females) were from the CPER. ~enty-nine of the foxes (34%) are dead including 8 (27%) G1 animals (5 females, 3 males) and 21 (38%) of the CPER foxes (13 males, 8 females). Coyotes were responsible for 72% of the mortality. Three animals were killed by automobiles, 3 were shot illegally, and one died after putting fore leg through its collar. a Table 2. Fox captures by night of capture July 1995-June for the 1995-96 field season. 1996 Night 1 2 3 4 5 Totals Captures 14 25 27 12 1 79 Trap Nights 596 596 596 475 40 2303 Catch/ 100 Traps 2.3 4.2 4.5 2.5 2.5 3.4 avg We located 6 pairs of foxes on the CPER in 1995 and 7 pairs in 1996. Six pairs of foxes were on G1 in 1995 and 5 pairs in 1996. In 1995 and 1996 we have counted a total of 41 pups from 16 litters (average = 2.6). We counted 8 pups from 4 litters on the CPER in 1995 and 17 pups in 5 litters in 1996. On G1, 6 pups were counted in 4 litters in 1995 and 10 pups in 3 litters in 1996. The smaller litter size (1.7 pups) and reduced production (14 pups) in 1995 compared to 1996 (3.4 pups/litter, 27 pups total) is attributed to the wet, cold spring that may have reduced prey abundance, particularly ground nesting birds and grasshoppers, and hampered hunting success of parents. During February, April,' and May we tested 47 infrared sensing camera units including a 22 camera g~id on the CPER. We staggered cameras at 1 or 2 mile intervals on the grid. We determined that optimum distance between the camera and sensor was 2.1 m with cameras working best at a height of 45 cm from the ground. These distances yielded pictures that typically showed the entire fox. Foxes were at stations an average of 2.1 minutes. We obtained pictures of numerous foxes, as well as a few badgers, skunks, jack-rabbits, and raccoons. Free ranging cattle were a constant problem. We will schedule population density estimation bouts with the cameras during late fall and winter of ~99697 to reduce interference from cattle. We characterized percent cover, vegetation height, and den parameters at 63 dens used by swift fox in 1994 and 1995. Fourty-eight were used as daytime' refuges the other 15 were whelping and pup rearing dens. Dens are clustered in shortgrass prairie as opposed to yucca, saltbush, or mesic bottomlands. Using 1:24,000 scale maps of primary and secondary vegetation we overlaid mylar locations of 29 dens used by foxes. For all dens the primary vegetation was typed as blue grama. Secondary vegetation typed as buffalo grass (24 dens), opuntia (3 dens), rabbitbrush (1) and tumbleweed (1). Visual estimation of �326 primary vegetation at the other 32 dens showed blue grama was the dominant grass. No active dens were found in saltbush communities. Den sites averaged 56% cover of perennial grasses (range 38-81) and 19% bare ground (range 9-40). Mean height of vegetation was 12.6 cm (range 5-27). Dens had from 1-3 entrances (mean=1.4) with 65% having a single opening. Two dens in which pups were reared had 2 entrances the others a single opening. The mean width of den openings was 20.6 cm (range 12-31, SD 3.5). Mean height was 18.6 cm (range 1133, SD 2.2). Twenty-two dens (35%) were on flat terrain. Only 12 (19%) were on slopes exceeding 4 degrees. Soil disturbance at dens varies. Some dens have soil deposits around the total entrance others have oval ramps extending outward from the den along the primary trail into the opening. Many dens show little or no soil accumulation at the entrance. Compass direction of den openings varied with 48% tending to open in a southerly direction. Table 3. captures on swift fox live-trapping plots by latilong block and county. July 1,1995-June 30,1996. SGP= short-grass prairie, SSP= sand sage pra~r~e, CRP= conservation reserve, C~= cropland, RIP= riparian, PJ= pinyonjuniper woodland, MXD= mixed crop/grassland. Latilong Block Northern Tier Eaton . FtMorgan Capture· Vegetation Total Plot # County M F 23 Weld SGP 0 0 o 68 Weld SGP 1 3 108 Weld SGP 1 2 4 3 13 Morgan SGP 0 1 1 Obs o Julesburg Sterling Tier Subtotals SGPIRIP/CL 0 0 o LoganlWeld SGP 1 1 5 Sedgwick 66 67 Weld SGP 1 0 2 1 104 Logan CUSGP 0 0 o 8 plots 4 counties 4 7 11 MXDIRIP 0 0 0 Washington SGP 0 1 1 YumalKit Carson . Bonny Res. 116 Last Chance 65 Limon 61 Elbert SGP 4 2 6 95 Lincoln SGP/CL 3 0 3 SGP/CL Tier Subtotals South-central 119 Lincoln 5 plots 5 counties 3 Cheyenne 6 8 14 13 11 24 1 0 1 Tier Cheyenne Wells SGP �327 Colorado Springs Karval 59 EIPaso SGP/SSP 1 1 72A 2 EI PasolLincoln SSP 0 0 0 25 ElbertlLincoln SGP 3 2 5 26 ElbertlLincoln SGP/CRP 0 0 32 Uncoln SGP/CL 4 .6 10 89 Uncoln SGP/SSP 8 5 13 108A Bent SGPIRIP 2 1 3 27 Pueblo SGPIPJ 9 plots 6 counties 0 Table 3. cont. Las Animas Pueblo Tier Subtotals Southern 0 0 0 19 15 34 Tier Kim La Junta Springfield 2 Las Animas MGP 4 2 51 Las Animas SGPIMGP 0 0 105S Bent SGPIRIP 0 0 0 65 Baca SGP/CRP 0 0 0 75 Baca MGP/CRP 0 0 0 6 1 1 111 Baca SGP 0 .0 0 129 Baca SSP 0 0 0 19 Las Animas SGP 1 3 4 Tier Subtotals 8 plots 3 counties 5 5 1 11 Totals 30 I;!lots 13 counties 41 38 1 79 Trinidad Line transect counts of dens were made in historically heavily grazed and lightly grazed pastures (Ashbyet al 1993). There was a significantly higher (x2 =21.3, 1. d.f., p=<.OI) number of dens (150) in the heavily grazed pasture than in the lightly grazed area (80 dens). In the lightly grazed pasture we noted fewer dens (34) with multiple openings compared to the more heavily grazed unit (60). Four radio-collared foxes used the heavily grazed area in 1994-95 compared to 2 foxes using the more lightly grazed area. Discussion Roell is completing analysis of population data for his M.A. Thesis. We will generate survivorship curves and discuss mortality and natality effects on the populations of the 2 sites in more detail at that time. Our den surveys are �328 similar in findings to several other studies. Fitzgerald et al. (1981) reported 59 of 69 dens to be in blue grama communities, 8 in crested wheatgrass, and 2 in mixed buffalo-grass - three awn pasture. Uresk and Sharps (1986) reported high percent frequency of occurrance of buffalograss (67%) and blue grama (53%) at 9 natal dens in South Dakota. Kilgore (1969) and Cutter (1958) reported respectively 18 of 35 and 2 of 25 dens to be in plowed fields. We have not observed use of such areas. Kilgore (1969) found swift fox dens in shortgrass prairie to be clustered on blow ridges (40%).or on flats (60%) whereas our dens are more varied in location but rarely on steep slopes. We report more dens with single openings and fewer total entrances than others. Kilgore (1969) reported an average of 4 openings (range 1-9). Cutter (1958) reported a range of 1-7 openings. Hillman and Sharps (1978) found an average of 5.8 (range 4-7) entrances for pup rearing dens and 2.6 (range 1-7) for other dens. Fitzgerald et al. (1981) reported few openings at dens with 48 of 69 having only a single entrance. We attribute the low number of dens with multiple entrances at our study sites to habitat conditions that provide a great number of sites suitable for denning and perhaps this leads to less philopatry of individuals to return to particular dens sites year after year. Prepared by: James P. Fitzgerald University of Northern Colorado Literature Cited Ashby, M. M., R. H. Hart, and J. R. Forwood. 1993. Plant community and cattle responses to fifty years of grazing on shortgrass pra~r~e. U.S.D.A., Agricultural Research Service, Rangland Resources Research.Unit Report 1:1-14. Beck, T. D. 1. 1995. Development of black bear i~ventory techniques. Annual report, Colorado Division of Wildlife, Mammals Research, 11pp Cameron, M. W. 1894. The swift fox (Vulpes velox) on the Pawnee National Grasslands: Its food habits, population dynamics and ecology. Thesis, University of Northern Colorado, Greeley 110 pp Cutter, W. L. 1958. Denning of the swift fox in northern Texas. J. Mammalogy 39:70-74. Fitzgerald, J. P~, R. R. Loy, and M. Cameron. 1981. Status of the swift fox on the Pawnee National Grassland, Colorado. Swift fox Symposium, Central Mountain and Plains Section, The Wildlife Society, Laramie 21pp Hillman, C. N. and J. C. Sharps. 1978. Return of swift fox to Northern Great Plains. Proc. south Dakota Academy of Science. 57:154-162. LOY, R. R. 1981. An ecological investigation of the swift fox (Vulpes velox) on the Pawnee National Grasslands. Thesis, University of Northern Colorado, Greeley. 64pp Uresk, D. W. and J. C. Sharps. 1986. Denning habitat and diet of the swift fox in western south Dakota. Great Basin Naturalist. 46:249-253. �329 Colorado Division wildlife Research July 1996 of wildlife Report JOB PROGRESS State of Project Colorado Mammals No. Work Plan No. 1 Covered: Author: July N. T. Hobbs, Research Multi-Species llA Job No. Period REPORT Investigations Predicting the Impacts of Environmental Change: Simulations of Genetic and Species Diversity at Landscape and Regional·Scales 1, 1995 - June 30, 1996 J. Miller, and J. A. Wiens Abstract Loss of intact, natural habita.t is the foremost threat to wildlife diversity in Colorado and the West. Historically, the prevailing source of habitat loss in the Rocky Mountain region has been harvest of natural resources (e.g., logging, mining, and agriculture). However, changing demographic and economic trends will drive a fundamental shift in the source of environmental change affecting wildlife habitat. As a result of these trends, residential development is likely to become the predominant human influence on the diversity of Colorado'S wildlife during the coming decade and beyond. It follows that protecting wildlife habitat will depend in a pivotal way on developing wise policy on land use in the places where people live. To foster such policy, we propose to develop a System for Conservation Planning (hereafter, SCoP). The initiating idea of SCoP is that the success of habitat protection will depend on offering wise alternatives for land use, alternatives that meet human needs for economic vitality as well as the needs of wildlife for natural landscapes. Providing information needed to develop these alternatives is the primary goal of the~SCoP project: ~be goal of SCoP is to obtain, assemble, and distribute state of tbe art infor.mation on effects of land use on wildlife diversity, particularly land use associated witb residential expansion in Colorado and tbe· West. Meeting this goal requires that we enhance access to current knowledge needed to support decisions on land use while we simultaneously strive to improve that knowledge. To that end, the SCoP has initiated pilot efforts in three Colorado counties, Larimer, Summit, and Boulder. In Larimer and summit Counties, we are working with local citizens and planners to design information systems that support land use decisions with regard to preserving and protecting wildlife habitat. In Boulder County, we have initiated studies of effects of residential development on avian communities in riparian zones. ��331 Predicting the Impacts of Environmental Change: Simulations of Genetic and Species Diversity at Landscape and Regional Scales P. N. Objective 1. Develop analytical tools to support decisions on management of wildlife diversity in Colorado. These tools will include simulation models and research results that predict consequences of changes in land cover types and land use for maintaining wildlife diversity. Segment 1. Determine and community Objectives effects of human population density composition in riparian habitats. on bird species 2. As part of the SCoP project, design and construct information support conservation planning in two Colorado Counties. 3. Publish Wildlife Protection diversity systems to Handbook 4. Develop standardized methodology accuracy of Landsat Thematic Mapper to estimate data. positional and classification 5. Conduct ground-truthing surveys to estimate positional and classification accuracy of vegetation maps of Summit and Larimer counties developed from Landsat Thematic Mapper data. Results Studies were conducted in Larimer and Boulder counties to determine effects of human population density on bird species diversity and community composition in riparian habitats. Seventeen study sites were censussed at 229 points on four sample dates during May-August. Fourteen sites were sampled using 900 artificial nests. ,Data are currently being analyzed. An information system was developed and Larimer counties (Appendix A) . to support conservation planning in summit We published -Managing Landscapes for Wildlife and People: A Habitat Protection Handbook.w This book will be distributed to local governments throughout the state and will be available on a World Wide Web site. We conducted ground-truthing surveys to estimate positional and classification accuracy of vegetation maps of Summit and Larimer counties developed from Landsat Thematic Mapper data (Appendix B) . ��333 Forecasting Impacts of Land Use Change on Wildlife Habitat: Collaborative Development of an Interactive GIS for Conservation Planning N. T. Hobbs "As anyone who lives in the region can attest, rapid population growth is changing the character of much of the West. " Chromartie 1994 Loss of intact, high quality habitat represents the foremost threat to maintaining vigorous and diverse populations of wildlife in the Rocky Mountain region. Demographic and economic trends . throughout the region suggest that residential development on private land is likely to become the predominant influence on wildlife habitat during the coming decade and beyond. Such development will occur to accommodate approximately 3 million new residents in the region during the 1990's alone (Chromartie 1994). These increments in the human population equate to an annual growth rate approaching 3% per annum, a rate that exceeds the tempo of population increases in most third world nations. Increases in the West's population will force fundamental changes in land use and land cover throughout the region. For example, about 90 thousand acres of agricultural land in Colorado are annually converted to other uses, primarily to residential development and associated . infrastructure (Colorado Department of Agriculture 1995). The area included in these conversions is roughly equivalent to a swath of land, 1 mile wide, extending 140.miles long=the distance from Fort Collins to Colorado Springs. Before it was developed, more than 60% of this agricultural land was native grassland that provided important wildlife habitat. Many of the high mountain valleys of the West are growing at a particularly rapid pace, primarily as a result of immigration. Immigrants are increasingly attracted to the valleys ofthe West because these areas offer environmental amenities found nowhere else. Of course, these' same valleys provide critical habitat for many species of wildlife. Careless development of these areas will undoubtedly cause fundamental, long term deterioration in the abundance and distribution of many species. Regional changes in land use are the result of many local decisions made one at a time---a ranch is converted to a subdivision; a mountainside is developed for skiing; a valley is dotted with vacation homes. Because these decisions are inherently local, the regional effects of growth on wildlife habitat are simply the collective outcome of many decisions made one at a time. This is problematic because there is often an unstated assumption in planning for such growth; we have assumed habitat lost in one place can be compensated byhabitat remaining undisturbed elsewhere. However, it is becoming clear thatthis assumption cannot holdforever=we have realized that �334 many small, seemingly benign impacts can accumulate to cause large, harmful effects on environmental values like wildlife habitat. As a society, we are challenged to anticipate and manage those effects. We have a timely opportunity to meet this challenge. In many counties in the West, geographic information systems (GIS) are being developed by counties to guide land-use planning. Such planning focuses on infrastructure (roads, water lines, schools etc.) as well as environmental values (open space, wildlife habitat). At.the same time, state and federal agencies are assembling spatial data on vegetation, hydrology, and current distributions of many wildlife species. As a result of these efforts, there is a large amount of data accumulating in county and agency offices throughout the West, data that could help citizens and decision makers foresee the way that small changes in land use will add up to cause large changes in the region's environmental values. However, although there is great potential in using such data to guide insightful land use planning and habitat protection efforts, the tools needed to provide such insights have not yet emerged. To provide such tools, we initiated a project in Colorado called a System for Conservation Planning, hereafter, ScoP (pronounced "scope'). The goal of the ScoP project is to support planning by local communities by providing them with readily accessible information on the consequences of development for wildlife. To meet that goal, we ptoposed to produce interactive· geographic information systems that would allow planners, decisions makers, and citizens to foresee how changes in land use land are likely to accumulate over time and space, and how these cumulative changes might affect the extent and distribution of habitat for wildlife. Our proposal was funded by the Great Outdoors Colorado Trust Fund during July of 1995 to support a pilot project in Summit County, Colorado. . Study Area Summit County extends across a high mountain valley approximately 100 km west of Denver. The county is one of the few in Colorado that embraces a complete watershed= the Blue River bisects the county along its north-south axis, extending its headwaters in the Ten Mile Range to a confluence with the Colorado River. The current population numbers about 1500 year-round residents, but the county is growing at greater than 2% per year and if this rate is sustained, will number 25-30,000 by the year 2020. The economy is based primarily on tourism attracted to several ski areas. As a result, the total population often exceeds the resident population by threefold. In addition, there are 1.3 dwelling units for each permanent resident. The county is predominantly public land, about 80% of the total area of the county is managed by the US Forest Service. Private land is concentrated in the valley bottoms, while the surrounding uplands and mountains are public: �335 3 System Development The Process of Collaborative Design From the outset, we were committed to designing a system reflecting the needs of users. In particular, we wanted to avoid developing an analytical tool influenced by the viewpoint of scientists alone. Instead, we believed that the utility of the system that we produced would . depend on the extent to which we could integrate a scientific view with the diverse views of information users, particularly those users who have a stake in the decisions that the information system would support. To assure that information system would meet the needs of a diverse set of clients, we assembled a team of users and experts to help us design the information system (Fig. 1). The users included a county commissioner, a planner, a developer, a land owner, environmental advocates, and a wildlife manager from the Colorado Division of Wildlife. The expert group consisted of a .GIS analyst, a land use attorney, an ecologist, a geographer, and a . scientific programmer. Collaborative Design environmental advocate. Users Iclentlfc I Experts I /! ~ programmer geographer ecologist Figure designed experts. attorney 1.. The ScoP information by a collaboration system was between users and We initiated the design process with a series of l-day "primers" on subject matter relevant to land use change and habitat conservation. Topics of these sessions . included principles of conservation biology and landscape ecology, land use law and planning, and human geography. The purpose of the primers was to provide a common understanding of some of the technical issues relevant to the information system. Following these primers, the team met to set goals for the system. We did this by asking the users several questions, including the following: • Imagine a situation (real or hypothetical) that is typical of your role in working with wildlife and development in Summit County. What insights about impacts of development on wildlife would help you play your role more effectively? • Describe a situation when you wanted to include wildlife concerns in land use planning or . in your work in development, but were unable to do so effectively. Be as specific as possible and try to choose an example that differs from the first one you discussed. �336 4 • The goal of the SCoP project is "To support land use planning by local communities by providing state-of-the-art information on the impacts of development on wildlife and by identifying actions that can be taken to minimize those impacts." What do you consider to be some important, tangible measures of success in meeting that goal? • Imagine yourself 10 years from now in Summit County. What specific things would you look for to indicate that Summit County was succeeding in meeting its goals for wildlife protection? The comments that emerged in response to these questions were drawn together into statements of goals for the information system. These goals focused on four general areas-education, screening, habitat value, and cumulative impacts. Goals for Education The users' groups felt strongly that one of the things that SCoP should do is to educate the citizens of Summit County about their wildlife. The information system should allow citizens to learn about the distribution of wildlife species and their habitats; it should inform about species status, their life histories, habitat requirements and sensitivities to impacts. The user should be able to locate an area on a map and find out what species have habitat in that area. He or she should also be able to identify a species and learn about the distribution of its habitat. Goals for Screening The developer and a citizen advocate on the design team asked that the system be able to offer a "coarse screening of impacts" of future development at a given site. "The developer wanted to be able to identify an area on a map that he might develop and learn about the potential concerns that would be raised about impacts on wildlife habitat at that site. This was important because the developer was willing to do his best to avoid sensitive areas, but he needed to know about such sensitivities ''up front", that is, before he had invested a great deal of time and money in preparing a formal development proposal for review by the county. The citizen advocate.asked for virtually the same function. He wanted to allocate his "advocacy time" in commenting on the develoment that had the greatest potential to impact wildlife. So, he wanted a tool to "screen" areas in urgent need of comment from those that were less urgent. He also wanted a sound, scientific source of information to use in formulating his comments. �337 5 Goals for Habitat Value All of the members of the design team felt that SCoP should map areas of the landscape according to their relative value as habitat-- areas that offer high value habitat as well as areas that are of lower value. Such a map would be useful in identifying alternative paths for development during master planning by offering a rationale for steering development away from valuable areas toward areas of lesser value. Goals for Cumulative Impacts The design team was unanimous in identifying the problem of "cumulative impacts" as a fundamental impediment to wise land use planning. It is clear that impacts of development . accummulate over time and space, and that the sum of these many impacts a the problem that must be overcome if planning is to be effective in conserving wildlife habitat. The impacts of a single development on the abundance and distribution of wilidfe is arguable, but the impacts of 100 developments are not. So, the design team asked that we work on a approach to forseeing how future development will spread in relation to exisitng widlife habitats. They asked that the system help them evaluate different approaches for managing population density (e.g., clustering, zoning, transfer of development rights, etc.). Implementation of System Available Data We built the system to make use of currently available data. Distribution maps were available for 30 species of vertebrates (mostly game mammals and fish) in Summit County. These maps show areas of the county known to be routinely used by the species. In addition, we developed habitat maps Habitat Maps for all of the county's 239 vertebrate species. Potentially suitable habitat for each species was 1 _\ .delineated (Figure 2) by identifying all 30 x 30 cells Vegetation 1\ in a raster map that met 3 criteria: Potentially Elevation 1) The cell must contain vegetative cover appropriate for the species. Distance to Water { ~ L\ Suitable Habitat } I" ) 2) It must be at an elevation within the upper and lower elevation limits for the species. 3) It must sufficiently close to water for· speices that require lakes or streams as part of their habitat. Figure 2. Habitat for each species was defined by its affinity for vegetative cover, its elevation range, and requirements for proximity to lakes and strearns. . �338 6 Vegetaton was mapped by supervised classification of a thematic mapper satellite image. The classification resulted in 14 vegetative cover types (Table 1). These were stored as an ARCINFO Grid coverage, with a 30x30 meter cell size. Species affinities for each type were derived from literature sources as well as by consultation with local experts. Table 1. Definition of land cover classes used to map Summit County, Colorado. Class Water Definition includes all types of open water: lakes, ponds and. reservoirs. Conifer (denser) medium to high density stand of single and mixed species, including lodgepole pine (Pinus contona), Engelmann spruce (Picea engelmannii), blue spruce (Picea pungens), sub-alpine fir (Abies lasiocarpa) and Douglas fir (Pseudotsuga menziesiii. Conifer (sparser) sparse to medium density stand of the above mentioned species. Sub-dorninant cover types range from bare soil and rock to willow, grass-forbs and, to a lesser degree aspen (Populus tremuloides). This class also includes a small amount of water edge. Conifer-Aspen stands of aspen (Populus tremuloides) mixed with various conifer species. Aspen primarily aspen (Populus tremuloides), but also includes cottonwood (Populus spp.). Irrigated primarily irrigated native meadows and hay fields. Willow primarily willows. Aspen- Irrigated-Willow all three cover types are represented. Grass-Forbs dominated by grasses and/or forbs; may also include sparse shrubs. Includes alpine meadows as well as lower elevation grasslands. Sage dominated by big sage (Anemisia tridentata). Sub-dominantcover types include grasses, bare soil and other shrub species (e.g., gray sage, bush cinquefoil, rabbit brush, and saltbrush). Mixed Shrub a mixed, general class consisting primarily of willows, sage, and limited amounts of serviceberry and chokecherry. Soil-dominated dominated by bare soil, often with sparse vegetation. Also includes areas of human development and a significant portion of Interstate 70. Rock includes talus slopes, mine tailings, dam faces and bare soil (including dry tailings ponds) Snow includes permanent and temporary snowfields. �339 7 Analysis Procedures: Habitat Value We developed 5 sepearate indices of habitat value (Table 2). The LOcal Diversity index showed the number of species that have potentially suitable habitat at each grid cell. User Defined Local Diversity was similar, except that the user could specify which species (ofthe 239 possible) would be included in the diversity calculation. Neighborhood Diversity was calcuated by centering a circular window (1 km radius) on each grid cell, determining the number of species that had potentially suitable habitat within the circle, and assigning that number to the grid cell. Table 2. Indices used to assess habitat value. Index Objective Local Diversity Indentify areas of the county that offer high levels of species diversity . resulting from overlap of habitats of individual species, Neighborhood Diversity Indentify areas of the county that, offer high levels of species diversity as a result of juxtaposition of vegetation types. User Defined Local Diversity Indentify areas of the county that offer high levels of species diversity as a reult of overlap of habitats of several individual species chosen by the user. Corridors Identifies areas of the county that facilitate movement of species among core areas of habitat. Patch Value Indentifies large intact patches of vegetation and weights them by relative rarity of species' habitat. . The Corridor index was calculated by first identifying groups of species using similar vegetation types through cluster analysis. The 5 largest, intact patches of habitat for each cluster were then delineated. We then calculated an impedence map assuming that resistance to movement was least among cells that included habitat for the cluster of species. We then summed the impedence map across the clusters of speices and normalized it to range between 0 and 1. High impedence was taken to mean low corridor value. The Patch Value index was calculated for all patches of vegetation containing ~ 50 continuous cells. For each patch we calculated the statistic V, s V = L Pi i = 1 where V is a realtive index of patch value, S is the number of species that have potentially suitable habitat within the patch, and P, is the proportion of the total habiat for the ith species within the county that is contributed by the patch. The value of V was then mapped using a relative color �340 8 scale. Large intact patches of vegetation offering habitat for specialist species receive high scores for patch value. Analysis Procedures: Simulating Land Use Change We developed a stochastic model forecasting the distribution of building units across the county. The model is based on historical data (from the tax assessor) on population density in each 1/4 section of the county. We analyzed these data using logistic regression to esimate a probablity that an undeveloped cell (i.e., population density = 0) will develop during any given year as a function of distance to existing development and to roads. We then fit exponential curves to describe the rate of growth of developing cells. The predictions of the model can be adjusted by changing assumptions on population growth rates and spatial restrictions (e.g., zoning) limiting the distribution of building units. The model assumes that each building is the center of acicular disturbance zone extending outward 50-500 m (Figure 4). The area of the disturbance zone is under the ccontrol of the user. It is assumed that the value of habitat is substantially reduced within the disturbance zone. Cumulative impacts are estimated by summing the total area contained within Disturbance Zone disturbance zones (Figure 5). • Changa.ln Vegatatlon Interface • Pradatlon The system runs a set of ARC-INFO AML's. A user interface provides point and click access to 5 main fucntions, Help, Education, Screening, Habitat Value, and Simulation (Figure 5). • The Help function provides context-sensitive access to an online manual desccribing how to use the sysytem. • Avoldanca of road. • Human/wlldllf. conflict. Figure 3. Simulations of cumulative impacts relies on an assumed radius of disturbance extending from each building. Simulating Figure 4. Impacts of Growth Cumulative impacts are simulated totaling the non-overlapping area included in disturbance. zones. by �341 9 • The Education function allows the user to obtain a variety of spatially referenced information on wildlife species. The user can locate himlherself on a map, specify an area of interest, and bring up a list of all of the speices that have potentially suitable habitat within the specified.area. After choosing an individual specices, the user can view habitat and range maps for the species and obtain a text description of its life history, sensitivities to managment practices, and habitat requirements (e.g., Figure 6) Education Function Screening Function Habitat Value Function ·Simulatlon Function Figure 5. • sees main menu. The Screening fucntion allows users to get information on potential impacts of development at a particular site. The user can identify an area to be developed using two .--.-1lJ"~ ••• Figure 6. species Data can be obtained for in the area chosen by the user. each approaches. Developments currently under review are show as points on a map of the county and the user can select an area, clicking on a point, the user can find out about impacts of the proposed development. Alternatively, the user can specifiy an area to be developed in the future by drawing a polygon on a map. In either case, a list of concerns about impacts on wildlife can be accessed after the development area has been identified (Figure 7). �342 10 These concerns include: •.. Habitats for listed species. The system reports all species of speical status that Figure 7. After identifying a development area, the user can view concerns on wildlife habitat. about impacts have potentially suitable habitat within the development area. Species of special status include those listed as threatened or endangered by the state or federal government, species that are listed as imperiled or crtically imperiled by the Colorado Natural Heritage Program, and indicator species listed as sensitive by the US Forest Service .. Once this list has been produced, the user can can view habitat and range maps for the species and obtain text description of its life history, sensitivities to managment practices, and habitat reqirements. •.. Impacts for listed species. By selecting this menu item, the user can find out if the proposed development will affect ~ 10% of the habitat for a listed species within the analysis area (i.e., county, planning basin, etc). Ranges of WRIS speties. Distribution data are available for WRIS species. If the development area overlaps ranges of these species, the system will show the overlap, and will describe patterns of seasonal use by the species. Heritage Program sites. If the development area overlaps areas that have been designated as conservation sites by the Colorado Natural Heritage Program inventories, then the system will list those sites and describe them Speties diversity. If the development area contains habitat for an unusually large number of species (relative to areas of similar size throughtout the county) the· system reports this 'finding. �343 11 • ". Rare vegetation types. If the development area overlaps rare vegetation types (those contributing < 15% of the total vegetative cover of the county), then the system shows the distribution of those types within the development area and reports statistics on the impact of the development on the amount of the type remaining. ". Edge sensitive species. The system reports changes in the realtionship between edge and interior of forested patches caused by the development. . The Habitat Value function shows a variety of maps shaded to reveal differences in habitat value indices described above (i.e., Table 1). These can be presented as panels, sequentially (i.e., like a slide show), or as an RGB color composite (Figure 8). Figure 8. The user can choose to map several indicators of habitat value. • The Simulation function allows users to examine potential effects of different land use control actions on cumulative impacts of development on wildlife habitat (Figure ). Such actions include incentives for clustering, conservation purchases and easements, phased development plans, site-level mitigation, transfer of development rights, and changes in zoning .. �344 12 Access to System The system resides on aSun Microsystems Spare 5 server at the Natural Resource Ecology Laboratory of Colorado State University. Citizens, planners, and decision makers will be able to access the information system in a couple of ways. We have provided a terminal available to the public in the Summit County government offices. This terminal is networked to the server via a 56K frame relay. We will also offer direct connections via telnet over the Internet. Such connections will requirea Unix terminal running X, or a PC with X emulation software. Evaluation of System The ultimate goal of the SCoP project is to help citizens and their governments make choices about land use that achieve a reasonable balance between the needs of wildlife for intact landscapes and the needs of people for economic vitality. A short term objective is to deliver information systems that users believe will contribute towards our ultimate goal. This short term objective is currently being evaluated in Summit County. A beta version is now being tested by the Design Team and other interested citizens. The system will be modified based on their suggestions. Many of the notable successes of bringing science to bear on environmental policy have involved a top down model where scientific information flows upward in a government hierarchy and regulations and policy flow downward through that hierarchy. This model does not serve in the case of land use change, because the political authority for decisions on land use reside at the local level. As a result, scientists are challenged to bring their knowledge to many local decisions, choices that are inherently diffuse in time and space. The System for Conservation .Planning project is working to meet that challenge. �345 1 APPENDIX CLASSIFICATION AND POSITIONAL A ACCURACY ASSESSMENT MAPPER DATA OF LANDSAT THEMATIC (TN) Tanya M. Shenk ABSTRACT A vegetation map of Summit County, Colorado was developed from Landsat Thematic Mapper (TM) imagery. This vegetation map will not only be used to make inferences about the abundance and distribution of each vegetation type within the county, but will also provide a foundational database for inferences about wildlife habitat. Therefore, it is critical to provide a rigorous, quantitative assessment of the accuracy of the map. A study plan was completed and preliminary work begun to provide the data necessary for such an accuracy assessment. Methodology was designed to involve voluntary citizen participation in ground-truthing of the vegetation map. Participation of citizen volunteers was incorporated to provide both cost-effectiveness and increased awareness and citizen participation in management of and research conducted for Colorado's wildlife. A pool of 75 randomly selected pixels (30m x 30m) for each of 13 vegetation classifications was generated. To date, 334 of these pixels have been· surveyed to determine dominant cover. A preliminary error matrix was constructed from these 334 ground-truthing surveys. Two measures of accuracy were determined from this error matrix: overall accuracy (60%) and errors of omission. Errors of omission suggest good accuracy (> 60%) for the vegetation classifications water, soildominated, and rock. Errors of omission performed most poorly for the vegetation classifications willow (0%) and grass-forb (23%). The greatest confusion occurred between irrigated and grass-forb, willow and irrigated, and a confusion of mixed shrub, grass-forb, and sage. Field work continues to complete vegetation surveys on the remaining 641 selected pixels. INTRODUCTION Remote sensing, image processing, and Geographic Information Systems (GIS) are increasingly being used to develop spatial databases (e.g., Scott et al. 1993, Rutchey and Vilcheck 1994). These databases, in turn, are used to support decisions on resource management and regulatory issues. The goal qf the CDOW's System for Conservation Planning (scoP) project (Hobbs et al. 1994) is to obtain, assemble, and distribute state of the art information on effects of land use on wildlife diversity. Meeting this goal will require developing an information system using geographic data, including a vegetation map constructed from Landsat Thematic Mapper (TM) imagery. This vegetation map will be of fundamental importance to the information system. Although there has been a tendency to treat such mapped-based data as errorfree (Bailey 1988, Monmonier 1991) it is now widely recognized that errors can be substantial (Cherrill and. McClean 1995, .Briggs 1995). Therefore, an accuracy assessment of the vegetation map is necessary to quantify confidence levels for any GIS data analyses conducted for the information system. �346 2 Reliable estimates of map accuracy will require appropriate statistical sampling (Card 1982) and data analysis (Kalkhan 1994). Therefore, we propose to (1) develop Qtandardized methodology to estimate positional and classification accur.acy of vegetation maps to be used in the SCoP information system and (2) conduct a field study using this standardized methodology to estimate accuracy of the vegetation map of Summit County, Colorado. Sources of error Error is inherent when constructing an information system based on GIS because of the techniques used to obtain and interpret the data. Remote sensing is the process of deriving information by means of systems that are not in direct contact with the objects or phenomena of interest (e.g., satellite sensors such as Landsat). Image processing refers· to the manipulation of the raw data produced by remote sensing systems to provide input data for measurement, mapping, monitoring, and modeling within a GIS context. It is clear that error is introduced, and thus accumulates, throughout this hierarchy of data acquisition, processing, and analysis. More specifically, three sources of error predominate (Burroughs 1986). The fiJi'stsource of error is related to basic features of the input data including: age of data, areai coverage (partial or complete), map scale, density of observations, relevance, format, and accessibility. Once the data have been collected however, scientists have tittle control over these sources of error although they must be kept in mind when inferences are drawn from any GIS results. The second major source of error results from natural variations or from original measurements including variation in data resulting from data entry or output faults, observer bias, natural variation, positional accuracy, and accuracy of classification. Assuming incorrect data handling has been controlled for and realizing natural variation is inherent, we will focus our effor~s on quantifying and providing information to improve positional and classification error. Positional and classification errors arise because of the methods used to interpret Landsat TM data. Thus, the last major source of error in GIS arises through processing of input data. Processing errors include numerical errors. in the computer, misuse of logic, problems associated with map overlay, and finally, classification and generalization problems including methodology, class interval definition, and interpolation. Computer limitations, logical errors, and map overlay problems can only be improved through verification of computer programming codes. Classification and generalization problems leading to error can, however, be improved through classification methods that incorporate information obtained from ground-truthing. Landsat TM uses passive sensors that measure the intensity of reflected radiation in seven spectral bands. From the digital satellite imagery, computer assisted classification is based on developing distinct spectral signatures that can be associated with specific land features. The spectral signature of an object is a repeatable set of reflected energy levels at specific wavelengths. The infrared radiation emanating from a scene is due to both self-emission from each object within the scene and reflected radiation· from the object as a result of illumination from natural sources such as the �347 3 sun, clouds, moon, and stars (Star and Estes 1990). Temperature and surface characteristics are the primary factors that govern the emission of infrared radiation. That is, the radiation emitted by a body at a given temperature is proportional to the characteristics of its surface (Star and Estes 1990). Thus, a rough surface emits more than a polished surface and dark-colored objects are usually better emitters than light-colored objects. ™ One of the primary methods used for classifying Landsat imagery is an unsupervised approach where the computer uses an algorithm to define groupings or clusters of pixels based on their associated spectral signatures (Croteau 1994). These. clusters are referred to as spectral classes and assigned a meaningful label (e.g., conifer) by the analyst working with site-specific information. The number of spectral classes is determined by the analyst. Alternatively, a supervised approach to classification can be used whereby the analyst develops multivariate descriptions of the different classes of interest (star and Estes 1990). The GIS is then used to assign all the regions in the data set to one of the class descriptions. Decision rules are established to make assignments between the different classes when there is no exact match between the available categories and the characteristics of a given pixel (Star and Estes 1990). Errors in data interpretation occur for both supervised and unsupervised classification methods. However, both unsupervised and supervised classification algorithms can be improved by incorporating information obtained from ground-truthing. Spatial resolution of TM data is approximately 30m2 (referred to as a pixel). A thematic map has all map pixels categorized into one and only one of a discrete number of categories (Green et al. 1993). Therefore, image classification methods, including minimum-distance-to-the-mean, maximumlikelihood (Jensen 1986, Mather 1987), and fuzzy classifiers (Fisher and Pathirana 1990, Wang 1990). are used to .assess the degree to which a pixel belongs to all cover types and then assigns a classification to each pixei. A thematic vegetation map is then constructed by assigning each pixel a vegetation classification based on interpretation of the reflectance data. Each classification methodology has associated errors based on their algorithms for interpreting the data. However, data collected to quantify current classification error could be used to further research on improving classification algorithms and thus, decrease classification error. ™ Accuracy assessment Ground-truthing of a vegetation map as large as that of Summit County will be time consuming and labor intensive. However, the resulting accuracy assessment will: (1) provide the necessary data to determine confidence in inferences drawn from further data analyses, (2) enhance the database through reclassification of misclassified pixels, and (3) provide data that may result in a better vegetation classification algorithm. In addition, the SCoP project was also designed to incorporate citizen input at all levels of the project including design, implementation, and enhancement. Ground-truthing techniques are repetitive, and can be conducted by persons minimally trained in sampling techniques, vegetation identification, and positional location. Therefore, we see this as an excellent opportunity for citizen involvement. We will model the citizen involvement program after CDOW's successful Rivers of Colorado Water Watch �348 4 Network (ROCWWN) program. This continued citizen involvement throughout should foster interest in and a sense of stewardship toward the natural resources of Summit county. Thus, citizen volunteers will be enlisted, trained, and organized to help conduct the accuracy assessment of the vegetation map. SCoP Specific objectives of this study include: (1) Ground-truth the Landsat Thematic Mapper (TM) vegetation map developed for Summit County, Colorado; (2) develop methodology to involve voluntary citizen participation in groundtruthing of the Summit County Landsat TM vegetation map; (3) construct an error matrix from the results of the ground-truthing surveys. Three measures of accuracy will be determined from this error matrix: overall accuracy, errors of omission, and errors of commission. Evaluation of the errors of omission and commission may provide information to define algorithms to better assign classifications to a given pixel and thus reduce classification error. The overall accuracy will be tested with pielou's index of segregation; (4) improve accuracy of the Summit county vegetation map database ·through reclassification of misclassified pixels1 and (5) develop standardized guidelines on the design and methods to access accuracy of Landsat TM imagery statewide. STUDY AREA Summit County, Colorado approximately 178,000 hectares is comprised primarily of the Blue River watershed bounded on the east by the Williams Fork Mountains and on the west by the Gore Range (Curdts 1994). The Blue River flows north from the Continental Divide near Hoosier Pass, and ends at its confluence with the Colorado River near the town of Kremmling. Summit County ranges in elevation from 2256 to 4350 meters above mean sea level. There is a substantial amount of alpine area above 3300m. Coniferous forests dominate the subalpine areas and consist primarily of lodgepole pine (Pinus con~or~a), Engelmann's and blue spruce (Picea engelmannii and P. Pungens), sub-alpine fir (Abies lasiocarpa), with smeHl amounts of Douglas fir (Pseudo~suga menziesii). Aspen (Populus ~remuloides) stands are found primarily in middle elevations. The lower elevations of the lower Blue River are. characterized by sagebrush (Ar~emisia spp) and pastured rangeland (CUrdts 1994). Landsat TM data were recorded on 5 July, 1989, are cloud-free, and cover TM scenes 3432 and 3433 (Curdts 1994). Rectangular subsets were created that contained the study area portions of each scene. Computer processing of the data was accomplished through the use ERDAS (1994) image processing software package on a SUN-IPX workstation (CUrdts 1994). The two subsets were merged (ERDAS:STITCH) into a single file, then geometrically rectified, using the cubic convolution algorithm. The hydo-unit boundary file was then used to "cookie-cut" (ERDAS:CUTTER) the data contained within the hydro-unit. ™ ™ METHODS Objective developed 1 - Ground-truth the Landsat Thematic for Summit County, Colorado. Mapper (TM) vegetation map �5 Two basic types of error are common in land cover maps generated from. Landsat TM.data. First, locational error, and second, the assignment of an incorrect vegetation classification to a given pixel (classification error). Data from ground-truthing sample surveys will reflect both sources of error as described below. A. Locational error Two sources of locational error are inherent in any Landsat TM imagery groundtruthing study. First, the vegetation classification map itself must be georectified to Universal Transverse Mercator (UTM) reference points. The process of geo-rectification is not error-free and results in positional error. The second source of inherent locational error occurs when locating a given pair of UTM coordinates taken from th~ geo-rectified vegetation map on the ground. 1. Positional accuracy Positional accuracy of remote sensing data refers to the accuracy of a geometrically rectified image. Rectification includes registration to a reference coordinate system together with a resampling procedure (Irish 1990). Positional accuracy of a geometrically rectified map may be reported as the root-mean-square error (RMSE) that is derived from the cubic convolution algorithm. The RMSE reflects the proportion or number of pixels that the image control points differ from the map or reference control points (Lunetta et ale 1991). Thus, the RMSE does not truly reflect the positional accuracy of all pixels within an image; the RMSE only addresses the control points and only with respect to the map (Lunetta et ale 1991). To geo-rectify the Landsat TM image for Summit county, 78 visible reference points were selected on the vegetation map throughout the county. Universal Transverse Mercator (UTM) coordinates for these 78 reference points were then obtained from 1:50,000 scale topographic maps. Of those 78 reference points, 54 had a RMSE ~ 1 pixel (Amy Cade, CDOW, personal communication). The .Landsat TM image was then geometrically rectified with these 54 points using the cubic convolution algorithm from the Earth Resource Data Analysis (ERDAS 1994) computer software program (Curdts 1994). The geo-rectification of the Landsat TM image has positional error (i.e., UTM coordinates from the ™ image will not identically match the UTM coordinates on either the topographic maps or on the ground). Therefore, when locating a single pixel to ground-truth the vegetation classification this positional error may cause us to determine a matched or mis-matched classification based on positional error and not on classification error. For example, such a situation could arise if we had 2 adjacent pixels (1 and 2) correctly classified into different vegetation classes (Pixel 1 = A classification, Pixel 2 = B classification). We wanted to ground-truth Pixel 1 but due to positional error we were actually ground-truthing pixel 2. We would assign a mia-classification to Pixel 1 because we would conclude it was really classification B and not the assigned A. Therefore, because positional error ia inherent in any Landsat TM image ground-truthing study, all inferences made from the accuracy assessment will refer to both positional and classification error. �6 2. Ground loca~ions Numerous techniques can be used to locate a given set of UTM coordinates on the ground. These techniques include topographic maps used with .a compass and altimeter, LORAN-C, or more recently, GPS. Each technique has its own sources of error. A GPS, however, provides the most accurate technique available for determining location. GPS measurements include error in the satellite clock, receiver error, geometric dilution of precision (GDOP) from angles of the satellites used to make the position measurement, and atmospheric and ionospheric conditions that slow the speed of light causing error. in interpretation of satellite location (Hurn 1989). These errors result in measurement accuracies of 18-60m. In addition, 'selective availability' or purposively degraded accuracy by the Department of Defense can result in measurement errors of 107m (Hurn 1989). The vegetation classifications for the summit County map have been made on 30x30m pixels. Therefore, this level of accuracy is unacceptable to locate individual pixels on the ground. Use of differential GPS, however, can provide measurement accuracies to within I-Sm. Differential GPS relies on information from a base receiver at a known location to correct for the above mentioned sources of error. Any errors in the satellite data received by the base are corrected and transmitted to any roving GPS receivers in the local area (up to 186km from the base receiver) to correct their position solutions. The base receiver at the United states Forest Service (USFS) in Fort Collins, Colorado will be used to provide satellite error corrections and provide ground accuracies of 1-Sm with a roving GPS. B. Classifica~ion error Accuracy testing of vegetation classification assignment (see Table 1 for vegetation classification types for Summit County) will'be accomplished by comparing the classification of randomly selected pixels on the map with reference data obtained from aerial photographs or ground checks for those same pixels. 1. Sampling design The method used in selecting a representative sample of ground reference data or remotely sensed data is considered an important part of any accuracy assessment (Card 1982). The selection of an appropriate sampling scheme is important in estimating the structure of the error matrix. Sampling techniques commonly used to assess the accuracy of a classification procedure include simple random sampling, systematic sampling, stratified random sampling, and cluster sampling (Congalton 1991) • .Simple random sampling is the most reliable sampling method to ensure unbiased estimates of an accuracy index and its associated variance (Congalton 1988, Kalkhan 1994). Simple random sampling has the advantage over other designs in that it is easy to apply (theoretically) and provides satisfactory results in evaluating the accuracy assessment of remotely sensed data. The disadvantage of simple random sampling is that the sample size within each thematic class is proportional to its area (Congalton 1991). Alternatively, stratified random sampling has the advantage over simple random sampling and systematic �351 7 1. Definition of land cover classes u.sedto map Summit County, Colorado (from Curdts 1994). Table Class Water Definition includes all types of open water: lakes, ponds and reservoirs. Conifer (denser) medium to high density stand of single and mixed species, including lodgepole pine (Pinus contorta), Engelmann spruce (Picea engelmannii), blue spruce (Picea ptingens), sub-alpine fir (Abies lasiocarpa) and Douglas fir (Pseudotsuga menziesii). Conifer (sparser)' sparse to medium density stand of the above mentioned species. Sub-dominant cover types range from bare soil and rock to willow, grass-forbs and, to a lesser degree aspen (Populus tremuloides). This class also includes a small amount of water edge. Conifer';"Aspen stands of aspen (Populus tremuloides) various conifer species~ mixed with Aspen primarily aspen (Populus tremuloides), includes cottonwood (Populus spp.). but also Irrigated primarily irrigated native meadows and hay fields. Willow primarily willows. Aspen-Irrigated-Willow all three cover types are represented. Grass-Forbs dominated by grasses and/or forbs; may also include sparse shrubs. Includes alpine meadows as well as lower elevation grasslands. Sage dominated by big sage (Artemisia tridentata). Subdominant cover types 'include grasses, bare soil and other.shrub species (e.g., gray sage, bush cinquefoil, rabbit brush, and saltbrush). Mixed Shrub a mixed, general class consisting primarily of willows, sage, and limited amounts of serviceberry and chokecherry. Soil-dominated dominated by bare soil, often with sparse vegetation. Also includes areas of human development and a significant portion of Interstate 70. Rock includes talus slopes, mine tailings, dam faces and bare soil (including dry tailings ponds) Sno!e' inCludeS permanent and temporary snowfields �352 8 sampling in that a small number of samples are selected from each category to assure reliable estimates. Therefore, because these ground-truthing data may also be .used to further improve the current supervised classification method (see Objective 4) stratified random sampling will be implemented to insure adequate coverage over all classifications. Thus, separate simple random samples .of pixels will be drawn from each vegetation classification on the map. UTM coordinates identifying the pixel (center and corner points) will be recorded for field location. Although elevation and slope may affect classification accuracy, 'not every vegetation classification occurs over a wide range of elevations or slopes (e.g., 'sage). Therefore, e1evationa1 and slope strata will not be designated. 2. Sample size To achieve a 95% confidence level, van Genderen and Lock (1977) recommend a sample size of 60 sampling units per strata, Hay (1979) recommends a minimum sample size of 50 or 100 per strata to insure valid error estimates, and Conga1ton (1988) recommends a 1% sample of the image for estimation of accuracy and its variance. We will ground-truth 75 randomly selected single pixels per 13 c1assific'ation strata (the snow classification will be eliminated from Table 1) as well as recording general vegetation information on the cluster of 8 pixels surrounding the randomly selected pixel. When field locations are differentially corrected we may find we did not capture the randomly selected pixe1'within the cluster of pixels ground-truthed. This may cause insufficient sample size for a given classification strata (more probable for the rarer classifications). If the sample size falls below 60 randomly selected pixels for a given strata we will continue the process of selecting random pixels within that strata and ground-truthing on a cluster of pixels around that randomly selected pixel until a minimum of 60 randomly selected pixels per strata are ground-truthed. Selection of random pixels by strata will be done using the IMAGINE option in ERDAS (1994). A total of 100 random pixels per strata will be identified. The first 75 will be the target survey sites. However, if those sites are completely inaccessible or have recently been converted to a new class (e.g., agriculture to housing development), new random sites will be taken from the· extra 15 randomly selected pixels. 3. Field loca~ion of randomly selec~ed pixels Ground-truthing teams will locate the UTM coordinates using first topographic maps and aerial photos to identify the general location of the sample site and then specific location of the UTM coordinates will be made with a GPS if the terrain permits. For the GPS to provide correct locations, 1ine-of-sight conditions with the satellites must exist. In forested areas where these criteria may not be met, GPS readings will be taken in the nearest open area. From the location of the open area, as determined by the GPS, tape measu,res and compasses will be used to locate the UTM coordinates of the sample point. If there are no open areas nearby, sample sites will be located using aerial photos, topographic maps, and compass readings. ' �353 9 4. Vegetation sampling Vegetation sampling will be conducted from late May through September to match vegetation phenology to that of when the Landsat imagery data were collected. This time coincides as well with easiest access to field sites and aids in vegetation identification. Vegetation survey crews will not be informed of the vegetation map classification for a given pixel to promote unbiased interpretation of ground-truthing data. Vegetation sampling will be conducted at each of the sample sites to provide information for both the accuracy assessment and a complementary study to further refine the classification algorithms used on the Landsat TM imagery. on-the~ground ~-E classification of the randomly S selected pixels, based on the results of the vegetation sampling, CLUSTER will be used to estimate map accuracy. Cluster sampling will be used to enhance understanding of PIXEL PIXEL PIXEL why certain pixels are misclassified based on the true vegetation classifications of adjacent pixels. General vegetation descriptions will be recorded conducted for the 8 pixels PIXEL PIXEL PIXEL adjacent to the randomly selected pixel (Fig. 1). From the vegetation sampling, ground classifications will be made for all 9 pixels surveyed. From the PIXEL PIXEL PIXEL series of 9 ground classifications, E we can test if a pixel surrounded by like pixels has a lower probability of being misclassified than the probability of being ~30 m---. misclassified when a pixel is adjacent to dissimilar vegetation Figure 1. Cluster sampling design for the classifications. Probabilities of randomly selected pixel (5) and the 8 misclassification may be higher for adjacent pixels. Each pixel is a 30m x given combinations of adjacent 30m square. pixel classifications. Identifying the combinations resulting in a high probability of misclassification can guide choices of areas for further ground-truthing to most efficiently improve map accuracy. 1 1 2 3 4 5 6 7 8 9 t Landsat TM imagery is based on the reflectance values of different canopy cover types. Cover is defined as the vertical projection of the crown or stem of a plant onto the ground surface and serves as a criterion for relative dominance within a community (Higgins et a1. 1994). Ground-truthing, then, must evaluate the dominant canopy cover type for a given pixel. To objectively assign a dominant canopy cover from the vegetation classifications in Table 1 to a sample pixel, the. following standard methodology will be applied. �354 10 1. If it is clear that a given pixel should. be classified into one of the vegetation classes defined in Table 1 the pixel will be assigned to that class without sampling the pixel. Examples of such clarity would include an agricultural field, dense urban development, or open water such as a lake or reservoir. 2. If the area is not homogenous, a systematic grid of points will be laid over the pixel (Fig. 2). Six north-south transects will be laid out over the pixel starting at 2.Sm from the west side and continuing at Sm-intervals. Along each of the six transects the cover type, as. defined by the classifications listed in Table 1, will be recorded at Sm-intervals, starting at 2.Sm from the start of the transect. 3. The vegetation classification with the highest determined as the dominant canopy cover. 4. A photograph will be taken of each pixel evaluated by a ground crew. To identify each photograph, a placard with the randomly selected UTM coordinates of the center pixel and the number of the position. of the. given pixel in the cluster ( e.g., randomly selected UTM coordinates x,y cluster position 1 would be displayed on the placard as x,y: 1) will be placed on the. ground. The photographs will be used as a quality control measure for the field crew as well as providing support for the ground-truthed classification of a given pixel. CLUSTER 123 456 7 8 1\9. PIXEL91\ \ ~'-----~---r----'---~~---r~1 27.p 22. ~ 17.f ..•... Q) • E 5. If the field visit shows that the pixel has clearly been recently converted to another class, the change will be noted. No vegetation surveys will be con~ucted on this site to avoid biasing the accuracy estimate. 6. If a randomly selected site is inaccessible, due to either terrain or landownership, data for the accuracy assessment will be made from low-level aerial photographs taken from a fixed-wing aircraft. frequency will be o 12.5 ('I') I 7.5 2.5 ._---- 30 meters --- ..• , • random UTM coordinates Figure 2. Systematic sampling scheme for recording canopy type at 36 points in each pixel of a cluster of 9. �.355 11 Field crews of trained citizen volunteers (see Objective 3) will conduct the vegetation sampling from sample sites. Assignment of sample sites will be made to minimize travel time for a given crew on a given day. A field test will be conducted by the principal investigator to evaluate and test equipment and sampling methods, evaluate and adjust sampling methods, and make estimates of the time required to complete a single sample. Objective 2 - Develop methodology to involve voluntary citizen participation in ground-truthing of the Summit County Landsat TM vegetation map. The primary requirement when incorporating citizen participation in the accuracy assessment of the vegetation map is that the resulting data must be defensible in both a. scientific and legal context. Therefore, the methods must be reliable, efficient, economical, and above all,. repeatable so that they may be appropriately implemented with minimal training by a number of citizen volunteers. At the same time, citizen volunteers will be introduced to SCOP, trained in scientifically rigorous ground-truthing techniques, and have the opportunity to actively participate in local wildlife management. Citizen volunteers will initially be sought from the CDOW Regional Volunteer Office and local groups concerned with the environment (e.g., high schools, Audubon and Sierra Club Chapters). All volunteers will be trained in the relevant data collection techniques. Necessary field equipment (compass, GPS unit, data collection forms, etc.) will be provided by the CDOW as well as technical support throughout the ground-truthing process. The CDOW will also assure that efforts are being conducted with standardized techniques (see Objective 1) and data are accurately recorded through a series of data checks. These data checks will include test survey plots and maintaining photographic records of all surveyed pixels. Test surveys will be conducted by having citizen crews locate and survey a series of test locations that have previously been surveyed by CDOW personnel. These sites will be randomly mixed in with assigned random selections and will not be known as test sites to the citizen vegetation crews. Evaluation of the test sites by CDOW personnel will be considered 'truth'. Evaluation of the same test plots will then be compared to 'truth' to quantify citizen crew accuracies. If a citizen crew's accuracy falls below 85% their survey data will not be used in the accuracy assessment or reclassification of the vegetation map. All citizen volunteers will receive a copy of the final report summarizing the results of their efforts. Objective 3 - An error matrix will be constructed from the results of the ground-truthing surveys and the Landsat TM imagery. Three measures of accuracy will be determined from the error matrix: overall accuracy, errors of omission, and errors of commission. Evaluation of the errors of omission and commission may provide information to define algorithms to better assign classifications to a given pixel and thus reduce classification error. The overall accuracy will be tested with Pielou's index of segregation. �356 12 Error~ of commission (user's accuracy) Errors of commission are defined as the probability of depicting a certain pixel to be in category i whereas the pixel is actually in category j (Green et al. 1993). The maximum-likelihood estimate for pr(AlICI) is nij nj and the maximum-likelihood estimate for the variance = v(Pr(AjlCi) A l i ·j nn•.+J2 A (1 - of pr(AjlCi) is nn~.j+) A where nIl is the number of pixels classified into category i and found to actually be in category j, nI+ is the total number of sample pixels in category i (from Green et al. 1993). Errors of omission (producer's accuracy) Errors of omission are defined as the probability of wrongly classifying a pixel into category i when it is actually in category i [pr(cI J Because the sample pixels were drawn from a stratified random sample, stratified over classification, the errors of omission are not multinomial probabilities but are conditional probabilities and must be estimated using Bayes Theorem (Green et al. 1993) as follows IA »). Pr(A.IC.) J ~ * [Pr(C.)/Pr(A.)] ~ " J where pr(Aj) = t [Pr(Ajl Ck) Pr(Ck) "k-l and pr(CilA) pr(AjlC) (ni/ni.l * Pr( Pr(ci)/pr(A) C) is the percentage of all pixels in the map classified into category i and known without error, Pr(Ck) is the percentage of all pixels in the map classified into category k and known without error, nkjis the number of pixels classified into category k and found to actually be in category j, m is the number of categories on the map, and all other parameters are defined as above. The" variance estimator for the pr(cIIAl) from a stratified random sample is given in Green et al. (1993). Pr(CI) �357 13 Assessment of overall accuracy ·Kalkhan (1994) found sampling schemes affect both bias of accuracy indices and estimates of their variance. Therefore, based on a stratified random sampling· scheme we will use Pielou's index of segregation (5) to estimate overall accuracy. Pielou (1961, 1977) introduced an index of segregation that measures the spatial association between two or more groups. Pielou's index of segregation, B, is defined as: i 'I' j where N is the sample size of classification categories of the thematic map, i is the row number, r is the number of rows, j is the column number, c is the number of columns, Xij is the number of counts of the classification category, Xi+ and x+j are the sum of the i-th row and j-th column respectively. If there is perfect agreement between the classification categories, 5 = 1, and 5 = 0 when there is no agreement'. Pielou's index of segregation follows a X2 distribution with (i-1) and (j-l) degrees of freedom, where i and j are the number of rows and columns in the error matrix, respectively (Kalkhan 1994). Although no analytical estimator is available to estimate the variance of Pielou's index of segregation an unbiased estimate can be obtained using bootstrapping (Efron 1979). These 3 estimates of map accuracy will be used to quantify confidence levels .for the vegetation ~ap. These estimates of map accuracy will also be used to provide levels of confidence in GIS metrics derived from the vegetation map layer. For example, assume species richness was to be estimated for each pixel as a function of the assigned vegetation classification of the pixel. The level of confidence on the species richness metric must incorporate the variance associated with vegetation classification of the pixel as well as its own associated variance. Therefore, it.is necessary to obtain an estimate of error in the vegetation map to provide confidence limits on estimated GIS metrics. Objective 4 - Improve accuracy of the Summit County vegetation map database through reclassification of misclassified pixels and improved supervised classification algorithms. Pixels misclassified on the Landsat TM vegetation map will be reclassified as the vegetation class determined from either ground sampling or aerial photography used to estimate map accuracy (see Objective 3). However, to encourage citizen participation and maintain volunteer enthusiasm for the ground-truthing effort, ground-truthing of sample pixels outside the stratified random sampling scheme will be included in the reclassification of the vegetation map. For example, citizens may be more likely to participate if they can ground-truth their own property •. The revised vegetation map, with all reclassified pixels, will be used in further GIS analyses performed in the SCoP information system. The result of using the revised map and the accuracy �358 14 errors estimated for the original map will be a conservative associated with any GIS data analyses. estimate of error Results from the confusion matrix and cluster analysis may provide information to improve the current, supervised classification algorithms. In particular, there may be enough information to separate out the vegetation classification of Aspen-Irrigated-Willow. If so, the vegetation map can be further revised to incorporate the more accurate classification scheme. However, the resulting map generated from the revised classification scheme would have to be evaluated with an independent ground-truthing study. Objective 5 - Develop standardized guidelines assess accuracy of Landsat TM imagery. on the design and methods to A handbook describing a standardized approach to the design and methods used to assess accuracy of vegetation maps constructed from Landsat imagery will be developed. The handbook will first provide a review of technical aspects of an accuracy assessment (e.g., sampling design, sample size, concepts of accuracy), and secondly, layout a .protocol for accuracy assessment that proceeds from how to contact volunteers, to field procedures, to data analysis. Any revisions from the above described approach to field techniques used in the ground-truthing, citizen recruitment, training, field crew logistics, field forms, or data analyses will be incorporated into the handbook. ™ RESULTS Cicizen AND DISCUSSION volunceers A total of 24 volunteers have been trained to date to collect data for the ground-truthing effort. The majority (14) of these volunteers were trained during a 2-day training session held May 4-5 in Frisco, Colorado. The remaining volunteers were enlisted and trained individually throughout the summer~ Of the 334 sites completed to date, volunteers conducted 42 of the vegetation surveys. Competency of the volunteers is excellent. However, the time .and energy demand to locate the random sites and conduct the vegetation surveys is high. Accuracy assessmenc A pool of 75 randomly selected pixels (30m x 30m) for each of 13 vegetation classifications was generated. To date, 334 of these pixels have been surveyed to determine dominant cover. A preliminary error matrix was constructed from these 334 ground-truthing surveys using only dominant vegetation cover. Although complete surveys were conducted at each site, for this first analysis only dominant cover type was analyzed. Thus, combination vegetation classifications including conifer-aspen, and aspen-irrigated-willow were not evaluated. Also, the two conifer classification, conifer dense and conifer sparse, were combined into a single conifer classification for these initial results. Two measures of accuracy were determined from the dominant cover error matrix: overall accuracy (60%) and errors of omission. A-preliminary estimate of the �359 15 overall map accuracy is 201/334 or 60.2%. However, very little useful information is contained in this estimate. Far more information concerning the accuracy of the map can be extracted from the error matrix. An error matrix was constructed from the 334 completed vegetation surveys (Fig. 3). From the error matrix, errors of omission were estimated (Table 2). Errors of omission suggest good accuracy (> 60%) for the vegetation classifications water, soil-dominated, and rock. Errors of omission performed most poorly for the vegetation classifications willow (0%) and grass-forb (23%). The greatest confusion occurred between irrigated and grass-forb, willow and irrigated, and a confusion of mixed shrub, grass-forb, and sage. Field work continues to complete vegetation surveys on the remaining selected pixels. Once completed, a full analysis will be conducted. * 641 CLASSIFICATION DATA (from vegetation map) W· CO CA A I WL AIW GF S MS SD R' ROW TOTAL W 72 0 0 0 0 0 0 0 0 0 0 0 '72 CO 0 35 1 0 0 0 0 4 0 0 1 0 41 CA 0 11 0 10 0 1 0 O. 0 0 0 0 22 A 0 1 0 23 2 2 1 3 0 0 0 0 32 I 0 0 0 2 10 0 0 8 0 0 0 0 20 WL 0 0 0 1 13 0 0 6 1 0 0 0 21 AIW 0 3 0 7 2 0 0 6 3 1 0 1 23 GF 0 6 0 1 2 0 0 12 2 0 0 1 24 S 0 0 0 0 1 0 0 2 17 0 0 4 24 MS 0 0 0 0 0 0 0 6 6 1 0 0 13 SD 0 2 0 0 0 0 0 4 2 0 19 0 27 R 1 1 0 0 0 0 0 1 0 0 0 12 15 COLUMN TOTAL 73 59 1 44 30 3 1 52 31 2 20 18 334 REFERENCE DATA (from ground-truthing) Classification codes are defined below. Descriptions of classification codes are given in Table 1. Classification will be eliminated from the ground-truthing surveys. W CO CA A Water Conifer (denser and sparser) Conifer-Aspen Aspen AIW GF S MS 'snow' Aspen-Irrigated-Willow Grass-Forbs Sage Mixed Shrub �360 16 Figure 3. Error matrix for the vegetation map of summit Classification descriptions are given in Table 1. Table 2. Errors of omission for the Summit vegetation ground-truthing surveys. Vegetation claesification County County. vegetation Errors of omission 72/73 98.6 Aspen 23/44 Irrigated 10/31 Willow 0/3 Grass-Forb 12/52 = = = = = = Sage 17/31 = 54.8 1 / 2 = = = 50.0 Water Conifer Mixed (denser and sparser combined) Shrub 35/59 soil-dominated 19/20 Rock 12/18 LITERATURE map based on 334 (%) 59.3 52.3 32.3 0.0 23.1 95.0 66.7 CITED Bailey, R. G. 1988. Problems with using overlay mapping for planning and their implications for geographic information systems. Environmental Management 12: 11-17. Briggs, D. 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Gap analysis: a geographic approach to protection of biological diversity. Wildl. Monogr. 41pp. Star, J. and J. Estes. 1990. Geographic Information systems: an introduction. Prentice Hall, Englewood Cliffs, New Jersey. 303pp. van Genderen, J. L. and B. F. Lock. 1977. Testing land-use map accuracy. Photogrammatic Engineering and Remote'sensing 43:1135-1137. Wang, F. 1990. Improving remote sensing image analysis through fuzzy information representation. Photogrammatic Engineering and Remote Sensing. 56:1163-1168. � https://cpw.cvlcollections.org/files/original/c9d884d5a0244a3566c9b957c9ec9c74.pdf 87694267a894c943ed2bf2b105e81f57 PDF Text Text JOB PROGRESS REPORT State of Colorado Project: W-15Q..R-9 ENDANGERED WILDLIFE INVESTIGATIONS Work Plan _.2.__ : Job _t__ Job Title: Peregrine Falcon Restoration PrQ~ Period Covered: 1 January - 31 December 1996 Personnel: G.R. Craig and C. Wightman, Colorado Division of Wildlife and J.H. Enderson, The Colorado College. ABSTRACT Activities conducted during this period have already been reported under W-150-R-8 (see attached report). ��3 JOB PROGRESS REPORT State of _----:,C04oW1o,wralDodl3.!o __ Project: _---'(W,..LL..- .•.• l50-""'-'R~-8""")~_ : Peregrine Falcon Restoration Program Period Covered: 1 July, 1995 - 30 June, 1996 Personnel: G.R. Craig and C. Wightman, Colorado Division of Wildlife and J .H. Fnderson, The Colorado College. ABSTRACT In the 1996 peregrine breeding season, 83 territories were occupied by 78 breeding pairs that fledged 140 young . . Productivity averaged 1.65 young fledged for those pairs that were monitored. Contents of 6 nonviable eggs were collected and have been preserved for future analysis. Shell fragments were collected at 18 sites and are awaiting measurement. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBUSHED OR QUOTED without permission of the author. ��5 PEREGRINE FALCON RESTORATION PROGRAM Gerald R. Craig SEGMENT OBJECTIVES 1. Ammally monitor the munber of breeding pairs of peregrines and their reproductive success in Colorado. 2. Ammally monitor organochlorine pesticide levels in Wild breeding peregrines in Colorado. 3. Monitor breeding J:qJUlation turnover through baIIl recoveries, presence of color markers, and telephotographic identification of individual breeding adults. 4. Augment poor wild production by placement of captive hatched Wild young and captive produced young into occupied Wild nests. 5. Release captive hatched and captive produced young at potential and vacant wild territories. 6. Monitor recruitment of reintroduced peregrines into the Wild breeding population of Colorado. METHODS AND MATERIALS 1. Visit all known peregrine breediog territories tbrougboot Colorado and observe them from a distance to establish the presence of breeding adults. Breeding pairs will be kept under surveillance to determine initiation of egg laying. Depending upon the individual female's reproductive history and eggshell condition (obtained through measurement of previous year's eggshell thicknesses) and availability of captive hatched young for release, breeding pairs either will monitored or manipulated as outlined in approach 4. Those pairs not designated to be manipulated will be revisited periodically throughout the nesting season to document reproductive success. When a pair's behavior indicates that egg laying has occurred and incubation is underway, the eyrie will be visited to document the mnnber of eggs produced. The. eggs will be candled to ascertain viability and approximate age. All nonviable eggs will be collected for chemical analysis. A second visit will be made to. determine productivity, band nestlings,. and collect eggshell fragments and unhatched eggs for thickness measurement and analysis under 2a and 2b. '2a. Eggsbell fragmeUs cu:ooIIered during eyrie visits described in approaches 1 and 4a will be measured for index to thickness following standardized procedures. 2b. Whole, nonviable eggs which are encountered during eyrie visits will be collected, preserved and submitted to the appropriate Fish and Wildlife service approved laboratory for pesticide analysis. Eggs collected from the wild in the coorse of Approaches 4a, 4b and 4c that are artificially at the Peregrine Fund's Boise, Idaho facility also will be submitted for shell thickness measurement and chemical analysis. 3. Peregrines presea at breediog territories will be examined to determine the presence of bands or color markers. Band confirmation will be accomplished through observation from a distance with telescopes and concealed remote controlled cameras. When banded falcons are encountered, every effort will be made to read band DJIDberswithoot trapping or baoUing the birds. It is possible this can be accomplished in most situations with a Questar field model telescope (80-13Ox). When band munbers cannot be discerned, attempts will be made to trap and examine the falcon at a time when capture will have least impact upon breeding activities. 4a. In accontance with an anwal release plan developed and approved by the State, U.S. Fish and Wildlife Service, Bureau of Land Management, National Park Service, and the Forest Service, a predetermined number of wild breeding pairs will be manipulated to augment natural productivity. Pairs with a history of reduced. clutch size, cracked eggs, or infertile or dead eggs will be candidates for fostering efforts. �6 4b. On occasion, it may be necessary to recycle several early breeding pairs in older to delay them until captive batcbed young of tile proper age are available for placement into wild sites. No later than 10 days after the last egg has been deposited, the eyrie will be visited and the entire clutch removed without replacement. Approximately 14 days after removal of the clutch, the pair will recycle, select another nest ledge, and deposit a secoed clutch of eggs. If the eggs are thin shelled, they may be replaced with plastic replicas and treated as outlined in approach 4a. This tecImique also worlcs well to augment captive production with wild produced eggs. 4c. At tinies, pairs will select inferior eyrie ledges that may compromise nest Success such as ledges that are too narrow to support a brood of large nestlings, the site may be vulnerable to predators, or it may be exposed to tile elelmn. If tile ledge cannot be mechanically improved, pairs can be relocated to other ledges through the recycling method described in approach 4b since they invariably relocate and select a new ledge when recycled. 5. In accon1aoce with an amml release plan develqx:d and approved by the State, U.S. Fish and Wildlife Service, Bureau of Latxl Mana~emeu. National Parle Service, and tile Forest Service, a predetermined DUIIlberof captive produced falcons will be released at unoccupied or potential sites through the technique of hacking. This teclmique is employed at locations that do .not have the benefit of protection or care from adults. Young falcons of aboot 35 days of age will be placed in a hack box on a suitable cliff ledge at the reintroduction site. They will be fed and cared for by attendants until they are flying and capable of fending for themselves. This technique assures that tile yooog become familiar with tbeir surroundings and hopefully will return to the site as adults and take up residency. Hacking requires constant attendance and observation to protect the vulnerable young and assure they have sufficient food while they are dependent upon the hack site. While the hack sites will be operated by the State, adual costs to operate the sites will be· borne by the appropriate land administering agency (Forest Service, Bureau of Land Management, and National Park Service). . 6. ConDrmed breeding territories and selected potential breeding sites will be surveyed ammally to document the presence of released falcons and ultimately determine the success of recovery efforts. RESULTS AND DISCUSSION Survey Effort In 1996, 4 teams comprised of 2 obsetvers each were assigned particular regions of the state to monitor breeding activities and survey potential cliffs as time permitted. Three teams were assigned regions west of the COntinental Divide and 1 team was located east of the Divide. Initially, 89 breeding territories were scheduled to be monitored by the teams, but three territories were not visited due to access difficulties and time constraints. An additional 43 potential nesting cliffs were examined for presence of nesting peregrines as time permitted and 14 new breeding territories were discovered. Territory {)cgJpancy Breeding territory occupancy increased from 71 sites in 1995 to 83 in 1996 (Fig.l). Seventy-eight of the sites were occupied by adult pairs that attempted to breed (laid eggs). Of the 5 pairs that did not breed (sites 44, 72, 88, 92 and 99) a menDer of ODepair (site 105) was comprised of an immature female and the adult female of another pair (site 72) was replaced by inunature female prior to egglaying. The rate of occupancy remained near the 80% level (Fig.2) and is generally considered appropriate for a stable population. an �7 Figure 1 NUMBER OF BREEDING PAIRS 80 ---------------------------------------------------------------------------------------------------- 70 ------------------------------------------------------------------------------.------------- 60 -------------------------------------------------------------------------------- 30 -------------------------------------------------------------- 10 o~-.-.""""--""'...-.,--..••••• -. T2 73 74 75 76 T1 78 79· 80 81 82 83 84 8S 86 87 88 89 90 91 92 93 94 95 96 YEARS Figure Z. RATE OF OCCUPANCY 80% -------------------- ~~-----------------------------------------------------------T2 73 74 75 76 T1 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 YEARS Reproduction The bigh proportion of of breeding pairs (94% )(Fig. 1) and a slight increase in their breeding success (Fig. 3) resulted in an improved productivity of 1.65 young fledged per total pair wbich wasthe bigbest since 1991 (Fig. 4) .. Eighteen of the breeding atteIq)(S fili1edwbile 60 pairs fledged at least 1 young. Actual fledging was not confirmed at only 3 sites (38,41 and 106) and for those sites with known outcome, the average fledged brood size (young fledged per successful pair) was 2.32. At least 160 young were hatched and 140 were confirmed to have fledged for an estimated mortality of 12.5 % for the nestling period. In addition. 8 young died between 3S days and fledge yielding a 5 % mortality for that age bracket. . �8 Figure 3. PERCENT OF TOTAL BREEDING PAIRS 1HA T WERE SUCCESSFUL 1()(J9.{, - - ~.; ':~:\ :;,;' .;.~: "~: ~" !;', ~.'.:.,~..~.:: ..: ::;:' 78 79 80 81 82 83 84 85 86 87 88 ,89 90 .. _----- ---_._-------- ;;', 91 92 ~f~J. 93 94 95 96 YEARS l~r21~kl Succ:e.sful PaiD; Figure 4. PRODUCTIVITY 2.5 - ~ 2 tn {31.5 u o ~ p.., 1\ ----r' \---------------- --- -----.---------------.~.r;a--'- -- - -- --"~--- .,.R......;! ~ 1 ---f----- --- z 8 I>- 0.5 o .____ ------ -- - -- _,,~ ------ _••• =sr=: -.----------------------------~---------------------------------------------------- cP -j-------:'-8/-~\;:.:-~:j\:.~:'--------j- -----.-----.---.--------.----.-------. --------.-! C!) »: ,..... 73 74 tY 75 76 77 78 79 ~ 80 81 82 Total Pairs 83 84 85 86 YEARS 87 88 89 90 91 92 93 94 95 96 --E3-. Unrnanipulated Pairs ~2shell Condition Six whole, nonviableeggs were encountered in the course of visits to 5 sites. These eggs have been preserved whole for contaminant analysis and shell thicknesses are not available at this time. Eggshell fragments were also collected from 18 additional nesting sites in the course of visits to band young. Thickness measurements have not been compiled for' these samples at this time, so DO coochIsions can be made about the 1996 thicknesses. Figure 5 shows the eggshell thickness trends through 1995. These values are highly variable due to small sample sizes, mixing of fragments from differeu eggs within the same cb.Jtch,and variation of thickness of fragments from the poles of the egg versus the waist. However, there appears to be a slight trend towaItl thicker eggs over the past 8 years with thickness averages less than �9 10% thin sioce 1992. Greater variability among eggs is also evident since 1988 with some eggs being thicker than preDDT era eggs and others 16% thinner. Figure 5 CUMULATIVE EGGSHELL TIllCKNESSES 0.39 - - - --- ---- --- -- -- - -- - -- -- - -- -- - -- -- - -- -- - ---- - -- -- - -- ---- - -- - -- --- -- -- - - - -_._----------,------------------------------------------------------------~ Pre-DDT Ere. Thi~e . M_ - - •• ~-----.-- 73 74 7S· 76 77 78 79 80 81 s::t I - -- -- - - - ------~ --- S3 8:4 8S .•.•••• and Minl1nuaI:L~ckn.""." Orpuochlorine St5 87 S8 89 90 91 9::i1 93 94 9.5 Years --=>- AT_age Thickn ••••• " Residue in Ea;a;s The 6 DOUViableeggs collected during the 1996 season have been preserved along with eggs encountered in 1994,.1993, 1992 mll991. This collection of 47 eggs is awaiting pesticide analysis by the Fish and Wildlife Service when funding is available. Release and Au~entation Efforts Remedial management efforts such as fostering or hacking have not been undertaken since 1989. Life Science Researcher N ��11 JOB PROGRESS REPORT State of Colorado Project: (W-151-R-8): Period Covered: Bald Eae:le Nest Site Protectiop apd Enhancement Proe:ram I July, 1995 - 30 June, 1996 Personnel: G.R. Craig, Colorado Division of Wildlife ABSTRACT Bakl eagles occupied 26 Colorado nesting territories in 1996. Five new territories was discovered and one that had been vacant in 1995 was reoccupied. Nineteen territories hatched young and all pairs fledged 32 young. Prodqctivity averaged 1.23 young per occupied territory. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBUSHED OR QUOTED without permission of the author. ��13 BAlD EAGLE NEST SITE PROTECTION AND ENHANCEMENT PROGRAM Gerald R. Craig SEGMENT OBJECTIVES 1. Monitor nest site occupancy and reproductive success. 2. Document survival rates and mortality factors. 3. Determine migration and wintering areas. 4. Determine if philopatry occurs in breeding eagles. 5. Determine nest site tenacity by individual breeding eagles. 6. Quamify nesting babitats and associated foraging areas in an effort to document nest site parameters conducive to improved reproduction. 7. Document pesticide contamination through eggshell measurement and chemical analysis of nonviable eggs. 8. Where necessary, implement actions to stabilize nests and majnrajn occupancy. MHfHODS AND MATERIALS This work will be a cooperative endeavor between the Division and Dr. Ricbard Knight of Colorado State University. 1. AImually visit all documented breeding sites to determine the presence of bald eagles. Pairs at territories will be documented by DWMs and other field personnel. Previously unrecorded pairs will probably be revealed in the course of aerial eagle and waterfowl flights. DWMs will confirm actual incubation from ground visits. 2. Occupied territories will be visited by DWMs periodically throughout the breeding season to determine hatch of young, nesting failures, etc. . 3. In May and June, a Utility Worker will observe breeding eagles from a distance and endeavor to follow their lOOVementsto locate important foraging areas. Responses of eagles to various buman activities and land uses will be recorded. 4. InJWJe, when the young are determined to be old enough to band, sites will be visited by Craig and Knight to place a federal band on one leg and a colored, alpha numeric marker on the other. The color markers will permit identification if the young return in subsequent years. During the same nest. visit the following will be recorded: . Physical parameters such as tree species, height, DBH, condition, and dominance. Nest condition, size, and location. Vegetative comnmnity and land use practices. In addition, Collect prey remains, nonviable. eggs and eggshell fragments. 5. ApproXimately Sec's ofblood will be collected from each nestling. The blood will be analyzed at the Savannah River Ecology Lab in Aiken, Sooth Carolina. Electrophoretic examination will permit genetic comparison with saIq)les collected from other populations in Saskatchewan, the Lake States and Arizona, as well as determine the heterogeneity of the Colorado birds. 6. When necessaty, remedial actions will be taken to stabilize nests that are threatened by wind throw. Should the �14 tree be decadent and in danger of falling, an artificial nest base may be placed in a suitable, adjacent tree. ActioIi will be taken only after it bas been deemed desirable to encourage the eagles to nest at the same location. RESULTS AND DISCUSSION Ierritozy Occupancy Ba1deagle nesting activities in Colorado are swnmarized in Figure 1 and Table 1 details the 1996 breeding season results. In 1996, 26 territories were occupied of which 5 (Jackson, Moffat IS, Rio Blanco 16 and 117, and Routt 112) were discovered in 1996. Nest size suggests that Rio Blanco 117 and Moffat 115were probably first nesting attempts. Although the Jackson site was discovered in 1996, the ranch caretaker indicated tbat it had been occupied at least the 2 previous years. Presence of an adult at Routt 112 in 1995 suggests tbat site was active the previous year. The Fremont site was occupied by adults early in the breeding season, rut did not produce young. Although the Grand site nest tree blew down shortly after the yooog fledged in 1995, the pair relocated to an artificial osprey nest platform and fledged 1 young. After the Rio Blaoco /14 rest slipped rut of the tree in 1994; the pair relocated upstream approximately .5 miles. The new nest was discovered in the course of s be1icopter inventory of big game. It is likely tbat the pair used the nest in 1995. Montezuma 112 and /13were occupied by geese. Figure 1 BREEDING PAIRS AND PRODUCTIVITY 35 -.----------------------------------------------------------, 30 - ------------------------------------------- -------- --------------------------------------- -- 25 20 - ------------------------- ------------------------------------------------ 1::5 - - - - -- - - - -- - -- --- - --- - - - -- - - - ---- ---- - - -- - -- --- - ---- - - -- - -- ---10 - --------------------- ---------------------------- o _'JM~~aL~JULJ~.c~ __ 74 75 76 77 78 79 80 81 82 83 84 8:5 86 87 88 89 90 91. 92 93 94 95 96 Yean; _____ Youn.g Proctuc:.d. _ Br •• d.U3.gPair. Reproductive efforts for 1996 are summarized in Table 2 and Figure 1 reviews the previous seasons' results. In 1996, 21 pairs laid eggs aOO32 yooog were produced by 19 successful pairs (1.68 young per successful pair) which yielded an overall productivity of 1.23 yooog per territory occupying pair. Seven pairs either did not produce eggs" or failed during incubation. Single unhatched eggs were encountered during banding visits at 2 nests (Moffat 111and 64). Sixteen young were banded and color marked at 11 locations (Adams, Jefferson, Grand, La Plata 111, Mineral, Moffat 61,112,63 and 64, Morgan and Routt 82) Fish and Wildlife Service bands were affixed to the nestlings' right legs and red alpha-numeric bands with yellow vinyl flags were affixed to their left legs. Culmen length and foot pad length measurements were obtained fromthe eaglets that were banded. Eaglets at 4 other sites were too old to band, and landowner permission could not be obtained at Rio Blanco 84. �15 Land Status The 5 new territories (Qackson, Moffat 115, Rio Blanco 116 and 117, and Routt 112) were on private property and livestock grazing is the primary land use. Banded Adults The adult male at Rio Blanco 115 was banded and color marked. He was produced at Rio Blanco 113 in 1991 which is approximately 16 miles downstream. The female was unbanded. The female at the Adams site, who had been banded on tbe Rocky MOOIlIainArsenal the winter of 1986 was replaced by an unbanded female in 1996. She was banded as an adult, so given at least 4 additional years to achieve adult plumage, she was at least 13 years old when she disappeared. Nest Stabjlization Efforts No rest stabilization effons were uWertaken in 1996. The nest at Rio Blanco 115 which blew out killing the chick in 1995 year, was recomtructed in a ome substantial position in 1996. The Rio Blanco 111 nest tree continued to remain upright in spite of its dead common. The pair did not frequent an artificial nest that had been constructed in an adjacent tree a wmber of years previously. After being vacant in 1995, Rio Blanco III was reoccupied and was successful despite the fire damage that killed the tree. The Archuleta site successfully fledged 2 young, although at time of fledging, the nest fell to tbe grwOO m1 tbe fledglings had to perch on neaIby limbs. An. artificial nest base will be placed at the site in the fall, The Jefferson rest is vulnerable to wind throw due to placement in the crotch of a dead limb. Since the limbs cannot be stabilized, consideration should be given to placement of an artificial base in an adjacent tree. The dead nest tree at the Adams site had its life extended when it was guyed up by REA~ Prepmdby: . C£. (1.... 6= oer.lid R. Craig Life Science Researclier IV �- Table 1. Bald Eagle Nesting Efforts in Colorado 0\ Site La Plata Co. #1 Moffat Co. #1 La Plata Co. #2 Grand Co. Montezuma Co. #1 Rio Blanco Co. #1 Rio Blanco Co. #3 Weld Co. #1 Montezuma Co. #2 Moffat Co. #2 Moffat Co. #3 Adams Co.. Archuleta Co. Montezuma Co. #3 Weld Co. #2 La Plata Co. #3 Rio Blanco Co. #4 Morgan Co. #1 Mesa Co. #1 Fremont Co. Routt Co. #1 Gunnison Co. Mineral Co. Weld Co. #3 Montezuma Co. #4 Jefferson Co. Mesa Co. #2 . Moffat Co. #4 Jackson Co. Rio Blanco #5 Moffat CO.#5 .Routt Co. #2 Rio Blanco #6 Rio Blanco #7 Total Young Total Pairs Young/Occ. Terr. IA - Inactive A 1974 1975 1976 1egg IA IA 1yng 2yng 2yng 1977 IA 2yng 2yng 1978 IA 2yng 2yng Oyng 1979 IA 1yng Oyng Oyng 1980 1981 1982 1983 ? ? ? ? -- 2yng 2yng -IA IA IA IA· IA IA IA IA A A A A 1yng 1yng ? 3yng 2yng 1984 ? 2yng IA IA IA eggs 2yng 2yng 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 ? eggs 2yng 1yng 2yng Oyng 1yng 1yng 2yng 3yng Oyng 1yng Oyng 1yng 2yng 1yng 3yng 2yng 2yng 2yng 2yng 2yng 2yng 1yng IA IA A IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA IA 1yng 1yng IA IA IA IA IA IA IA IA IA IA IA IA Oyng 2yng 2yng 2yng.2yng 2yng 2yng 1yng 2yng 2yng IA 3yng 2yng Oyng 1yng A .2yng 1yng 3yng 3yng 2yng 1yng 2yng 2yng 2yng eggs IA IA IA Oyng IA IA IA IA IA IA 2yng 1yng 1yng 1yng 1yng 1yng Oyng 1yng IA IA IA IA 1yng Oyng 2yng 3yng 2yng 2yng 3yng Oyng Oyng 3yng 2yng 1egg IA ? eggs IA Oyng Oyng 2yng 2yng 1yng 2yng eggs 1egg eggs 2yng 2yng 3yng 3yng 2yng 3yng 3yng 1yng eggs 2yng IA IA IA Oyng Oyng eggs IA 1yng 2yng 1yng 1yng 1yng 2yng 1yng 2yng 1yng 2yng IA eggs IA IA IA IA IA IA IA 2yng 2yng ? ? IA ·lyng Oyng 2yng 1yng 2yng 3yng Oyng Oyng 2ng 2yng 3yng 2yng Oyng 3yng 3yng Oyng eggs eggs Oyng IA IA 2yng IA Oyng Oyng 2yng Oyng 2yng Oyng 1yng 2yng Oyng LAD A ? Oyng 2yng A Oyng Oyng Oyng Oyng IA IA IA Oyng Oyng Oyng 1yng 2yng 2yng 2yng 1yng 1yng 1yng ?yng 2yng 2yng Oyng 1yng OYng 1yng 2yng Oyng o 1 4 4 4 1 0 3 6 2 6 6 5 10 8 16 13. 19 20 24 19. 25 32 1 1 2 2 3 3 1 3 4 2 4 5 10 9 8 10 10 13 14 20 17 21 26 0.00 1.00 2.00 2.00 1.33 0.33 0.00 1.00 1.50 1.00 1.50 1.20 0.50 1.11 1.00 1.60 1.30 1.46 1.43 1.20 1.12 1.19 1.23 LAD - Lone Adult - Active ~- �17 Table 2. Colorado Bald Eagle Nesting Efforts - 19% Site Age of Birds Male Female Comments Young Produced Central Region Adams Co. Jefferson Co. Adult Adult Adult Adult 1 1 Northeast Region Jackson Co. Morgan Co. Weld County #3 Adult Adult Adult Adult Adult Adult 2 3 0 Adults present early, eggs not confirmed, Southeast Region Freniont Co. Adult Adult ? Adult perched by nest early in season. 2 Fledged, nest fell out at that time. Adult female only bird observed. Southwest Region Archuleta Co. Gunnison Co. La Plata Co #1 La Plata Co. #3 Mineral Co. Montezuma Co. #2 Montezuma Co. #3 Northwest Region Grand Co. Mesa Co. #2 Moffat Co.#1 Moffat Co #2 Moffat Co. #3 Moffat Co. #4 Moffat Co. #5 Rio Blanco Co.#1 Rio Blanco Co. #3 Rio Blanco Co. #4 Rio Blanco Co. #5 Rio Blanco Co. #6 Rio Blanco Co. #7 Routt Co, #1 Routt Co. #2 Total Total pairs: 26 Egg laying pairs: 21 Productive pairs: 19 IA = Inactive Adult Adult ? Adult SubadAdult Adult Adult Adult Adult 0 1 0 2 Goose in nest Goose in nest Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult SubadAdult Adult Adult 23 26 1 2 1 2 2 1 0 3 2 2 1 2 0 0 1 32 Unhatched egg in nest. Unhatched egg in nest. Female in incubating posture, failed. Relocated to new nest, probably used in 95. Adult male was hatched at Rio Blanco #3 in 1991. Observed from aircraft. Female in incubating posture, failed. ��19 Colorado Division of wildlife Wildlife Research Report July 2, 1997 JOB PROGRESS REPORT State of project: Work Plan Colorado W-164-R-2 2 Laboratory Investigations Job 4 Job Title: WILDLIFE FORENSIC TRAINING SLIDE SERIES FOR FIELD AND ·LABORATORY INVESTIGATIONS. Period Covered: 1 July, 1996 - 30 June, 1997. Personnel: W.J. Adrian. ABSTRACT wildlife forensics is a rapidly changing science with very few methods of disseminating research findings to field and laboratory personnel in a timely manner. Publication of research findings is a necessary part of any research project, but getting that information on line in the field and other forensic laboratories is subject to great time delays. This goal of this project is' to obtain existing slides and exhibits used to teaching wildlife forensics to field and laboratory personnel, produce slides, title slides and exhibits that are not currently available to forensic laboratories and field personnel and to make this material available to all contributing forensic scientists. The scope of this project is cover all laboratory forensic techniques both at the professional laboratory personnel level and the wildlife officer level, how to submit all types of evidence to the laboratory and cover all field forensic techniques. Segment Objectives "1. Obtain existing slides and exhibits used in teaching wildlife forensics to field and laboratory personnel. 2. Produce slides, title slides and exhibits necessary for teaching wildlife forensics to field and laboratory personnel. 3. To make duplicates of all slides and exhibits all contributing forensic scientists. for �20 METHODS AND MATERIALS Obtain existing slides and exhibits used in teaching wildlife forensics to field and laboratory personnel, produce those slides, title slides and exhibits not currently available, and to make duplicates of all slides and exhibits for all contributing forensic scientists. RESULTS AND DISCUSSION Existing slides have been obtained for Nebraska, Wyoming, Tennessee and California that are used in teaching wildlife forensics to field and laboratory personnel. Title slides are being made to update these new slides. This project is approximately 60% complete and includes (to date) 6 new sections on laboratory and field forensics techniques. Prepared �21 JOB PROGRESS REPORT State of . Project Job Title: Colorado W-166-R : Mjgratoty Game Bird Investigations Work Plan _L: Job _2L Preseason monitor banding of Mallards in Colorado Period Covered: 01 January through 31 December 1996 Author: Michael R. Szymczak Personnel: Pat Medina, U. S. Forest Service; J. Broderick, R. Caskey, P. Creeden, R. Del Piccolo, J. Ellenberger, V. Graham, J. Gray, 1. Gumber, T. Mathieson, J. Miller, M. Szymczak, and S. Yamashita, Colorado Division of Wildlife. ABSTRACT Ducks were trapped in modified Salt Plains bait traps and banded at 1 wetland location near Grand Junction and 1 location near Yampa, in western Colorado in August and September 1996. Three hundred and eighty-two mallards (Anas platyrhnchos) were banded; 362 near Grand Junction and 20 near Yampa. ��23 PRESEASON MONITOR BANDING OF MALLARDS IN COLORADO 1. Band mallards in Colorado's portion of Banding Reference Areas in the Pacific Flyway that will contribute information on harvest rates, survival rates, and distribution of harvest for use in Adaptive Harvest Management of western mallard populations. SEGMENT OBJECTIVES 1. Trap and band mallards in the Grand IunctionlDeltalOlathe area in late August-early September using salt plains bait traps (Szymczak and Corey 1976). Banded ducks will be classified according to age and sex using accepted techniques (Carney 1964, Weller 1976: 35). Banding schedules and recapture reports will be submitted to the U. S. Fish and Wildlife Services' Bird Banding Laboratory. Band return reports will be summarized and remain on file with the Colorado Division of Wildlife. INTRODUCTION In 1990, the Pacific Flyway Study Conunittee formulated a 5-year cooperative mallard and northern pintail (Anas acuta) preseason banding program that was endorsed by the Pacific Flyway Council. This program was designed to address banding needs throughoutthe western U. S., including Alaska; and in the provinces of British Columbia and Alberta. Through the first 5 years, about 9,000 ducks were banded in Colorado under this program. Following the 5th year of banding, an analysis of recoveries of all mallards banded during the 5. year period in the western U. S. and Canada showed that cohorts of mallards banded in southeastern Idaho, western Wyoming, northern Utah, and western Colorado had similar recovery distribution properties. Since trapping and banding efforts in the 4-state region had been most successful in southeast Idaho and western Colorado, those 2 .areas were selected for continued banding in relation to the Pacific Flyway Council efforts to establish a western mallard management unit. METHODS Trap Area Selection The selection of wetland locations for continued banding were based on number of birds banded per unit of effort during 1991 through 1995, and on the availability of personnel to operate the banding stations. The Walker State Wildlife Area (SWA) near Grand Junction and Markley's Pond near Olathe were selected sites. In addition, birds were also banded at Gardner Park Reservoir near Yampa. Gardner Park was the only high elevation site where birds were �24 trapped during the S-year period and few birds were banded at that location. Additional banding and resulting recoveries would add to and strenghthen the distribution information collected for birds breeding in that area. . Trapping Ducks were trapped and banded near Grand Junction from 26 August through S September, and at Gardner Park Reservoir from 12 September through 18 September. All birds were trapped in modified Salt Plains bait traps (Szymczak and Corey 1976) using whole shelled com for bait. Traps were visited daily. Mallards and pintails were the target species. Banded birds were recorded by wetland site. Band numbers of all birds captured that were banded in previous years or outside the specific area of trapping were recorded. RESULTS Trapping, banding and record keeping Trapping occurred only on the Walker SWA and at Gardner Park Reservoir (Table 1). A trapping crew was not assembled to trap in the Uncomphragre River valley. At Walker, 3 riverine sites were selected for trap sites. A total of382 mallards was banded during trapping in western Colorado in 1996 (Table 2) with a reduced trapping effort compared to the previous S years. Immatures comprised 76 % of the sample on the Walker SWA, but only 20010of those banded at Gardner Park. Percent immatures at Walker was similar to that during the initialS-years of banding. Only mallards were. banded at Walker, but 1 adult female blue-winged teal (Anas discors )/cinnamon teal (Anas cyanoptera)was banded at Gardner Park. Band Reporting and Record Keeping All band numbers of newly banded birds and recaptures were submitted to the U. S. Fish and Wildlife Service's Bird Banding Laboratory on standard forms. Computer files containing the number of birds banded by area, site, day, age and sex were constructed at the Colorado Division of Wildlife's Research Center. �25 Table 1. Trapping locations during preseason banding in western Colorado, August- September, 1996. Wetland Name Location Colorado River WalkerSWA TIN, R2W, Sec 36, SWII4 Upper Yampa River Gardner Park Reservoir TIN, R86W, Sec 22, NE1I4 Table 2. Number of Mallards banded by area; site, age and sex during pre-season trapping in western Colorado, 1996 Age/Sex Area Site . AM AF 1M IF Totals Colorado River Walker SWA 68 18 143 133 362 Upper Yampa River Gardner Park Reservoir 2 14 3 1 20 70 32 146 134 382 Totals LITERATURE CITED Carney, S. M. 1964. Preliminary keys to waterfowl age and sex identification by means of wing plumage. U. S. Dep. Inter., Fish and Wildt. ServoSpec. Sci. Rep. - Wildt. 82. 47pp. Szymczak, M.R, and 1. F. Corey. 1976. Construction and use of the Salt Plains duck trap in Colorado. Colo. Div. Wildl., Div. Rep. 6. 13pp. . Weller; M W. 1976. Molts and plumages of waterfowl. Pages 34-38 in F. C. Bellrose. Ducks, geese and swans of North America. Stackpole Books, Harrisburg, Pa. Prepared by: ?n.....R_. L ~ ~ MichaclRSzymczak Researcher/Scientist IV ��27 Colorado Division Wildlife Research June 1997 of Wildlife Report JOB state of Colorado Project W-166-R Work Plan 1__: Job Title: Period Author: Avian Job Integrated Covered: PROGRESS Research REPORT - Migratory Game Bird Investigations 24 Waterbird 1 January Management 1996 through Studies 31 December 1996 James H. Gamrnonley Personnel: J. H. Gamrnonley, J. K. Ringelman, M. R. Szymczak, C. E. Braun, Claussen, M. A. Reddy, D. Pavlacky, B. st. George, Colorado Division of Wildlife; M. K. Laubhan, National Biological Service. M. ABSTRACT We studied foraging and nesting ecology of breeding waterbirds from 15 April to 3 July 1996 at Russell Lakes State Wildlife Area (RLSWA). During 4 15-day sampling periods, we measured habitat variables (water depth, conductivity~ and temperature; vegetation height, density, and species composition) in 330 randomly located plots within short emergent (SE), tall emergent (TE), shallow open water (SW), semipermanent open water (SPOW), saltgrass (Distichlis spicata) (SG), and upland shrub (US) cover types. We collected mallards (Anas platyrhynchos) (n = 8); redheads (Aythya americana) (4), cinnamon teal (Anas cyanoptera) (5), and American avocets (Recurvirostra americana) (2) for analysis of food habits and body cOhdition in relation to reproductive status; food samples were also taken at the site where each bird was collected to compare food use to availability at the scale of individual foraging locations. We also collected time-activity data for each of the above species and killdeer (Charadrius vociferus) and Wilson's phalarope (Phalaropus tricolor), and measured habitat variables at sites where birds foraged during each time budget bout. We conducted nest searches on all habitat plots and in other selected portions of RLSWA; habitat variables were measured at 268 nest sites and-nests were revisited to determine nest fates. Processing of food samples and data entry continued through December 1996. ��29 INTEGRATED WATERBIRD MANAGEMENT STUDIES P. N. OBJECTIVES 1. Map location of wetlands and wetland 2. Document hydrologic regime and water, characteristics of each wetland type. 3. Identify aquatic invertebrates associated with each wetland community, and document seasonal trends in invertebrate diversity, abundance and biomass. 4. Quantify the abundance, spatial and temporal use patterns, behaviors, and food habits of waterbirds in different wetland types. Relate the dynamics of endogenous lipid and protein reserves to food habits and migration and breeding ecology. 5. Determine seasonal wetland habitat requirements for all waterbirds, and consolidate these needs into a conceptual design for an optimum wetland community. 6. Develop the water management protocol and wetland development guidelines needed to produce the optimum wetland community. Prepare wetland development and water management plan for the RLSWA. SEGMENT communities on the RLSWA. soil and vegetation a OBJECTIVES 1a. Map cover types (short emergent, tall emergent, shallow open water, semi-permanent open water, saltgrass (Distichlis spicata), and upland shrub) and other landscape features (roads, ditches, gravel pits, and parking areas) at RLSWA, and digitize into a GIS format. lb. Randomly overlay a grid representing 0.25-ha plots on the GIS map. Select a stratified random sample of plots in each cover type (total n 330) . 2a. Randomly select a subset (n = 15) of plots in each cover type for invertebrate and seed sampling. During a one-week period, collect seed and invertebrate satnples and determine density and biomass of food items at random sites in each plot. Collect 3 replicate samples each from the benthos, water column, and vegetation at each site. Repeat sampling every 3 weeks from April to July. 2b. During a 2-week period, use focal sampling (Altmann 1974, Tacha et al. 1985) to determine activities of mallards (Anas platyrhynchos), redheads (Aythya americana), cinnamon teal (Anas cyanoptera), American avocets (Recurvirostra americana), killdeer (charadrius vociferus), and Wilson's phalaropes (Phalaropus tricolor). Repeat sampling with I-week breaks between sampling periods, 15 April to 5 July. 2c. During each 2-week sampling period, search each selected plot (1b) for waterbird nests. Map the location of each nest on aerial photos and return at the expected hatch date to determine nest success. Obtain GPS coordinates of each nest and enter on GIS map (la). �30 3. On the first and last day of each 2-week sampling period, use linetransect sampling (Buckland et al. 1993, Laake et al. 1994) to estimate densities of each species in 2b within each cover type. Fourteen northsouth transects spaced at 440 m intervals across RLSWA will be traversed during each count; total transect length i~ 38.4 km. 4. During each 2-week sampling period, determine food habits of species in 2b by collecting actively foraging birds and measuring the numbers and biomass of food in esophageal contents. Determine reproductive status (pre-laying, laying, incubating, post-breeding), molt intensity, and nutrient composition of each collected bird. Collect food samples from collection sites using procedures in 2a. 5. At collection sites (4), nest sites (2c), focal time-budget sites (2b), food sampling sites (2a), and random sites in each selected 0.25-ha plot (1b), measure the following habitat variables, where appropriate, during each sampling period: water depth, conductivity, and temperature; cover type; plant species; and vegetation height and visual obstruction (Robel et al. 1970). Use habitat variable measures to estimate the availability of each cover type for a) foraging and b) nesting waterbirds of each species listed in 2b. 6a. Determine food selection by species listed in 2b by comparing (density, biomass) esophageal contents to foods at collection sites (4), and at random sites in each cover type (2a). 6b. Determine foraging habitat selection by species listed in 2b by comparing habitat variables at collection sites and focal observation sites to habitat variables at random sites (5). Compare food density and biomass between collection sites and random sampling sites, and among cover types (2a and 4). 6c. Determine nesting habitat selection by species listed in 2b by comparing habitat variables at nest sites to habitat variables at random sites (5), and by comparing nest densities in each cover type to the availability of each cover type (2c and 5). Compare nest success among cover types (2c), and in relation to habitat variables (2c and 5). INTRODUCTION The San Luis Valley (SLV) is one of the most important breeding areas for waterbirds in Colorado (Ryder et ale 1979, CDOW 1989, Nelson and Carter 1990, Gilbert et ale 1996). A wetland ecosystem can be managed for habitats that maximize requirements for a narrow group of avian species, or for. more diverse habitats that optimize resources for a variety of avian species. The latter approach of integrated waterbird management better fits the philosophy of increased emphasis on managing landscapes for species diversity. One goal of the Colorado State Waterfowl Management Plan is to provide habitat of sufficient quality to maintain duck and goose populations at desired levels for maximum recreational opportunities (CDOW 19.89). In addition, the SLV draft management plan for waterbirds recommends maintenance of diverse wetland habitats with 25% of the actively managed habitat on public lands managed for nongame waterbirds (Olterman 1993). In 1994, CDOW, in cooperation with NBS, initiated a study to examine resource use by both game and nongame waterbird species breeding at Russell Lakes State Wildlife Area (RLSWA). STUDY AREA We studied Saguache County. waterbird ecology at RLSWA, a wetland complex in the SLY in We categorized habitats (cover types) at RLSWA according to �31 hydrology (flooding depth and duration) and vegetation structure as follows: short emergent (SE), tall emergent (TE), shallowly flooded open sites with no emergent vegetation (SW), semi-permanently flooded sites with no emergent vegetation (SP), saltgrass (SG), and upland shrub (US); we also delineated alkali flats and gravel areas. We created a GIS map of RLSWA based on these cover types, using aerial photos (scale = 1:4,000) taken on 5.May 1994 (cover type designations were ground-checked during summer 1994). The 1,239 ha included in the study area were apportioned as follows: SE, 296 ha (24%); TE, 185 ha (15%); SW, 51 ha (4%); SPOW, 192 ha (15%); SG, 58 ha (5%); US, 446 ha (36%); alkali flat, 4 ha (0.3%); gravel, 7 ha (0.6%). Our study focused ori mallards, redheads, cinnamon teal, American avocets, killdeer, and Wilson's phalaropes. Other waterbird species were included in line-transect counts and nest monitoring. METHODS Cover types were deline~ted and mapped in a GIS. A grid of 0.25-ha square plots was randomly placed over the GIS map. Each plot was categorized according to its dominant (>50%) cover type, and a random sample of plots classified as SE, TE, SW, SPOW, SG, and US was selected for habitat sampling (total n = 330). Field work was conducted during 4 15-day sampling periods; 1 week separated the last day of a sampling period and the first day of the next sampling period. During each sampling period, we measured water depth, conductivity, and temperature, and vegetation height, density (Robel et al. 1970), and species compo~ition (hereafter referred to as habitat variables) at random locations in each 0.25-ha plot. On the first and last day of each sampling period, we used 14 northsouth line transects located 400 m apart, to census waterbirds on RLSWA. As observers walked each transect line, they recorded the species, sex (when possible), flock size, perpendicular distance from the transect line, and the cover type from which waterbirds flushed. We used program DISTANCE (Buckland et al. 1993, Laake et al. 1994) to estimate densities of all waterbird species for which we had adequate sample sizes. We shot foraging waterbirds during each sampling period. We observed foraging birds for >20 minutes prior to collection, to ensure that individuals contained food items obtained at the collection site, and to determine the pair status of collected birds. Immediately following collection, the esophagus of each bird was removed and stored iri 95% ethanol. Birds were weighed and stored in a freezer for later processing .. In the laboratory, we identified and sorted food items contained in the esophageal contents of each bird. Each food item was dried (60 C) and measured to the nearest 0.1 mg. We also collected food samples in the water column, vegetation, and benthos, and measured habitat variables at the site where each bird was collected to compare food and foraging habitat use to availability at the scale of individual foraging locations. We used focal sampling (Altmann 1974, Tacha et al. 1985) to determine the diurnal activities of selected waterbird species in each cover type. During each 10-minute focal session, observers recorded each time the subject changed its activity (resting, preening, locomotion, foraging, alert, or aggression) or cover type. We measured habitat variables at locations where birds foraged during time-budget bouts. During .each sampling period, we searched all selected plots for waterbird nests. We also searched other selected portions of RLSWA to increase sample sizes. Habitat variables were measured, and the location of each nest was marked on an aerial photo and later digitized on the GIS map. We revisited each nest near its expected hatch date to determine its fate. A nest was classified as successful if >1 egg hatched. Unsuccessful nests were categorized as deserted, avian predation, mammalian predation, unknown predation, or other. �32 RESULTS Seasonal hydrologic patterns at RLSWA were similar between 1995 and 1996 (Table I), with the area more extensively flooded during April 1996 than in 1995 and generally drier during June 1996 than in 1995 (Fig. 1). Most cover types provided a diversity of water depths; percent of the total area in each habitat that was flooded at desirable foraging depths (1-15 cm) for most of our study species ranged from 37-54% in SG, 54-65% in SE, 19-22% in SPOW, 4958% in SW, 51-55% in TE, to 31-37% in US (Table 1). Based on these estimates, the availability of potential foraging habitat (based only on water depth) was greatest each year in SE (160-192 hal, followed by US (138-165 hal, TE (94-102 hal, SPOW (36-42 hal, SW (25-30 hal, and SG (21-31 ha). Each year, 32-52% of SG and 59-65% of US habitats were not flooded, and therefore not available for most foraging waterbirds except killdeer (Table 1). Waterbird abundance varied seasonally and among years. For individual species, peak densities on the study area were about 1.2 mallards/ha, 2.2 cinnamon teal/ha, 0.6 gadwalls (Anas strepera)/ha, 0.25 redheads/ha, 0.5 American avocets/ha, 0.4 killdeer/ha, 0.3 Wilson's phalaropes/ha, and 1.2 white-faced ibis (Plegadis chihi)!ha (Fig.2). Densities of most of these species were generally highest in SW habitats (Fig. 2). We collected 9 mallards, ·4 redheads, 5 cinnamon teal, and 2 American avocets in 1996; emphasis was placed on collecting birds foraging in SE habitats, to increase sample sizes for this cover type. In general, gut contents varied widely among individuals of each species. Total sample of individuals used for food habits analyses (i.e., individuals with >0.1 mg [dry wt.] of food in the esophagus) 1994-96 and the mean (SO) percent dry weight of esophageal contents comprised of invertebrates was as follows: 36 mallards, 38.4 (30.8); 30 redheads, 10.4 (15.6); 53 cinnamon teal, 53.4 (39.4); 51 American avocets, 95.8 (10.9); 44 killdeer, 99.3 (3.5); 50 Wilson's phalaropes, 95.8 (18.2). Waterbird dissections and carcass analyses will be completed in 1997. Food habits will be analyzed in relation to species, sex, reproductive status, and body condition. Diet composition of collected waterbirds will be compared to composition of 1) foods available at the collection site and 2) foods available at random locations in foraging habitats used by each waterbird species, to examine food selection by species, sex, and reproductive status. Carcass lipid, protein, and mineral content will be compared among birds in different reproductive categories for each species. External body measurements will be used to correct for variation in nutrient composition due to individual differences in structural size. Food items collected from random locations in habitat plots and from waterbird collection sites were identified, sorted, and weighed in 1996. Data from random sites will be analyzed to examine differences among habitat types and collection periods in taxonomic composition and biomass of foods consumed by foraging waterbirds. We collected 442 focal sessions (73.7 hr) of time-activity data in 1996. Time budget data will be analyzed to determine the effects of cover type, month, time of day, and social st.a't.us on acti vi ties of each species. Percent time spent foraging in each cover type will be combined with line-transect data to rank the relative use of each cover type as foraging habitat by each species. We monitored 241 nests of 13 'spec i es in 1995. Most waterbird species nested earlier in 1996 than in 1995; average nest initiation dates differed significantly (t-tests) between years for American avocets (P = 0.009), American coots (P = 0.01), gadwall (P = 0.003), and mallards (P = 0.007), but not for cinnamon teal (P = 0.11), killdeer (P = 0.08) r . and redheads (P = 0.98) (Fig. 3). Mayfield estimates of nest success and daily survival rates of nests will be calculated. Habitat measurements were recorded and nest locations have been plotted on the GIS map; nest success will be examined in �33 relation nests. to habitat variables at the nest PLANS site and the spatial location of FOR 1997 Analyses of data collected during 1994-96 will continue. Habitat manipulations will be conducted on selected areas at RLSWA to 1) identify the water management capabilities at RLSWA, and 2) test whether waterbirds respond to changes in conditions as predicted based on results from 1994-96. Similar habitat experiments will continue on RLSWA and other managed wetland areas in Colorado through 1999. LITERATURE Altmann, J. 1974. Observational Behaviour 49:227-267. study CITED of behaviour: sampling methods. Buckland, S. T., D. R. Anderson, K. P. Burnham, and J. L. Laake. 1993. Distance sampling: estimating abundance of biological popu.Lat.i.ons . Chapman and Hall, London, U.K. 327pp. Colorado Division of Wildlife. 1989. Colorado statewide waterfowl management plan 198·9 - 2003. Colorado Div. Wildl., Terrestrial Wildl. Sect., Migratory Game Bird Program Unit. 97pp. Gilbert, D. W., D. R. Anderson, Response of nesting ducks National Wildlife Refuge. Laake, J. K. Ringelman, and M. R. Szymczak. 1996. to habitat and management on the Monte Vista Wildl. Monogr. 131. 44pp. J. L~, S.T. Buckland, D. R. Anderson, and K. P. Burnham. 1994. DISTANCE user's guide, version 2.1. Colorado Coop. Fish and Wildl. Unit, Colorado State Univ., Fort Collins. 84pp. Nelson, D. L., and M. F. Carter. 1990. Birds of selected wetlands Luis Valley. Colorado Div. Wildl., unpubl. Rep. 40pp. Olterman, J., ed. 1993. The San Luis valley Wildl., Unpubl. Rep. Robel, R~ J., J. N. Briggs, Relationships between grassland vegetation. waterbird of .the San Colorado A. D. Dayton, and L. C. Hulbert. 1970. visual obstruction measurements and weight J. Range Manage. 23: 29.5-297. Ryder, R. A., W. D. Graul, and G. C. Miller. movements of ciconiforms in ColQrado. Conf. 3:49-58. Tacha, T. C., P. A. Vohs, and G. C. Iverson. 1985. and continuous sampling methods for behavioral Ornithol. 56:258-264. Prepared plan. by: ~~ /James H. GammOnley Researcher/Scientist III Res. Div. of 1979. Status, distribution, and Proc. Colonial Waterbird Group A comparison of interval observations. J. Field �34 Table 1. that were Percent flooded of total area (ha) of each 6 cover in different depth categories. Percent Cover type SG Month Year Dry <5 5-15 April 1995 50.9 21.4 18.1 1996 32.2 33.3 1995 43.6 1996 (58 hal June SE April (296 hal June SPOW April (192 hal June SW April (51 hal -June TE April (185 hal June US hectares April (446 hal June by water types depth at RLSWA category (cm) 30-46 46-76 6.8 2.3 0.4 0.0 20.5 10.2 3.1 0.6 0.0 25.5 22.3 6.1 1.9 0.5 0.1 51. 6 18.9 18.8 8.8 1.3 0.5 0.0 1995 26.0 37.3 22.6 9.8 2.8 1.4 0.1 1996 21.2 33.5 29.2 11. 4 3.7 0.9 0.1 1995 22.8 38.0 27.4 8.6 2.1 0.9 0.1 1996 30.5 30.1 23.8 10.9 3.5 1.3 0.0 1995 11. 4 9.6 10.5 7.3 5.4 15.5 39.9 1996 9.4 9.0 10.2 8.4 7.5 15.5 40.0 1995 9.2 11.3 11.1 7.8 5.5 15.5 39.6 1996 12.3 8.6 10.7 7.6 6.1 15.7 30.0 1995 28.4 28.5 25.0 11.9 2.9 3.3 0.0 1996 27.5 24.9 22.6 15.7 5.9 3.2 0.2 1995 27.1 31.7 27.0 9.1 2.9 2.1 0.0 1996 30.8 24.2 25.1 14.4 2.9 2.6 0.0 1995 20.6 24.8 27.8 13.8 3.5 6.7 2.7 1996 16.0 21.5 30.4 15.7 6.5 7.0 2.9 1995 17.7 24.2 30.9 14.5 3.7 6.3 2.7 1996 21.6 24.6 27.0 13.6 4.3 6.2 2.7 1995 61.2 24.8 10.4 2.9 0.5 0.1 0.0 1996 60.0 22.0 12.9 4.0 1.0 0.1 0.0 1995 58.8 24.8 12.3 3.0 0.8 0.3 0.0 1996 65.0 19.3 11. 8 3.2 5.9 0.1 0.0 15-30 >76 �35 Figure legends Fig. 1. Distribution of different April and June, 1995 and 1996. water depth categories at RLSWA during Fig. 2. Densities of waterbirds at RLSWA estimated from line-transect counts during 1994-96. For each species, density estimates and standard error bars are displayed for the entire study area (all cover types), and for each cover type. Fig. 3. Box plots of nest initiation dates for American avocets (AMAV), American coots (Fulica americana) (AMCO), cinnamon teal (CITE), gadwall (GADW), killdeer (KILL), mallards (MALL), redheads (REDH), and Wilson's phalaropes (WIPH) at RLSWA, 1995 and 1996. �Russell Lakes SWA hj ,.... t.O s:: i'i CD . I-' Water Depth II >75 em (> 30 in) April 1995 June 1995 . • 45-75 em (18-30 in) • 30-45 em (12-18 in) • 15-30 em (6-12 in)· III 5-15 em (2-6 in) . D >0-5 em (>0-2 in) IINot Flooded April 1996 June 1996 \H 0\ �00000 ..•...•...•...•. 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Job Mi2ratocy Game Birds Investigations _j_ Job Title: Cooperative Mana2emeot Pro2rams Period Covered: 01 January through 31 December 1996 Author: Michael R. Szymczak Personnel: James H. Gammonley and Michael R. Szymczak, Colorado Division of Wildlife. ABSTRACT Recommendations for wetland habitat improvements and/or management were provided for public and private land managers across the state. Proposals for funding projects with Duck Stamp monies were evaluated and rated. Presentations on wetland ecology in relation to waterfowl were given at workshops. Work with the Wetland Focus Area Committees continued in relation to wetland project development proposals for the Great Outdoors Colorado Wetland Legacy grant, the Intermountain West Joint Venture, and the Playa Lakes Joint Venture. Responsibilities as Colorado's representative on Pacific Flyway Study Committee and Council and Central Flyway Technical Committee, including Pacific Flyway Consultant to the U. S. Fish and Wildlife Service's Regulation Committee were fulfilled. A 3:..5 year cooperative post-breeding goose banding program was continued in the Durango areas and the upper Gunnison Basin. ��59 COOPERATIVE MIGRATORY BIRD MANAGEMENT PROGRAMS In 1988, the Colorado Division of Wildlife (CDOW) created the Migratory Game Bird Program Unit (MBPUYwithin the Terrestrial Wildlife Section. This administrative change combined all individuals having statewide responsibilities for research and management of migratory game birds. Members of the MBPU worked in concert to improve migratory bird management in Colorado. This job was created to allow team members to participate in these management programs. In November 1993, project personnel assumed additional ' responsibility for leading and administering the Duck Stamp wetland development program. Since 1993, personnel of the MBPU have taken on additional responsibilities with wetland programs in Colorado. In July 1996, the MBPU was dissolved with most of the responsibilities being transferred to the Avian Program. This report covers activities of the migratory bird segment of the Avian Program. P.N.DBJECTI\TES 1. Continue to aid the formation of Wetland Focus AreaCommittees in the state, monitor the functioning of these committees, and aid in obtaining funds for proposed projects by serving as chairman of the state-wide oversight committee and state coordinator for the Intermountain West Joint Venture. 2. . Advise public land management agency personnel, including area biologists and district wildlife managers, on the potential benefits to migratory birds of acquisition and/or development of wetland areas. Activities may include on-site inspection, formulating plans for the collection of biological data, recommending wetland enhancement developments, or review of development plans. Provide information on wetland habitat development to private land managers. 3. 'Prepare and present information programs on the principles of migratory bird management. Preparation may include literature review, construction of charts, graphs, . and tables as photographic slides or posters, and rehearsal of presentations. 4. Attend flyway Technical Committee and Council meetings and flyway special workshops as assigned. Compile information on current status of Colorado's migratory bird population and hunting season results through consultation with CDOW biologists and managers. Prepare reports for presentation at meetings and workshops. Serve on committees as assigned by flyway Technical Committee and/or Council Chairman. Attend selected meetings in Colorado that address migratory bird management programs, at which in-depth biological expertise would be of value. 5. Provide methodology to migratory birds and wetland managers for sampling the biological parameters of interest. Literature review may be required to develop appropriate methodology. Parameters of interest may include breeding pairs, nesting densities, nesting success, fledging success, vegetation composition and density, invertebrate composition and density, or population survival. . . �60 6. Assist in collecting information that will enable waterfowl and wetland managers to make decisions on population and wetland management. SEGMENT 0BJECI1VES 1. Use the knowledge and skills of the federal aid supported members of the Migratory Bird Unit to facilitate wetland and waterfowl management and informational programs within Colorado. a. Continue to aid the formation of Wetland Focus Area Committees in the state, monitor the functioning of these committees, and aid in obtaining funds for proposed projects by serving as chairman of the state-wide oversight committee and state coordinator for the Intermountain West Joint Venture. b. Advise public land management agency personnel, including area biologists and district wildlife managers, on the potential benefits to migratory birds of acquisition and/or development of wetland areas. Activities may include on-site inspection, formulating plans for the collection of biological data, recommending wetland enhancement developments, or review of development plans. Provide information on wetland habitat development to private land managers. c. Prepare and present information programs· on the principles of migratory bird management. Preparation may include literature review, construction of charts, graphs, and tables as photographic slides or Posters, and rehearsal of presentations. d. Attend flyway Technical Committee and Council meetings and flyway special workshops as assigned. Compile information on current status of Colorado's migratory bird population and hunting season results through consultation with CDOW biologists and managers. Prepare reports for presentation at meetings and workshops. Serve on committees as assigned by flyway Technical Committee and/or Council Chairman. Attend selected meetings in Colorado that address migratory bird management programs, at which in-depth biological expertise would be of value. e. Provide methodology to migratory birds and wetland managers for sampling the biological parameters of interest. Literature review may be required to develop appropriate methodology. Parameters of interest may include breeding pairs, nesting densities, nesting success, fledging success, vegetation composition and density, invertebrate composition and density, or population survival. f. Assist with Canada goose trapping and banding in the Durango area, and the upper Gunnison Basin. �61 RESULTS The Wetland Initiatiye, Wetland Focus Area Committees, and Waterfowl Habitat PrQject Reyiew Committee (WHPRC) Actiyjties Through the state-wide Wetland Initiative Planning Grant over 200 wetland projects were identified. Project specific evaluation forms were completed for some projects, including some preliminary engineering information. As member of the Wetland Initiative project planning team, migratory bird researchers assisted in preparing the fmal grant application and developing project selection criteria. As chairman of the WHPRC, Szymczak chaired 1 committee meeting for ranking and funding proposals submitted for the 1996-97 funding year; informed proposal submittees of the outcome of their funding request; periodically monitored progress of project planning, construction, and money.for new and previous years funded projects; coordinated Site Specific Agreements and fund reimbursement with the Ducks Unlimited Inc. MARSH program; served as the Project Officer for wetland development contracts formulated with Ducks Unlimited, the Bureau of Land Management, and the U. S. Fish and Wildlife Service. Specific projects were funded along with non-project specific funding support for the USFWS Partners For Wildlife program. In addition, Szymczak initiated the formation of Wetland Focus Committees in northeast Colorado (South Platte River), southeast Colorado (Arkansas River), and along the Front Range of Colorado and attended Wetland Focus Committee meetings in the San Luis Valley (2), North Park (2), Southeast Colorado (2) Lower Colorado/Gunnison (2), Front Range (2) and Upper Gunnison (1). Szymczak visited existing and potential wetland sites and recommendations for development and/or management were made for: Roots Reservoir near Loma (private), Fort Collins Water Treatment Facility near Windsor, the CDOW Frank Easement near Windsor, and a Colorado Department of Transportation site near LOveland. State Wildlife Areas in South Park were also visited. Gammonley met with Rick Schnaderbeck (USFWS) and Murray Laubhan (USGS) to visit and discuss water management at the White Ranch, a new holding of the Alamosa-Monte Vista NWR complex. Recommendations were made to use available well water to emulate a ephemeral/temporary flooding regime that should enhance and maintain the saltgrass dominated wetland site for use by migrating shorebirds, cranes, and waterfowl. Gammonley, Laubhan, and Schnaderbeck also visited sites on the Alamosa and Monte Vista NWRs, the Chiles property, the Rocky Mountain Bison Ranch, and several other wetland developments on private lands in the San Luis Valley to discuss management practices and recommendations for future developments and management scenarios. Informational Proerams A wetland workshop was organized and conducted for about 100 people. Presentations were directed at members of Wetland Focus Area Committees who were considering development or enhancing projects. A meeting agenda is included (Appendix A).. Gammonly participated in: (1) a work session to develop management objectives for the Alamosa-Monte Vista National Wildlife Refuge complex and provided information on waterbird ecology and wetland plant management principles in the San Luis Valley, and (2) a �62 planning session for continued ecological monitoring of the Hebron Ponds area (Bureau of Land Management) in North Park, discussing monitoring techniques for wetland vegetation and waterbird abundance and productivity. Waterfowl Technical Committee and Council Meetin2s The Iuly 1996 Technical/Study Committee and Council meetings for all Flyways were held jointly. Project personnel attended independent Central Flyway Waterfowl Technical Committee (CFWTC) and Pacific FlyWay Study Committee (PFSC) and Council sessions as well as all joint sessions. Waterfowl population status was reviewed along with characteristics of the. 1995-96 waterfowl hunting season harvest, and proposed 1996-97 hunting season recommendations were formulated and forwarded through the Council's to the USFWS Regulation Committee. In addition to the normal activities for Iuly Flyway meetings, the joint session enabled Technicians and Council members to discuss waterfowl issues that transcend flyway boundaries. At the winter meeting of the CFWTC in December attended by Gammonley, major items of direct relevance to Colorado included a review of analyses of neck collar observations of short-grass prairie population Canada geese, work on a revision of the management plan for the high-line population of Canada geese, and development of Central Flyway . recommendations for duck harvest regulation packages under the Adaptive Harvest Management (AHM) approach. . In addition to flyway meetings, Gammonley completed statistical analyses and began writing the final draft of the Evaluation of the High Plains Mallard Management Unit in the Central Flyway, Part B - Duck Wintering Populations, Harvest, and Hunting Effort. Completion of this report in 1997 is required by the u.s. Fish 'and Wildlife Service. He also reviewed and provided comments on the .Mourning Dove Management Plan, and served on a CMUTC subcommittee to review and rank proposals submitted for Migratory Shore and Upland Game Bird research grants. . In addition to PFSC duties, Szymczak was assigned as the Colorado representative on the Pacific Flyway Council. He attended the March 1996 Council meeting as Colorado's representative and was also responsible for representing the Pacific Flyway Council as a Consultant to the USFWS's Regulation Committee for the June and Iuly meetings of the .Service Regulations Committee. Council Consultants also serve on the AHM Task Force and Szymczak attended 1 Task Force Meeting .. Written reports to the Council were prepared for the Task Force and SRC meetings. C~ Canada Goose Bandin2 Canada geese were banded in the Durango-Bayfield area for the third consecutive year and in the upper Gunnison basin for the second consecutive year. The trapping operations were conducted with the cooperation and personnel of the CDOW West Region. A total of 97 goslings and 45 adults was banded at 5 locations in the Durango-Bayfield area, and 75 goslings and 50 adults were banded in the Gunnison area at 2 locations in late June 1996 (Appendix B). The total number banded in four years in the Durango-Bayfield area has been 517, with 220 banded in the Gunnison area. �63 . DISCUSSION Project personnel provide useful information in planning and evaluating waterfowl management and habitat enhancement programs in Colorado and educating bind management agency personnel about the habitat requirements of waterfowl. With increasing emphasis on wetland habitat in Colorado, and the initiation of new programs with expanded responsibilities for project personnel, wetland-related objectives of this job will receive more emphasis in the near future. Colorado will shortly have 11 Wetland Focus Area Committees functioning in the state that will require coordination and expertise in wetland project planning. The resources provided by project personnel will insure that money raised through the Colorado Duck Stamp program or any other funding initiative will be spent in accordance with the objectives of the program. , COnducting and/or formulating surveys and banding efforts and informing management agency personnel about various aspects of waterfowl and wetland ecology provides a valuable service to management agencies, the waterfowl resource and, in some cases, the hunting public. . Continued participation on.Pacific and Central Flyway committees ensures that Colorado will remain informed on migratory bird matters, have input in migratory bird hunting regulations, and influence habitat programs affecting migratory game birds. Prepared by: ?rJ.:.klt?~ Michael R. Szymczak Researcher/Scientist N ~ James H. Gammonley Researcher/Scientist ill �64 Table 1. Age, sex, number and band numbers of Canada geese banded on wetlands in the Durango-Bayfield area, June 1996. Age and sex Wetland LF LM AM AF Totals Band Numbers James Ranch 19 14 10 11 54 858-44501-521 858-11668-700 Rainbow Springs 11" 9 6 6 32 858-11522-533 Vallecito 6 13 5 1 25 858-1154-578 Tay-Col Cattle Company 3 "9 4 1 17 858-11579-595 Harpers Pond 7 6 "1 0 14 858-11596-600 858-11701-709 46 51 26 19 Totals 142 Table 2. Age,"sex, number and banding years of Canada geese banded on wetlands in the Durango-Bayfield area. Age and sex " Wetland AM LF LM AF Totals Years banded James Ranch 49 57 27 24 157 1994-96 O'Neal 31 41 29 27 " 128 1994-95 Vallecito 17 28 24 26 95 1994-96 Tay-Col Cattle Company 22 36 15 18 91 1994-96 Dove Ridge Subdivision 7 5 1 1 14 1994-95 11 9 6 6 32 1996 137 176 102 102 517 Rainbow Springs Totals �65 Table 3. Canada geese trapped and banded, by location, inthe Gunnison area, 1996. Age and sex Wetland LM AM LF AF Totals Band Numbers Wilson 29 31 16 12 88 858-11747-800 858-11801-834 Oleson 8 7 11 11 37 858-11710-746 Totals 37 38 27 23 125 Table 4. Age, sex, number and banding years of Canada geese banded on wetlands in the Gunnison area. Age and sex Wetland LM LF AM AF Totals Years banded Wilson 63 58 34 28 183 1995-96 Oleson 8 7 11 11 37 1996 71 65 45 39 220 Totals �ATTACHMENT ~ 66 WETLANDS WORKSHOP COLORADO WETLAND PARTNERSHIP May 22, 1996 8:30 AM - 4:30 PM Hunter Education Building Colorado Division of Wildlife 6060 Broadway Denver, CO 8:30 - 8:45 Welcome and Introductions - Jim Ringelman, Colorado Division of Wildlife (CDOW) 8:45 - 9: 15 Functions, Values, and Types of Wetlands in Colorado - John Sanderson, Colorado Natural Heritage Program 9: 15 - 9:45 Threatened and Endangered Species and the Clean Water Act ~ Bill Noonan, U.S. Fish and Wildlife Service/CDOW 9:45 - 10:05 Break 10:05 - 10:35 Conservation Easements and Wetland Protection - Heidi Sherk, The Nature Conservancy 10:35 - 11:20 Colorado Water Law - Grady McNeill, CDOW and Steve Sims, Colorado Attorney Generals Office . 11:20 - 11:50 Seasonal Requirements of Waterfowl in Colorado - Jim Ringelman 11:50 - 1:20 Lunch 1:20 - 1:50 Ecology and Management of Wetland Wildlife Species - Pat Magee, CDOW 1:50 - 2:20 Wetland Creation and Enhancement Techniques - Rick Schnaderbeck, U.S. Fish and Wildlife Service and Bob Sanders, CDOW 2:20 - 2:40 Break 2:40 - 3:10 Funding Sources for Wetland Creation and Enhancement - Jim Ringelman 3: 10 - 3:40 Combining Resources, Partnerships, and the Process of Delivering Wetland Projects - Mike Szymczak, CDOW 3:40 - 4:30 Discussion and Closing Remarks �67 JOB PROGRESS state of Colorado Project W-166-R . Job REPORT Migratory Game Birds Investigations Work Plan 22 Job Title: Migratory Game Bird Publications Period Covered: Author: 2 01 January through 31 December 1996 Michael R. Szymczak Personnel: James H. Gammonley and Michael Colorado Division of wildlife R. ABSTRACT No articles were published during this segment. Prepared by: ~P?~~ Michael R. Szymczak Researcher/Scientist IV Szymczak, ��69 JOB PROGRESS REPORT State of: Colorado Project: W-167-R Work Plan: __j_: Job _2i_ Avian Research Job Title: Evaluation of Habitat Deyelopment for Ring-necked Pheasants in Eastern Colorado Period Covered: 01 January through 31 December 1996 Authors: Thomas E. Remington and Warren D. Snyder Personnel: C. E. Braun, T. J. Davis, M. J. Emerson, M. A Et~ K. M. Giesen, E. T. Gorman, L. K. Haynes, R W. Hoffinan, K. D. Johnson, J. L. Mekelburg, K. L. Martin, M. L. Molarsky, T. E. Remington, W. D. Snyder, M. L. Trujillo, 1. D. Wieland, B. T. Weinmister, J. A Yost, D. 1. Younkin; Colorado Division of WIldlife. ABSTRACT Expenditures under the Pheasant Habitat Improvement Program (PHIP) increased from about .$299,000 in 1995 to $318,000 in 1996. Most habitat developments were either plum thickets (225 of 509), or sorghum food plots (175), although, as in past years, most (84%) PHIP expenditures went toward establishment of plum thickets. PHIP has now resulted in the establishment of754 plum thickets in 5 years. Above average moisture throughout the area improved plum and juniper seedling survival and improved the quality of sorghum food plots (vertical obstruction of 5.1 dm, and a vertical obstruction reading [VORl of 1.6-3.5). Average counts of crowing male ring-necked pheasants (phasianus colchicua) did not differ (f = 0.38) between treatment and control blocks. Counts averaged about 14 calls per station in 1993, 17 in 1994, 13 in 1995, and 14 in 1996. Hunter pressure was similar to 1995, while harvest rates increased about 78% to 0.14 birds per hour in 1996. Hunters were most successful when hunting sorghum food plots, where harvest rates were over 3 times higher than the average for other cover types. One hundred and seven hens were captured and radiomarked, while 45·hens radiomarked in 1994 or 1995 were recaptured and fitted with new radios. Sample size of birds available for survival estimates was 203. Annual survival of radio-marked hens (1 Nov - 31 Oct) was 23.8%, an increase from the 15% in 1995 but still substantially below the 41% survival in 1993-94. The intermediate survival was attributed to intermediate height and quality of wheat stubble and declining quality in Conservation Reserve Program (CRP) fields. No differences in survival between treatment and control blocks were apparent. Most mortality was due to avian (51%) or mammalian (41%) predation. About 12% of wheat in Phillips County was combined with stripper headers. Most stripped-wheat stubble was treated with herbicides (74%)· or mechanically cultivated which decreased its cover value to pheasants. About half of conventionally-harvested wheat stubble was sprayed with herbicides, undercut, or disced in the fall which essentially eliminated these fields as pheasant habitat. ��71 EVALUATION OF HABITAT DEVELOPMENT FOR RING-NECKED PHEASANTS IN EASTERN COLORADO Thomas E. Remington and Warren D. Snyder INTRODUCTION Pheasants are pursued by more hunters than any other small game species in Colorado (83-88% of small game license buyers). In a recent survey, 74% of pheasant hunters rated their hunting trips in Colorado as poor (45%) or fair (29010), while only 10% rated their trips as very good or excellent. Lack of birds and places to hunt were identified as the most significant reasons why some hunters did not hunt pheasants in Colorado. Small game license sales in Colorado have declined by about 90,000 (45%) in the last 10 years. It is apparent that if the Division ofWlldlife is going to tum this decline around, pheasants will be a key species. Presumably, recruitment and retention of hunters will increase if the quality of pheasant hunting is improved, i.e., increases in pheasant numbers and places to hunt. Previous research has indicated that over-winter survival of pheasants is the most critical factor limiting pheasant populations. The Pheasant Habitat Improvement Program (pIllP) was created to establish over-winter survival cover within historically good pheasant range in eastern Colorado. The program was conceptually designed to overcome significant obstacles to developing habitat, mainly a lack of . manpower and a burdensome contractual system (costs of administering contracts exceeded costs of developments). Under PmP, the Division ofWlldlife contracts with individual Pheasants Forever chapters in eastern Colorado to contact landowners and develop habitat on private lands following specific guidelines. Each chapter develops contracts with individual landowners and pays them when the habitat work is completed and verified. Division ofWlldlife personnel inspect habitat developments and verify completion and compliance with guidelines. A new method of combining wheat has potential to increase pheasant survival. Stripper headers strip the grain from the stalks rather than cutting and thrashing the stalks to separate the grain, which leaves much taller stubble. Pheasant survival from July through April is largely dependent upon the height and cover value of wheat stubble (Snyder 1985). We measured the height and cover value of paired blocks of stripped and conventionally-cut wheat to assess the value of stripped wheat for pheasants, and trapped and radiomarked a sample of hens in each area to evaluate impact of stripped wheat on pheasant survival. P. N. OBJECTIVES To determine ifhabitat developments offered through the Pheasant Habitat Improvement Program increase pheasant survival, breeding density, and pheasant harvest within selected northeast Colorado study areas. . �72 SEGMENT OBJECTIVES 1. Work with Pheasant Forever chapters, management personnel, and landowners to develop annual sorghum plantings, disturbance tillage, switch grass, or plum thickets and to develop stripped-wheat test plots in other locations within the primary pheasant range in northeast Colorado. 2. Monitor hen pheasants previously radiomarked with mortality-sensing transmitters within treatment and control blocks to compare survival, nesting success, and use of habitats through 1996. Trap and radiomark additional hens as necessary to obtain a sample of 100 hens respectively within stripped wheat and controls in fall 1996. 3. Conduct pheasant crowing counts within all treatment and control blocks during AprilMay 1996. 4. .Monitor hunting pressure, hunting success; and pheasant harvest within treatment and control sites and within stripped-wheat fields. 5. Conduct evaluations of the quality of annual plantings as survival cover. 6. Evaluate Pheasants Forever chapter and landowner acceptance of the program and consider modifications as suggested or needed. 7. Prepare an annual progress report. METHODS Procedures used during this work segment were described in previous progress reports (Remington and Snyder 1994, 1995, 1996). The PHIP Habitat Project Guidelines for participating Pheasants Forever chapters were modified slightly from 1995 and are attached (Appendix A). The most significant changes were requiring pre-approval of shrub thicket sites by CDOW personel, the addition of a payment rate for the use of2x12 inch staples to pin down weed barrier fabric in the tree-planting furrow in place of using dirt, and standardization of payment rates for thickets/windbreaks at $0. 561 linear ft. We shifted emphasis from evaluating impacts offorage sorghum on hen pheasant survival to evaluating impact of combining wheat using stripper headers (rather than conventional headers which cut the wheat stalk) on hen pheasant survival. Complexes of stripped-wheat fields were identified for use as treatment areas. Following wheat harvest in July 1996, we mapped all the wheat stubble in Phillips and Sedgwick counties where stripper headers were prevalent. Stubble was classified along road-side transects as to header type used and how the stubble was treated post-harvest; i.e., untreated, sprayed with herbicides, or undercut. We designed and build a walkin trap that could be used to trap hen pheasants in stripped-wheat fields because the stubble height was judged too tall to trap by conventional nightlighting techniques. Although the trap proved �73 quite successful when tested with game farm hens, wild hens would flush rather than run through the trap entrance. An alternate means to evaluate survival in stripped wheat fields was used. We trapped hens from CRP and conventionally-harvested stubble in areas we had trapped in previous years, and moved about half of them into stripped-wheat fields in stripped-wheat study areas. Research in Missouri (WIlson et al. 1992) indicated most. translocated hen pheasants stayed within 1-2 miles of their release point. Study areas and release points were established so that hens moving 1-2 miles would still have access to stripped-wheat fields. To reduce the potential for an age-related bias in survival estimates, we also moved almost half of hens surviving from 1994 (1 of 4) and 1995 (17 of 41) trapping efforts. RESULTS Rainfall and weather patterns The survival and quality of habitat developments and the quality of wheat stubble, CRP, and other cover types is ultimately dependant on the amount and timing of precipitation. Precipitation was deficient during the early part of 1996 and considerable wind erosion of winter wheat fields occurred, primarily in Phillips County in late winter (Table 1). However, rainfall was both adequate and well distributed through the growing season and into mid-October. No significant precipitation was received in November and December 1996. Annual precipitation averaged about 50 em (19.5 inches) among the 4 stations with a considerable variance among stations (Table 1). Most of the region received above average precipitation in 1996. Wheat harvest was delayed in many areas due to wet conditions in July. Planting of winter wheat was also delayed by wet field conditions in September. Nearly all ofPhilIips County was devastated by severe hail either in late Mayor in early August. Several radio-marked hens were killed by the August hail storm northeast ofPaoIi in Phillips County and in northwest Yuma County. Presumably mortality of broods also occurred in these areas. Table 1. Monthly precipitation (cm) received at 4 U.S. Weather Service stations in the Pheasant Habitat Improvement Program area, 1996. Yuma Holyoke Month 1.25 0.94 Jan o 0.28 Feb 2.67 2.44 Mar 2.82 3.12 Apr May 8.84 14.83 10.11 6.40 Jun S.21 12.90 Jul 7.26 14.45 Aug 9.45 8.13 Sep 0.61 0.69 Oct Nov No Significant Precipitation Received Dec No Significant Precipitation Received Totals 64.13 48.27 Akron4E Sterling 0.64 0.25 2.87 1.19 10.44 7.34 7.75 7.47· 8.66 0.99 2.26 47.60 38.05 o 1.70 1.27 8.00 6.20 5.39 5.21 7.26 0.76 �74 Seven Pheasants Foreverchapters in eastern Colorado participated in the Pheasant Habitat Improvement Program (plllP) in 1996 and received funds from the Division of Wildlife for habitat development. The RingneckRednecks (Morgan Co.) Chapter requested and received funds for the first time. The 7 chapters began the 1996 growing season with $381,480.64 in PlllP funds. They used $317,853.23 (83%) of that amount in habitat work (Table 2), leaving a balance of $63,627.41 to be carried to 1997 .. Expenditures have risen each year since the Program was initiated in 1992 (Table 3), illustrating continued popularity ofPlllP with Pheasants Forever chapters and landowners. Shrub thickets and windbreaks Shrub thicket plantings continued to be the most popular and most expensive habitat component; 225 were planted in 1996 bringing the 5-year total to 754 (Table 4). Most were planted by volunteers from Pheasants Forever chapters. All Washington County plantings were subcontracted to local Future Farmers of America (FF A) and Young Farmer chapters. Yuma County plantings were completed by an FF A Chapter and Yuma County Soil Conservation District personnel. About 30 sites within Phillips and Logan counties were sub-contracted to a professional tree planter. An Explorer Scout Troop from Sterling assisted the Northeastern Colorado (Sterling) Chapter. Survival and growth of seedlings on most sites has been good to excellent. Plumsin many sites that were planted during 1992 and 1993 are root sprouting between the rows to begin forming closed-canopy thickets. Survival of Rocky Mountain junipers has been even higher than that of plums at most sites. Over 200 shrub sites were inspected during fall/winter of 1995-96 to assess where fair to poor survival mandated replanting. Replanting of junipers was completed on most sites where it was needed; but time and manpower constraints prevented replanting plums on all sites. Replanting will be intensified in 1997 to improve sites while they are still young. To facilitate this replanting effort, a letter and evaluation form was sent to all landowners with thickets on their property asking them to inspect their thicket/windbreak for replanting needs or fabric repair, etc., and return the evaluation form to us. A d-base file was created of all cooperating landowners which detailed all habitat developments. Replanting needs were added to this database. Shrub survival was good to excellent on nearly all the 1996 sites. Rocky Mountain juniper seedlings apparently froze before being delivered and planted in Kit Carson County and will be replaced by the Colorado State Forest Service in 1997. A small percentage of plums died after planting or were dead when planted on most other sites. Heavy rains, reportedly as high as 20 em, fell within a few hours in mid-September and caused extensive flooding along several intermittent drainages, primarily in southeastern Logan County. Flood waters washed over at least a dozen shrub sites and washed out part of the weed-barrier fabric on a few. Repairs were completed on most in fall 1996; the remainder will be completed in spring 1997. Simazine herbicide (Princep") was applied by personnel hired by the Phillips County Chapter of Pheasants Forever in early spring 1996 to about 100 new and previously planted shrub sites in Phillips County and to 3 plantings in eastern Logan County. This persistent herbicide was applied at a rate of 4 quarts of the 4L formulation/acre (9.3 Vha) to 2, 3-dm wide bands along the outside �75 edges of the weed barrier fabric to suppress annual weeds. Weed growth was retarded on most sites but was not eliminated. A higher application rate might more effectively suppress weed growth and should be evaluated. Weeds along fabric edges were mowed on over 50 shrub sites, primarily 1996 sites, in mid- to late-summer, within Phillips, Sedgwick, Logan, and northern Washington counties. Glyphosate (RoundupR) was applied with a wick applicator(I:7 mixture; sponge applicator from H.G. Hank Schnider, Inc., Hugo, MN) to weeds and grasses growing through the slit in the weed-barrier around plums and junipers in numerous shrub sites, primarily in Logan and Phillips counties. Wicking with Roundup was effective at killing weeds and grasses and suppressing competition with seedlings. The wick applicators used had plastic valves and other plastic components which did not hold up to sustained use. We should develop or find a commercial source for a more durable wick applicator. Sor:bum food/coyer plots Sorghum food/cover plots were planted on 175 sites totaling 660 acres in 1996 (Table 5). Division personnel planted about 70 of these within the Mailander, Kurtzer, Fleming, Pauli, and Clarkville treatment blocks. Pelletized nitrogen (46% a.i.) was applied prior to the final tillage and. disking and these tracts were planted using surface planters with 30-inch row spacings. All sites were subsequently cultivated to reduce grass and weed competition. Germination and early growth was good to excellent in all plots. Hail severely impacted plantings on the Mailander block (for the second consecutive year), and moderately impacted plots in the Kurtzer block in early August. The other 3 blocks received less severe hail which caused little damage. Late summer rains stimulated extensive regrowth of hailed sorghum but the stalks were not strong enough to withstand high (near 100 km/hr) winds that occurred one day in October. Those winds broke the stems on much of the taller sorghum but most, especially the grain sorghum, remained standing to provide fair to good cover and food for pheasants. Plots in areas that were not impacted by hail, especially in the Fleming, Pauli, and Clarkville areas, grew well and provided good cover for pheasants (Table 5). Sorghum plots.planted by landowners using drills with a 12inch row spacing could not be cultivated with our equipment. These plots were invaded by annual grasses, and most developed into dense stands of stunted sorghum. Cover value was poor, particularly since much of the sorghum in these stunted plots blew over during the October wind storm. Only 10 sites were retained in disturbance tillage (annual weeds), however most of these developed into excellent stands of tall annual weeds. �76 Table 2. Pheasant habitat planted and/or contracted by Pheasants Forever chapters during 1996 in northeastern and east-central Colorado through the Pheasant Habitat Improvement Program. Habitat/Contract SarPum & Oth~c Eaad :flats Washington County Phillips County .Frenchman Creek (Fleming) Yuma County N.E. Colorado (Sterling) Ringneck Rednecks (Morgan) Subtotal SwitchgmSS :flantin~s Washington County N. E. Colorado (Sterling) Phillips County Frenchman Creek (Fleming) Yuma County Subtotal Disturhan~ Iillag~ (Annual Earhs) Phillips County Yuma County N. E. Colorado (Sterling) Subtotal Plantin~s 12 42 12 85 13 11 Numh~ Acres 32.5 135.0 43.8 380.0 54.5 .l.S..Q 175 660.8 7 2 44 9 25.5 6.0 126.8 23.6 ~ .l2Q..Q. 102 301.9 1 6 10.9 43.0 .a .l1.Q 10 70.9 52 54 40 32 19 17 31.5 28.5 19.5 16.9 9.4 9.3 Shrub Thi~kets and Windbceaks Washington County Phillips County Yuma County Frenchman Creek (Fleming) N. E. Colorado (Sterling) Ringneck Rednecks (Morgan) East Central (Kit Carson) Subtotal 225 123.1 Total Plantings/acres 512 1,156.7 11 __8jl Replantin~ Seedlings, Custam Work, Eertilizer, etc. Washington County N. E. Colorado (Sterling) Phillips County Ringneck Rednecks (Morgan) Subtotal Total Expenditures Payment ($) 1,300~00 5,257.00 1,752.00 14,220.00 2,075.00 559.00 25,163.00 1,275.00 240.00 5,332.50 1,150.00 . 4,800.00 12,797.50 377.00 1,720.00 470.00 2,567.00 67,309.81 62,087.55 50,457.20 36,075.21 19,059.37 20,866.51 10,J20AO 266,176.05 3,791.31 516.65 6,679.72 162.00 11,149.68 $317,853.23 �77 Tabl. 3. Ph••• ant Habitat Improvement Program expenditure. ($) by habitat type and Pheasant. Forever Chapter, northeastern COlorado, 1992-96. Habitat/Chapter Plum Thickets/Windbreaks Phillips Yuma Washington Sterling Fleming Burlington Horgan Subtotal 1992 1993 1994 1995 1996 Totals 23,454 8,730 59,661 27,525 41,760 11,992 52,982 36,992 58,030 23,263 5,556 12,319 77,866 47,083 65,513 16,642 9,888 17,738 146,494 183,586 234,730 62,088 50,457 67,513 19,059 36,075 10,320 20,867 266,176 276,051 170,787 232,613 77,079 45,963 45,933 20,867 869,293 2,800 1,676 20,030 76 754 1,692 5,160 150 .700 888 5,230 21,795 6,898 5,333 4,800 1,275 240 1.150 12,798 33,323 6,699 1,975 3,574 1.150 46,721 7,388 17,385 22,102 700 9,472 14,640 30,322 1,100 7,075 8,218 29,377 560 8,856 11,278 49,659 53,137 47,011 1,170 1,610 3,615 5,250 300 1.200 10,365 590 6,640 360 3,520 180 1.072 5,132 6,123 38,307 Switchgrass Plantings Phillips Yuma Washington. Sterling Fleming Subtotal Sorghum Plantings Phillips Yuma Washington Sterling Fleming Horgan Subtotal 360 3,530 pisturbance Tillage Phillips Yuma Wa.hington Sterling Subtotal Tall Wheat/No-Till Phillips Yuma Washington Sterling Subtotal 520 3,300 377 1,720 470 2,567 45,860 99,551 3,660 34,866 1,752 559 186,248 6,112 18,740 950 3,797 29,599 Wheat 1,069 1,000 2,069 CUstom Worka Phillips Yuma Washington Sterling Horgan Subtotal Totals 1.005 8,235 5,257 14,220 1,300 2,075 1·,752 559 25,163 $54,954 aReplanting shrubs, repairing 300 400 60 300 1,979 150 1,400 3,829 150 90 400 640 360 1,591 3,290 1,257 2,382 6,264 1,253 3,020 90 1,313 2,983 163 8,520 10,537 4,549 3,791 517 162 11,150 $221,028 $277,930 $298,680 $317,854 760 fabric, applying 360 fertilizer, 6,680 discing, etc. 14,625 5,856 11,051 3,062 162 34,756 $1,170,446 �78 Table 4. Number and type of plantings under the Pheasant Habitat Improvement Habitat/ Chapter Plum Thickets/Windbreaks· Phillips Yuma Washington Sterling Fleming Burlington Morgan Subtotal 1992 1993 1994 1995 1996 26 51 7 30 63 38 5 31 20 41 36 46 25 6 14 17 54 40 52 19 32 11 138 162 191 Tall Wheat/No-Till Phillips Yuma Washington Sterling Subtotal 15 _l1. 38 Totals 235 151 179 84 40 48 ~ 225 754 44 40 140 49 7 63 1 22 1 2 7 9 6 9 4 2 21 24 73 29 ~ 102 ~ 228 15 113 131 110 169 171 4 8 5 38 52 46 64 42 85 12 13 12 322 571 29 213 12 55 300 333 295 6 24 22 11 2 Disturbance T.illage Phillips Yuma Washington Ste.rlinq Subtotal 50 8 switchgrass Plantingsb Phillips . Yuma Washington Sterling Fleming Subtotal Sorghum Plantings Phillips Yuma Washington Sterling Fleming Morgan .Subtotal completed per Pheasants Forever chapter Proqram, northeastern COlorado, 1992-96. 5 9 19 14 2 7 55 5 ...ll _ll 175 1,158 1 6 1 -2. ___:J_ ___a _J 16 42 45 31 8 2 36 75 10 _2Q 7 141 Wheat 5 5 1 1 3 4 _J ~ 13 8 8 12 1 ~ 1 30 -Woody plantings consist of 0.1 to 0.3 acre plum thickets and most are accompanied by three-row juniper or juniper/plum windbreaks. ~any of the 1996 switchgrass plots had previously been sorghum plots that were seeded back to grass within CRP fields. �79 Table 5. Vegetation characteristics of sorghum plots planted within treatment blocks in 1996 and sampled in January-February 1997, northeastern Colorado. B~ight ~QB Treatment block H Kurtzer Pauli Fleming Clarkville Mailander 5 6 6 4 3 Means (dm) S.D. (dm) 1.29 2.49 3.47 2.15 1.58 0.30 0.71 0.71 0.42 0.09 2.88 4.98 5.90 5.10 3.23 0.52 0.98 0.76 0.62 0.40 2.20 0.85 4.42 1.30 S.D. QADg~ sorghum (l) Cg:i:~1: S.D. Forbs 26.8 33.8 33.4 38.0 36.0 15.3 9.0 4.2 12.4 0.8 29.5 20.4 25.5 20.5 22.3 15.0 8.8 7.2 10.8 12.2 33.6 4.2 23.6 3.9 S.D. Swjtch&rass Switchgrass was planted on numerous plots that had been planted to sorghum in previous years within Conservation Reserve Program (CRP) fields; Satisfactory stands were attained on nearly all of these, but herbicides were not used to reduce weed competition so growth was marginal. Switchgrass wasplanted in several other locations, with generally excellent establishment and first-year growth. Switchgrass was replanted on several tracts where satisfactory stands had not been attained in previous years. The Phillips County Chapter of Pheasants Forever contracted a private applicator to treat some previously-seeded switchgrass plots with herbicide to reduce weed competition. Coyer Yalue of Wheat Stubble Stripper headers were first used in Phillips County in 1992. In 1995, 12.4% of wheat harvested in Phillips County was combined with a stripper header. While stripped wheat has the potential to provide tremendous cover for pheasants (Remington and Snyder 1996), only 16.6% of stripped. wheat fields were left untreated after harvest. Most stripped-wheat fields were treated postharvest with either a herbicide application (74%), or by undercutting (9.3%) to control weeds. Landowners in this area purchased stripper headers primarily to facilitate a crop rotation where 2 crops are grown in 3 years. In this cropping system dryland corn or sunflowers are planted into the wheat stubble. A fall application of herbicides is used to conserve moisture for the spring crop. Herbicide application and undercutting both significantly reduce the cover value of stripped wheat for pheasants (Fig. 1), although both treatments still leave stubble with higher VORs and . height than does conventionally-combined stubble. The cover value of conventionally-cut wheat stubble has declined as semi-dwarfvarieties of wheat are planted almost exclusively, and as fall herbicide use or mechanical cultivation become more prevalent throughout northeastern Colorado. In Phillips County, about 52% of wheat combined with conventional headers was left untreated, while 28% was sprayed with herbicides and 20% was undercut or disced in fall. Semidwarf varieties of wheat seldom yield adequate stubble height for pheasant survival when harvested by modern combines; this problem is exacerbated when herbicides control weed growth that can mitigate short stubble height. The vertical obstruction reading (VOR) for chemicalfallowed, conventionally-cut stubble was less than half that of untreated stubble, and only onefifth that of untreated stripped-wheat stubble (Fig. 1). �80 4 a. 3.2 3 f- E "C 0:: 2 STRIPPED 0 2.1 1.9 w f-- ~ w ~ a:: 0 w I- 1 Z :::::> I- ~ 0... en CUT I:::::> 1.4 w w z w o a:: 0 :::::> 0 ~ a:: 06 I- Z :::::> a ~ D.. en 7 b. 6 f-- ........•. 5 f-- 5.5 E -0 ....._,. I- 4.8 4.3 4 f- :r: C> W :r: 3 - 2 f- 1 - 2.6 2.4 o Fig. 1. Visual obstruction readings (al VOR-dm) and height (bl dm) of wheat stubble harvested by stripper or conventional headers and subsequently treated herbicides, undercut, or untreated, February-March, 1997. with �81 Pheasant Crowim: Census Average counts of crowing males did not differ between treatment and control blocks (Table 6; f = 0.95 and 0.50 using high count of replicate counts and average of replicate counts, respectively). Counts, on average, increased slightly from about 13 calls per station in 1995 to about 14 in 1996., Table 6. Pheasant crowinq census data amonq treatment northeastern COlorado, sprinq 1996. and. control blocks, Count 1 Block ,2 3 Average of counce" Hiqhest count High countl station· Treatments Holyoke SE Mailander Kurtzer Clarkville Pauli Y-W Co. Line Fleming Kuntz otis CUrve 21.5 7.7 20.5 6.8 7.1 14.7 15.0 7.9 6.3 13.3 10.1 16.6 Means 30.0 21.5 7.7 16.9 8.5 7.-1 14.7 20.5 7.9 6.3 1~.3 ± 4.1 21.5 7.7 20.5 10.1 7.1 14.7 30.0 7.9 6.3 21.5 7.7 21.7 10.9 7.1 14.7 30.4 7.9 6.3 14.0 ± 4.4 14.2 ± 8~5 10.9 13.2. 16.1 12.6 24.8 10.9 15.1 10.3 13.8 10.9 13.2 16.9 12~6 31.2 10.9 15.2 10.3 13.8 controls Paoli HE Haxtun HE Paoli South St. Pete .Kelly Yuma CO. Lonestar Platner Washington W. 10.9 13.2 13.7 24.8 10.9 15.1 10.3 13.8 16.1 12.6 22.7 9.9 13.7 Means A Obtained 10.9 13.2 14.9 12.6 23.8 10.9 12.5 10.3 13.8 using the highest ± 4 1 count per station e. 14.2 ± 4.4 among counts before 15.0 ± 6.4 averaging. Bupter Pressure and Success During opening weekend 299 hunters were contacted and interviewed to ascertain their success. While this was a slight (7%) decline from 1995, we had 7 rather than 9 people contacting hunters, a 22% decline in sampling effort. Harvest rates improved by 78% over 1995, from 0.076 to 0.136 birds harvested per hour of hunting effort (Table 7). Harvest rates have almost tripled since 1994. Hunters expended 7.3 hours of effort to harvest a rooster, or on average 1.1 roosters were bagged per 8-hour hunting day. This is the first time since we began collecting harvest information in 1992 that harvest has exceeded a bird per hunter per day. Hunter effort and harvest were recorded and summarized by habitat �82 type (Table 7). Hunters were most successful in sorghum food plots and stripped wheat and least successful in com and wheat stubble. Harvest rates in sorghum food plots were 2-4 times higher than harvest rates in other, more commonly hunted cover types such as creek bottoms, CRP, or wheat stubble. This confirmed earlier hypotheses that sorghum food plots could concentrate pheasants and make them more vulnerable to hunters (Remington and Snyder 1996). Table 7. Pheasant hunter 9-10 November 1996. Cover effort, Hours lAssumes a 8-hour 100 58 374 311 347 7.2 4.2 26.9 22.4 25.0 125 lilO hunting rates in northeastern , crippled Colorado, Flush/hr Bag/hr 1.35 0.62 0.80 0.53 0.41 0.46. 0.21 0.14 0.13 0.10 13.2 7.7 14.5 18.4 15.0 3.7 1.7 1.1 1.0 0.8 . OliO 0102 1015 012 0.56 0.14 15.5 1.1 Daily bag , Total Sorghum Stripped wheat Creekbeds, weeds CRP Wheat stubble Corn stubble Weighted average flush and harvest day Pheasant Trappin~ and Suryjyal Annual mortality (1 Nov - 31 Oct) mortality of hen pheasants trapped and radio- marked between October and December 1995 was relatively low. Mortality was less than 10% of the hens alive at the beginning of the month for 7 of 12 months, compared to 3 of 12 for the previous year's cohort (Fig. 2). The pattern of mortality observed was typical of patterns identified over the last 4 years (Fig. 3); increasing mortality from November through January, low mortality in February through April followed by peaks in May and July. 3 of 4 years the spike in mortality in July has run counter to a strong declining trend in mortality from May through August. This may reflect a short-term impact on survival following wheat harvest in early to mid-July during a period when cover is generally adequate and improving. Predation accounted foralmost all mortality (119 of 132 mortalities; 90%). Avian predators accounted for 51% of mortalities; great homed owls (BWm virgioianus) and prairie falcons ~ mexicauus) were responsible for 84 and 10010of avian kills, respectively, while 6% could not be attributed to a particular avian predator. Most (5 of6) prairie falcon kills occurred between November and February, suggesting winter residents rather than breeding birds were responsible. Mammalian predation accounted for 41% of mortalities; coyotes (Canis latrans) were responsible for 45 of 49 (92%) mammalian kills. Red fox (YuJpes wIpes) are uncommon, but do occur in our study area. It is unlikely unless tracks are present that we could differentiate between coyote and red fox kills, so some of the predation attributed to coyotes could be due to red foxes. Other mammals killing pheasants included long-tailed weasels (MusteJa frenata; 2 kills), domestic cat (1), and domestic dog (1). Other birds were killed by hail (4), shot by hunters illegally (3) hit by vehicles (2), collided with a fence, and 1 died of disease or stress. Cause of mortality could not be determined in 2 cases. Mammalian predators appeared to be more successful than avian predators between 1 April and 15 July when green wheat was actively growing and providing cover (14 of22 or 64% of mortalities versus 26 of67 [39%] mortalities during In �83 the rest of the year). Pheasants may be less vulnerable to avian predators (which locate prey primarily by vision) when they are hidden by green wheat. The concealment value of green wheat should not impact mammalian predators as much, because they can rely on scent to locate prey. Night-trapping began on 22 October and concluded on 12 December. We captured and radio marked 107 new hens, of which 55 were moved to stripped-wheat study areas and released. Through a combination of day and night trapping we also captured and replaced radio transmitters on 45 hens surviving from trapping efforts in 1994 and 1995, of which 18 were moved to stripped-wheat areas. Thus, 152 radiomarked hens were available after 1 October for survival estimates in 1996-97; 73 in stripped wheat and 79 released where they were trapped in conventional stubble or CRP. Translocated hens were moved a distance offrom 5 to 75 km and released into stripped- wheat fields. Most translocated hens (62 of73) stayed within 1-2 km of their release point. Mortality of this radio-marked group was average in the October through December period (Fig. 2). �25~--------------------------------~~ 00 ~ 20 rz 15 w o c::: W 0. 10 5 o NOV JAN MAR MAY JUL SEP NOV JAN MAR MAY JUL SEP NOV JAN MAR MAY JUL SEP NOV DEC FEB APR JUN AUG OCT DEC FEB APR JUN AUG OCT DEC FEB APR JUN AUG OCT DEC Fig. 2. Monthly mortality of radio-marked hen pheasants, November 1993 through December 1996. �16~~--------------------------------~-----' 14 12 t- z w o 0:: W a.. 10 8 6 4 2 o NOV DEC JAN FEB MAR APR MAY JUN JUL AUG SEP OCT Fig. 3. Monthly mortality of radio-marked hen pheasants, average for 1993-96. 00 VI �86 LITERAUJRE CITED Remington, T. E., and W. D. Snyder. 1994. Evaluation of habitat development for ring-necked pheasants in eastern Colorado. Colorado Div. Wildl., Prog. Rep., Fed. Aid Proj. W-167-R. Apr. 1-19. . ____ , and Colorado. __ . 1995. Evaluation of habitat development for ring-necked pheasants in eastern Colorado Div. Wildl., Prog. Rep., Fed. Aid Proj. W-167-R. Apr. 1-13. and . 1996. Evaluation of habitat development for ring-necked pheasants in eastern Colorado. Colorado Div. Wlldl., Prog. Rep., Fed. Aid Proj. W-167-R. Apr. 45-66. Snyder, W. D. 1985. Survival of radio-marked hen ring-necked pheasants in Colorado. J. Wildl. Manage. 49:1044-1050. Wilson, R. 1., RD. Drobney, and D. L. Hallett. 1992. Survival, dispersal, and site fidelity of wild female ring-necked pheasants following translocation. J. Wildl. Manage. 56:79-85. Prepared by -rL C &i:Jnj_ Thomas E. Remingt LSSRN Prepared by ~ 1). ~&.uu Warren D. Snyder LSSRN [Jo) �87 Appendix A - 1996 PHIP SPECIFICATIONS SHRUB THICKETS AND SUPPLEMENTAL WINDBREAKS A shrub (plum) thicket may be planted with or without a windbreak, but a windbreak will not be funded if planted alone. Plantings will be eligible for funding only in farmed areas (dominated by cropland) within less than 0.1 mUe of cultivated cropland and ereater than 0,1 mile away from occupied dwellings. If placed in pastured grass they must be at the pasture edge adjacent to farmed cropland, and the windbreak and thicket must be fenced as one unit (using permanent fencing materials) to exclude livestock. PIllP does not pay for fencing. If placed in CRP, thickets must be within 0.1 mile of cultivated cropland. Plantings must not be disturbed for 10 years. Plantina sites must be inspected and approved by a Colorado Division of Wildlife person before being prepared and planted. CDOW personnel are instructed to not approve plantings in very marginal pheasant ran~ where woody plantings alone will do little to increase pheasant populations or in low areas where they miibt be flooded. . . Maximum Funded: No more than 1 thicket (with or without wind barrier) can be planted per 80 acres. Each thicket/windbreak must be at least 114mile from another thicket/windbreak. Siu: Shrub thickets must be at least 1/IOth acre (4,300 ft2) in size, and contain at least 8 rows (excluding windbreak rows). Twelve hundred (1,200) feet is the maximum linear feet offabric funded per thicket. Supplemental windbreaks, if planted, must be placed on the north and west side of the thicket and will be funded up to 900 linear feet total if straight and ·1,200 linear feet total ifL-shaped. They must include at least 3 rows, one of which must be juniper/cedar. Spacing between the thicket and windbreak should be 100 feet (range 60 ft. min.; 120 ft. max.). Mulcbine: Woven polypropylene fabric such as Dewitt's Sunbelt or Eartbmat is required on all plantings .. Minimum fabric width is 6 ft: (3 ft. on each side of the row). Drip Systems will not be cost shared. Payment Rate: Payment will be SO.56l1inear ft. offabric to the maximums listed above (900 & 1200 lin. ft.) completed by PF Chapters, subcontracted to approved groups, or to approved private contractors. Supplemental payment ratesllinear foot will be: SO.Ol/ft for use of 10 to 12-inch (8 to 12-gauge) wire staples to replace dirt in center offabric, and SO.OI/ft. for band application of an approved herbicide along the exterior edge of the fabric, preferably with a wick applicator. Fertilizer will not be funded. The payment rate may be reduced up to SO.12l1inear ft. where seedlings are not planted properly or on time, the site is poorly prepared, or the fabric is not laid properly. These plantings are expensive and longterm, therefore, quality is emphasized. A SO.Ol/ linear ft. incentive payment may be made for sites where quality in site selection and planting (staples must be used) has been emphasized. Plantjn& Dates: Between March 20 and May 15. �88 Appendix A - 1996 PHIP SPECIFICATIONS Pre-llapt Treatmept: Sites must be tilled, preferably in fall and then rototilled in spring. Tillage must be to bare soil with little residue remaining and must be deep enough to kill existing vegetation. Approved Species: American Plum (bare root), Rocky Mt. Juniper, E. Red Cedar (potted). Golden willow may be used in sites previously inundated by water. One or two rows (maximum) may be planted to choke cherry or sumac (quail bush) within thickets. Between-row Spacin&: A maximum of 10ft. spacing will be permitted (8 ft. recommended) for shrub thickets. A maximum of 15 feet will be permitted for wind breaks. In-row Spacjn&: A maximum of8 ft. (6 ft recommended) will be permitted for shrub thickets, and 12 feet for evergreens within wind barriers. Replacement Plantjn&: Payment will be at the actual cost of seedlings (State Forest Service price less quantity discount) plus $O.2S per seedling for labor, and SO.l1 per wire staple (lor 2 per seedling). Labor will not be funded for landowners replanting their own sites. SUPPLEMENTAL Purpose: PAYMENTS FOR CUSTOM SITE PREPARATION To prepare planting sites when the landowner does not have the proper equipment. Treatment: Breaking out small tracts within CRP or sodded waste areas with a mold-board plow, sweep plow, heavy disc, rototiller, or combinations of these, to completely destroy existing vegetation for reseeding to switchgrass, planting sorghums, or site preparation for thickets and windbreaks. This does not include previously farmed (planted) areas. Tillage must be to a depth of at least 6 inches. Payment Rate: Payment rate will be SlS.OO/acre for preparation of sites greater than 1 acre in size, which may involve two to three treatments. The payment rate will be S20.00/acre for preparing small sites (less than 1 acre) for shrub thickets/windbreaks where a rototiller is used because of the extra time needed per acre. Equipment transportation to and between will be paid atl SlS.OO/hour. PERENNIAL GRASS AND GRASS-LEGUME PLANTINGS Switchgrass provides tall cover that stands well over winter. Small, unfarmed tracts containing short, sodded grasses, are recommended for revegetation to switchgrass. Other shorter, cool-season grasslegume mixtures may be used in roadsides where snowdrift is a problem. This practice is funded only in farmland (not rangeland) settings. Payment Rate: SSO.OO/acre as a one-time payment for sites up to 10 acres. For each additional acre (in sites larger than 10 acres [40 acres maximum]) the rate is S3S.00/acre. An additional SlS.OO/acre will be paid for breaking out sod in heavily sodded sites and supplemental discing prior to planting switchgrass (this does not apply to roadsides). �89 Appendix A - 1996 PHIP SPECIFICATIONS Preplant Soil Preparatjop: Adequate tillage to completely destroy existing perennial vegetation and to establish a firm, weed-free seed bed is required. Interseeding is not approved. Roundup herbicide can be applied to kill weeds just before switch grass seedlings emerge. Planting a tall-sorghum mix (for which payment is available) is recommended the first year. Switchgrass can be seeded into the residual sorghum without tillage during the subsequent spring. Plantjn& Procedures: Planting procedures outlined in the Division's Game Information Leaflet #113 should be considered when planting switchgrass. In general, about 20 pure live seeds/ff (2 - 3 lbslacre) should be planted using-a drill with double-disk furrow openers, l-inch depth bands, and packer wheels. If a herbicide is not used, up to 1 lb/acre of an adapted dryland alfalfa and up to Y2 lb of sweet clover should be added. Approved Species: In plots, switchgrass should comprise at least 75% of the live seed (alfalfa and sweet clover are approved additions). Within roadsides, switchgrass is the priority species where snowdrift is not a problem. Other approved warm-season grasses include bluestems and Indian grass. Where these can not be used the tallest wheatgrasses (tall or intermediate) the roadside site will allow should be used in combination with alfalfa (1 to 2 lbslacre). Plantjn& Dates: Warm-season grasses including switchgrass: March 15 to May 25: Cool-Season Grasslegume Mixtures: March 15 - July 15 Plot Duratjon: Grass and grass-legume plantings must remain ungrazed and undisturbed for at least 7 years. Roadsides should remain unmowed unless essential to reduce snowdrift. If essential; mowing should be delayed until 1 August and restricted to the road shoulder. Prescribed burning, thinning tillage, or other rejuvenation treatments may be applied after 7 years. Grass stands that are relatively thin provide taller, better cover for pheasants. Legumes provide nitrogen and increase growth and quality when added to grass mixtures. DISTURBANCE TILLAGE AND TALL WILD ANNUALS Wild sunflowers, kochia, pigweed, and other tall annuals which attain 4 to 6 ft. height stand better through winter than other herbaceous vegetation, and provide excellent cover for broods, protection from blizzards and predators, and supplemental food. This is the most effective and least expensive approach for increasing pheasants and other upland game birds. Fallow land that is left idle usually converts to annual grasses or dog-hair stands of weeds by the 2nd year following tillage. Therefore, at least one tillage in mid-spring is usually needed to promote growth of tall annuals and a second thinning tillage is sometimes needed. Maximum Funded: 14 acreslO.25 section, 28 acres/landowner and section. Plots larger than 3 ac. should be at least 0.25 mi. apart. Fundin& Rate: S30.00/year for patches 0.1 to 0.5 acres in size, patches larger than 0.5 acre are considered 1 acre. S40.00/acre/year for sites up to 5 acres (7 ac in pivot comers). S30.00/acre/year for �90 Appendix A - 1996 PHIP SPECIFICATIONS additional acres up to 10 (6th-IO acres).Seeding wild sunflower or other approved wild annuals at 2 to 4 Ibs. per acre will be funded at direct seed costs (see seed sources below). Plot Dimensions: preferred. Short, relatively wide patches, which will not be easily inundated by drifting snow, are . Placement: Adjacent to woody cover when possible. Draw bottoms that already contain weeds and above average moisture are ideal. Sites containing noxious perennials should be avoided. Specifications: Initial tillage with a disk plow or mold-board plow is needed in sites containing perennial grass to destroy all perennial cover, preferably, immediately after the ground has thawed in early March. Large clod size is preferred to retain thin stands of annual forbs. Initial tillage in subsequent years should be conducted prior to May I A second thinning tillage may be used prior to the 1st of June. Spring tillage is needed each year to retain tall annuals. Annual grasses usually dominate if tillage is not used each spring. WIld sunflowers can be drilled or broadcast and harrowed at low rates to help establish tall annuals, if they are not already present. Known sources in Colorado include the Arkansas Valley Seed CompanyDenver & Longmont, and Sharp Bros. Seed Company - Greeley. Retentiop: Tall annuals must remain undisturbed through March of the following year. Sites should be prepared for the next year's growth during mid to late April if weedy cover exists. ANNUAL SURVIVAL PLANTINGS - SORGHUMS & DRYLAND CORN APPLICATION: On CRP, annual Set Aside, and other cropland or tilled wasteland. When applied within CRP fields, SCS specifications for CP-12 must be used (see supplement). Landowners must obtain approval from the ASCS before planting grains as WIldlife Food Plots on annual Set Aside acres and these acres can not be harvested. Dryland com is primarily applicable in center-pivot comers next to irrigated com. MAXIMUM FUNDED: at least 114mile apart. I plot/80-acre field, 2 plotsll60 acres, 4 plots (28 ac.)/section. Plots must be PAYMENT RATE: S40.00/acrelyearfor I - 5 acres (7 acres within center pivot comers) and S2S.00/acrelyear for additional acreages in tracts larger than 5 acres (12-acre maximum). S30.00/acrelyear if planted after June 20th. S7.S0/acre for application of30 lbs ofnitrogenlacre. S15.00/acre will be paid for breaking out sod in CRP or heavily sodded sites and supplemental discing prior to planting. Landowners can not be paid for labor/tillage on their own land other than for breaking out sod. . PLACEMENT: . . Plots should be within or near cropland and placed crosswise to prevailing winds. �91 Appendix A - 1996 PHIP SPECIFICATIONS SPECIFICATIONS: Preplant Soil Preparation: Initial treatment: vegetation in early' spring prior to annual growth. Adequate tillage to destroy existing perennial Subsequent years: Preferably minimum tillage shredding of old materials as needed prior to April 25. Annual application of nitrogen at 30 - 40 lbs./ac. is recommended. Plot Dimensions: Minimum total plot width shall be 150 feet. Wider strips are preferred to reduce impacts of drifting snow. (See restrictions on dimensions in CRP). Row Spacin&: Sorghums - 15 to 30 inches; Dryland com - 30 to 36. Seed Specificatjons: Sorghum Patches - At least 60% (75% preferred) of an adapted tall forage sorghum that will stand well with minimal lodging and will mature before frost. Up to 40% can be adapted varieties of grain sorghum. These can be mixed or planted in separate rows (i.e., 2 rows of grain sorghum to 6 rows offorage sorghum. These sorghums should equal a minimum of75% of the total weight. Maximum . amounts for other grains include: Dryland com (25%), sunflowers (10%) and proso millet (10%). Addition of 1 to 2Ibs./ac. of wild sunflower seed is recommended (Source: Arkansas Valley Seed Company - Denver). Dryland Com Plots - Early maturing dryland varieties adapted to NE Colorado. Seed, from these varieties, that is one year removed from purchased hybrid can be used to reduce seed cost. Plantjn& Dates & Rates: Sorghums - Between April 25 and June 15; Mid- to late-May is recommended. Plantings conducted after June 15 may be assessed a $10/acre payment reduction. Sorghums should be planted at 4 - 8 lbs./acre (30-inch rows) and at higher rates if drilled. Dryland com - Between April 25 and May 15. Plantings after June 1 will not be accepted for payment. Seeding for dryland varieties should be from 10,000 to 13,000 seeds/acre. At least one cultivation is needed for com and fertilizer should be used. Plot Duratiop: 1 year. Sorghum and com plantings must remain undisturbed through March of the following year. Dryland com must be left standing through March unless harvested. Harvesting may be conducted after March 15 of the following year if approved by ASCS. �92 Appen~ix A - 1996 PHIP SPECIFICATIONS SUPPLEMENT FOR SORGHUM PLANTINGS WITHIN CRP SCS Notification: The CRP contract must be amended at the local NRCS office prior to implementing CP-12 and breaking out food plots within CRP. This requires filling out a one-page form at yourNRCS office. The ASCS must be advised of the change for their records. Dryland com is not approved for plots within CRP. Winter cover-food plots within CRP must be replicated until the end of the CRP contract. If the farmer wishes to discontinue this practice he must reestablish grass (required by the ASCS). Payments will be made annually. Reimbursement will be at S40.00/acre to cover reseeding grass unless Division of WIldlife personnel do the work. Maximum Funded: I plot/80-acre field, 2 plotsll60 acres. Plots must be at least 114 mile apart. Maximum Size: The maximum size is 3 acres per site. CRP fields must contain at least 40 acres to be eligible for a CP-12 food plot. Plot Dimensions: Plantings may be up to 200 feet wide (100 ft in sandy soils). TypiCal3-acre plots measure 198 x 660 feet. Where a 100 ft maximum is required a 30 ft wide buffer of untilled grass is left between two 99 x 660 ft parallel strips to obtain a 3 acre plot. Smaller plots should have reduced length to retain at least the ISO ft. minimum width. For example, a plot 99 ft Wide x 440 ft. long equals 1 acre and two adjacent plots will exceed the minimum width requirement. Placement: Preferably within 50-100 yds of edge and near cropland, but location can vary depending on soil, wind, and moisture, and location of other winter covers if they occur. Sorghum plantings are not permitted in soils containing free lime (shows effervescence), or soils that are deep sands or choppy sands. �93 JOB PROGRESS REPORT State of: Project: Work Plan: Job Title: Colorado _~WI.L.;.i-l~6,,:!:ry.;;;.l-R~~~ :, Upland Bird Research 3 :Job 19 Implications of Habitat Loss and Fra2IDentation on Conservation Strate~es for Gunnison SaKe Grouse Period Covered: 01 January through 31 December 1996 Author: Personnel: Sam J. Qyler-McCance Clait E. Braun, Colorado Division of Wildlife; Sara J. Oyler-McCance, Colorado State University ABSTRACT Data were compiled and a parsimonious model was developed to predict sage grouse (Centrocercus minimus) occupancy using the combination of variables which most successfully predicted occupancy without overfitting the data. Blood and feather samples were collected from 49 Gunnison sage grouse in southwestern Colorado between April and September 1996. Samples were obtained from 4 distinct populations. DNA was successfully extracted from all samples. A subset of the samples was used to screen microsatellite primers. Primers which looked promising were used in polymerase chain reaction (PeR) reactions and parameters in those reactions were optimized to produce clear products. Screening and optimization will continue in 1997-98. In addition, DNA will be extracted from single feather samples for which there are no blood samples. A geographic information system (GIS) coverage of all the areas used in the habitat-based model and a coverage of paved roads in southwestern Colorado were created. A user-friendly model was developed which allows a user to investigate the effects on habitat quality of changing habitat patch size and distance to the nearest paved road. Additional coverages will be added and development of conservation strategies will continue in 1997-98. ��95 IMPliCATIONS OF HABITAT WSS AND FRAGMENTATION ON CONSERVATION STRATEGIES FOR GUNNISON SAGE GROUSE . Sara J. Oyler-McCance INTRODUCTION There are serious knowledge gaps which impede development of a conservation plan for Gunnison sage grouse. first, it is not known how much sage grouse habitat has already been lost and how much might be lost in the future given human population growth and land development This is essential information if a balance between human growth and sage grouse conservation is to be achieved. Second, little is known about landscape level habitat requirements of sage grouse living in fragmented habitats. Questions such as: how large must a habitat patch be to support sage grouse?, can a well-connected network of small patches support a sage grouse population?, and are some interpatch matrices more detrimental to sage grouse than others? need to be answered before a landscape level conservation plan can be developed. Third, little is known of the movement of young sage grouse, as only one study has addressed this issue. Dunn and Braun (1985) measured natal dispersal of sage grouse in contiguous but altered habitats of northwestern Colorado and found average dispersal distances of 8.8 km for juvenile females and 7.4 km for juvenile males. It is not known, however, whether Gunnison sage grouse move among fragmented habitats (across distances up to 300 km) or whether some populations in southwestern Colorado are truly isolated. The amount of inbreeding in small, isolated populations is also unknown. Young (1994) measured inbreeding within the Gunnison population, yet that population is large compared to other populations ·of this species. Knowledge of movement among patches and the amount of inbreeding would provide essential information about potential inbreeding effects and aid in any conservation plan which addresses reintroductions. P. N. OBJECTIVES The objectives of this study are to (1) develop a habitat-based model which can be used to predict sage grouse occupancy of patches in southwestern Colorado, (2) investigate gene flow among isolated populations using microsatellites as a molecular marker as well as sequencing the control region of the mitochondrial genome, (3) determine the level of inbreeding within populations using the aforementioned techniques, (4) document the loss of sagebrush-steppe habitat using aerial photography and satellite imagery, and (5) develop a spatially explicit model which integrates information gained in the previous portions of the study to be used to assess potential conservation strategies of Gunnison sage grouse. SEGMENT OBJECIlVES 1. Review literature pertinent to the objectives of this study. 2. Develop a habitat-based model to identify what habitat or landscape features may be manipulated to improve occupied areas and to identify any unoccupied areas which might be suitable for sage grouse habitation. 3. Collect blood and feather samples from 4 different populations in southwestern Colorado. 4. Isolate DNA and begin to screen micro satellite primers and optimize PCR reactions. 5. Begin to construct GIS coverages to be used in the final spatially explicit model. Such coverages include a digitized coverage of areas included in the habitat-based model, and a coverage of paved roads associated with those areas. . �96 6. Begin to develop a computer program which interfaces with the GIS coverages such that the effects of manipulating specific variables (e.g., patch size) can be seen visually in a GIS map coverage. . 7. Prepare annual report. MEfHQDS Habitat-based Model A suite of micro-scale and landscape level variables were chosen which were thought to be important to sage grouse to develop the habitat-based model. The micro-scale variables were used to describe the quality of the habitat in each patch. These variables included the percent cover of : live sagebrush, dead sagebrush, oakbrush, other brush, pinion/juniper, forbs, and grass - height of: live sagebrush, dead sagebrush, oakbrush, other brush, pinion/juniper, forbs, and grass density of sagebrush bushes - presence/absence of fence posts, wet meadow, pinion/juniper, and oakbrush. Landscape level variables describe the overall patch and its relation to landscape surrounding it These variables included the area of each patch, the area/perimeter ratio, distance to the nearest paved road (from the centroid of the patch), the presence of powerlines, and the habitat type of the surrounding matrix. Twenty-five patches in southwestern Colorado (12 occupied and 13 unoccupied) were chosen for inclusion in this study. A patch was defined as a discrete expanse of sagebrush-steppe habitat with boundaries consisting of either a paved road or non sagebrush steppe habitat, the only exception being agricultural fields. In agricultural fields, there were often small islands of sagebrush surrounded by either plowed or planted fields. Because sage grouse could readily move among these small islands, I considered all islands of sagebrush in the midst of agricultural fields as belonging to the same patch as long as they were not separated by paved roads or non sagebrushsteppe habitat (excluding agricultural fields). Data were collected by walking transects and taking measurements in a 1 m2 plot frame every 200 meters. Patches greater than 2 km2 were sub sampled such that grids 1 km2 were sampled yielding measurements taken at 25 points. Landscape level data were determined from satellite images and maps. The number of variables had to be reduced because there were more variables than actual data points (patches). Thus, the variables considered to be most important (e.g., patch size) were included as well as biologically meaningful composite variables (e.g., percent of all brush that was sagebrush). The list of variables included in model selection were: patch area, distance to the nearest paved road, presence/absence of oakbrush, presence/absence of pinion/juniper, percent cover of all brush, percent of all brush that was live sagebrush, average brush height, percent cover of forbs, and height of forbs. A logistic regression framework was used to predict the probability of occupancy based on certain variables. Most variables have a linear relationship between the value of the variable and the probability of occupancy (e.g., the larger the patch the higher the probability of occupancy), however some variables have a quadratic relationship between the value of the variable and the probability of occupancy (e.g., the percent of all brush that was live sagebrush). For those variables with quadratic relationships, the variables were transformed by using the square of the deviation from the mean. Once the variables were selected and transformed, model selection was conducted with model selection criterion being Akaike's Information Criterion (AlC) corrected for small sample sizes. �97 Genetic Analysis Blood samples from as many individuals as possible (max. = SO) were obtained from each of the following populations: Dove Creek, Dry Creek/Miramonte, Crawford, and Gunnison. Blood samples were obtained by capturing sage grouse using spotlight trapping (Giesen et al. 1982), clipping a toe nail and extracting 2-3 drops of blood. Trapped birds were banded and released. Blood samples were placed in sterile 15 ml Eppendorftubes in a STE buffer and frozen, Feather samples were also obtained from each captured bird and frozen along with the blood samples. Samples were then transported to Dr. Tom Quinn's molecular genetics lab at the University of Denver and stored in a -70 C freezer until DNA extractions were performed. DNA was extracted from blood samples using the protocol for DNA extractions from low volume blood samples. The blood samples were thawed and up to SOpl of blood was removed and used for extraction. Any unused portion of blood was re-frozen and stored for later extractions. DNA was extracted using established protocols (below). 1. Place SOOplTE (10mM Tris, ImM EDTA pH 8.0) in each 1.5 ml tube. 2. Add 40pl proteinase K solution (2.25 mg/ml) in each tube. 3. Add SOpl blood, close cap, and mix. 4. While vortexing, add 26pl of 10% SDS. 5. Rotate for 3 hours at SOC. 6. Perform a phenol extraction 3 times, after first extraction transfer upper layer contents to a new microfuge tube, for every extraction afterwards, remove phenol from underneath with pipette. 7. Extract 1 time with chloroform:isoamyl alcohol mix (24:1). 8. Add 100pilOmb/ml 9. Add 26pl of 10% SDS, vortex, add 4Opl2.5mglml 10. Rotate at SO C for 3 hours. 11. Extract with phenol 2 times; transfer to a new tube, leaving blob behind after first transfer. 12. Extract 1 time with chloroform:isoamyl alcohol (24:1). 13. Ethanol precipitate. 14. Conduct spectrophotometer reading: determine concentration using 5pl DNA in 995pl H20. RNase, vortex incubate for 3 hours at 37 C. proteinase K, vortex. After extraction, 5pl of each DNA sample was diluted (1: 10) and 'used as a working sample. The remaining volume of each sample was stored in the -70 C freezer. Microsatellite primers were obtained from two sources. The first group of primers were those designed for use with poultry chickens and were obtained from Hans Cheng at the Poultry Research Group. The second group of primers were designed for use with red grouse (l&2RPUSla20pus scoticus) and wereobtained �98 from Stuart Piertney at the University of Aberdeen. Primers were radioactively labeled for later visualization on autoradiography film, The T4 Polynucleotide Kinase (PNK) Labeling procedure was used. In this procedure, T4 PNK catalyzes the transfer of the y-phosphate from ATP to the 5' terminus of polynucleotides or to mononucleotides bearing a 3' phosphate group. One primer (either the forward or reverse primer) was chosen and radioactively labelled using established procedures (below). 1. In a O.5pl Eppendorf tube mix: 1pl lOpM primer, 1}l1lOX Buffer, 0.25pl T4 PNK; 0.25pl p33 y-ATP, and 7.5pl H20. 2. Incubate at 37 C for 15 minutes. 3. Stop reaction by heating to 70 C for 10 minutes. Screening microsatellite primers consisted of choosing 4 - 12 DNA samples (representing at least one individual from each population, plus at least one individual from Cold Springs Mountain to use as an outgroup) and amplifying the micro satellite region using PCR. The components of a PCR reaction for each individual were: 1. 2. 3. 4. 5. 6. 7. 1pl diluted DNA, 2.5pl Taq buffer (lOx), 2.5pl dNTP mix (O.25mM), l.25pl forward primer, l.25pl reverse primer, 16.375pl H20, and O.l25pl Taq polymerase. Once all components of the PCR reaction were added, 2 large drops of ultra-clean mineral oil was added to each tube (to prevent evaporation) and the tubes were placed in a thermal cycler.' The PeR reaction typically consists of a denaturing step (-94 C), an annealing step (variable temperature), and an extension step (,..,72C). These three steps were repeated for a number of cycles. The number of cycles, exact temperatures, and time at each step were variable and depended on the primers used. At the completion of the PeR, samples were then run for 2.5 hours on a 6% polyacrylamide gel using electrophoresis. The gel was then dried and exposed to film for 1.5 - 3 days. The film was then developed and analyzed. Spatially Explicit Model GIS coverages of the areas sampled in the habitat-based model were created by overlaying a blank coverage over a satellite image which had been classified as to habitat type .. Patches of sagebrush were digitized into the blank coverage in ARCIINFO. A coverage of paved roads was created by downloading digital line graph files containing transportation information from the EROS data center. These files were then linked together in ARCIINFO and the appropriate information was accessed and saved as a separate coverage. Additional information (e.g., % of all brush that was sagebrush) was added to the sagebrush area coverage so that an interactive program could be developed allowing the user to investigate the effects of changing certain variables on the quality of the habitat. A program was written in C language to interface with the GIS coverages. �99 RESULTS AND DISCUSSION Habitat-Based Model AlC corrected for small sample sizes (AlCc) was used as the model selection technique. Given an established set of models (as defined by the subset of variables used in model selection) AlCc uses maximum likelihood to optimize the trade-offs between bias and variance using the following formula: -210~(L) + 2(K) + 2(K(K+ 1»/(N-K-l) where K is the number of parameters and N is the sample size. The best model (Table 1) included the variables patch size, distance to the nearest paved road, and the percent of all brush that was sagebrush. The logistic regression equation for this model is 1 Prob(occupancy)= 1+ _ e-(-15.4 +4.1(distance to road) +O.35(patch area) - 317.6(% of all brush that was live sagebrush» The parameter estimates and standard errors for this model varied (Table 2). The only other variable which proved to be somewhat important (by its inclusion in the top four models) in predicting sage grouse occupancy was percent cover of forbs. The best model to make inferences from the data includes patch area, distance to the nearest paved road (from the centroid of the patch), and the percent of all brush that was live sagebrush. This model can be used to predict sage grouse use of patches in southwestern Colorado. This model can also be used to rank the existing patches and identify the suitability of patches which are candidates for transplants. Additional data to be used in the validation of this model will be collected in 1998. Genetic Analysis Of the 49 blood samples for which DNA extraction was attempted, DNA was successfully extracted from all samples. When screening primers, 6 outcomes were possible, not including the outcome of no product (below). 1. 2. 3. 4. 5. 6. Clean bands, polymorphic. Clean bands, not polymorphic. Questionable if a product exists. Fuzzy bands with shadowing bands above, possibly polymorphic. Fuzzy bands with no shadowing bands, possibly polymorphic. Fuzzy bands, not polymorphic. Thirty-three chicken primers were labeled and screened. Of those, 11 showed no product and were discarded. The results of the first screen of the remaining 22 primers varied (Table 3). Twelve red grouse primers were labeled and screened. All primers produced some product. The results of the first screen of the red grouse primers varied (fable 4). For those primers in which the products were not clear and easy to read (outcomes 3 - 6), PCR optimization (changing parameters in the PCR reaction) was and will continue to be attempted. DNA from feather only samples will attempt to be extracted and another molecular marker will be investigated in 1997. Spatially Explicit Model GIS layers will continue to be developed as the data becomes available (e.g., genetics data and data from analysis of habitat loss). �100 Table 1. Four best habitat-based models chosen with the AlCc model selection technique. K AlCc Delta AlCc Cbi-SQuare GOFdf Parea, roaddis, lsbpcnt 4 18.57 0.00 8.574 21 Parea, roaddis 3 19.03 0.46 11.89 22 Parea, roaddis, lsbpcnt, forbcov 5 21.07 2.50 7.91 Parea,roaddis,forbcov 4 21.67 3.10 11.67 21 Model = = Parea patch area, roaddis distance to the nearest paved road from the centroid of the patch, lsbpcnt live sagebrush, forbcov % cover of forbs. = 20 = % of all brush that is Table 2. Parameter estimates for the best model. Parameter Estimate Standard Error Intercept -15.40 Roaddis 4.10 3.13 Parea 0.35 0.25 -317.62 Lsbpcnt 11.15 278.03 Parea = patch area, roaddis = distance to the nearest paved road from the centroid of the patch, lsbpcnt live sagebrush, forbcov % cover of forbs. = = % of all brush that is �101 Table 3. Results of the first screen of 33 chicken primers. Primer Identifier Product Code Adl24 Adll23 Adl171 Adl22 Adl118 Adll06 Adl145 Adllli Adl155 Adll20 Adl19 Adl181 Adl171 Adl210 4 2 2 4 3 4 3 2 1 2· 4 4 3 5 5 2 2 5 2 Adl136 Adll58 Adl172 Adl231 Adll54 Adl157 Ad1228 Adll73 1 2 ·3 = clean 4 5 4 bands, polymorphic. = clean bands, not polymorphic: = questionable products. 4 5 6 = fuzzy bands with shadowing - possibly polymorphic. = fuzzy bands with no shadowing - possibly polymorphic. = fuzzy bands - not polymorphic. �· 102 Table 4. Results of the first screen of 12 red grouse primers. Product Code ~D1erldentidier Rg2 Rg3 RgTl RgT2 Rg4 Rg5 Rg7 Rg8 Rg9 RgT3 RgT4 RgT5 = = 1 clean bands, polymorphic. 2 clean bands, not polymorphic. 3 = questionable products. 6 4 1 4 4 3 2 1 4 1 3 3 4 5 6 = fuzzy bands with shadowing - possibly polymorphic. = fuzzy bands with no shadowing - possibly polymorphic. = fuzzy bands - Dotpolymorphic. LITERATURE cUED Dunn, P.O., and C. E. Braun. 1985. Natal dispersal and lek fidelity of sage grouse. Auk 102:621-627. . Giesen, K. M., T. J. Schoenberg, and C. E. Braun. 1982. Methods for trapping sage grouse in Colorado. Wildl. Soc. Bull. 10:224-231. Young, J. R. 1~. Sexual selection of sage grouse. Ph.D. Diss., Purdue Univ., West Lafayette, IN. 123 pp. �103 JOB PROGRESS REPORT State of: Colorado Project: W-l67-R Work Plan: 12 : Avian Research Job 18 Job Title: Genetic Diversity Among Populations of Merriam's Wild Dlrkeys in Colorado Period Covered: 01 January through 31 December 1996 Author: Richard C Dujay Personnel: Richard W. Hoffman, Colorado Division of Wildlife; Richard C. Dujay, Colorado State University ABSTRACT Blood samples were collected from 315 Merriam's wild turkeys (Meleagris ga)]opavo merriami) trapped (n = 292) anellor harvested (n = 23) in Colorado. Samples were obtained from 3 East Slope and 5 West Slope flocks. In addition, samples were obtained from the Sacramento Mountains in New Mexico (n = 5) and the Mogollon Rim in Arizona (n = 32). DNA was successfully extracted from 309 of the 315 samples processed. All samples were assayed using spectrophotometry and DNA concentrations were quantified. A protocol for Non-isotopic Restriction Fragment Length Polymorphism (RFLP) analysis was developed using the M13mp18 phage probe. Twenty randomly selected samples from 2 Colorado counties (Montezuma and Montrose) along with 20 randomly selected samples from Arizona were subjected to restriction digest by the Pst-I restriction enzyme. The restricted samples were electrophoresed through 0.8% argarose gel and Southern transferred to nylon membranes. The DNA fingerprints were digitized to ascertain band sizes (nucleotide bases per band) -. Polymorphismsare readily apparent and easily identifiable among these 3 populations. ��105 GENETIC DIVERSITY AMONG POPULATIONS OF MERRIAM'S WILD TURKEYS Richard C. Dujay INTRODIICTION Until recently, genetic considerations have been largely ignored in wild turkey restoration and range expansion programs due to lack of cost-effective techniques for examining genetic attributes of populations (Leberg 1990, Stangel et al. 1992)~ Establishing high genetic diversity in introduced populations is considered desirable because the level of genetic diversity influences the population's growth rate and ability to adapt to new environmental . conditions (Leberg 1990, Vrijenhoek and Leberg i991). Practices common to wild turkey transplant programs that may contribute to low genetic diversity in new populations include releasing more females than males, releasing birds captured from the same flock, and obtaining release stock from the same population (Leberg 1990, 199i). This lack of genetic diversity in the released stock may explain why some wild. turkey populations experience consistently low reproductive success and fail to increase even when habitat conditions appear suitable. Supplemental releases into genetically stressed populations may be one way to enhance genetic vigor and stimulate reproductive performance. However, before any further manipulations are attempted, as much genetic information as possible must be gathered from existing populations (Leberg et al. 1994). With advancements in DNA technology, large scale surveys can now be economically conducted over broad geographic areas to assess the current genetic condition of populations and to evaluate the genetic consequences of past management activities. These genetic data are essential in prescribing supplemental releases and in directing management efforts towards other factors, such as habitat quality, if the data reveal no genetic problems. p. N. OBJECTIVES The objectives of this study are to (1) compile a database of genetic structure and variation among populations of Merriam's wild turkeys in Colorado, (2) compare genetic diversity of introduced/reestablished populations with the original source population, (3) compare genetic diversity of Merriam's wild turkey populations in Colorado with non-manipulated (pure) populations of Merriam's turkeys in Arizona and New Mexico, and (4) identify populations that are genetically stressed and develop management recommendations to enhance the genetic attributes of these populations. SEGMENT OBJECTIVES 1. Review literature pertinent to the objectives of this study. �106 . 2. Collect blood samples from Memam's wild turkeys in Colorado. 3. Arrange for blood samples to be collected from Merriam's wild turkeys in Arizona and New Mexico. 4. Extract and purify DNA from collected blood samples. 5. Quantify DNA concentrations in extracted samples. 6. Expose samples to restriction digest. 7. Perform electrophoreses on digested samples. 8. Compile data, analyze results, and prepare progress reports. STImv AREA AND MEmons The following' populations/geographic areas were selected for sampling: 1. Larimer County - between the Poudre and Big Thompson drainages. This area is classified as nonhistoric turkey range. The population originated from the release of 15 birds (8 males, 7 females) captured in 1957 on what is now the Spanish Peaks State Wildlife Area. 2. Boulder County - between Lefthand and Saint Vrain Creeks. There are no records of any releases in Boulder County. This population occurs at the northern periphery of the native distribution of Merriam's wild turkeys and probably originated from the northward expansion of native populations. However, it is also possible that birds from the introduced population in Larimer County expanded south into Boulder County. 3. Las Animas County - on and surrounding the Spanish Peaks State Wildlife Area. Spanish Peaks and Devil' s Creek (Archuleta County) have been the primary sources of birds for transplant programs elsewhere in the state. Located within the core of the native distribution of Merriam I s turkeys, Las Animas County supports some of the highest densities of turkeys in Colorado and is the leading harvest area in the state. No recent transplants have been made into Las Animas County. However, in the 1940ls and 501s, male turkeys were interchanged between Las Animas and Archuleta counties as part of a breeder exchange program. 4. Archuleta County - including Devil's Creek State Wildlife Area and Cat Creek (Southern Ute Indian Reservation). This area also is within the core of the native distribution of Merriam I s wild turkeys. Supplemental releases from Las Animas county were made into Archuleta County (specifically at Devil's Creek) during the 1940ls and SOlS as part of a breeder exchange program. �107 5. Montezuma County - including Boggy Draw and Hartman Draw. This area has a history of population declines followed by reintroduction programs. The first decline occurred prior to 1940. Reintroductions were made during the early 1940's using birds trapped at the Devil's Creek State Wildlife Area. By the early 1960's, turkeys were presumed extirpated from the area. Reintroduction efforts were initiated in 1983 using birds trapped from Las Animas and Pueblo counties. The population has substantially increased in recent years and has provided a source of transplant stock for other areas in southwestern Colorado. 6. Montrose County - between the Dave Wood and Delta-Nucla roads. Historical records suggest wild turkeys are native to Montrose County, but disappeared from the area by 1900. Restoration attempts were first made in 1934 using birds obtained from private sources in Texas, Oklahoma, and New Mexico. These may have been captive birds, and some were eastern (M. g. silzestris) or more likely Rio Grande (M. g. lntermedia) wild turkeys. Few, if any birds, remained by 1940. New attempts to restore the population began in 1944 and continued through 1949 using birds trapped in Archuleta County. The reintroduced population increased and expanded its range through the early 1960' s, when another crash occurred. A third restoration attempt was begun during the early 1980's using source stock from Las Animas and Pueblo counties. Some turkeys still remained in the area when the third restoration program was initiated. The restored population has increased and expanded into formerly occupied . habitats. 7. Mesa County - within the Plateau .Valley. There are no records of turkey releases in the Plateau Valley. However, releases were made in the 1950's and 1980's near Rifle along Divide and Beaver creeks. Turkeys also were released near Cedaredge on the south side of the Grand Mesa. It is possible birds from these releases expanded their range into the Plateau Valley. 8. Garfield County - including Beaver Creek, Divide Creek, and Rifle Creek. This area, along with the Plateau Valley, is considered non-historic turkey range. Releases made in the 1950's were comprised of birds trapped in southwestern Colorado, primarily from Archuleta County. Releases in the 1980's were from birds trapped in Las Animas County. More recently, birds have been trapped and moved within the Garfield County area. 9. Mogollon Rim - Chevelon Ranger District, southwest of Flagstaff, Arizona. This area was selected because it supports one of the few remaining unmanipulated populations of Merriam's wild turkeys within the native distribution of the subspecies. Turkeys have been trapped and moved from the Mogollon Rim, but no birds from other populations have been released into the area. 10. Sacramento Mountains - Cloudcroft Ranger District, southeastern New Mexico. An unmanipulated population of Merriam's wild turkeys that is isolated from other populations. �108 Turkeys were baited with oat hay and com and live-trapped using cannon nets, drop nets, or box traps. Captured turkeys were classified as to age and sex, and banded with seriallynumbered aluminum leg bands. Ages were recorded as subadult (8-10 months) or adult (> 18 months). Blood was collected by jugular venipuncture. At least 1.5 mls of blood were collected from each bird and placed in EDTA purple top vials for DNA analysis. Samples collected for DNA analysis were stored in a-20° C freezer until extractions were performed. DNA extractions used the Analytical Genetic Testing Center's Quik Gene Kit with adapted protocol for avian blood. The whole blood samples were thawed and a volume of 0.250 mls was drawn from each vial for DNA extraction. The unused portion of whole blood was re-frozen and stored as a backup sample for later extraction. The extraction protocol was as follows: 1. Place 0.250 JlI of whole avian blood into a graduated conical centrifuge tube. 2. Add 2.5 ml of ice cold red blood cell (RBC) lysis buffer OX), shake to mix, and vortex for 30 seconds. 3. Let stand on ice for 10 minutes. 4. Centrifuge 25 minutes at 3500 rpm. 5." Decant and discard supernatant (if still lumpy, repeat steps 2-5). 6. Add 1 ml ofRBC lysis buffer (1X) and rinse pellet (do not rinse pellet if it was necessary to pipette supernatant). 7. Add 4 ml of white blood cell (WBC) lysis buffer (IX), shake until viscous, and vortex for 5-10 seconds. 8. Incubate for 30-45 minutes at 55° C in a shaker water bath. 9. Add 200 JlI10% SDS, 500 JlI protein precipitant solution, and 1 ml of ultra pure water, and vortex for 1 minute. 10. Incubate in 55° C water bath for 20 minutes. 11. Centrifuge at 3500 rpms for 25 minutes. If not clear after spin, add 112 volume of ultra pure water and 200 JlI of SDS, vortex 30 seconds, and repeat steps 10 and 11. 12. Pipette clear supernatant into clean test tube. 13. Add 2-3 volumes ice cold absolute alcohol to precipitate DNA. �109 14. Remove DNA with sterile glass hook, dry slightly, and dissolve in a microcentrifuge tube containing 700 III ofTE Buffer (EDTA TRIS at 7.4 pH). 15. Place in refrigerator at 40 C for 24-48 hours, then into -200 C freezer for short term storage (3-6 months) or -700 C for long term storage (indefinitely). The washing and purification procedure involved thawing the extracted DNA samples and placing the solution into a 15-ml·conical centrifuge tube. The volume of solution was determined, and 10% SDS and protein precipitant were added at 50 III and 90 ILlper 1.0 mlof DNA solution, respectively. Each sample was mixed thoroughly by vortexing and allowed to stand at room temperature for 10 minutes. Samples were then centrifuged at 3500 rpms for 25 minutes and the supernatant was decanted or pipetted into a clean centrifuge tube, .where the DNA was precipitated using absolute ethanol. The DNA was collected and dissolved again in TE buffer and frozen at -200 C. Genomic turkey DNA was subjected to endonuclease restriction by the enzyme Pst-L, Electrophoresis of the restricted DNA was performed in a 0.8% agarose gel and then Southern transferred to a nylon membrane, and prepared for hybridization. Conditions of the hybridization and stringency washes were varied individually until the desired results were obtained. After many trials, the optimal hybridization and wash conditions were determined as follows: Hybridization - high temperature hybridization (650 C) with the M13mp18 probe Washes moderate detergent concentration (0.1 % SDS) moderate salt concentration (2X SSC) both high and low temperature washes (2X @ 250 C; IX @ 650 C) RFSIIl.TS AND DISCIJSSION Blood samples were collected from 315 Merriam's wild turkeys trapped (n =292) or harvested (n = 23) in Colorado, Arizona, and New Mexico (fable 1). Within Colorado, samples were obtained from 3 East Slope and 5 West Slope populations. DNA was extracted from 309 of the 315 samples processed. All samples failing to yield any DNA were from hunter-harvested birds. All samples were subjected to spectrophotometric analysis and showed moderate to high concentrations (0.50-1.01, Ilg/lll) of high molecular weight (absorbency 260 nm 0.100-0.202) DNA. Power calculations were performed using SAS statistical software in a completely randomized design .. Caiculations that compared each one of the 10 populations under investigation (n, nlO) to the pooled sample (N) indicated that a power> 0.80 was obtained when any n = 20 and N > 300. To assure confidence in statistical analysis, .the minimum sample size from any population should be > 20. This indicates that additional samples are needed from Larimer County (n = 11) arid the Sacramento Mountains in New Mexico (n = 5). �llO Table 1. Location and number of blood samples collected from Merriam's wild turkeys in Colorado, Arizona, and New Mexico. Location Boulder County Larimer County Las Animas County Montezuma County Archuleta County Montrose County Mesa County Garfield County Sacramento Mountains, NM Mogollon Rim, AZ Total n 20 11 27 61 33 38 51 37 5 32 315 Digitization of the fingerprints from 3 locations was completed (fable 2). The fingerprints of the samples from Montezuma County, Montrose County, and Arizona showed distinct banding patterns along with substantial polymorphisms. Band sizes range from < 1,000 nucleotide bases to > 28,000 nucleotide bases. The number of bands per individual varied from 7 to 21. Band frequencies were determined for flocks and the population at large (fable 3). Chi square analysis was performed on the band frequencies found in the 5,000 to 10,000 nucleotide base range (fable 4). No significant differences were detected. With additional data still needing to be compiled, differences are likely to be identified. The presence and/or absence of particular bands within certain flocks indicates different evolutionary or manipulation histories. �TabJc2 Bands IlCCUalogio 1M Doge from S,OOO 10 JO,ooo ol"'~tidc blllca (± - p"""""'cc oftbc baod io ao iadhlidllal) KILO-BASBS Individual 9.6 8.8 8.2 6.8 5.8 7.7 7.2 6.3 5.4 Montezuma County, CO on + + 0/3 + + + 0/4 0/5 0/6 + + + on 0/8 0/9 0/10 0/11 0/13 0/14 0115 0/16 0/17 0/18 0/19 0120 onl 0122 Montrose County. CO Mn M/3 + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + MI20 Mnl MI22 Arizona An AI3 + + + + + + + + + + + + + + + + + + + + + + + + + + + + AI7 + + + + + + + + + + + + + + AIlS Al16 All7 All8 All9 AnO Al2l AI22 + + + + + + AllO AI 11 Al13 Al14 + + + + + Mn Al8 Al9 + + + + + Al4 AlS Al6 5 + + + + + + + + + M/4 MIS M/6 M/8 M/9 MilO Mill M/13 M/14 MilS M/16 M/17 M/18 M/19 + III + + + + + + + + + + + + + �112 Table 3. Band frequencies among wild turkey flocks .and the population at large. Flock 9.6 8.8 8.2 KILO-BASES 7.7 7.2 6.8 Montezuma 0.60 0.60 0.00 0.15 0.00 0.55 0.00 0.20 0.00 0.45 Arizona 0.50 0.40 0.35 0.60 0.50 0.05 0.00 0.00 0.00 0.30 Montrose 0.30 0.30 0.45 0.20 0.00 0.40 0.05 0.05 0.40 0.00 Population 0.47 0.43 0.27 0.32 0.17 0.33 0.02 0.08 0.13 0.25 6.3 5.8 5.4 5.0 Table 4. Chi square analysis on band frequencies of wild turkeys against the population. Flock Chi square Montezuma County 0.998667 > 0.5 Montrose County 0.998311 > 0.5 Arizona 0.997599 > 0.5 IJTERATIIRE CITED Leberg, P. L. 1990. Genetic considerations in the design of.introduction programs. Trans. North Am. Wildl. and Nat. Resour. Conf. 55:609-619. __ . 1991. Influence of fragmentation and bottlenecks on genetic divergence of wild turkey populations. Conserv. BioI. 5:522-530. __ , P. W. Stangel, H. O. Hillestad, R. L. Marchinton, and M. H. Smith. 1994. Genetic structure of reintroduced wild turkey and white-tailed deer populations-. J. Wildl. Manage. 58:698-711. Stangel, P. W., P. L. Leberg, and J. 1. Smith. 1992. Systematics and population genetics. Pages 18-28 in J. G. Dickson, ed. The wild turkey, biology and management. Stackpole Books, Harrisburg, Pa. �113 Vrijenhoek, R. C., and P. L. Leberg. 1991. Lets not throw the baby out With the bath water: a comment on management for MHC diversity in captive populations. Conserv. Bioi. . 5:252-254. Prepared by: 1\,\ e.hancLC. D~ Richard C. Dujay Research Tech. I (ll» ��115 JOB PROGRESS REPOe state ofz Colorado Projectz W-167-R-5 Work Planz Job Titlez 13 Job: Moyements. Plains ShahP-tailed Period COveredz Authorz Upland Bird Research Reproductiye Success. and Habitat Use by Introduced Grouse 01 January Kenneth 10 . throuqh 31 December 1996 M. Giesen Personnelz Jim Aragon, Clait E. Braun, Kenneth M. Giesen, COlorado Division of Wildlife Chuck Loeffler, ABSTRACT A total of 33 plains sharp-tailed grouse (tympanuchus phasianellus jamesi) was trapped in southeastern Wyoming and transplanted onto Raton Mesa i.nLas Animas COunty, COlorado in April 1996. Ten of 20 males and 10 of 13 females were fitted with radio transmitters prior to release. Weather conditions at the time of release were more moderate than in 1995 with cool temperatures and less than 30\ snow cover on the Mesa. Documented mortality of radio-marked birds was 35\ (7 of 20) within 60 days postrelease and signals from another 8 birds (4 males, 4 females) were lost due to lonq distance dispersal or radio failure. Maximum individual dispersal from the release site ranged from 1.3 to 41.6 km with 7 of 13 birds movinq ~ 5.0 km. Home ranges of 3 birds surviving at least 60 days ranged from 0.32 to 117.12 km2• Height density indices within grasslands used by radiomarked birds ranged from 1.24 ± 0.66 dm to 3.45 ± 1.35 dm. Siqhtings of unmarked birds and survival of radio-marked birds indicates that spring through fall habitat on Raton Mesa meets the minimum requirements for this species. ��117 NOVEME!r.rS, REPRODUCTIVE SUCCESS, AND HABIDT USE BY INTRODUCEDPLAINS SBARP-~AILED GROUSE Kenneth M. Giesen IHmOWCTIOH Plains sharp-tailed grouse historically occurred in suitable foothill and riparian habitats along the Front Range of Colorado. Sharp-tailed grouse populations declined with human settlement and were exti;pated from most of their range in eastern Colorado by the late 1800's. Although the historical breeding population of sharp-tailed grouse in Douglas County continue to decline, small breeding populations and winter migrants or transients have been reported in recent years from Yuma, Logan, and Weld counties (Haag and Braun 1990, Braun unpubl. data). However, the total breeding population of plains sharp-tailed grouse in Colorado remains small «300 birds) and most populations are associated with privateiy-owned lands and, therefore, subject to land management activities which may have detrimental consequences. Plans to increase distribution and popu~ations of plains sharp-tailed grouse in Colorado will rely primarily on transplants (Braun et ale 1992). While numerous transplants of prairie grouse have been attempted in North America, few have been successful (Toepfer et ale 1990, Rodgers 1992, Hoffman et ale 1992). A previous transplant of sharp-tailed grouse into Las Animas County near Raton Mesa was attempted with a total of 85 males and 83 hens being . released over a 3-year period (1987-89). While this transplant was not thought to be successful in establishing a breeding population, little followup of released birds was conducted and important data on survival and movements are lacking. Thus, it is.desirable to document responses of sharptailed grouse to experimental transplants and evaluate parameters potentially affecting success including movements, habitat use, mortality, and reproduction. P. N. OBJECTIVES The objectives of this project are to assist with trapping and transplanting of plains sharp-tailed grouse into selected sites along the Front Range of Colorado and evaluate transplant success. Population characteristics of the transplanted population including movements and home range size, timing and causes of mortality, habitat use, and nest success will be compared to those described in the literature for native and transplanted prairie grouse. Results of this study will assist in developing transplant protocols for future transplants of prairie grouse. SEGMENTOBJECTIVES 1. Review literature habitat use. on prairie grouse introductions, movements, and 2. Coordinate landowners for plains 3. Transplant up to 50 plains sharp-tailed grouse from southeastern into suitable habitats along the Front Range of COlorado. efforts with Wyoming Game.and Fish personnel and affected in southeastern Wyoming to locate potential trapping sites sharp-tailed grouse. Wyoming �118 4. Radiomark up to 25 sharp-tailed grous~ in the transplanted population and monitor movements, habitat use, reproduction, and mortality. 5. Conduct a pre-release evaluation of the habitat at the selected release site. 6. Prepare annual progress report. METHODS Contact was made with personnel of the Wyoming Game and Fish Department to obtain the necessary- permits for trapping plains sharp-tailed grouse in southeastern Wyoming for transplant into Colorado. Active dancing grounds in southeastern Wyoming were located as potential trapping sites and permission from affected landowners was obtained. Walk-in funnel traps (Toepfer et ale 1988, Schroeder and Bra~n 1991) were used to capture male and female sharptailed grouse on dancing grounds. captured birds were classified to age and sex and fitted with serially-numbered aluminum ban4s on the right leg and yellow colored plastic bandettes on both legs. Battery-powered transmitters (weight 12-13 gmS) were attached with a necklace (Amstrup 1980) to selected birds to facilitate monitoring of movements and survival after release .on Raton Mesa. A portable Global Positioning Satellite (GPS) receiver was used to record locations of birds and minimum convex polygon home ranges were calculated using the McPaal software package (M. Stuwe and E. E. Blohowiak, Conserv. Res. Cent., Natl. Zool. Park, Smithsonian Inst., Front Royal, Virginia, 1985). Vegetative cover (height-density indices) in the release area was measured using a Robel pole (Robel et ale 1970). BESm.TS Transplant o£ sharp-tailed grouse A total of 33 (20 males, 13 females) sharp-tailed grouse was captured in Platte and Goshen counties, Wyoming and released onto Raton Mesa in Las Animas County, Colorado in April 1996. One additional male died during the trapping process. Birds were trapped on 4 dancing grounds (Baker Swale c 1 male, 8 females, Kennedys'-=10 males, 5 females, Thomas Jeffersons -=4 males; Grange c 5 males). captured birds were held in captivity up to 4 days prior to release. Birds were transported by truck and helicopter to the release site. At the release site-birds placed in holding boxes for 5-10 minutes before the doors were opened to allow escape. Twenty birds (16 males, 4 females) were released on 5 April, 3 hens were released on 12 April, and 4 males and 6 hens were released on 19 April. Ten male and 10 female sharp-tailed grouse were fitted with radio transmitters prior to release. In contrast to 1995, weather conditions at the time of release and during the following month were characterized by mild temperatures and average precipitation. Snow cover on Raton Mesa was <30 percent, compared to 100 percent in 1995, which allowed better access for monitoring birds. Mortality Depredation of 6 released birds (3 males, 3 females) was documented from 5 to 35 days post-release (Table 1). One-hen was_found freshly killed by vehicle traffic 61 days post~release, 5 birds (2 males, 3 females) dispersed off Raton Mesa and could not be located, and signals from 2 birds (1 male, 1 female) �119 were not heard following their release. Mortalities were documented from 1.31 km to 41.62 km from the release site with most occurring within 30 days of release. Several attempts to locate radio signals from missing birds using aircraft were unsuccessful. Known mortality and dispersal from Raton Mesa resulted .in high loss of radiomarked birds following release (at least 14 of 20 birds). While depredation by avian arid mammalian predators was high, dispersal from the release site was likely a greater cause of mortality following release of translocated birds. No sharp-tailed grouse known to have dispersed from Raton Mesa was known to return and the habitat surrounding Raton Mesa was likely lacked food resources or adequate cover. Mortality attributed to depredation may have been influenced by the condition of the birds following capture, captivity (1-4 days), and lack of familiarity with food resources and escape cover following release. Table 1. Fates of radio-marked sharp-tailed Las Animas County, Colorado, 1996. Bird # Age 2+ 2+ 2+ 2+ 2+ 2+ 12+ 2+ 2+ 2+ 2+ 2+ 12+ 2+ 2+ 12+ 2+ 1383 1497 1566 1649 1779 1819 1889 0455 1799 1195 1958 1698 1751 1589 1518 1539 1395 1859 1343 1678 Sex MaxDist& (m) M M M M M F F M F M F F F F F F F M M M 9,550 n.d.b n.d. 4,670 2,350 n.d. 6,619 3,984 2,350. n.d. 7,680 n.d. n.d. n.d. 1,310 n.d. 41,620 7,590 2,390 2,204 grouse released Home Range (km2) 2.39 0.31 117.12 0.32 & Maximum distance from release site. b No data, bird not located after release Movement. on Raton Mesa, Fate Raptor kill, 10 May Signal from off Mesa Signal from off Mesa Radio fell off, 23 Apr Depredation, 1 May Signal from off Mesa Radio fell off, 28 Aug Raptor kill, 2 May Mortality, 1 May No signal postrelease .Last signal 3 July Signal from off Mesa No signal postrelease No signal postrelease Mammal kill, 9 May Signal from off Mesa Roadkill, 19 June Radio fell off, 28 Aug Raptor kill, 24 Apr Last signal 6 June onto Raton Mesa. and Home Range The distribution of movements from the release site was bimodal with some birds staying relatively close to the release site while others dispersed off Raton Mesa where none were relocated alive (Table 1). High mortality of some birds Boon after release may have skewed innate movement patterns, although some birds dispersed> 6.0 km.but eventually returned to the vicinity of the �120 release site. Some of these birds were not visually located or the radios recovered although mortality signals were received for >30 days. Although sample sizes were small, it did not appear that mortality or post-release movements were related to sex of the grouse, with both males and hens showing long dispersal movements. Minimum convex polygon home ranges were calculated for 4 grouse (1 male, 3 females) which were located periodically for> 60 days after release. Home ranges of 3 birds were relatively small (0.31 - 2.39 Jcm2,Table 1) and may have been affected by the few locations of each bird. The largest home range resulted from a long dispersal movement from the. release site into New Mexico (where the bird was struck by a vehicle and killed). COnsecutive locations at 1-2 week intervals showed that birds were usually within 200-400 m of their previous locations. One hen initially dispersed >6.6 Jan.south from the release site but returned after 5 weeks to within 1.1 Jan of the release site where she nested. She incubated a clutch of 11 eggs of which 5 eventually hatched in late June. Her movements for the remainder of the summer were within 400m of the nest site, and at least·3 chicks survived to 60 days-of-age. Habitat Height-density of vegetation was measured during June-August at 200 points within 4 pastures (50 measurements/pasture) where radio-marked sharp-t~iled grouse were regularly located. The greatest height-density was measured at the release site pasture (3.45 ± 1.35 dm). Overall the two pastures in COlorado had 58 of 100 VOR measurements ~ 3.0 dm and 23 of 100 VOR measurements ~ 4.0 dm. Pastures in New Mexico where sharp-:-tailed grouse occurred during May-September had mean height-density measurements Of 1.59 ± 0.54 dm and 1.24 ± 0.66 dm, reflecting greater livestock grazing. However, height-density was quite variable in these pastures with 21\ having heightdensity meas.urements ~ 2.25 dm. The average height-density measured in both COlor~do pastures in 1996 was slightly less than in 1995 and likely refle·cted the high precipitation the areas received during March-June of 1995 and the drier conditions in 1996. DISCUSSION Results from the second years t~ansplant indicate high ~ortality of released birds and dispersal from the release site. The long movements may result from birds seeking suitable habitats or trying to return to their native area. Hoffman et ale (1992) recorded post-release movements up to 29 Jan for greater prairie-chickens (Tympanuchus cupido)transplanted in spring. High winds and the location of the release site relative to the edge of the mesa likely resulted in some birds flushing frein the mesa top and landing in forested habitats. Because the elevation on top of Raton Mesa is 300-600 m above the surrounding terrain any birds leaving the mesa top may have perished before finding their way back. No birds were known to leave the top of the mesa and return. The west, north, and east sides of Raton Mesa are surrounded by forests and steep cliffs; ~onditions which may hinder return movements to the mesa top. Lack of familiarity with food resources and escape cover combined with several'days of captivity, may have weakened the birds and increased their susceptibility to predation. �121 It is not known whether the radio-marked birds suffered higher mortality than those not marked as reported in other studies (Marks and Marks 1987). Sharptailed grouse without radios were observed regularly following the release indicating actual survival may have been higher than indicated by radio-marked birds. While no nesting attempts were documented, it is possible that reproduction may have occurred by unmarked birds and was not detected. The success of this project may depend upon the successful release of additional sharp-tailed grouse in 1996. If weather conditions are favorable, mortality and dispersal movements of released birds may be reduced and chances for reproduction enhanced. One factor not studied is the availability of winter food for this population. Waste grain from wheat fields was available in Wyoming where these grouse were captured but is lacking on Raton Mesa and adjacent areas. If the grouse released onto Raton Mesa need to disperse into New Mexico to find winter food, they may not return to breed near their release site. LITERATURE CITED AmstrUP, S. C. 1980. A radio-collar 217. for game birds. J. Wildl. Manage. 44:214- Braun, C. E., R. B. Davies, J. R. Dennis, K. A. Green, and J. L. Sheppard. 1992. Plains sharp-tailed grouse recovery plan. COlorado Div. Wildl., Denver. 33 pp. Hoag, T. W., and C. E. Braun. 1990. Status and distribution tailed grouse in COlorado. ·Prairie Nat. 22:97-102.' of plains sharp- Hoffman, R. W., W. D. Snyder, G. C. Miller, and C. E. Braun. 1992. Reintroduction of greater prairie-chickens in northeastern COlorado. Prairie Nat. 24:197-204. Marks, J. S., and V. S. Marks. 1987. Irifluence of radio-collars sharp-tailed grouse. J. Wildl. Manage. 51:468-471. on survival Robel, R. J., Briggs, J. N., Dayton, A. D., and Hulbert., L. C. 1970. Relationships between visual obstruction measurements and weight grassland vegetation. J. Range Manage. 23:295-297. Rodgers, R. D. 1992. A technique for establishing sharp-tailed unoccupied range. Wildl. Soc. Bull. 20:101-106. grouse of of in Schroeder, M. A., and C. E. Braun. 1991. Walk-in traps for capturing prairie-chickens on leks. J. Field Ornithol. 62:378-385. Toepfer, J. E.; R. L. Eng, and R. K. Anderson. 1990. Transplanting prairie grouse: what have we learned? Trans. North Am. Wildl. and Nat. Resour. COnf. 55:569-579. _____ , J. A. Newel, arid J. Monarch. 1988. A method for trapping grouse hens on display grounds. Pages 21-23 in A~ D. Bjugstad, prairie Tech. �122 Coord. Prairie chickens on the Sheyenne National Agric •• For. Servo Gen. Tech. Rep. RM-159. Prepared by Kenneth M. Giesen Wildlife Researcher C Grasslands. u.S. Dep. �123 JOB PROGRESS REPORT State of: Colorado Project: Work Plan: W-167-R 13 :Job Job Title: Avian Research 11 Eya1uation of Columbian Sharp-tailed Grouse Reintroduction Opportunities in Western Colorado Period Covered: 01 January through 31 December 1996 Author: Richard W. Hoffman Personnel: Richard W. Hoffman, Colorado Division of Wildlife ABSTRACT Efforts for this reporting period focused on reviewing literature, conducting 1ek surveys, and gathering information to assist in the formation of working groups. ��125 EVALUATION OF COLUMBIAN SHARP-TAILED REINTRODUCTION OPPORTUNITIES IN WESTERN COLORADO Richard W. Hoffman "INTRODUCTION Use of common names and misidentification of blue grouse (Dendragapus obscurus) and sage grouse (Centrocercus urophasianus) by" early explorers have made it difficult to ascertain the precise distribution of Columbian sharptailed grouse (Iympartuchus phasianellus columbianus) in Colorado (Rogers 1969, Giesen and Braun 1993). However, historical records suggest this subspecies may have occurred in at least 22 counties in western Colorado (Bailey and Niedrach 1965, Rogers 1969). Recent surveys indicate viable populations are restricted to Moffat, Routt, and Rio Blanco counties, with possible remnant populations in Mesa and Montrose counties (Giesen and Braun 1993). Similar reductions in the distribution of Columbian sharp-tailed grouse have occurred throughout western North America (Miller and Graul 1980). This decrease in distribution resulted in "designation as a Category 2 species (U. S. Dep. Inter. 1989). Factors responsible for the reduction in distribution include conversion of native rangeland to cropland, excessive grazing by livestock, vegetative succession due to fire suppression, herbicide treatments, mineral exploitation, and urban development (Meints et al. 1992, Giesen and Connelly 1993). These factors have had the most pronounced impact on nesting, brood rearing, and winter cover through loss of native grasses and deciduous shrubs (Giesen and Braun 1993). Cover types used by Columbian sharp-tailed grouse tend to be structurally and vegetatively diverse with an extensive deciduous shrub component (Meints et al. 1992, Giesen and Connelly 1993). In Colorado, �126 Columbian sharp-tailed grouse occur in mountain shrub communities interspersed with grasslands, small aspen (Populus tremuloides) stands, and riparian zones (Giesen 1987). Serviceberry (Amelanchier spp.) is an essential element of these communities and usually grows in association with one or more of the following deciduous shrubs: Gambel oak (Quercus gambelii), common chokecherry (Prunus virginiana), snowberry (Symporicarpos spp.), and sagebrush (Artemisia spp.) (Giesen 1987). Wheat is the primary agricultural crop within the range of sharptails in western Colorado. Wheatfields may be used during late summer and fall after the wheat has been harvested. These fields are usually snow- covered and unavailable during winter. Much of what is known about Columbian sharp-tailed grouse in western Colorado has resulted from studies in the northwest portion of the state (Dargan et al. 1942, Rogers 1969, .Giesen 1987). Little is known about sharp- tailed grouse in southwestern Colorado other than they once occurred there and may still exist in low densities on the north end of the Uncompahgre Plateau (Rogers 1969, Giesen 1985). It has been 10 years since the last intensive effort to conduct lek surveys for Columbian sharp-tailed grouse in western Colorado. Another intensive effort is needed because changes in land use practices have occurred since then including implementation of the Conservation Reserve Program, additional mining· and development activities, and alteration of grazing practices. Perhaps the most important action in the last 10 years affecting the need for current population and distribution data has been the petition to list Columbian sharp-tailed grouse as "threatened" or "endangered" in the lower 48 conterminous United States pursuant to the Endangered Species Act (Carlton 1995). This action is of special significance in Colorado because Idaho and Colorado are the only states that allow hunting of Columbian sharp-tailed grouse and that still have adequate populations to �127 provide transplant stock for future restoration programs. Opportunities for management of sharptai1s in western Colorado may be limited because much of the occupied habitat occurs on private lands. The· most extensive areas of public lands within the historic distribution of Columbian sharp-tailed grouse are in southwest Colorado. The last confirmed sighting of sharptai1s on these lands was in 1985 (Giesen 1985). Before a reintroduction program can be implemented, current habitat conditions and status of sharp tails on these lands must be evaluated and management strategies formulated based on the outcome of the evaluation. It is likely that any effort to restore sharptai1s in western Colorado will require a commensurate effort to restore and protect the habitat. P. N. OBJECTIVES Objectives of this project are to (1) form a sharp-tailed grouse working group with broad citizen, community, and agency representation, and in cooperation with this group, prepare a conservation plan for Columbian sharptailed grouse in Colorado, (2) conduct intensive 1ek surveys of Columbian sharp-tailed grouse in northwest Colorado, (3) ascertain presence or absence of sharp tails on historic range in southwest Colorado, (4) identify potential reintroduction sites within the historic range of Columbian sharp-tailed grouse, (5) evaluate existing habitat conditions on these sites based on the habitat suitability index model describe&by Meints et a1. (1992), and (6) cooperate with other western states in preparing conservation strategies for Columbian sharp-tailed grouse. SEGMENT OBJECTIVES 1. Review literature pertinent to the objectives of this study. �128 2. Identify participants interested in preparing conservation plan. 3. Organize and conduct public meetings. 4. Form working group and conduct regularly scheduled meetings to develop management strategies and prepare conservation plan. 5. Prepare conservation plan in collaboration with working group. 6. Conduct leks surveys in Moffat, Routt, and Rio Blanco counties. 7. Conduct surveys to ascertain presence or absence of sharptails in Mesa (north end of Uncompahgre Plateau) and Montrose (Horsefly Creek) counties. 8. Select study sites for habitat suitability analysis. 9. Estimate percent winter cover and nestingfbrood habitat within 6.5 and 2.5 km radius, respectively, of study leks using aerial photographs. 10. Obtain height/density measurements along randomly located transects in nest/brood habitat. 11. Within each winter cover type, estimate distance to nearest nest/brood habitat. Within each nestfbrood habitat type measure the distance to the nearest winter cover. 12. Calculate habitat suitability indices for winter and nest/brood habitat and rank lek sites. 13. Compile data, analyze resul~s, and prepare progress report. RESULTS AND DISCUSSION Segment objective 1 - Literature on all aspects of the biology and ecology of sharp-tailed grouse was reviewed. Literature searches were conducted through Current Contents and the Fish and Wildlife Reference Service. Efforts were made to review draft and final conservation plans and strategies prepared by other states and to talk with the people involved in �129 preparing these documents. Efforts also were made to review all documents pertaining to the petition to list the Columbian Sharp-tailed grouse as threatened or endangered. Segment objective 2-4 - Discussions were held with the Human Dimensions section of the Colorado Division of Wildlife and with other CDOW personnel with experience in organizing and conducting public meetings. In addition, stakeholder documents (i.e., manuals, plans, and strategies for effective stakeholder involvement) obtained from the Human Dimensions section were reviewed. The CDOW maintains a database of external publics and key contacts in other agencies that are routinely invited to participate in stakeholder meetings. This list was reviewed and names were added or deleted as deemed necessary. Meetings were held with CDOW field personnel to determine their interest in participating in the working groups and to obtain names of local landowners, businesses, and organizations that should be invited to participate. A letter was drafted to organize informational meetings at several locations in western Colorado. Too date, no official working group has been formed. Attempts were made to join existing sage grouse working groups in northwest and southwest Colorado, but members of these groups were opposed to the idea. This has created problems in terms of generating interest from other agencies because their personnel are already involved with other working groups and there is only so much time they can devote to this activity. Another problem was the failure of the U. S. Fish and Wildlife Service to act on the petition to list the Columbian sharp-tailed as threatened or endangered. listing, interest in sharptails has diminished. Without the threat of However, this should change in the near future as the Service is required by law to act on the petition �130 and should do so sometime in 1998. This can be used as leverage to generate interest in developing conservation plans to prevent listing. Segment objective 5 - Information (iiterature, unpublished data ..etc.) has been gathered to use in the preparation of conservation plans but preparation of the plan will not begin until 1998. Segment objective 6 - Lek surveys were conducted during March-June 1997 and results will be reported in the 1997 federal aid reports. efforts focused on locating new leks. the status of existing leks. Briefly,· Area personnel were asked to document There was 'insuffient manpower and time to search Moffat, Routt, and Rio Blanco counties; thus, searches were confined to Routt County with plans to search Moffat and Rio Blanco counties during spring 1998. Thirty-one new leks were located in Routt county during 1997; 61% were in CRP fields, 32% in mine reclamation, and 7% in grassy openings within the mountain shrub community. In addition, 8 historic leks listed as inactive were relocated within 0.5 km of the original lek site. In all cases the leks moved into nearby CRP fields. Segment objective 7 - No sharptails were located in Mesa (north end of Uncompahgre Plateau and Pinon Mesa), Montrose (Horsefly Creek), o~ Dolores (Salter Y) counties. The last confirmed sightings of sharptails in southwestern Colorado occurred in 1985 near the Cold Springs Ranger Station in Mesa County. Segment objectives 8-12 - Habitat suitability analyses were not performed because there was insuffient time to do both habitat work and lek surveys. The habitat work should be done in mid- to late May, however, there are still opportunities .to conduct lek surveys at this time. northwestern Colorado are not accessible until mid Mayor Many areas in later and it was considered more important to survey these areas for leks than to collect �131 habitat data. Efforts will continue to focus on lek surveys in spring 1998. However, attempts will be made to collect some habitat·data in 1998 from 3 active leks in Moffat and Routt counties. The Habitat Suitability Model developed for sharp-tailed grouse in Idaho (Meints et al. 1992) was ·applied to the western portion of Mesa Verde National Park during spring 1996. Sharp-tailed grouse have been identified in archeological zecords from Mesa Verde NP and were known to occupy habitats northwest of the Park until the early 1960's. Approximately 2,886 ha of potentially suitable habitat was identified within the Park; 2,078 ha (72%) was identified as potential winter habitat and 808 ha (28%) as nestingfbrood rearing habitat. Most of the suitable habitat was created within the past 20 years by a series of intense fires that decreased the amount of oakbrush, juniper, and pinyon and stimulated the growth of serviceberry, snowberry, and grasses. The mean distance from nestfbrood cover to the nearest winter cover was 241 m, whereas the mean distance from winter cover to the nearest nestfbrood cover was 504 m. Visual obstruction measurements (Robel 1970) averaged 3.4 dm (13 in) for nestfbrood areas and 5.0 dm (20 in) for winter areas. The suitability indices for winter and nestfbrood cover were 0.4 and 0.72, respectively. Meints et ale (1992) advised against releasing birds into areas where the HSI is < 0.75 for one or both of the habitat components. In .the case of Mesa Verde NP, the habitat is suitable for sharptails but the quantity and dispersion of winter and nestfbrood rearing habitats are below recommended levels for introducing sharptails. �132 LITERATURE CITED Bailey, A. M., and R. J. Niedrach. Mus. Nat. Hist., Denver, Co. Carlton. J. C,. 1995. 1965. Birds of Colorado, Vol. 1. Denver 454pp. Petition for a ~ule to list the Columbian sharp-tailed grouse, Tympanuchus phasiane11us columbianus, as "threatened" or "endangered" in the conterminous United States under the Endangered Species Act, 16 U.S.C. Sec. 1531 et seq. (1973) as amended. Biodiversity Legal Foundation, Boulder, CO. 52pp. Dargan, L. M., H. R. Shepherd, and R. N. Randall. 1942. grouse in Moffat and Routt counties. Sage Grouse Survey, Vol 4, Denver. Giesen, K. M. 1985. Colorado. Giesen, K. M. Data on sharp-tailed Colo. Game, Fish, and Parks Dep, 28pp. Inventory of Columbian sharp-tailed grouse in western Colo. Div. Wildl., Unpubl. Rep. Fort Collins. 1987. 6pp. Population characteristics and habitat use by Columbian sharp-tailed grouse in northwest Colorado. Res. Rep., Part 2. Pages 251-279 in Wildlife Colo. Div. Wildl., Fed Aid Proj. W-152-R, Apr. 1987. Giesen, K. M., and C. E. Braun. 1993. sharp-tailed grouse in Colorado. Giesen, K. M., and J. W. Connelly. Status and distribution of Columbian Prairie Nat. 25:237-242. 1993. Guidelines for management of Columbian sharp-tailed grouse habitats. Wildl. Soc. Bull. 21:325-333. Meints, D. R., J. W. Connelly, K. P. Reese, A. R. Sands, and T. P. Hemker. 1992. Habitat suitability index procedure for Columbian sharp-tailed grouse. Idaho For .• Wildl .• and Range Exp. Stn. Bull. 55, Moscow. 27pp. Miller, G. C., and W. D. Graul. America. 1980. Status of sharp-tailed grouse in North Pages 18-28 in P. A. Vohs and F. L. Knopf, eds. of the prairie grouse symposium. Proceedings Oklahoma State Univ., Stillwater. �133 Rogers~ G. E. 1969. The sharp-tailed grouse in Colorado. and Parks Tech. Publ. 23, Denver. Colo. Game, Fish, 94pp. Robel, R. J., J. N. Briggs, A. D. Dayton, and L. C. Hulbert. 1970. Relationships between visual obstruction and weight of grassland vegetation. J. Range Manage. 23:295-297. U. S. Department of the Interior. 1989. Endangered and threatened wildlife and plants; annual notice of review; proposed rules. 54:560. Prepared by Researcher/Scientist IV Fed. Register ��135 JOB PROGRESS REPORT State of: Colorado Project: W-167-R Work Plan: Job Title: 22 Upland Bird Research : Job _1_ Upland Bird Research Publications Period Covered: 01 January through 31 December 1996 Author: Clait E. Braun Personnel: Clait E. Braun, K. M. Giesen, R. W. Hoffinan, T. E. Remington, and W. D. Snyder, Colorado Division of Wildlife ABSTRACT The following articles were published in 1996: Braun, C.E. 1996. Status of prairie grouse in North America. 7th Int. Grouse Symp., Fort Collins, CO. Abstract. W-37-R, Work Plan 3, many jobs; W-167-R, Work Plan 3, Jobs . IS and 19. ___ -" and T.D.I. Beck. 1996. Effects of research on sage grouse management. Trans. North Am. Wildt. and Nat. Resour. Conf. 61:429-436. W-37-R, Work Plan 3, Jobs 3-17; W-167-R, Work Plan 3, Jobs IS and 19. __ ---' J.H. Enderson, M.R Fuller, Y.B. Linhart, and C.D. Marti. 1996. Northern goshawk and forest management in the southwestern United States. The Wildt. Soc. Tech. Rev. 96-2. 19pp. W-SS-R, Work Plan 4, Jobs 4-S. .. Commons, M.L., RK. Baydack, and C.E. Braun. 1996. Gunnison sage grouse use of fragmented habitats in southwestern Colorado. 7th Int. Grouse Symp., Fort Collins, CO. Abstract. W-167-R, Work Plan 26, Job 1. ___ .:» ---' and . 1996. Sage grouse movement and habitat use in southwestern Colorado. Proc. Annual Conf., The Wildt. Soc. 3:62. W-167-R, Work Plan 26, Job 1. Connelly, Jr., J.W., and C.E. Braun. 1996. Long-term changes in sage grouse populations in the western United States. 7th Int. Grouse Symp., Fort Colliris, CO. Abstract. W-167-R, Work Plan 3, Job 19; Work Plan 26, Job 1. Giesen, K.M. 1996. Grouse habits and habitats. Colorado Outdoors Hunting Guide 1996: 3133. W-37-R and W-167-R, Work Plan 17, Jobs 1-7 and Work Plan 13, Jobs 9 and 10. �136 ___ -> and G.D. Kobriger. 1996. Status and management of sharp-tailed grouse in North America. 7th Int. Grouse Symp., Fort Collins, CO. Abstract. W-167-R, Work Plan 13, Job 10. Hoffinan, RW. 1996. Where the birds are - the sport of ptarmigan hunting. Colorado Outdoors Hunting Guide 1996: 26-29. W-37-R, Work Plan 17, Jobs 1-7 and W-167-R Work Plan 17, Job 7. ___ and G.M. Beauprez. 1996. Reintroduction 'of greater prairie-chickens in northeastern Colorado. 7th Int. Grouse Symp., Fort Collins, CO. Abstract. W-167-R, Work Plan 14, Job 4. -> Lancia, RA., C.E. Braun, M.W. Collopy, R.D. Dueser, r.o. Kie, c.i Martinka, lD. Nichols, . T.D. Nudds, W.R Porath, and N.G. Tilghman. 1996. ARM! For the future: adaptive resource management in the wildlife profession. Wildl. Soc. Bull. 24: 436-442. W-167R, Work Plan 26, Job 1. Martin, K., P.B. Stacey, and C.E. Braun. 1996. Demographic rescue and the maintenance of population stability in grouse-beyond metapopulations. 7th Int. Grouse Symp., Fort Collins, CO. Abstract. W-167-R, Work Plan 17, Job 7. Melcher, C., and K. Giesen. 1996. Blue grosbeaks: a response to habitat changes and a late nesting record. Colorado Field Ornithol. Jour. 30: 158-161. W-167-R, Work Plan 13, Job 10. Oyler, S.J., C.E. Braun, and K.P. Burnham. 1996. Use ofa habitat-based model to predict sage grouse occupancy of patches in southwestern Colorado. 7th Int. Grouse Symp., Fort Collins, CO. Abstract. W-167-R, Work Plan 3, Job 19. Quinn, T.W., N.W. Kahn, J.R Young, N. Benedict, S. Wood, D. Mata, and C.E. Braun. 1996. Probing the evolutionary history of sage grouse populations using mitochondrial DNA sequence and nuclear micro satellites. 7th Int. Grouse Symp., Fort Collins, CO. Abstract. W-167-R, Work Plan 3, Job 19. Remington, T.E. 1996. Bright future for pheasants. Colorado Outdoors Hunting Guide 1996: 20-25. W-167-R, Work Plan 1, Job 24. ___ ___ and RW. Hoffinan. 1996. Costs of detoxification ofxenobiotics in conifer needles to blue grouse. 7th Int. Grouse Syrnp.jFort Collins, CO. Abstract. W-167-R, Work Plan 9, Job 8. -> -' and . 1996. Food habits and preferences of blue grouse during winter. J. Wildl. Manage .. 60: 808-817. W-167-R, Work Plan 9, Job 8. Snyder, W.O. 1996. Habitat management for upland game birds on eastern Colorado sandhill rangeland. Colorado Div. Wildl., Wildl. Info. Leafl. 115. 4pp. W-167-R, Work Plan 21, Job 3. Prepared by Clait E. Braun Avian Program Manager �137 JOB PROGRESS REPORT State of: Colorado Project: W-167-R Upland Bird Research Work Plan: 26 : Job _1_ Job Title: Analysis of Upland Bird Population Trends Period Covered: 01 January through 31 December 1996 Author: Clait E. Braun Personnel: Clait E. Braun, Kenneth M. Giesen, Richard W. Hoffinan, Thomas E. Remington, and Warren D. Snyder, Colorado Division of Wildlife ABSTRACT The following reports were published in 1996: Braun, C.E. 1996. Sage grouse counts, Blue Mountain, 1996. 1996. Sage grouse counts, Cold Spring Mountain, 1996. 1996. Sage grouse counts, Gunnison Basin, 1996. 1996. Sage grouse counts, Lower Moffat County, 1996. 1996. Sage grouse counts, North Park, 1996. Commons; M.L., and C.E. Braun. Sage grouse investigations, Crawford Area, Montrose County, 1996. Giesen, K.M. 1996. Columbian sharp-tailed grouse harvest data, northwest Colorado, 1976-96. Hoffinan, RW. 1996. Analyses of statewide blue grouse wing collections for 1996. Remington, T.E. 1996. Analysis of small game harvest surveys conducted by telephone and by conventional post-card surveys. ___ .. 1996. Analysis of pheasant program in relation to LRP goals. Snyder, W.D. 1996. Planting shrub thickets - a review. Prepared by Clait E. Braun Avian Program Manager ��139 JOB PROGRESS REPORT State of Colorado Project: W'-171-R-l: WO.rk Plan Bald Ea21e Nest Site Protection and Enhancement Pro2ram _2_ : Job _2_ Job Title: Bald Eaele Period Covered: Nest Site Protection and Enbapcement Proeram 1 January - 31 December 1996 Personnel: G.R. Craig, Colorado Division of Wildlife ABSTRACT Activities conducted during this period have already been reported under W-151-R-8 (see attached report). � https://cpw.cvlcollections.org/files/original/50dabd9a65b36e2d00d4d051cd18ead6.pdf 2481996d13f652ad0ac201a1232afd07 PDF Text Text Colorado Division Wildlife Research July 1997 of Wildlife Report JOB PROGRESS State of Project REPORT Colorado No. ~W~-~1~5~3,-~R~-_.1~0~ _ Mammals Research Work Plan No. Multispecies Job No. consulting Services Mammals Research Period Covered: Author: July Investigations for 1, 1996 - June 30, 1997 G. C. White Personnel: R. M. Bartmann, R. B. Gill, T. D. I. Beck, D. J. Freddy ABSTRACT Progress towards the objectives of this job include: 1. Development of a mule deer population model that allows the user to sample the modeled population to mimic current CDOW procedures. The purpose of this model is to assLst biologists in evaluating their big game modeling procedures. 2. A study of compensatory effects of harvest on the Piceance Basin mule deer population was continued as part of Federal Aid Project W-153-R Work Plan 2 Job 15, entitled Compensatory Effects of Harvest in a Mule Deer Population. Experimental harvests have been conducted in December, 1989, 1990 and 1991. Radio collars to monitor over-winter survival of fawns were placed on the animals during November, 1989-95. The results of this study have been accepted for publication in the Journal of Wildlife Management. 3. Preliminary cooperation 4. A World-Wide-Web page at http://www.cnr.colostate.edu/-gwhite/software.html has been maintained to distribute software developed contract • . 5. analysis of elk sightability with David Freddy. models was performed under in this Assistance was provided in the design and analysis estimate black bear abundance in western Colorado. of a study to 6. Assistance was provided in the design estimate kit fox abundance in western and analysis Colorado. of a study to 7. Assistance was provided in the design and analysis estimate swift fox abundance in eastern Colorado. of a study to ��3 CONSULTING SERVICES FOR MARK-RECAPTURE ANALYSES G. C. White P. N. OBJECTIVES Develop a model of mule deer populations monitoring procedures. SEGMENT 1. based on current and proposed OBJECTIVES Summarize philosophy and methodology in the development of a mule model based on current and proposed CDOW monitoring procedures. RESULTS CDOW deer AND PISCUSSION Introduction. The Colorado Division of Wildlife (CDOW) has been a leader in development of methods for monitoring the status of mule deer populations. Quadrat counts (Kufeld et al. 1980, Bartmann et al. 1986) conducted from helicopters during December-January provide population estimates, and December age and sex ratios, again determined from helicopters, provide estimates of recruitment and herd composition. Although annual estimates of these parameters would be desirable, costs are prohibitive, so population size is estimated every 3-5+ years and age ratios estimated every 1-2 years for major management units. Harvest estimates are obtained annually from phone surveys (White 1993, Steinert et al. 1994). From these data, population models are developed to project the population and establish harvest objectives for the coming year. Unfortunately, the 1 variable to which the model is most sensitive is survival, and no estimates of survival are routinely taken as part of monitoring procedures. This paper has 2 objectives: 1) to present reasons why monitoring of survival is es.sential to project the trajectory of deer populations, and 2) to describe a monitoring system that includes estimates of survival and is within current budget constraints for state-wide deer monitoring. To implement these objectives, we will first describe a simple population model. Then, the importance of the sensitivity of the model to parameter values and the imPortance of temporal variation to model predictions will be explained. Finally, the need for a more complex "planning model" currently under development will be described. The crucial philosophy underlying this paper is that management decisions must be based on data. In other words, the management of mule deer in Colorado should not be based on model predictions where the model inputs are not provided from measurements made in the field. Complex models of mule deer dynamics may capture most of our knowledge of this system, but such models do not provide reliable predictions of year-to-year dynamics because of the lack of annual information on required inputs. The issue of model complexity is better comprehended with an analogy to an auto trip from New York City to Los Angeles. No reasonable driver would start this trip with 7.5 minute USGS topographic quadrangles as his/her model. Certainly the topographic quadrangles contain all the necessary information, but the detail is considerably more than needed. A simpler model will suffice, such as state road maps, and is more likely to result in success. An even simpler model of just a single map of the Interstate highways would suffice, but would not provide all the details we might like. Unfortunately, costs usually limit the amount of information available, even though we may desire more. �4 The second crucial philosophy underlying this paper is that good data on a few mule deer herds are better than poor data on all the herds in Colorado. In other words, rigorous monitoring of a few herds provides better inferences to the herds not monitored than does inadequate monitoring on all the herds. Colorado's mule deer populations are managed as Data Analysis Units (DAUs) within which are 1 or more Game Management Units (GMUs). GMUs typically represent mule deer populations or a subset thereof. Population modeling and population objectives are conducted at the DAU level, whereas most monitoring and harvest estimation takes place at the GMU level. MULE DEER POPULATION MODEL To make this presentation explicit, a model of mule deer population dynamics is necessary. This model provides the framework to justify any population monitoring scheme, i.e., the model establishes what population parameters must be measured. The model is simple to economize the amount of input data necessary to use it. Yet, the model must adhere to biological authenticity so that it is useful in projecting mule deer population status. Mule deer population dynamics are much more complicated than the model portrays. However, routine measurement of a wider array of inputs required for a more complicated model is unrealistic. Thus, the model presented here is a reasonable trade-off between what can be measured practically and what is needed to predict mule deer populations for management purposes. The model has only 2 age classes: fawns and adults. The gender of fawns will not be differentiated until they are I-year old. Thus, we define 3 categories in the population: fawns (labeled Juveniles or J), females (F), and males (M). Fawns are recruited into the population in early December when the ratio of fawns to females is estimated. The number (N) of fawns on December 1 is computed as where R(t) is the estimated ratio of fawns to yearling sampled in the population in year t. Total population December in year t is thus and adult females size (NT) in early NT(t) = NJ(t) Total population size prior to the next hunting season is determined by multiplying December fawn and female population segments by over-winter survival rates followed by spring to fall female survival. Estimates of spring to fall survival rates are usually close to 1 so, for simplicity, we will ignore the small amount of mortality during that period. Further, we will assume a constant 50:50 sex ratio for fawns. The equations to project the population from December of year t forward to December of year t+l and after harvest (H) in year t+l are: NF(t+l) SJ(t) 0.50 NJ(t) + SF(t) NF(t) - HF(t+l) NM(t+l) = = SJ(t) 0.50 NJ(t) + SM(t) NM(t) - HM(t+l) NJ(t+l) = R(t+l) and NF(t+l) . The fawn" age class is the observed recruitment discussed above. The model contains 4 parameters that are year-specific: recruitment, juvenile survival, female survival, and male survival. Estimates of harvest could be inflated to account for wounding loss. other assumptions implicit in this model are that males and yearling females have the same survival as 2+-year-old females. We chose to not distinguish yearlings from older animals because data are not collected to �5 support this additional complication. A more elaborate data collection operation would justify a more elaborate model. Given the insufficiency of current data collected by CDOW on mule deer, we opted for the simplest model possible. WHY SURVIVAL ESTIMATES ARE CRITICAL TO MODELING MULE DEER POPULATIONS The relative importance of a parameter in a mule deer population model must be evaluated from 2 perspectives. First is sensitivity of the model to the parameter. Second is how much variation from year to year takes place for each parameter. Sensitivity Sensitivity is defined as the amount of change of the model's output compared to the amount of change of the parameter, referred to as parameter sensitivity (Innis 1979).. Thus, suppose the output from the model is rate of population change defined as A Nt+1/Nt• If adult doe survival (SF) is increased 10% from 0.85 to 0.935, the change in A for SF = 0.85 to the new value of A computed for SF = 0.935 relative to the change in SF is a measure of the sensitivity of A to SF' Technically, sensitivity is defined as the partial derivative of A with respect to the parameter of interest. If SF is increased by amount ~, then = Sensitivity aA = -- asF ' and is often presented as a percentage by multiplying by 100. The proportional sensitivity, or elasticity (Caswell 1989), of 2 or more parameters can be compared by multiplying the sensitivity of a parameter by the parameter value divided by A. Elasticity gives the proportional change in A resulting from a proportional change in the parameter. For SF' the elasticity would be Elasticity aA = = ---SF A aS F Any ungulate model will have a very high sensitivity to adult female survival rates, while sensitivity for recruitment and juvenile survival is similar but considerably less than for adult survival rates. Intuitively, this is because adult survival occurs in the model multiple times for a single cohort of animals, whereas recruitment and juvenile survival only occur once per cohort. For the model described above, an analytical expression can be derived for the rate of population change (A Nt+11 Nt) as a function of the survival and recruitment parameters from a Leslie matrix (Leslie 1945, Caswell 1989) formulation. The Leslie matrix for the above difference equations is = SJ RRSF 0 2 SJ -2 SJ -2 with the dominant eigenvalue SF 0 0 SM of this matrix , A, so that �6 h = RSJ + 2SF 2 where the value 2 is the result of the even sex ratio. Note that the adult male survival rate does not affect population growth rate (and does not appear in this equation), as only females give birth. With this equation, we can compute sensitivity directly, as described above, plus we can compute sensitivity analytically by taking the partial of h with respect to .each of the parameters (Le., R, SJ' and SF). Taking the numerical values of R = 0.64, SJ = 0.40, and SF = 0.85 (Table 1), the resulting value of h is 0.978. When a 10% increase is made in each of the 3 parameters, 1 at a time, the estimates of elasticity are 0.1309, 0.1309, and 0.8691, respectively, for R, SJ' and SF. That is, a 10% increase in either R or SJ results in a 1.309% increase in h, whereas a 10% increase in female survival results in an 8.691% increase in h. The resulting values of hare 0.9908 for Rand SJ' and 10063 for SF. These results suggest a precise estimate of female survival must be used in the model, or else population projections will be seriously biased. Much more bias (about 6.6 times) will result in projections from a 10% error in SF than from a 10% error in either R or S J. Although the model used to obtain these results is not complex, conclusions will be essentially the same regardless of how much more complex the model is structured. Adult survival will always be the most sensitive parameter in a reasonable mule deer population model. Recruitment and overwinter fawn survival will have identical sensitivities (unless sex ratio or sex-specific survival rates are used) and be much lower than adult survival. Temporal Variation The second perspective on the relative importance of parameters in the model is year-to-year variability of the parameters, often labeled temporal variation or environmental variation. How much do each of the 3 parameters vary from year to year? Although computing the variance of a series of estimates of recruitment or survivals would seem appropriate, such is not the case. Variation of the true, but unknown, population parameters is of interest. True survival or recruitment rates are not observed. Rather, we make estimates of these parameters. Thus, the total variance of the series of estimates includes both sampling variance (because only estimates are available) and temporal variation of the true process. To properly estimate temporal variation of the series, the sampling variance of the estimates must be removed. To further understand this concept, consider 2 studies to compute juvenile survival over a 10-year period on the same study area •• One study uses only 10 radios/year, whereas the other uses 100/year. The study with the small sample size will have considerably more variation in the series of estimates because of larger sampling variation, while temporal variation for both studies is identical. Thus, to estimate temporal variation properly, we must remove the sampling variation. In this section, we describe a procedure to remove sampling variance from a series of estimates to obtain an estimate of the underlying process variation (which might be temporal or spatial variation). The procedure is explained in Burnham et al. (1987:260-278). Consider the example of estimating over-winter survival rates for a deer population annually for 10 years. Each year, the true survival rate is different from the overall mean because of snow depth, cold weather, etc. Let the true, but unknown, overall mean be S. Then the survival rate for each year can be considered to be S plus some deviation attributable to temporal variation, with the expected value of the ei equal to zero: �7 Environmental The true population Variation i i Mean 1 8 8 + e1 81 2 8 8 + e2 82 3 8 8 + e3 83 4 8 8 + e4 84 5 8 8 + e5 85 6 8 8 + e6 86 7 8 8 + e7 87 8 8 8 + e8 88 9 8 8 + e9 89 10 8 8 + elO 810 Mean 8 Year i S S mean 8 is computed Year S: as 10 S with the variance of the = S.~ computed LSi i=1 , 10 as: 10 02 = L ( Si - S) 2 i=1 10 where the random variables e. are selected from a distribution with mean 0 and variance 02• In reality~ we are never able to observe the annual rates because of sampling variation or demographic variation. For example, even if we observed all the members of a population, we would still not be able to say the observed survival rate was 8j because of demographic variation. Consider flipping 10 coins. We know the true probability of a head is 0.5, but we will not always observe that value exactly, i.e., get 5 heads from 10 flips. Further, imagine if you flip 11 coins -- the true value is not even in the set of possible estimates. That is, the only possible estimates are 0/11, 1/11, •••, 11/11, with none of the estimates equal to 0.5. The same process operates in a population as demographic variation. Even though the true probability of survival is 0.5, we would not necessarily see exactly ~ of the population survive on any given year. Hence, what we actually observe are the quantities: �8 Environmental Variation + SamEling Truth Year i Variation Observed Year i i Mean 1 S S + el + fl 51 2 S S + e2 + f2 52 3 S S + e3 + f3 53 4 S S + e4 + f4 54 5 S S + es + fs 55 6 S S + e6 + f6 56 7 S S + e7 + f7 57 8 S S + es + fs 5a 9 S S + e9 + f9 59 10 S + elO + flO 510 Mean S S A 5 5 where the ej are as before, but we also have additional sampling variation, or demographic variation, or both, variation from in the fj. The usual approach to estimating sampling variance separately from temporal variance is to take replicate observations within each year so within-cell replicates can be used to estimate sampling variance; whereas the between cell variance is used to estimate the environmental variation. Years are assumed a random effect, and mixed model analysis of variance procedures are used. This approach assumes that each cell has the same sampling variance. Classical analysis of variance methodology assumes the variance within cells is constant across a variety of treatment effects. This assumption is often not true, i.e., the sampling variance of a binomial distribution is a function of the binomial probability. Thus, as the probability changes across cells, so does the variance. Another common violation of this assumption is caused by the variable of interest being distributed lognormally, so that the coefficient of variation is constant across cells and the cell variance is a function of the cell mean. Further, the empirical estimation of the variance from replicate measurements may not be the most efficient procedure. Therefore, the remainder of this section describes methods that can be viewed as extensions of the usual variance component analysis based on replicate meaeuzemenea within cells. An estimator of the temporal variation is provided for the situation where the within cell variance is not estimated by the method of moments estimator based on replicate observations. Assu~e that we can estimate the sampling variance for each year, given a value of 5.~ for the year. For example, an estimate of the sampling variation for a binomial is va r ( S ~.1 5~.) = �9 where ni is the number of animals monitored to see if they survived. Then, can we estimate the variance term due to environmental variation, given that we have estimates of the sampling variance for each year? If we assume all the sampling variances are equal, the estimate of the overall mean is still just the mean of the 10 estimates: 10 with the theoretical variance var LSi i=l = S 10 being = (5) 2 + E[var 0 (SiS) ] 10 i.e., the total variance is the sum of the environmental variance expected sampling variance. This total variance can be estimated plus the as 10 = va r (5) We can estimate variances the expected L(Si-S)2 i=l 10(10 - 1) sampling variance as the mean of the sampling 10 = E(var(SIS)] so that the estimate L var (SiIS) i=l 10 of the environmental variance obtained by solving for d 10 (10 - 1) However, sampling variances are usually not all equal, so we have to weight them to obtain an unbiased estimate of 02• The general theory says to use a weight, Wi w.~ 1 = 2 0 + so that by replacing var (5.~ IS.) with ~ tor of the weighted mean is S with theoretical the estimates) variance var (5.1~ S ~. ) it!:!estimator var (5..z IS.)~ , the estima- = (i.e., sum of the theoretical variances for each of �10 var and the empirical = (S) variance 1 = var estimator 10 L w. (s. - va r i=l = (S) ~ i=l Wj are the true 2 10 Lw. When the S) ~ (but unknown) (10 - 1) ~ weights, we have 10' _fuWi(Si-S)2 1 10 Lw. i=l giving (10 - 1) ~ the following 10 "L.J W i i=l (S i - S) 2 = (10 - 1) 1 Hence, all we have to do is manipulate this equation with a value of 02 to obtain an estimator of 02• To obtain a confidence interval on the estimator of 02, we can substitute the appropriate chi-square values in the above relationship. To find the upper confidence interval value, solve the equation 0;, 10 " _ A A L.J Wi (S i: - S) 2 i=l = (10 - 1) and for the lower confidence interval 10 10 - 1,aL 10 - 1 A2 0L' solve value, the equation _ L.J Wi (Si " 2 X A - S) i=l (10 - 1) A 2 2 X 10 - 1,au 10 - 1 As an example, consider over-winter fawn survival data from mule deer fawns in Piceance Basin in northwest Colorado (Table 1). Survival rates are from the staggered entry Kaplan-Meier estimator (Pollock et ale 1989). The standard deviation of the 14 survival estimates is 0.22. When sampling errors are removed (mean SE = 0.07), the standard deviation of temporal variation is estimated as 0 = 0.21 (95% confidence interval 0.15 to 0.35). This confidence interval represents the uncertainty of the estimate of temporal variation, i.e., the sampling variation of the estimate of temporal variation. �11 Note that the temporal variation estimate is only slightly smaller than the overall standard deviation, as the sampling variation of the estimates is relatively small. Similar results are shown for adult survival and recruitment (Table 2). For mule deer in DAU D-7, which includes Piceance Basin, in northwestern Colorado, the relative variability of recruitment rates, and juvenile and female survival have been measured with the coefficient of variation, defined as the standard deviation of temporal variation (0) divided by the mean of the parameter estimates. From Table 2, we see there is much more variation of over-winter fawn survival than of either recruitment or adult survival. Ev.en though the model is most sensitive to adult survival, this parameter varies little from year to year. Thus, we conclude a precise estimate of SF must be obtained. In contrast, the model is not terribly sensitive to SJ' but this parameter varies considerably from year to year and thus must be estimated each year. Recruitment (R) is not particularly variable, nor is the model particularly sensitive to R. Thus, we don't need to put nearly as much effort (dollars) into estimating recruitment as into estimating survival. PROPOSED MONITORING SCHEME Current CDOW monitoring places all effort into measuring recruitment and occasionally population density, and none into estimating juvenile or female survival rates. Thus, we conclude current monitoring efforts are wasteful because the variable being measured most often is likely the least important to measure annually. As a result, CDOW lacks the necessary information to properly monitor mule deer populations (R. M. Bartmann, Colo. Div. Wildl., unpubl. rep.). In this section, a monitoring scheme that shifts emphasis from monitoring recruitment to monitoring survival is developed. An obvious reason why survival is not monitored is that it is more expensive to measure than recruitment. To rigorously estimate age-specific survival, the fate of a sample of marked animals must be determined. The most direct approach is via radio-tracking, but mark-resight or banding analysis methods are also possible (van Hensbergen and White 1995). However, mark~ resight or mark-recapture (e.g., Burnham et ale 1987, Lebreton et ale 1992» and banding methods (Brownie et ale 1985) are indirect in that additional parameters (resighting probability or band recovery probability) must be estimated. These parameters are nuisance parameters in the sense that they are not the real parameters of interest. However, precision of survival estimates is greatly affected by the precision with which nuisance parameters are estimated. As a result of the increased number of parameters, a larger sample size is required with indirect methods than with radio-tracking methods. For example, White and Bartmann (1983) estimated survival of ~ule deer banded during winter. Even though 1,923 animals were banded over a 5year period, annual survival estimates had coefficients of variation averaging over 32% for juvenile survival and over 19% for female survival. Had radio collars been used, the average coefficients of variation would have been approximately 14% and 5% for juvenile and female annual survival rates, respectively. However, an even bigger problem with using banding or mark-resight methods for monitoring annual survival rates is that estimates are not obtained for the current year, but only for intervals prior to the current year. This phenomenon occurs because the survival parameter and the recovery or resight parameter are confounded for the last survival interval of the data set. This confounding is removed only by adding another year of marking and recovery or resighting data. Thus, these methods are not useful for monitor- �12 ing because estimates of survival will not be available until after the current year's harvest. White et al. (1996) developed a method to estimate adult and juvenile over-winter survival based on age ratios of the population prior to and after winter, and the age ratio of animals dying during the winter. However, the assumptions of this method are unlikely to be met and the potential for biased estimates is considerable. They suggest radio-tracking is generally more appropriate for estimating survival of juvenile and adult female cohorts unless special circumstances exist. Therefore, we conclude radio-tracking is the most economical method to estimate survival even though initial costs are high. Additional benefits of radio-tracking are that cause of death can be determined so insights into the mechanisms affecting population dynamics may be gained. Considering the standard error of the survival estimate as a function of the number of radioed animals (n) for various survival rates, approximately 50 animals must be marked to achieve survival estimates with reasonable precision (Fig. 1). The variance of a survival estimate is a function of both sample size and true survival. Variance is symmetrical about 0.5, with the maximum variance at 0.5 [see White and Garrott (1990) for a review of estimating survival with radioed animals]. This requirement is regardless of the size of the unit or the density or number of deer, because the fraction of the population sampled with radios is too small to affect finite population correction. As shown in Fig. 1, the magnitude of the survival rate does affect the standard error of the estimate. If a sample of 50 radios are needed to estimate over-winter fawn survival adequately, approximate costs can be determined. Assuming $350/fawn for capture (helicopter netgunning) and $200/radio, $27,500 will be needed to initiate monitoring. Additional costs are incurred for monitoring. Assuming $160/hour for tracking via fixed-wing aircraft and 10 flights of 4-hours duration each to determine live/dead status of each animal, an additional $6,400 is required. Thus, approximately $34,000 (exclusive of personnel costs) is required to monitor over-winter fawn survival for a single population or DAU. Obviously, monitoring fawn survival in all 53 DAUs in Colorado is impractical. Instead, we suggest the CDOW annually monitor over-winter fawn survival in a subset of DAUs around the state, labeled core DAUs here. This core subset should represent larger mule deer populations and different habitats. Presumably, estimates from core DAUs would be representative of surrounding, or satellite, DAUs, thus providing estimates of survival in DAUs similar in nature to 1 of the core DAUs. An objective approach to deciding on which DAUs to include in the core subset would be to perform a cluster analysis of DAUs based on available information such as harvest rates, recruitment, habitat, and elevation. However, using estimates from a core DAU to manage a satellite DAUs is risky and this approach should be evaluated. A random sample of satellite DAUs could be selected each year for monitoring along with the core units. Through time, a correlation between the core DAUs will be developed with each satellite DAU. The validity of inferences from core units to any satellite DAU will thus be able to be tested over time. Instead of DAUs, more effective and efficient monitoring might be provided by GMUs. In the past, CDOW biologists have not consistently collected monitoring data for entire DAUs. Instead, some subset of GMUs within DAUs may be sampled. This practice leads to estimates of population parameters-that are not comparable across years because different portions of a DAU are sampled in different years (R. M. Bartmann, Colo. Div. Wildl., unpubl. rep.). The reason for monitoring 1-2 GMUs within a DAU is that the GMUs �13 generally represent distinct mule deer populations or population subsets which is unlike a DAU where a potpourri of populations may be represented. Thus, GMUs may provide more practical and useful data than DAUs. So far, we have focused on over-winter fawn survival. This is because over-winter fawn survival was found highly variable from year to year and necessitated annual monitoring. Adult survival is also critical in that the model is most sensitive to this parameter. However, because of little annual variation in adult survival, this parameter can be estimated with data collected across a series of years. Thus, we propose that core units have an initial sample of adults included in the monitoring program. Female fawns could be fitted with expandable collars so that survivors of their first winter would contribute to estimating adult female survival rates during ensuing years. The annual effort needed to monitor adult survival can be considerably less than for fawns because data can be pooled across years. A sample of at least 20 adults in each core unit, as well as any satellite units, should be maintained. Ideally, recruitment and density should probably be monitored annually in core units and in each randomly selected satellite unit. However, we have not determined the optimal allocation of effort between monitoring over-winter fawn survival, adult survival, recruitment, and population size, or the costs associated with each scenario. Based on the analysis presented in this paper, we assume that an adequate monitoring system requires annual survival information on fawns. Information on recruitment and density will also be required, but how often and what quality of information will be needed in the core units? How much effort (meaning dollars) should be diverted from monitoring survival to monitoring recruitment and/or density? An objective approach to determine monitoring intensities and intervals for fawn and adult survival, recruitment, and density is to develop a simulation model of a mule deer population that includes random temporal variation. The model we have developed allows the user to sample the modeled population to mimic monitoring procedures. The optimal strategy for monitoring requires allocating effort to the monitoring of the various parameters as a function of the cost of collecting data and the temporal and sampling variability of each parameter. Estimates of cost for the various monitoring procedures are used to set the amount of data collected for each parameter monitored. From these data, harvest levels are set to maintain the population at a herd objective ·as is currently done for real populations. Because the true population is known, an evaluation of performance can be made. For a fixed cost, different allocations of monitoring effort can be compared relative to the mean squared error between the true.population and the herd objective population size, Le., minimize ~ ~ (NT rue - NOb'Ject'Lve ) 2 where the summation is over years. Specifically, we would want find the relative amount of effort for sampling age ratios and population density across time versus the relative amount of effort for sampling survival with radio-tracking. Can better management be achieved for the same cost by monitoring survival frequently and recruitment and population size intermittently than by monitoring all 3 at the same level annually? Alternatively, instead of a optimizing results from a computer model, possibly analytical solutions can be developed to allocate effort optimally to monitoring the different population parameters. However, at this time, we do not understand how to derive such analytical relations. Our model to develop an optimal sampling strategy assumes that December herd composition (and thus recruitment) and population density can be sampled simultaneously during the same helicopter survey. Randomly selected quadrats are counted and classified to provide the data. The biological parameters in �14 the model are taken from Table 2, except that fawn survival was increased to 0.6 and recruitment increased to 0.691 so that the population has A>l, requiring harvest to maintain the population at a specified objective. An initial population of 10,000 animals was assumed, with the population objective of 5,300 adult females. Costs associated with monitoring are $600/hr of helicopter time, with 1/4 hr required to count and classify a quadrat, and $600/animal to capture and radio an animal to determine its fate. The hypothetical DAU sampled contains 665 quadrats. The budget for sampling is assumed to be $30,000/yr. Radios on adult females were assumed to last 4 years, thus, most adult radios provide data beyond the year in which the radio was put on the animal. Fawn radios were assumed to drop off after 1 year. Based on these inputs, the optimal sampling strategy to minimize the squared deviation of the true population size from the desired objective is to spend approximately 18 hours of helicopter time each year performing herd composition and population counts, and split the remaining $19,200 evenly between collaring fawns and adult females to measure survival (Fig. 2). Note that changes in the input values will change these results somewhat. However, the optimal allocation of radios between fawns and adults generally is close to 50:50. The final step to evaluate the proposed change in monitoring strategy is to demonstrate that adequate correlations exist in over-winter survival between the core units and the satellite units. These correlations must be estimated from field data, so this evaluation will take many years to complete. Without reasonably good correlations, the lack of monitoring in the satellite units would lead to inadequate information for management. Thus, the proposed monitoring procedure can be considered adaptive management (Walters 1986, Hilborn and Walters 1992) in that the validity of using core units to manage satellite units will be evaluated through time. Likely, the set of core units may change as we gain more information on the similarity of parameters across units. A final caveat must be offered. Monitoring mule deer populations does not provide cause and effect relationships that govern the population dynamics. Monitoring will suggest that the population is changing. However, to understand the mechanisms that are causing the change, designed experiments must be conducted. Thus, a sound monitoring program does not remove the need for a sound research program. In summary, the following steps must be taken to implement the proposed monitoring scheme. 1. Select a set of core units for monitoring that are representative of the mule deer populations in Colorado. 2. Determine the optimal allocation of effort for monitoring of over-winter fawn survival, adult female survival, recruitment, and population size on core units. This allocation of effort will likely change as more data become available, and will vary depending on costs for the particular unit being monitored. 3. Monitor core units annually, always including over-winter fawn and adult female survival as part of this monitoring. 4. Monitor a randomly selected subset of satellite units annually so correlations between satellite units and core units can be developed to evaluate the effectiveness of the monitoring scheme. 5. Evaluate the effectiveness of the monitoring scheme annually to determine if a more efficient scheme can be developed. �15 CONCLUSION Current CDOW monitoring procedures for mule deer populations are inadequate, because the parameters most important in projecting mule deer population status are not measured. A monitoring scheme that includes over-winter fawn survival and adult female survival is proposed. To evaluate this monitoring scheme, and to initiate it objectively, a simulation model of mule deer management has been developed. Results from this model suggest that annual helicopter surveys of herd composition and population density and overwinter fawn and adult female survival are required. Further, a key assumption of the proposed monitoring scheme is that correlations exist in basic population parameters between similar units. This assumption can only be tested with field data, not through simulation. 1. Preliminary analysis of elk sightability models was performed in cooperation with David Freddy. Model selection with AIC was performed for a list of candidate models. 2. Assistance was provided in the design and analysis of a study to estimate black bear abundance in western Colorado. Program NOREMARK has been updated to perform the necessary analyses. 3. Assistance was provided in the design and analysis of a study to estimate kit fox abundance in western Colorado. Alternative sampling strategies were discussed. 4. Assistance was provided in the design and analysis of a study to estimate swift fox abundance in eastern Colorado. Alternative sampling strategies were discussed. 5. A paper summarizing the effect of experimental harvest on overwinter survival of mule deer fawns has been accepted for publication by the Journal of Wildlife Management. LITERATURE CITED Bartmann, R. M., L. H. Carpenter, R. A. Garrott, and D. o. Bowden. 1986. Accuracy of helicopter counts of mule deer in pinyon-juniper woodland. Wildl. Soc. Bull. 14:356-363. Brownie, C., D. R. Anderson, K. P. Burnham, and D. S. Robson. 1985. statistical inference from band recovery data -- A handbook. Second ed. U. S. Fish and Wildl. Servo Resour. Publ. 156. 305 pp. Burnham, K.P., D. R. Anderson, G. C. White, C. Brownie, and K. H. Pollock. 1987. Design and analysis methods for fish survival experiments based on release-recapture. Am. Fisheries Soc. Monogr. 5. 437 pp. Caswell, H. 1989. Matrix population models. Sinauer Assoc. Inc., Sunderland, Mass. 328 pp. Hilborn, R. and C. J. Walters. 1992. Quantitative fisheries stock assessment -- choice, dynamics and uncertainty. Chapman and Hall, New York, N. Y. 570 pp. Innis, G. S. 1979. A spiral approach to ecosystem simulation, I. Pages 211386 in Systems Analysis of Ecosystems. G. S. Innis and R. B. O'Neil, eds. International Co-operative Publishing House, Fairland, MA. Kufeld, R. C., J. H. Olterman, and D. C. Bowden. 1980. A helicopter quadrat census for mule deer on Uncompahgre Plateau, Colorado. J. Wildl. Manage. 44:632-639. �16 Lebreton, J.-D., K. P. Burnham, J. Clobert, and D. R. Anderson. 1992. Modeling survival and testing biological hypotheses using marked animals: a unified approach with case studies. Ecol. Monogr. 62:67-118. Leslie, P. H. 1945. On the use of matrices in certain population mathematics. Biometrika 33:183-212. Pollock, K. H., S. R. Winterstein, C. M. Bunck, and P. D. Curtis. 1989. Survival analysis in telemetry studies: the staggered entry design. J. Wildl. Manage. 53:7-15. Steinert, S. F., H. D. Riffel, and G. C. White. 1994. Comparison of big game harvest estimates from check station and telephone surveys. J. Wildl. Manage. 57:336-341. Van Hensbergen, H. J., and G. C. White. 1995. Review of methods for monitoring vertebrate population parameters. Pages 489-508 in J. A. Bissonette and P. R. Krausman, eds. Integrating people and wildlife for a sustainable future. Proc. first International Wildlife Management Congress. The Wildl. Soc., Bethesda, Md. Walters, C. J. 1986. Adaptive management of renewable resources. Macmillian, New York, N.Y. 374 pp. White, G. C. and R. M. Bartmann. 1983. Estimation of survival rates from band recoveries of mule deer in Colorado. J. Wildl. Manage. 47:506-511. White, G. C. and R. A. Garrott. 1990. Analysis of Wildlife Radio-tracking Data. Academic Press, New York, NY. 383 pp. White, G. C. 1993. Precision of harvest estimates obtained from incomplete responses. J. Wildl. Manage. 57:129-134. White, G. C., A. F. Reeve, F. G. Lindzey, and K. P. Burnham. 1996. Estimation of mule deer winter mortality from age ratios. J. Wildl. Manage. 60:37-44. Table 1. survival, Year 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 Mean SD fawn Estimates of recruitment (fawns/100 adult females), over-winter and annual adult female survival in DAU D-7 in northwestern Colorado. Adult Female Survival Recruitment Fawn Survival Estimate Estimate SE SE Estimate SE 76.4 72.6 72.9 78.6 78.1 75.5 62.9 58.7 73.4 61.5 2.73 2.30 2.17 2.78 2.04 1.96 1.53 1. 71 1.86 1.53 63.0 63.9 62.1 55.6 49.7 47.1 54.9 47.6 64.14 10.55 1.20 0.97 1.33 1.12 1.38 1.40 1.38 3.04 1.80 0.32 0.07 0.20 0.54 0.43 0.24 0.27 0.78 0.32 0.46 0.11 0.55 0.70 0.62 0.40 0.22 0.09 0.05 0.08 0.07 0.06 0.08 0.08 0.08 0.09 0.11 0.04 0.06 0.05 0.06 0.07 0.80 0.78 1.00 0.92 0.72 0.82 0.86 0.83 0.97 0.72 0.72 0.90 0.85 0.97 0.85 0.10 0.13 0.11 0.00 0.05 0.09 0.12 0.13 0.08 0.03 0.07 0.07 0.04 0.05 0.03 0.07 �17 Table 2. females), northwest Parameter Estimates of temporal variation in recruitment (fawns/100 adult over-winter fawn survival, and adult female survival in DAU D-7 in Colorado. 95% Confidence Coefficient Mean Temporal Interval of Variation Variation Recruitment 64.1 Over-winter Adult Fawn Survival Female Survival 10.3 0.40 0.21 0.85 0.078 7.6 - 15.7 16% 0.15 - 0.35 52% 0 - 0.14 9% 0.16 -------0.1 - - 0.2 - -- - 0.3 ------0.4 - 0.14 L.. 0 L.. L.. w "E CO 0.5 0.12 0.1 0.08 'U c: CO 0.06 .•.... en , , .... 0.04 .... _'- ...• _--- - __ ------ 0.02 ---------- ---------- 0 0 100 200 300 400 Sample Size (n) Figure 1. Standard error of the estimate of survival (S) for a radiotracking study with n radio-marked animals. The 5 lines portray the SE forS = 0.5, 0.4, 0.3, 0.2, and 0.1 from highest to lowest. �18 -w ..c L- a> a. o o .CD :::c 1·~.~~~~~~~~~~~~~~~~~~ 0.00 0.25 0.50 0.75 1.00 Proportion of Radios on Fawns MSE 420000 500000 580000 460000 540000 620000 2. contour plot of the mean squared difference of true population size and the desired population size as a function of allocation of effort between helicopter surveys, and fawn and adult female radio collars to monitor survival. Figure �19 colorado Division Wildlife Research July 1997 of Wildlife Report JOB PROGRESS State of Project REPORT Colorado No. W-153-R-10 Mammals Research Work Plan No. Multispecies Job No. Mammals Library Period Covered: Author: July Jacqueline Personnel: Inyestigations Publication, Services Editing, and 1, 1996 - June 30, 1997 A. Boss Jacqueline A. Boss and Nancy W. McEwen ABSTRACT During the Segment * the following were accomplished: 36 pUblications were purchased at the request of Mammals Researcher personnel and placed into the Colorado Division Wildlife Research Center Library collection. of * 12 free reports and short publications from state or federal agencies or from private sources were located, ordered, and obtained for use by mammals Research personnel. * 54 theses or books were obtained on Interlibrary for use by Mammals Research personnel. * 924 individual use by Mammals * 10 manuscripts by Mammals accepted for publication. * 0 manuscripts were prepared personnel for peer review. articles Research were located personnel. Research Loan or as gifts and delivered personnel and submitted on request were published by Mammals or Research for ��21 MAMMALS PUBLICATION, EDITING Jacqueline AND LIBRARY SERVICES A. Boss P. N. OBJECTIVE To provide a centralized support program for manuscript editing services to facilitate publishing results of research conducted Federal Aid Project W-153-R. SEGMENT and library by staff of OBJECTIVE To provide a centralized support program for Mammals Research editing, library, and publishing services so that Mammals Research personnel can be most efficient in publishing results of their research. SUMMARY or SERVICES Publications Purchased with Mammals Research Funds and Placed in the Research Center Library Andrew, R. and R. Righter. 1992. Colorado birds : a reference to their distribution and habitat. Denver, CO : Denver Museum of Natural History. 442pp. Bass, R. 1995. The lost grizzlies: a search for survivors of Colorado. New York : Houghton Mifflin Co. 239pp. in the wilderness Boyce, W. M., P. R. Crosbie, and R. M. Lee, eds. [1993]. Desert Bighorn Council : 1993 transactions : a compilation of papers presented and submitted at the 37th annual meeting, 7-8 April 1993, Mesquite, Nevada. Las Vegas, NV : Desert Bighorn Council. 60pp. Boyce, W. M., P. R. Crosbie, and R. M. Lee, eds. [1994]. Desert Bighorn Council : 1994 transactions : a compilation of papers presented and submitted at the 38th annual meeting, 6-8 April 1994, Moab, Utah. 5.1. : [Desert Bighorn Council]. 36pp. Boyd, R. J., J. A Armentrout, and W. R. Brigham, eds. [1995]. Desert Bighorn Council : 1995 transactions : a compilation of papers presented and submitted at the 39th annual meeting, 5-7 April 1995, Alpine, Texas. 5.1. : [Desert Bighorn Council]. 108pp. e . Chase, A. 1986. Playing God in Yellowstone: the destruction first national park. New York : Atlantic Monthly Press. Colorado Counties, Denver, CO Inc. 1995. Colorado's public lands: Colorado Counties, Inc. 102 leaves. a county Crawford, W. and M. Gorman. 1995. Future libraries : dreams reality. Chicago: American Library Assoc. 198pp. Cvancara, V., ed. 5.1. : s.n. 1996. Current references var. pagination. of America's 446pp. in fish research profile. madness vol. & 21, 1996. �22 Doughty, R. W. 1989. of Texas Press. Return 182pp. of the whooping crane. Austin, TX University Douglas, C. A., T. D. Bunch, P. R. Krausman, D. M. Leslie, Jr., and J. J. Spillett, eds. [1981]. Desert Bighorn Council: 1981 transactions: a compilation of papers presented at the 25th annual meeting, April 8-10, 1981, Kerrville, Texas. Las Vegas, NV : Univer. of Nevada. 70pp. Drummond, A. 1995. Enos Mills: Press of Colorado. 433pp. citizen of nature. Niwot, CO : University Ehrlich, P. R., D. S. Dobkin, and D~ Wheye. 1988. The birder's handbook: a field guide to the natural history of North American birds : including all species that regularly breed north of Mexico. New York : Simon & Schuster, Inc. 785pp. Eversole, A. G., ed. [1996]. Proceedings of the forty-eighth annual conference : Southeastern Association of Fish and Wildlife Agencies : October 23-26 1994 : Biloxi, Mississippi. S.l. : Southeast. Assoc. of Fish and Wildl. Agencies. 669pp. Eversole, A. G., ed. [1997]. Proceedings of the forty-ninth annual conference : Southeastern Association of Fish and Wildlife Agencies : September 23- . 27, 1995 : Nashville, Tennessee. S.l. : Southeast. Assoc. of Fish and Wildl. Agencies. 736pp. Geist, V. 1991. Elk country. Geist, V. 1990. Mule Minocqua, deer country. WI Minocqua, Gittleman, J. L. 1996. Carnivore behavior, Ithaca, NY : Cornell University Press. Gunnel, G. K. 1995. CO : Westcliffe : NorthWord WI : Northword ecology, 644pp. Guide to Colorado wildflowers, Publishers. 2 vols. l75pp. Press. and evolution, vols~ Commercialization and wildlife Hawley, A. W. L. 1993. Malabar, FL : Krieger. 124pp. with the devil. Hoyt, Press. 1 & 2. management 176pp. vol. 2. Englewood, dancing J. A. 1994. Animals in peril: how "sustainable use" is wiping out the world's wildlife. Garden City Park, NJ : Avery Publishing Group. 257pp. Krausman, P. R., W. M. Boyce, M. C. Wallace, and R. M. Lee, eds. [1992]. Desert Bighorn Council : 1992 transactions : a compilation of papers presented and submitted at the 36th annual meeting, 8-11 April 1992, Bullhead City, Arizona. Las Vegas, NV : Desert Bighorn Council. 87pp. Lankester, M. W. and H. R. Timmermann, eds. [1997]. Alces: devoted to the biology and management of moose: volume Thunder Bay, Ontario: Lakehead University. 197pp. a journal 32, 1996. Lankester, M. W. and H. R. Timmermann, eds. [1997]. Alces devoted to the biology and management of moose: volume Thunder Bay, Ontario: Lakehead University. 217pp. a journal 33, 1997. �23 Lynch, W •. 1993. Bears: The Mountaineers. monarchs 242pp. of the northern wilderness. Seattle, WA Miller, B., R. P. Reading, and S. Forrest. 1996. Prairie night: blackfooted ferrets and the recovery of endangered species. Washington, Smithsonian Institution Press. 254pp. Murray, J. A., ed. 1992. grizzly. Anchorage, The great bear: contemporary AK : Alaska Northwest Books. writings 245pp. on the Nesbitt, WID. H., ed. 1996. Proceedings of the 81st Convention; 1991 : International Association of Fish and Wildlife Agencies : September 11, 1991 : Hot Springs, Arkansas. Washington, D.C. : International Association of Fish & Wildlife Agencies. 422pp. Perrin, P. G. 1972. William Morrow Reference & Co., Inc. handbook 297pp. of grammar Petersen, D., ed. 1996. A hunter's heart New York : Henry Holt & Co. 331pp. Rising, J. D. sparrows 365pp. honest & usage. essays 7- New York on blood sport. 1996. A guide to the identification and natural history of the of the United States and Canada. New York: Academic Press. Schullery, P. 1992. The bears of Yellowstone. Publishing Company. 318pp. Worland, Wassink, J. L. 1993. Mammals of the central Rockies. Mountain Press Publishing Company. 161pp. Webb, DC WY High Plains Missoula, MT J. W., ed. 1959. Proceedings of the eleventh annual conference: Southeastern Association of Game and Fish Commissioners : October 1958 : Mobile, Alabama. Columbia, SC : Southeast. Assoc. of Game Fish Commissioners. 393pp. 20-23, and Webb, J. W., ed. [1959]. Proceedings of the thirteenth annual conference: Southeastern Association of Game and Fish Commissioners: October 25-27, 1959 : Baltimore, Maryland. Columbia, SC : Southeast. Assoc. of Game and Fish Commissioners. 407pp. Webb, J. W., ed. 1960. Proceedings of the fourteenth annual conference: Southeastern Association of Game and Fish Commissioners : October 23-26, 1960 : Biloxi, Mississippi. Columbia, SC : Southeast. Assoc. of Game and Fish Commissioners. 284pp. ~heses and Books Loan or as Gifts Alderton, D. 192pp. Obtained on Interlibrary for Use by Researchers 1993. Wild cats of the world. New York Facts On File, Inc. Alvarez, K. 1993. Twilight of the panther: biology, bureaucracy and failure in an endangered species program. Sarasota, FL : Myakka River Publishing. 501pp. �24 Baker, J. M. 1992. Habitat use and spatial organization of pine marten of southern Vancouver Island, British Columbia. M.S. Thesis, Simon Fraser University, Burnaby, BC. 119pp. Baker, R. D., R. S. Maxwell, V. H. Treat, & H. C. Dethloff. 1988. Timeless heritage : a history of the Forest Service in the southwest. [Washington, D.C.] : U.S. Forest Service. U.S.D.A. Forest Service publication; FS-409. 208pp. Botkin, D. B. 1995. Our natural history : the lessons New York: G. P. Putnam's Sons. 300pp. Bowles, M. L. and C. J. Christopher. : conceptual issues, planning, University Press. 394pp. of Lewis and Clark. 1994. Restoration of endangered species and implementation. New York : Cambridge Callicot, J. B., ed. 1987. Companion : University of Wisconsin Press. to a Sand County 308pp. Cameron, J. 1929, The Bureau of Biological Survey and organization. Baltimore, MD : John Hopkins graph of the United States government; No. 54. Almanac. Madison, WI its history, activities Press. Service mono339pp. Carbyn, L. N., S. H. Fritts, and D. R. Seip, eds. 1995. Ecology and conservation of wolves in a changing world. Edmonton, Alberta : Canadian Circumpolar Institute. Occasional publication series (Canadian Circumpolar Institute); no 35. 620pp. Caughley, G. and A. Gunn. 1996. Conservation biology Cambridge, MS : Blackwell Science. 459pp in theory Copeland, J. P. 1996. The biology of the wolverine in central Thesis, University of Idaho, Moscow, ID. 138pp. Crisler, L. 1958. Arctic wild. New York: Harper and practice. Idaho. & Brothers, Pub. M.S. 301pp. Doan-Crider, D. L. 1995. Population characteristics and home range dynamics of black bears in northern Coahuila, Mexico. M.S. Thesis, Texas A & M University - Kingsville, TX. 117pp. Driver, B. L., D. Dustin, T. Baltic, G. Elsner, and G. Peterson, eds. Nature and the human spirit : toward an expanded land management State College, PA: Venture Publishing, Inc. 467pp. 1996. ethic. Elias, S. A. 1996. The ice-age history of national parks in the Rocky Mountains. Washington, D.C. : Smithsonian Institution Press. 170pp. Evans, H. E. Chicago 1984. Press. Life on a little-known 318pp. planet. Fleharty, E. D. 1995. Wild animals and settlers Norman, OK : University of Oklahoma Press. Greenwood, M. 1932. Epidemiology: MD : The Johns Hopkins Press. historical 80pp. Chicago : University on the Great 316pp. of Plains. and experimental. Baltimore, �25 Greenwood, M. 1935. Epidemics and crowd-diseases : an introduction to the study of epidemiology. London: Williams & Norgate, Ltd. 409pp. Greenwood, M., A. Bradford Hill, W. W. C. Topley, and J. Wilson. 1936. Experimental epidemiology. Medical Research Council (Great Britain) Special report series; no. 209. London: H. M. Stationery Off. 204pp. Grinnell, G. B. New York 1892. Blackfoot lodge tales: the story of a prairie people. Charles Scribner's Sons. 310pp. Harwood, R. F. 1979. Entomology in human.and animal health. Macmillan Publishing Co. p. New York Humphrey, S. R. 1992. Rare and endangered biota of Florida Gainesville, FL : University Press of Florida. 392pp. mammals vol. I. IUCN : the World Conservation Union. 1995. Publications Services Unit. 33pp. Wild cats. Cambridge, UK IUCN Idaho Dept. of Fish and Game. n.d. Mountain lion : species management plan 1991 - 1995. [Boise, ID : Idaho Dept. of Fish & Game]. no pagination. Kitchener, A. 1991. The natural history of the wild cats. Comstock Publishing Assoc. 280pp. Ithaca, NY : Kucera, Thomas E. 1988. Ecology and population dynamics of mule deer in the eastern Sierra Nevada, California. Ph.D. Dissertation. University of California - Berkeley. 207pp. Lawton, J. H. 233pp. 1995. Extinction rates. New York Oxford University Press. Livingston, J. A. 1981. The fallacy of wildlife conservation. : MCClelland & Stewart, Inc. 117pp. Livingston, J. A. 1994. tion. Boulder, CO Luce, A. A. 1993. 191pp. Toronto, Onto Rogue primate : an exploration of human domesticaRoberts Rinehart Pub. 229pp. Fishing and thinking. Camden, ME : Ragged Mountain Press. McCullough, D. R. and R. H. Barrett, eds. 1992. Wildlife 2001 New York : Elsevier Applied Science. 1,163pp. populations. McIvor, D. E., J. A. Bissonette, and G. S. Drew. 1994. A critical review of the status of the Yuma mountain lion, Felis concolor browni. Logan, UT : Utah Coop. Fish and Wildl. Res. Unit (U.S. National BioI. Surv.) Rep. 94-1. 155pp. Mares, M. A. and H. H. Genoways, eds. 1982. Mammalian biology in South America: a symposium held at the pymatuning Laboratory of Ecology, May 10 - 14, 1981. Linesville, PA : University of Pittsburgh. Special publication series (pymatuning Laboratory of Ecology); vol. 6. 539pp. Mares, M. A. and D. J. Schmidly, eds. 1991. Latin American mamma logy : history, biodiversity, and conservation. Norman, OK : University of Oklahoma Press. 468pp. �26 Miller, B., R. P. Reading, & S. Forrest. 1996. Prairie night: black-footed ferrets and the recovery of endangered species. Washington, D.C. : Smithsonian Institution Press. 254pp. Nilsson, G. 1983. The endangered species handbook. Animal Welfare Institute. 245pp. Washington, D.C. The Powell, R. A., J. W. Zimmerman, and D. E. Seaman. 1997. Ecology and behaviour of North American black bears : home ranges, habitat and social organization. New York: Chapman & Hall. 203pp. Preece, R. and L. Chamberlain. 1993. Animal welfare & human values. Waterloo, Onto : Wilfrid Laurier University Press. 334pp. Prescott-Allen, R. and C. Prescott-Allen. 1982. What's wildlife worth? Washington, D. C. : Inter'l. Institute for Environment & Development. 92pp. Price, M. F. and D. I. Heywood. 1994. Mountain .environments and geographic information systems. Bristol, PA : Taylor & Francis, Inc. 309pp. Remmert, H. ed. 1994. Minimum animal populations. New York : SpringerVerlag. Ecological studies; vol. 106. 156pp. Ricklefs, R. E. and D. Schluter, eds. 1993. Species diversity in ecological communities : historical and geographical perspectives. Chicago: University of Chicago Press. 414pp. Savage, C. 1993. Wild cats: lynx: Sierra Club Books. 136pp. bobcats mountain lions. San Francisco Schwerdtfeger, W. ed. 1976. Climates of Central and South America. New York : Elsevier Scientific Publishing Company. World Survey of Climatology; Vol. 12. 532pp. Taylor, P. W. 1986. Respect for nature: a theory of environmental ethics. Princeton, NJ : Princeton University Press. 329pp. Tucker, E. A. & G. Fitzpatrick. 1972. Men who matched the mountains : the Forest Service in the Southwest. [Albuquerque, NM] : U.S.D.A. Forest Service. Southwestern Region. 293pp. Turbak, G. 1986. 77pp. America's great cats. Flagstaff, AZ Northland Press. Turner, D. C. and P. Bateson. 1988. The domestic cat : the biology of its behaviour. New York: Cambridge University Press. 222pp. Van Sickle, W. D. 1990. Methods for estimating cougar numbers in southern Utah. M.S. Thesis, University of Wyoming, Laramie, WY. 73pp. Vander Wall, S. B. 1990. Food hoarding in animals. Chicago Press. 445pp. West, L. S. 1951. and control. Chicago University of The housefly : its natural history, medical importance, New York : Comstock Publishing Company. 541pp. �27 White, E. and J. Losco, eds. 1986. Biology and bureaucracy: public administration and public policy from the perspective of evolutionary, genetic and neurobiological theory. Lanham, MD : University Press of America. 633pp. Young, S. P. & E. A. Goldman. 1944. The wolves of North America : part I. their history, life habits, economic status, and control : part II. Senior biologists, section of Biological Surveys, Division of Wildlife Research, Fish and Wildlife Service, Department of the Interior. Washington, D.C. : American Wildlife Institute. 636pp. Reference Document Location and Deliyery The Research Center Library staff also located and delivered individual articles or free documents on request for Mammals personnel during this segment. Manuscripts Job Progress Published Reports; FY approximately Researcher 942 1996-97 Federal Aid. All studies. Freddy, D. J. and G. C. White. 1997. Implications Colorado. Proc. Western States and Provinces Workshop. Tucson, AZ. (abstract in press). of elk survival rates Joint Deer and Elk in Fulton, D. C., M. J. Manfredo, and J. Lipscomb. 1996. Wildlife value orientations : a conceptual and measurement approach. Human Dimen. Wildl. 1(2):24-47. Kufeld, R. C. and D. C. Bowden. 1996. Survival shirasi) in Colorado. Alces 32:9-13. rates of Shiras Jessup, D. A., T. E. Thorne, M. W. Miller, and D. L. Hunter. and translocation of wild ungulates in North America. Selvaggina XXIV:355-365. moose 1996. Suppl. (Alces Capture Ric. Biol. Shenk, T. M. and A. B. Franklin. 1996. Accuracy assessment of a Landsat ™ data vegetation map. page 152 in The Wildlife Society : Third Annual Conference : Cincinnati 96 : program and abstracts. Bethesda, MD : The Wildlife Society. (abstract). Shenk, T. M., A. B. Franklin, and K. R. Wilson. 1996. A model to estimate the annual rate of golden eagle population change at the Altamont Pass Wind Resource Area. pages 47~56 in the Proceedings of National AvianWind Power Planning Meeting II. King City, Ontario: LGL Ltd. Shenk, T. M., N. T. Hobbs, and D. M. Theobald. 1997. Estimating accuracy of Landsat TM vegetation classifications using fuzzy logic: a case study. in The Wildlife Society : Fourth Annual Conference : Snowmass, co (abstract in press). Shenk, T. M., N. T. Hobbs, and D. M. Theobald. 1997. Use of fuzzy set theory to deal with error in spatial databases : a case study. Ecological Society of America: 1997 Annual Meeting (abstract in press). �28 Shenk, T. M. and G. C. White. 1996. Detecting temporal trends in demographic parameters Ecology. (in review). Shenk, T. M., G. C. White and K. P. Burnham. 1996. Effects variance on detecting density dependence from temporal populations. Ecology. (in review). Manuscripts in Review density dependence from using logistic regression. of sampling trends in natural FY 1996-97 At the end of FY 1994-95 all manuscripts were either 'in preparation' press.' None were in the stage of 'in review' by periodicals. Prepared by Jacqueline Librarian A. Boss or 'in �29 Colorado Division Wildlife Research July 1997 of Wildlife Report JOB PROGRESS state of Project REPORT Colorado No. ~W~-~l~5~3~-~R~-~l~0~ _ Mammals Research Work Plan No. Hultispecies Job No. Mammals 1 Research Administration Period Author: Covered: July 1, 1996-June Inyestigations 30, 1997 R. Bruce Gill Personnel: R. Bruce Gill and Diane K. Haerter ASTRACT Plans were developed, approved and budgeted for 6 projects relating to deer, elk, or moose research. All projects were successfully completed, but encumbered more resources than were allocated because budget allocations were finalized too late to adjust project spending. Work of all Mammals 1 Research Staff members was evaluated and plans for fiscal year 1997-98 were prepared and submitted for approval and bugeting. Support staff continued to provide library and publications services as requested by Mammals 1 Research staff members. ��31 Mammals Research 1 Administration R. Bruce Gill P.N. Objective 1. Supervise and administer research Mammals 1 Research Section. Segment 1. Supervise and administer 1 Research Section. on deer, elk, and moose with the elk, and moose in the Mammals Objectives research on deer, RESULTS Research and technical activities of six full-time employees were supervised during the segment. The segment objective was only partially fulfilled for reasons described below. • An audit of the Division of Wildlife's performance under the Application for Federal Assistance revealed that previous Job Progress and Job Final Reports had failed to address all of the objectives in the project agreements satisfactorily. Consequently, the Division's application for fiscal year 1997-98 remained unfunded until reports were submitted addressing the unreported objectives. Amended Job Progress and Job Progress reports were submitted and accepted by the U.S. Fish and wildlife Service. • Annual work plans were prepared for all Mammals 1 staff members. Work was satisfactorily completed as outlined in the plans. However, allocated budgets were over-encumbered because final budget allocations and work authorizations from the Division's Planning, Budgeting, and Evaluation Unit were not finalized and formalized until late April 1997. The final allocations were reduced by 10% from the previous year's allocation, but notification of the reduction occurred too late to make adjustments in project spending. • Library services provided to the Mammals 1 and Mammals 2 Research staffs included the purchase of 36 requested publications; 12 free reports or short publications were obtained from other state or federal agencies; 54 theses or books were obtained on Interlibrary Loan or other sources; 924 technical or scientific articles were located and delivered to research staff upon request. • Publication services were provided to Mammals 1 and Mammals 2 Research staff and resulted in the publication or acceptance for publication of 10 scientific, technical, or professional publciations. In addition, over 500 35mm slides were produced upon request from research staff for use in professional, technical, and/or scientific talks. �32 • Work performance evaluations were completed for all Mammals Research staff members and included evaluations of federally sponsored project activities. • Project staff provided technical input into several continuing and ongoing policy issues, including management and mitigation of crop and property damage from big game mammals; deer, elk, and moose hunting regulations; human dimensions of big game hunting satisfication and opportunity; assessment of potential factors contributing to a decline in Colorado deer numbers. Prepared by: R. Bruce Gill Mammals Program Leader �33 Colorado Division of Wildlife Wildlife Research Report July 1997 JOB FINAL REPORT state of Colorado Project No. W-1S3-R-10 Mammals Research Work Plan No. Deer Inyestigations Job No. Compensatory Effects of Harvest in a Mule Deer Population Period Covered: Author: July 1, 1996 - June 30, 1997. R. M. Bartmann, G. C. White. Personnel: R. M. Bartmann and G. C. White ABSTRACT Field data collection was terminated as of 30 June 1996 with data analysis continuing into 1996-97 (Segment 10). A publication, "Effect of Density Reduction on Overwinter Survival of Free-Ranging Mule Deer Fawns", was submitted to "The Journal of Wildlife Management" and accepted for publication, probably in the April 1998 issue. ��35 COMPENSATORY EFFECTS Richard OF HARVEST M. Bartmann P. IN A MULE DEER POPULATION and Gary C. White N. OBJECTIVES 1. Increase the winter survival rate of mule deer fawns by lowering deer density to reduce competition for forage during winter. 2. Increase the harvest rate of deer through increased productivity of adult does and decreased natural mortality of fawns resulting from closer alignment of population size with carrying capacity. SEGMENT annual survival rates total OBJECTIVES 1. Estimate units. of adult females on control and treatment 2. Analyze data, prepare manuscript for publication, and submit "The Journal of Wildlife Management" for publication. manuscript to Adult female mule deer were not monitored for survival after 30 June 1996. Annual survival rates for adult females have been consistently high so continued expenditure of time and money to monitor survival until 1 December 1996 was deemed not necessary for successful conclusion of the study. Rather, data analyses were conducted to enable preparation of a final report in the form of a publication. A manuscript, "Effect of Density Reduction on OVerwinter Survival of FreeRanging Mule Deer Fawns", was submitted to "The Journal of Wildlife Management" and accepted for publication, probably in the April 1998 issue. The abstract of that publication follows. Abstract: Understanding how Overwinter survival of fawns changes as a function of density is a critical relation for managing mule deer (Odocoileus hemionus) populations. We examined change in Overwinter survival of fawns in response to intentional density reduction by radiotracking fawns on control and treatment areas. Deer density on the treatment area was lowered -75%, mostly from antlerless harvests in December. There were 7 years of pretreatment data, 4. years of harvest, and 3 more years of posttreatment monitoring. Fawn survival rate on the treatment area the 3 winters after density was lowered averaged 0.16 higher (P = 0.001) than the control area. After density was lowered, body mass of fawns on the treatment area in November-December averaged 0.8 kg more than the control (P < 0.001). A parallel decline in deer density began on the control area 2 years after initiation of the intentional density reduction on the treatment area. This decline was unexpected and the cause unknown leaving unanswered what differences in fawn survival and body size between the 2 areas would have been had the control population remained high. Prepared by _ Richard M. Bartmann, LSSR III Dr. Gary C. White, Professor ��37 Colorado Division Wildlife Research July 1997 of Wildlife Report JOB PROGRESS state of Project REPORT Colorado No. W-153-R-I0 Mammals Research Work Plan No. Multispecies Job No. Monitoring and Managing Chronic Wasting Disease in Deer and Elk Period Covered: July Inyestigations 1, 1996 - June 30, 1997 Authors: M. W. Miller Personnel: A. L. Case, T. R. Spraker, and E. Zimmerman S. Tracy, M. A. Wild, E. S. Williams, ABSTRACT Deer and elk from throughout Colorado were examined for occurrence of chronic wasting disease using a combination of targeted surveys and harvest or roadkill surveys. We continued to develop and modify a statewide targeted surveillance program for acquiring, examining, and reporting on CWO suspects submitted from Colorado. Between June 1996 and May 1997, 22 chronic wasting disease (CWO) cases (16 deer, 6 elk) were diagnosed among 52 "suspects" (34 deer, 18 elk) submitted from northeastern Colorado; CWO was not diagnosed in any of 12 additional "suspects" (6 deer, 6 elk) submitted from elsewhere in Colorado. Four CWO cases originated in game management units (GMUs) where the disease had not been detected previously; these included an elk case in GMU 7 and deer cases in GMUs 29, 93, and 95. Harvest and road-kill surveys were used to estimate CWO prevalence in enzootic management units. About 6.3% of deer and 1.4% of elk harvested in Larimer County data analysis units (DAUs) (D4/E4 or DI0/E9) tested positive for CWO via immunostaining; CWO was not detected in any of the harvested deer or elk submitted from outside Larimer County. Requiring hunters to participate in harvest surveys appeared to effectively increase the number of samples submitted for CWO examination; submission rates were 3 to 5 times higher in units and seasons where regulations required submissions of heads from harvested deer or elk. About 11% of road-killed deer from D4 tested positive for CWO; none of the road-killed deer or elk from other DAUs were positive. Data from both targeted surveillance and surveys indicate that Larimer County remains the most significant focus of CWO in Colorado, although some natural spread may be occurring both southward and eastward. Targeted surveillance of clinical suspects appears to be the most sensitive approach for initially detecting CWO in deer and elk populations throughout Colorado, and should be �continued statewide. Once detected, combinations of harvest and road-kill surveys should be employed to estimate prevalence, monitor prevalence trends, and compare prevalence among DAUs. [Note: Some of the research activities described under this Work Plan/Job are a continuation of work initiated under Work Plan la, Job 6 (Monitoring and Managing Wildlife Health in Colorado).] �39 MONITORING AND MANAGING CHRONIC WASTING DISEASE IN COLORADo M.W. Miller P. N. OBJECTIVES (1) Design, conduct, and report results of surveys to estimate and detect changes in prevalence of chronic wasting disease (CWO) in wild deer and elk populations. (2) Develop and refine simulation models of CWO dynamics to predict potential impacts of CWO on deer and elk populations. (3) Design, captive (4) Design, conduct, and report on experiments transmission of CWO in deer and elk. (5) Develop conduct, and report results deer and elk herds infected adaptive resource management AGREEMENT of epizootiological with CWO. related studies to diagnosis of and plan. for CWO in deer and elk. OBJECTIVES (1) Design, changes conduct, and report results of surveys to estimate and detect in prevalence of CWO in wild deer and elk populations. (2) Design, captive conduct, and report results deer and elk herds infected (3) Design, conduct, and report on experiments transmission of CWO in deer and elk. of epizootiological with CWO. related studies to diagnosis of and Chronic wasting disease (CWO) affects native deer and elk, causing behavioral changes and progressive loss of body condition that invariably lead to the death of affected animals (Williams and Young 1992). Neither the causative agent nor its mode of transmission have been identified. There are no tests currently available for diagnosing CWO in live animals, and pos~mor~em tests require microscopic examination of brain tissue. There are no known treatments for CWO. Previous attempts to eradicate CWO from research facilities failed on at least 2 occasions (Williams and Young 1992; Miller et al., 1997). Although similar in some respects to other transmissible spongiform encephalopathies that affect domestic sheep (scrapie) and cattle (bovine spongiform encephalopathy; "mad cow disease"), there is no evidence suggesting CWO can be naturally transmitted to domestic livestock, or that scrapie or BSE can be transmitted to native cervids. Moreover, there is no evidence suggesting that CWO presents a threat to human health. "Chronic wasting disease" was first recognized by biologists in the 1960's as a disease syndrome of captive deer held in wildlife research facilities in Ft. Collins, CO, and was subsequently recognized in captive deer, and later in captive elk, in wildlife research facilities near Ft. Collins, Kremmling, and Meeker, CO and Wheatland, WY (Williams and Young 1980, 1982). Since 1981, CWO has also been diagnosed in free-ranging mule deer, white-tailed deer, and elk from northcentral Colorado; most of these diagnoses have been made since 1990 �40 (Spraker et al. 1997). Although CWO was first diagnosed in captive cervids, the original source of CWO is unknown; whether CWO in captive cervids really preceded CWO in wild cervids, or vice versa, is equally uncertain (Spraker et a1. 1997). At present, the known world-wide distribution of CWO in wild cervids appears to be limited to northeastern Colorado and southeastern Wyoming. In Colorado, free-ranging CWO cases have primarily originated from along the Front Range near Estes Park and west of Ft. Collins-Loveland. Cases were diagnosed in 6 different game management units (GMUs)(191, 9, 19, 20, 94, and 96) prior to 1996, but two GMUs (19, 20) yielded about 85% of the documented cases; the affected GMUs comprise portions of 3 deer (04, 010, 044) and 2 elk (E4, E9) data analysis units (DAUs). There is no evidence that wild deer or elk outside northeastern Colorado are infected with CWO. The significance of CWO and its impacts on native deer and. elk populations are unclear. Simulation models of CWO dynamics predict that this disease could cause significant declines in affected deer and elk populations (Miller and McCarty, unpublished data). In light of CWO's potential impacts on wildlife resources and the difficulties inherent in eliminating CWO from captive or wild cervid populations once established, it seems most prudent to assume CWO could adversely affect native deer and elk populations and manage to reduce its occurrence· and prevent its further spread. A more complete understanding of CWO is fundamental to developing a comprehensive management program. In 1996, ongoing surveillance efforts were enhanced to provide a better tool for estimating and monitoring changes in CWO prevalence in enzootic areas and for potentially detecting emergence of CWO in new areas. Reliable estimates of CWO prevalence are particularly critical to detecting trends, predicting potential impacts of disease on long-term population performance, and assessing efficacy of management interventions; moreover, such data are needed to guide policy decisions and to provide information to hunters and other publics. Ultimately, surveillance data will be the foundation of an adaptive resource management plan for CWO in deer and elk; that plan will provide a mechanism for incorporating new knowledge gained through surveys, modeling, and experimental studies into a continuously evolving management program formulated to reduce the occurrence of CWO and minimize the risk of its spread to other native deer and elk populations in Colorado. MATERIALS AND METHODS Surveillance We monitored deer and elk populations throughout Colorado for occurrence of CWO using a combination of targeted surveillance and harvest or road-kill surveys. These were organized and conducted as follows: Targeted (= clinical disease) surveillance: Deer and elk showing clinical signs consistent with those seen in chronic wasting disease were collected by field personnel statewide and brain tissues examined for evidence of spongiform encephalopathy. The "suspect case" profile was defined as follows: • Species: mule deer white-tailed elk deer �41 • Age: ~ 18 months • Signs: emaciated and abnormal behavior &/or indifference to human activity &/or increased salivation &/or tremor, stumbling, incoordination &/or difficulty or inefficiency in chewing/swallowing &/or increased drinking and urination Where possible, submissions were subjected to complete necropsy; in some situations, only heads were available for examination and sampling. In all cases, histopathology of brain tissue (Williams and Young 1993) was used to diagnose CWO; in some cases, immunohistochemistry or other ancillary tests were used to confirm or support diagnoses. Harvest surveys: In order to obtain reliable estimates of CWO prevalence that will serve as a basis for monitoring responses to management interventions, we continued conducting harvest surveys on select deer and elk populations. During the 1997-1997 hunting seasons, fresh brain and select lymphatic tissues were collected from 262 deer and 277 elk harvested in enzootic GMUs; 92 deer and 25 elk harvested in other GMUs throughout Colorado were also sampled. Brain tissues were examined at ,the Colorado State University Diagnostic Laboratory for histopathological lesions or anti-PrP immunostaining reactions consistent with CWO infection. Because sample sizes for most individual GMUs were too small to provide reliable prevalence estimates by GMU, we pooled data by DAU for comparisons within and among species. Small sample sizes also precluded meaningful analysis of data from deer or elk harvested outside known enzootic areas. Road-kill surveys: In addition to targeted surveillance and harvest surveys, we also began collecting road-killed deer and elk in select areas during 19961997 to augment harvest survey efforts. Thirty-seven deer and 12 elk were sampled from enzootic DAUs in Larimer County; 5 deer from D44 and 18 deer from Middle Park (D9) were also sampled. Epizootiological Studies We summarized epizootiological data gathered over 21 yrs from CWO-infected captive elk populations. A draft manuscript was submitted to the Journal of Wildlife Diseases. Diagnostic Approaches Immunohistochemistry (IHC) techniques were applied to samples collected during targeted surveillance and harvest/road-kill surveys. Additional evaluation of IHC, as well as development of Western blot techniques, is planned for next segment. Transmission Studies A study plan outlining experiments designed to evaluate susceptibility of cattle to chronic wasting disease was drafted by Williams et ale No other transmission studies were planned or conducted this segment. �42 RESULTS AND DISCUSSION Surveillance Targeted (= clinical disease) surveillance: Between June 1996 and May 1997, 22 CWO cases (16 deer, 6 elk) were diagnosed among 52 "suspects" (34 deer, 18 elk) submitted from northeastern colorado; CWO was not diagnosed in any of 12 additional "suspects" (6 deer, 6 elk) submitted from elsewhere in Colorado. Ten of the 22 new CWO cases originated in GMU 20. Four cases originated in GMUs where CWO had not been detected previously; these included an elk case in GMU 7 and deer cases in GMUs 29, 93, and 95 (Fig. 1). As in past years, most (77%) CWO cases were observed and submitted during October-April. Among deer cases, males and females were represented equally; in elk, females cases outnumbered males 5:1. All 4 of the cases representing apparent extensions of CWO distribution involved female animals. Although the number of cases detected during 1996-1997 is greater than in previous years, this is likely a function of increased surveillance efforts statewide. Similarly, the 4 newly identified GMUs represent predictable range extensions of CWO distribution in northeastern Colorado. These results also indicate that, afforded sufficient effort, our targeted surveillance approach should be effective in detecting CWO outside known enzootic areas. Harvest surveys: About 6.3% of deer (11/176) and 1.4% of elk (4/277) harvested in Larimer County DAUs (D4/E4 or D10/E9) tested positive for CWO via immunostaining; CWO was not detected in any of 86 deer harvested in DAU D44, or in any of the deer or elk harvested outside known enzootic areas. Among deer populations, prevalence did not differ (P = 1.0) between D4 (6.5%) and D10 (5.9%); prevalence in D44 (0%) was lower (P ~ 0.036) than in D4 or D10. Deer survey data revealed no evidence of sex-related bias in CWO prevalence: although comparatively few· does were harvested in D4 and D10, prevalence among does (10%) and bucks (5.5%) did not differ (P = 0.413). Estimated prevalence among harvested elk did not differ (P = 0.328) between E4 (0%) and E9 (2%), and prevalence also did not differ (P = 0.552) between cows (1.8%) and bulls (2.5%) harvested in E9. Overall, CWO prevalence among elk harvested in Larimer County was lower (P = 0.007) than in sympatric deer populations. The foregoing prevalence estimates may be somewhat liberal because the definition of "positive" included subclinical cases where either histopathological lesions or anti-PrP immunostaining reactions in brain tissue were observed. Of the 15 cases identified, only one harvested deer appeared to be suffering from clinical CWO. Six of the 11 positive deer showed both histopathological lesions and immunostaining; the other 5 deer and all 4 elk were classified as positive solely on the basis of immunostaining reactions. Although no known "false positives" were identified among the 92 deer and 25 elk examined from outside known enzootic DAUs, further evaluation of both sensitivity and specificity of existing diagnostic techniques still appears warranted. Observations based on harvest survey data agreed with those based on targeted surveillance data in two respects, but differed in a third. Using either data set, Larimer County appears to be the most significant focus of CWO in northeastern Colorado; although 4 clinical CWO cases have been submitted from GMUs east of I-25, harvest survey data demonstrate that prevalence is lower in D44 than in D4 or D10. Similarly, CWO prevalence is higher in Larimer County deer populations than in sympatric elk populations; these species differences are also reflected in the predominance of deer cases detected via targeted �43 surveillance. However, targeted surveillance data have historically suggested that mule deer bucks were affected by CWO more frequently than does; harvest survey data, although somewhat limited for does, do not support that observation. Larger sample sizes provided by doe harvests in 04 and 010 during 1997-1998 should provide sufficient power to more reliably test hypotheses about sex-related differences in CWO prevalence. During the 1996-1997 rifle seasons, harvest survey participation was required of successful antlerless elk hunters in E4, all elk hunters in E9, and all deer hunters in GMUs 94 and 951. In E9, about 70% of the successful elk hunters submitted heads as required. In E4, submission rates (r~) were higher (P < 0.001) for successful antlerless elk hunters (r~ = 37%) than for successful antlered elk hunters (r~ = 7%) who were expected to comply voluntarily. Similarly, submission rates in GMUs 94 and 951 (r~ = 31%) tended to be higher (P = 0.13) than in GMU 96 (r~ = 24%), where a letter sent to hunters requested their voluntary participation; submission rates were substantially lower (P < 0.001) in 04 and 010 (combined r~ = 11%), where deer hunters were expected to participate in the absence of any direct requirement or request to do so. In general, it appears compelling participation in harvest surveys via regulation is the most effective method for increasing sample sizes. Three of 27 the deer or road-killed survey (P = evidence of between the (11%) road-killed deer from 04 tested positive for CWO; none of elk from other DAUs were positive. Prevalence estimated from deer in 04 did not differ from prevalence estimated via harvest 0.418). As with harvest survey data, road-kill data revealed no sex-related bias in CWO prevalence: prevalence did not differ 18 does (11%) and 9 bucks (11%) sampled from 04. Although sample sizes provided by road-kill collections were far too small to use in monitoring trends or comparing prevalence among DAUs, this approach could be used to augment survey efforts in urban areas (e.g., Boulder County, Estes Park). Based on 1996-1997 data, sampling road-killed deer appears at least as sensitive as harvest sampling in detecting CWO; however, both survey approaches are about an order of magnitude less sensitive in detecting CWO than targeted surveillance of clinical suspects. Epizootiological Studies Epizootiology of chronic wasting disease in captive Rocky Mountain elk (Ceryus elaobus nelsoni) (Miller, Wild, and Williams): Between June 1986 and May 1997, chronic wasting disease (CWO) was the only natural.cause of adult mortality among captive Rocky Mountain elk (Cervus elapbus nelsoni) held at a wildlife research facility near Fort Collins, Colorado, USA. Of 23 elk that remained in this herd> 15 mo, four (17%) developed CWO. All affected elk were unrelated females from the founding cohort, captured as neonates and raised in 1986. The index case was diagnosed in 1989; time intervals between subsequent cases averaged about 21 mo (range 13 to 32 mo). Initial age at onset of clinical signs ranged from about 2.9 to 8.1 yr; duration of clinical disease averaged about 7.5 mo (range 5 to 12 mo) prior to euthanasia. Intraspecific lateral transmission of CWO provided the most plausible explanation for the epizootic pattern observed; neither periparturient nor maternal transmission were necessary to sustain this epizootic. Early detection and elimination of incubating or clinical individuals may have aided in reducing infection rates during the epizootic. Transmission routes and rates, pathogenesis, antemortem diagnostic tools, and the potential role of reservoirs or environmental contamination in perpetuating CWO epizootics warrant further investigation. �44 Diagnostic Approaches Immunohistochemistry (IHC) appeared to enhance CWO diagnosis in evaluating both CWO suspects and samples collected via ·surveys. Sensitivity and specificity of IHC remain to be fully determined; in particular, apparent false positive results from stained llymphoid tissue warrant further investigation. Additional evaluation of IHC, as well as development of western blot techniques, should greatly enhance efficiency and reliability of CWO surveillance activities. Transmission Studies A draft plan describing studies Those studies will be initiated of CWO transmission next segment. to cattle is appended. Acknowledgments The statewide CWO monitoring and surveillance program described here relies heavily on efforts of dedicated field personnel throughout the Colorado Division of Wildlife, and truly represents a division-wide effort to improve our understanding and management of this important disease problems. In addition to those specifically listed, we collectively thank all of those regional and area biologists, district and area wildlife managers, volunteers, deer and elk hunters, and others who assisted by submitting suspect cases, harvested animals, or road-killed animals throughout the year. Literature cited Miller, M. W., M. A. Wild, and E. S. Williams. 1997. Epizootiology of chronic wasting disease in captive Rocky Montain elk. J. Wildl. Dis. 34: in review. Spraker, T. R., M. W. Miller, E. S. Williams, D. M. Getzy, W. J. Adrian, G. G. Schoonveld, R. A. Spowart, K. I. O'Rourke, J. M. Miller, and P. A. Merz. 1997. Spongiform encephalopathy in free-ranging mule deer (Odocoileus hemionus), white-tailed deer (Odocoileus virginianus), and Rocky Mountain elk (Cervus elaphus nelsoni) in northcentral Colorado. J. Wildl. Dis. 33:1-6. Williams, E. S., and S. Young. 1980. Chronic Wasting disease of captive mule deer: A spongiform encephalopathy. Journal of Wildlife Diseases 16: 89-98. _____ , and 1982. Spongiform encephalopathy of Rocky Mountain elk. Journal of Wildlife Diseases 18: 465-471. _____ , and 1992. Spongiform encephalopathies in Cervidae. Revue Scientifique et Technique Office International des Epizooties 11: 551-567. _____ , and 1993. Neuropathology of chronic wasting disease in mule deer (Odocoileus hemionus) and elk (Cervus elaphus nelsoni). Veterinary Pathology 30: 36-45. Prepared by Wildlife Research Veterinarian �45 CWO-endemic GMUs prior to 1996 other GMUs In CWO-endemic OAUs Figure 1. PriQr to 1996, all of Colorado's free-ranging CWO cases had come from 1 of 6 different northeastern Colorado game management units (GMUs); during 1996-1997, cases were detected in 4 additional GMUs (7, 29, 93, 95). Despite these apparent extensions of CWO distribution, most 1996-1997 cases originated from Larimer County GMUs considered collectively to be the major focus of CWO in Colorado. �46 UNITED STATES DEPARTMENT OF AGRICULTURE OMB Approved OS24-OO33 COOPERATIVE STATE RESEARCH, EDUCATION, AND EXTENSION SERVICE Expires 6/97 NATIONAL RESEARCH INITIATIVE COMPETITIVE GRANTS PROGRAM PROPOSAL TYPE Principal Investlgator(s): PI #1 Elizabeth o Standard S. Williams Institution University Institution Colorado Division Game and Fish Dept. PI #2 ______________________ Michael W. Miller PI #3 T_e_rry-=--J_._Kr __e_e.::.g_er Institution Wyoming PI #4 H_a_n_a_V_a_n __C_a_m_,p'-e_n Institution University ProjectTitle: " - of Wyoming of Wildlife of Wyoming Research Proposal o Conference o AREA Award a Postdoctoral o New Investigator Strengthening; " 0 Career Enhancement o Equipment o Seed Grant Standard Strengthening o S::..u::..s::.;c:..:e:Lp:..:ti::;:b~iH:.:.·tyt.....=.0f:._ca=:;"::.:t:.::tl.::.e...:.to~c.::.e:....:rv:..:.;id:..".:s.E::.po::,;n:..:J9:2,!i!,:fo:.:.nn::..:.!..e::,;n:.:,;ce=p:.:.h _ Key Words: _ ___;s;:Jp:..:o:..:n.:;;gz.:,:if:.,::o:.:.:nn:..:..:._;e::.:n..:=ce:.::.r::p:.:.:h:::a.:.::I0:.r::p:::a:.::th.!JY"'-'..::c:!,!h!.:ro~n,!!;ic:::....:.!w~a~s.!!tin~g::z...:d::.::is:..:e B=S.=E"._ . .::.e ••• xp~e=:n .•.•. im.•.•. e~n"",t""a,,-I ~i"-ra"""",ns:::Jm ..•..•.•.• is""s:!.>io"""n .••. ,_"ca.,,,...,ttwleo<- PROJECT SUMMARY (Aproximately 150 words) The emergence of bovine spongiform encephalopathy (BSE) in Great Britain has shaken animal agriculture throughout the wortd. The cattle industries in Great Britain and other affected countries have been severely impacted. International trade in bovine products among many countries and trading blocks has been disrupted. Perhaps more importantly, there has been loss of confidence by some of the public in governmental enJities responsible for food safety, and, in particular, safety of beef and beef products. It is important, therefore, to understand host range and effects of spongiform encephalopathies endemic in the United States. Chroni wasting disease (CWO), a naturally occul)ing spongiform encephalopathy of cervids (members of the dee family), affects captive and free-ranging mule deer, white-tailed deer, and Rocky Mountain elk in northcentral Colorado and southeast Wyoming. Range cattle and CWO-affected cervids are sympatric in these areas. CWO is laterally transmitted among three cervid species and cross-species lateral transmission from sheep to goats occurs in scrapie, the prototype animal spongiform encephalopathy. This suggests cattle could become infected with the prion of CWO by contact with affected cervids or from contaminated ranges. The long-term goal of the proposed research is to determine if cattle are susceptible to CWO. To test hypotheses that domestic cattle are susceptible to CWO we propose two approaches, In the contact exposure study, cattle and mule deer will be housed in CWD-endemic wildlife facilities and exposed to " clinical cases of cWO as they occur. Mule deer will serve as controls for exposure to infectivity in the facilities and from CWO affected deer. In the oral exposure study, cattle will be exposed to CWO-affected mule deer brain given orally at a higher dose than likely to occur under natural conditions. Tonsillgr and lymph node biopsies Will be collected from principals for immunohistochemistry and Western blots to detect pfPSc as. a method to monito for extraneural replication of the agent If experimental cattle and deer develop CWO, clinical" disease, neurops,th0logy, and_distribution of pfPSc in the anirt_lSls will be documented. We believe exposure via contact in e~e1t)iQ (i;l~lities and by exposure would represent a much greater exposure than would be expected on range, but witr provide direct evidenCe of the potential for CWO to cross species lines to cattle. In the absence of these proposed trials, national and lnternafional perceptions of the risk of a new spongiform encephalopathy in catHe couid adversely affect United States cattle industries well into the future. o~ �47 Colorado Division Wildlife Research July 1997 of Wildlife Report JOB PROGRESS State of Project REPORT Colorado No. W-153-R-10 Mammals Research Work Plan No. Elk Inyestigations Job No. Estimating Developing Population Period Author: Covered:; July Survival Rates of Elk Techniques to Estimate Size 1, 1996 - June 30, 1997 D. J. Freddy Personnel: J. Broderick, G. Byrne, J. Ellenberger, D. FOx, J. Frothingham, Graham, D. Masden, C. Mehaffy, M. Okata, J. Ritchie, P. Will, R. Winn, CDOW; D. Bowden, G. White, CSU; Diagnostic Laboratory, CSU; N. Miers, D. OUren, volunteers; BLM Glenwood Springs, CO, NBS Ft. Collins, CO, USFS Rifle, CO, Rocky Mountain Elk Foundation, cooperating. v. Abstract We captured and radio-collared 70 calf elk (Cervus elaphus nelsoni) 6-months old in December 1996 to estimate survival rates during winter 1996-97 and to increase numbers of radiocollared elk available for experiments utilizing mark-resight models to estimate population density. Elk were captured using helicopter net-gunning and portable corral traps. Survival rates (± 95% CI) for 6-11 month old calves during winter-spring averaged 0.89 ± 0.04 with yearly rates ranging from 0.86 to 0.92 between 1993-94 and 1996-97. Survival was similar among years (P > 0.50) and sexes (P > 0.50). Primary causes of death for calves were suspected predation (58%) and suspected malnutrition (26%). Survival of adult females (~ 12 months old) during winter-spring averaged 0.96 ± 0.02 and was similar among years (P > 0.60) with yearly rates ranging from 0.94 to 0.98 between 1993-94 and 1996-97. Hunting was the primary cause of death (59%.) for adult females during winter-spring. Survival of elk 18-23 -months of age during winter-spring averaged 0.97 ± 0.03 for males and 0.97 ± 0.04 for females. Annual survival (1 December-30 November) of adult females averaged 0.81 ± 0.06 from 1993-94 through 1995-96 and survival was similar among years (P > 0.50) with hunting accounting for 81% of the deaths. Survival during summer-fall for adult females averaged 0.90 ± 0.03 with hunting accounting for 95% of the deaths. Survival during summerfall for elk 12-17 -months of age averaged 0.87 ± 0.07 for males and 0.94 ± 0.05 for females with all deaths attributed to hunting. Survival during summer-fall for male elk 24-months old averaged 0.15 ± 0.10 with all deaths due to hunting. Elk population size in 1997 was 1,854 ± 194 (95% CI) based on random quadrat sampling and 3,610 ± 641 based on the JHE pooled mark-resight estimator. These estimates were different (P < 0.05) and continued the disturbing trend of quadrats estimating only 51-93% of the numbers of elk estimated by mark-resight estimators. Adjusting quadrat estimates with �sighting bias formulas did not meaningfully narrow differences observed between quadrats and mark-resight estimators. Discrepancies in estimators reflect the possibility that we may be violating major assumptions(s) underlying either mark-resight or quadrat theory. We believe further field testing to evaluate potential sources of bias will be necessary to evaluate these techniques to estimate population size. �49 ESTIMATING SURVIVAL JOB PROGRESS REPORT RATES OF ELK AND DEVELOPING ESTIMATE POPULATION SIZE David TECHNIQUES TO J. Freddy P. N. OBJECTIVE Estimate estimate survival rates of adult population size. female SEGMENT 35 male and calf elk and develop techniques to OBJECTIVES 1. Radio-collar and 35 female calf elk in December 1996 in GMU-42. 2. Estimate winter radiocollared radiocollared 3. Estimate density of elk in a portion of GMU-42 using 4 helicopter flights involving 2 nonrandom mark-resight flights and 2 flights using a random quadrat sampling system. Apply sighting bias corrections to adjust numbers of elk counted on quadrat flights. Compare bias adjusted quadrat estimates with mark-resight estimates based on numbers of marked elk seen during all 4 flights. 4. Analyze 5. Prepare draft of manuscript on sighting collected in 1994 and 1995. 6. Continue and annual survival rates of calf and previously adult female and male elk from known fates of animals. data and summarize to monitor annually locations in Federal Aid Job Progress bias models and movements developed of selected reports. from data radiocollared elk. INTRODUCTION OUr objectives are to provide reliable estimates of survival rates for calves during winter and for adult females and males throughout the year for the period 1993-94 through 1996-97. For adults, we are especially interested in survival rates inclusive of hunting mortalities which reflects human-induced rates of removal and exclusive of hunting mortalities which reflects natural rates of survival. Additionally, we will develop and test a system for estimating population size that will incorporate estimates of sighting bias in conjunction with a random sampling system using search quadrats as sample units. OUr winter study area encompasses about 839 km2 (324 mi2) in the eastern half of Game Management Unit 42 south and east of Rifle, Colorado. Elk winter range vegetation types include juniper-pinyon woodland (Juniperus osteosperma-Pinus edulis), oakbrush-mountain shrub (Quercus gambeliiAmelanchier alnifolia), aspen (Populus tremuloides), sagebrush (Artemisia tridentata), and agricultural fields (Freddy 1993, 1994). METHODS Capture and Marking We placed radio collars (172-176MHz) having mortality sensors on 70 6-month old calves, of which 35 were males and 35 were females. Calves were trapped �50 from 7-10 December 1996 using helicopter net-gunning. Helicopter capture occurred at 10 remote sites located primarily on public lands. Trapping effort was allocated among 8 geographic trap zones to assure that radioed elk were representative of most if not all segments of the population (Table 1). Radio collars were of the same type used in previous years (Freddy 1994). Calves captured by net-gunning were ferried by helicopter to processing points usually within 1.6 km of capture sites. At processing points, body weight, total body length, hind foot length, and rectal body temperature (F) were measured anq calves were then radio-collared and released. Body measurements for calves were compared between sexes and years using Proc FREQ and GLM (SAS 1988). Survival We monitored life or death status of radioed elk during daily ground surveys and .aeria1 surveys conducted at 2-4 week intervals from December 1996 through April 1997 and via aerial surveys at 2-4 week intervals from July to November 1996 and May to June 1997. Life or death status of all calves radioed in December 1996 (70) was known for the period 7 December 1996 through 14 June 1997 which is the time period over which survival rates for calves, age 6-11 months, are calculated. On 15 June, calves by definition become 12-month old yearlings. Life or death status of 151 female and male elk previously collared in December 1993-1995 that had survived to 14 June 1996 was known for 148 of these elk through 15 June 1997. Survival rates (S) of radioed elk were calculated using the binomial estimator with a variance, VAR(S) = S(l-S)/n (White and Garrott 1990). Survival rates are expressed as the mean estimate ± the 95% confidence interval. We used x2 -contingency tests to compare survival rates. We defined 4 major time intervals for survival analyses: winter-spring was 1 December to 14 June, summer-fall was 15 June to 30 November, annual was 1 December to 30 November to coincide with timing of capture and radioca11aring, and yearly, only for yearling elk aged 12-23 months, was 15 June to subsequent 14 June. Causes of death were estimated from multiple sources of evidence including: presence or absence of gunshot wounds, presence or absence of bite wounds on carcass and predator tracks or scat at carcass site, physical positioning of carcass remains whether buried, covered, scattered, or consolidated, relative amount of internal fat and marrow fat if present with carcass, and results of histopathology and marrow fat analyses (Wad~ and Browns 1982, Halfpenny and Biesiot 1986). Fat content (percent dry matter) of bone marrow and estimates of age based on dental cementum were obtained for dead elk by the Colorado Division of Wildlife Laboratory while histopathology analyses were provided by the Colorado State University veterinary Diagnostic Laboratory. Photographs were taken of nearly all mortalities so that physical evidence could be reviewed and judged by outside experts (pers. comm. A. Ande~son, T. Beck, W. Ande1t). Population Estimates We continued to evaluate methods to estimate elk population size or density during winter in that portion of GMU 42 from East Alkali Creek west to Grass Mesa, south of Rifle, CO. In previous years, 95% confidence intervals about population estimates based on quadrat sampling exceeded ± 30% of the estimated population size. We addressed this problem by refining strata boundaries and �51 by sampling 100% of those quadrats in high density strata, effectively reducing sampling error to 0 in these high density strata. About 30% of the quadrats in low density strata were randomonly sampled which was similar to previous years. With this strategy, we selected 72 quadrats in 15 strata representing a 53% sample of the 137 mi2 (351 km2) winter range (Table 14). As in winter 1996, each potential quadrat was rated as high or low in expected density, individual.quadrat boundaries remained unchanged from previous years and were based on topographic features, and quadrats were about 1 mi2 (2.6km2) in size. We felt this strategy to increase numbers of quadrats flown during 1 flight, instead of flying 2 repl~cates of a smaller sample of quadrats per flight as originally planned, would be more cost-effective in addressing problems of precision. We counted elk during 3 flights using a Bell-Soloy helicopter having a pilot, observer, and navigator-observer, all of whom could potentially detect elk during surveys. One flight per day was flown on 10 and 13 February 1997 to count as many marked and unmarked elk as possible encountered along a nonrandom route within the 137 mi2 area with flying time limited to about 7.5 hours per flight. On 11 and 12 February, marked and unmarked elk were counted on quadrats. Two helicopters were used each day and assigned to different strata to shorten the total time interval over which quadrats were completed. Total flight time for both helicopters was 29 hours. Pilots were common to all 3 flights. The primary observer on nonrandom flights did not participate in quadrat flights. The navigator-observer for the nonrandom flights was 1 of 3 primary observers used on quadrat flights while navigator-observers for quadrat flights only flew quadrats. On February 11 and 12, fixed-wing flights were conducted to confirm locations of. 223 radiocollared elk to determine whether these elk were within or outside of the 137 mi2 sample area. We generated estimates of population size for each individual flight and 3 flights pooled using several mark-resight estimators in program NOREMARK (White 1996). We considered estimates based on all 3 flights pooled to be our best estimate of population size and the benchmark against which estimates based on quadrat sampling would be compared. Estimates of population size based on stratified random quadrat sampling were computed using program DEAMAN (Colo. Div. Wildl. software). We then applied a sighting bias correction formula to each group of elk counted on each quadrat, resulting in adjusted counts per quadrat, and recalculated populations estimates based on quadrat sampling. To correct for negative sighting bias, we used a simple 2 parameter formula to correct for the probability of detecting an individual group: Yccorrecte<lqroupsizel = 1.1619(Log Group Sizeccounte<ld + 0.0806 (Freddy 1995). MOYements We continued to locate selected radioed elk at least once per month since capture to document seasonal movements via telemetry using a Cessna 185 • .These elk were originally selected at random from within trap zones and equalized by age class in January 1994. Elk from the original sample that died were replaced each subsequent January primarily with randomly selected 6 month-old calves of the same sex from the same trap zone(s) as elk that died. As of 1 January 1997, these 44 elk were classified as 23 adult females, 4 yearling females, 5 female calves, 1 adult male, 5 yearling males, and 6 male calves. During June 1997, we again located an additional 28 adult females, that were originally selected at randQm in 1995, to document locations during the calving period. Females from the original sample that died were replaced each subsequent June with adult females randomly selected from the same trap �52 zones of elk that died. movements. As needed, we located other elk to document unusual RESULTS AND DISCUSSION capture There were no acute deaths of calves during capture. One male calf (174.619/96) died within 14 days of capture due to the effects of capture myopathy (histopathology, CSU Diagnostic Lab) and was subsequently censored from survival analyses (Table 3). Survival Between 1 December 1993 and 14 June 1997, 146 radiocollared elk died (Table 2, Appendix 1). Hunting was apparently involved in 71% of the deaths. For adults ~12 months old, hunting accounted for 90% of 115 deaths. There were 2 periods of mortality during the year. Adults died primarily from September to January when hunting seasons occurred (Fig. 1) while calves died primarily from February to May (Fig. 2). Calves Survival for calves during winter-spring averaged 0.89 ± 0.04 (n = 280) with yearly rates ranging from 0.86 to 0.92 (Table 3). We failed to detect differences in survival of calves among years (X23 = 1.5, P >0.50), between sexes pooled among years (X21 = 0.43, P >0.50), and between sexes within each yearly cohort (X21 ~ 0.79, P > 0.40) (Tables 3, 5, 6, 7). Sex of dead calves for all years was 17 male and 14 female (Table 2). These survival rates were associated with winters considered mild in temperature. Snow was usually low or moderate in depth except in 1995 when considerable snow fell during March and April but usually melted rapidly at lower elevations, and in 1997 when a storm in January delivered at least 40 cm of snow on lower elevation winter ranges. Rain in January 1997 subsequently settled this snow to about 25 cm and caused severe crusting of snow. Elk in some segments of the lower winter range moved in excess of 24 km to areas at even lower elevations along the Colorado River near Rulison and Parachute, CO soon after this storm occurred. Survival rates of elk, however, remained high and apparently were not greatly affected by this weather event. Primary causes of death for all calves during winter-spring were malnutrition (13%), suspected malnutrition (13%), mountain lion predation (35%), suspected predation (23%), .and unknown cause (10%) (Tables 2, 4). Timing of deaths peaked in March and April for both males and females (Fig. 3). Deaths attributed to predation occurred from January into June and those attributed to malnutrition occurred from February to May (Fig. 2) but both presumed causes of mortality peaked in March and April suggesting a functional change in vulnerability of calves during these months. However, percent fat content in marrow of calves lost to predation and malnutrition suggested that malnutrition may not have predisposed calves to predation. Percent fat in bone marrow of calves suspected of dying from predation was 69 for males (range 2-95, n = 10) and 36 for females (range 8-63, n = 8) and for calves dying from suspected malnutrition, 11 for males (range 0.2-41, n = 4) and 18 for females (range 8-31, n = 3) (Table 4). Possibly elk become more sedentary during March and April to conserve energy reserves and thus more predictable in behavior allowing predators, especially mountain lions, to hunt calves efficiently. �53 Calf Body Size Averaged over all years, male calves had larger body weights, longer total body length, longer hind leg lengths, and higher condition indexes (P ~ 0.023) than female calves (Tables 11, 12). Differences in body size were greatest for weight, as males (115 kg) were about 8% larger in mass than females (106 kg). Differences between sexes in other body dimensions were <3%. This advantage in mass, however, did not translate into higher rates of survival for male calves. Body weights were largest for males in 1995 (119 kg), for females in 1996 (110 kg), and for sexes combined in 1995 (113 kg) (Table 12), but we failed to detect differences in body weights among years (P = 0.15) and there was no interaction between year and sex (P = 0.34). Increases in weights in 1995 were not unexpected as the locally moist summer was favorable to forage production on summer ranges. Weights measured at capture in December for calves suspected of dying from predation (Table 4) averaged 116 kg for males (range 72-140 kg, n = 9) and 96 kg for females (range 78-120, n = 10). Predators thus appeared to take males of average and females of smaller than average size (Table 12). Weights of calves suspected of dying from malnutrition (Table 4) averaged 93 kg for males (range 77-127, n = 5) and 101 kg for females (range 100-102, n = 3). Malnutrition thus appeared to affect smaller than average sized males and females (Table 12). Weights of calves dying from all causes averaged 108 kg for males (range 72-140, n = 16) and 97 kg for females (range 78-120, n = 14) (Table 4), which was about 7% below average weights at capture for both sexes. Yearlings Yearly survival for elk 12-23 -months of age was 0.84 ± 0.08 for males (n 90) and 0.91 ± 0.06 for females (n = 97) when averaged among years with hunting deaths included (Tables 5, 6, 7). Survival among years for males ranged from 0.79 to 0.88 and for females from 0.81 to 0.97. We failed to detect differences in yearly survival rates among years for males (X22 = 0.88, P > 0.50) but survival was lower (0.81) for females in 1996-97 (X22 = 5.75, P = 0.06). Yearling males were subjected to about the same rate of illegal hunting during all years which accounted for most of the male mortality, while in 1996-97, females had a lower survival rate because 5 of 6 deaths were hunting related (Tables 5, 6, 7). With hunting deaths included, we failed to detect differences in survival between sexes within (X21 ~ 2.25, P > 0.15) and 2 among years (X 1 = 1.71, P > 0.25). Of 23 deaths, 19 (83%) were hunting related. Hunting deaths involved 12 m~les of which 9 (75%) were classified as illegal kills (Table 2). Yearly survival for elk 12-23 -months of age was 0.97 ± 0.03 for males (n 79) and 0.98 ± 0.02 for females (n = 91) when averaged among years with hunting deaths censored (Tables 5, 6, 7). Survival among years for males ranged from 0.96 to 1.00 and for females from 0.97 to 1.00. We failed to detect differences in yearly survival rates among years for males (X22 = 1.14, P > 0.50) and for females (X22 = 1.23, P > 0.50), and between sexes within and among years (X\ ~ 0.007, P > 0.90). Survival during summer-fall for yearling elk 12-17 -months of age was 0.87 ± 0.07 for males (n = 91) and 0.94 ± 0.05 for females (n = 98) when averaged among years and inclusive of hunting deaths, and 1.00 for both males (n = 79) and females (n = 92) with hunting deaths censored (Tables 5, 6, 7). Survival among years for males ranged from 0.83 to 0.90 and for females from 0.87 to �54 0.97 inclusive of hunting mortalities. We failed to detect differences in survival rates. among years for both males (X22 = 0.70, P > 0.50) and females (X22 = 3.63, P > 0.20) inclusive or exclusive of hunting deaths. Survival rates of females (0.94) were higher than males (0.87) when averaged among years and inclusive of hunting deaths (X21 = 2.73, P = 0.10). There were no non-hunting deaths during summer-fall for either males or females (Table 2). survival during winter-spring for yearling elk 18-23 -months of age was 0.97 ± 0.03 for males (n = 78) and 0.97 ± 0.0.04 for females (n = 91) when averaged among years and inclusive of hunting deaths, and 0.97 ± 0.03 for males (n = 78) and 0.98 ± 0.03 for females (n = 90) with hunting deaths censored (Tables 5, 6, 7). We failed to detect differences in survival rates among years for both males (X22 = 1.16 P > 0.50) and females (X22 < 2.59, P > 0.30) inclusive or exclusive of hunting deaths. We also failed to detect differences in survival rates between sexes within and among years inclusive or exclusive of hunting deaths (X\ < 0.24 P > 0.50). During winter-spring, there were 2 nonhunting deaths each for males and females and 1 illegal hunting death for a female (Table 2). Yearling spike-antlered males were generally not legal quarry during hunting seasons. In 1994, 32 yearling males from the 1993-94 calf cohort entered the hunting seasons presumably as spike-antlered males and 4 (13%) were illegally taken during rifle seasons. In 1995, 30 yearling males from the 1994-95 calf cohort entered the hunting season and 2 (7%) were illegally taken and 1 (3%) was fatally wounded during archery season in an area where the elk was legal quarry. In 1996, 29 yearling males from the 1995-96 calf cohort entered the hunting seasons and 4 (14%) were illegally taken during rifle seasons and 1 (3%) was an assumed wounding loss during rifle seasons because the animal had a 3 x 3 antler point configuration that may have resembled a legal branchantlered male (Tables 2, 5, 6, 7). Adult Females Survival during winter-spring for all adult females ~12-months of age was 0.96 ± 0.02 (n = 409) when averaged among years and inclusive of hunting deaths and 0.98 ± 0.01 (n = 399) with hunting deaths censored. Survival among years ranged from 0.94 to 0.98 and from 0.97 to 0.99, inclusive and exclusive of hunting mortalities, respectively. We failed to detect differences in survival among years inclusive (X23 = 1.93, P >= 0.50) or exclusive (X23 = 2.85, P > 0.60) of hunting mortalities (Table 8). There were 17 winter-spring deaths, of which 10 (59%) involved hunting: 5 were killed and 3 were wounded during rifle late-seasons and 2 were illegally shot out of hunting seasons (Table 2). Seven natural deaths were attributed to mountain lion predation (1), suspected predation (2), complications while calving (1), and unknown cause (3) (Table 2). survival during summer-fall for all adult females ~12-months of age was 0.90 ± 0.03 (n = 372) when averaged among years and inclusive of hunting deaths and 0.99 ± 0.01 (n = 337) with hunting deaths censored. Survival among years ranged from 0.87 to 0.94 and from 0.99 to 1.00, inclusive and exclusive of hunting mortalities, respectively. We failed to detect differences in survival among years inclusive (X22 = 3.31, P > 0.15) or exclusive (X22 = 1.24, P > 0.50) of hunting mortalities (Table 8). Hunting was involved in 35 (95%) of 37 summer-fall deaths: 24 were killed, 8 were wounded, and 3 disappeared during archery, muzzleloading, and rifle seasons. Natural deaths were attributed to suspected predation (1) and complications while calving (1) (Table 2). At the beginning of hunting seasons in fall 1994, 1995, and 1996 �55 there were 99, 129, and 142 radiocollared adult females (~12-months of age), respectively, available to hunters (Table 8, non-hunting deaths censored). Percent of marked elk removed by all types of hunting-related mortalities averaged 10% annually (range = 6-12). Annual survival for adult females ~12-months of age marked as a cohort in December 1993 was 0.81 ± 0.06 (n = 163) when averaged among years and inclusive of hunting deaths and 0.96 ± 0.04 (n = 138) with hunting deaths censored. Survival among years ranged from 0.78 to 0.88 and from 0.92 to 0.98, inclusive and exclusive of hunting mortalities, respectively. We failed to detect differences in survival among years inclusive (X21 = 0.19, P > 0.50) or exclusive (X21 = 0.15, P > 0.50) of hunting mortalities (Table 9). For this cohort of adult females, hunting was involved in 25 (81%) of 31 deaths during all years: 13 were killed, 9 were wounded, and 2 disappeared during archery, muzzleloading, and rifle seasons and 1 was illegally killed out of hunting seasons. Natural deaths were attributed to mountain lion predation (1), suspected predation (1), and complications while calving (2). Adult Males Survival during summer-fall for all adult males 24-months of age was 0.15 ± 0.10 (n = 53) when averaged among years and inclusive of hunting deaths and 1.00 (n = 8) with hunting deaths censored. Survival among years ranged from 0.08 to 0.21 inclusive of hunting mortalities. We failed to detect differences in survival among years (x\ = 1.86, P > 0.20) (Table 5, 6). All deaths were attributed to hunting. At 24-months of age, males are branchantlered and therefore become legal quarry for hunters. Two of 4 males that lived to be 36-months old survived their second hunting season as legal quarry. Survival during winter-spring for adult males >30-months of age was 1.00 (n = 8) during all years (Tables 5, 6). There were no non-hunting deaths of adult males. Hunting was the cause of mortality among male elk ~24-months of age. From the 1993-94 cohort of 36 male calves, 28 (78%) lived to become 24-month old branch-antlered males and enter the 1995 hunting seasons. Of these 28, 22 (79%) were harvested in 1995 inclusive of 2 that disappeared during hunting seasons and 2 wounding losses (9% of 22). From the 1994-95 cohort of 33 male calves, 25 lived to 24-months and entered the 1996 hunting seasons. Of these 25, 23 (92%) were harvested in 1996 inclusive of 2 that disappeared during hunting seasons, 1 illegally taken during rifle season, and 1 wounding loss (4% of 23). Regular rifle seasons accounted for 33 (70%) of the hunting mortalities (Table 2). Most 24-month old males were harvested in the Grand Mesa DAU or in adjacent GMU 43. Exceptions were 3 males killed in GMUs 63 and 52 about 160 km south of where these males were captured as calves in GMU 42. The differential impact of hunting on survival and recruitment of males and females to young adult age classes is demonstrated by net survival rates of calf cohorts. For 1993-94 and 1994-95 calf cohorts, net survival from 6 to 36 months of age ave~aged 0.09 for males and 0.80 for females (Tables 5, 6). Survival Rates and Population Modeling We incorporated estimates of survival rates, population size, and composition into a simple spreadsheet model to assess potential for population growth or decline (Table 13). We used natural survival rates (hunting deaths censored) �56 for summer-fall and winter-spring time periods for adult males, yearling males, adult females, yearling females, and calves. Hunting deaths were incorporated'as harvest removal rates for each sex/age class. We assumed survival of calves 0-5 months of age during summer was 1.0 because the model uses post-season calf:cow ratios (measured in January) as an estimate of net calf recruitment to the beginning of winter (December). We also assumed that the recruited sex ratio calves to December is 1:1. Modeling suggests that if harvest rates of antlered elk remained as measured and harvest rates of antlerless elk were reduced to 0, the population could grow at an annual rate of 17%. Current rates of antlerless harvest allow annual growth rates of 7%. To stabilize population growth, harvest rates on antlerless age classes must increase nearly two-fold. Obtaining this level of harvest could be a formidable management challenge. Population Estimates Elk population size in 1997 was 1,854 ± 194 (95% CI) based on quadrat sampling and 3,610 ± 641 based on the JHE pooled mark-resight estimator (Tables 14, 15, 16). The quadrat sample was lower than the JHE estimate (P ~ 0.05). We achieved our primary goal of improving precision of the quadrat estimate to facilitate detecting dif·ferences in quadrat and mark-resight estimators (Table 16). A disturbing trend has emerged in estimates of population size during the last 3 years. Quadrats provided estimates of size that represented only 51 to 93% of the numbers of elk estimated by pooled mark-resight estimators, excluding initial and preliminary quadrat sampling in 1995 when the quadrat estimate was 39% of the mark-resight estimate. Mark-resight estimates have ranged from 3,177-3,924 elk based on pooled or individual flights while quadrat estimates have ranged from 1,854-3,387 elk (Table 16). The consistent magnitude in discrepancy among mark-resight and quadrat estimates was demonstrated well in 1997 as the 3, one-sample Lincoln-Peterson mark-resight estimates, whether based on marked and unmarked elk counted on nonrandom flights or on quadrats, were 3,289-3,924 elk while the estimate based on stratified quadrat sampling was 1,854 elk (Table 16). The 3 mark-resight estimates were based on 3 flights that were independent in methodology, strongly suggesting that the probability of detecting marked elk is consistent among different combinations of observers and types of flight patterns. A similar trend among estimates occurred in 1996 (Table 16). We adjusted quadrat estimates for sighting bias using a simple 2 parameter sighting bias model. Adjusting for sighting bias increased quadrat estimates only 7-14% (Table 16). Using more complex sighting models did not meaningfully narrow the differences observed between quadrat-and mark~resight estimates. Discrepancies in estimators reflect the real possibility that we may be violating a major assumption(s) underlying either mark-resight estimators or quadrat sampling with sighting bias corrections. Three important potential sources of bias are: 1) detecting and accurately counting groups of elk but missing marked elk within detected groups would inflate mark-resight estimates, 2) overestimating numbers of marked elk within the sample area at the time of flights would inflate mark-resight estimates, and 3) detecting a lower than expected percentage of elk groups on quadrats wo.uld deflate quadrat estimates. If bias 1) is true, we suggest that behavior of marked elk when approached by a helicopter would be different than other elk in the same group �57 with the possibility that marked elk either do not move with the group when the group is initially disturbed or they move away from the group reducing the chances of observers seeing marks. Another possibility is that some fraction of the white collars used as marks were not visible due to discoloration but we tend to discount this possibility because collars that were returned by hunters or were removed from other recovered mortalities were still white. If bias 2) is true, accuracy in determining elk locations during specific census location flights was much poorer than our measured accuracy of ~300 m. We cannot eliminate this source of bias, but each marked elk was located and the estimated GPS location verified on maps delineating the sample area. If bias 3) is true, our estimated sightability of elk groups on quadrats which was 82% and developed during 199 test trials (Freddy 1995) greatly overestimates the true detection rate achieved during management application of quadrat sampling, even though observers used during testing and application remained the same individuals. The magnitude of discrepancy between estimators would further suggest that we not only missed groups on quadrats, but groups relatively large in size, even though sightability of groups ~7 in size exceeded 0.90 (Freddy 1995). We are currently trying to evaluate the most likely source(s) of bias and then develop procedures to test the appropriate hypothesis during winter 1997-98. Elk Movements In December 1995, 35 male and 34 female calves were radiocollared and of these elk, 24 males and 27 females survived to become IS-months old on 1 December 1996 (Table 7). By the end of winter 1996-97, 21 (7M, 14F) or 41% had dispersed to a winter range outside of the intensive winter range study area in GMU 42 where these elk were initially captured (Table 17). Rates of dispersal were 29% for males and 52% for females. Calves trapped in zones E, F, and G which encompass Alkali, Dry Hollow, and the Mamm creeks, comprised 43% (9) of the dispersing yearlings and .these elk moved primarily west to winter ranges near the towns of Collbran and Parachute-Rulison. Calves trapped in zones C and D which encompass West Divide Creek, also comprised 43% of the dispersing juveniles and these elk moved south and southwest to winter ranges near Paonia Reservoir and Hotchkiss. We completed initial draft maps (1:100,000 scale) of a GIS land database for the area frequented by radiocollared elk as part of a cooperative effort through the National Biological Service, Rocky Mountain Elk Foundation, USFS, and BLM. Layers of GIS information include topography, based on 1:24,000 scale USGS quadrangles, land ownership, hydrology, transportation, transportation management zones, winter trapzones for elk, all elk locations, locations of female elk in June, and locations of all elk mortalities. Maps will be reviewed for accuracy by cooperating agencies. We will acquire vegetation-type coverage for the area and develop a relational database between elk locations and GIS layers to allow descriptive analyses of areas frequented by elk. Sighting Bias Draft Manuscript Discrepancies in estimates of population size based on quadrats, sighting-bias adjusted quadrats, and mark-resight estimators prompted us to re-evaluate potential sources of bias in all of these estimators. We therefore, have delayed drafting and submitting for peer review a manuscript evaluating sighting bias adjusted estimates of population size. �58 CONCLUSIONS We obtained acceptably precise estimates of calf and adult survival rates and recommend monitoring survival of remaining radiocollared adult male and female elk during the next 2 years. We recommend conducting aerial surveys in 199798 to estimate bias in detecting and counting marked elk on sample quadrats and mark-resight surveys to complete our evaluation of techniques to estimate elk density. LITERATURE CITED Freddy, D. J. 1993. Estimating survival rates of elk and developing techniques to estimate population size. Colo. Div. Wildl. Game Res. Rep. July: 83-117. Freddy, D. J. 1994. Estimating survival rates of elk and developing techniques to estimate population size. Colo. Div. Wildl. Game Res. Rep. July: 27-42. Freddy, D. J. 1995. Estimating survival rates of elk and developing techniques to estimate population size. Colo. Div. Wildl. Game Res. Rep. July. Halfpenny, J. C., and E. A. Biesiot. 1986. A field guide to mammal in North America. Johnson Books, Boulder, co. 161pp. SAS Institute Inc. 1988. SAS/STAT Cary, NC. 1028pp. User's Guide, 1982. Procedures Texas Agric. Expt. 6.03. SAS Institute, D. A., and J. E. Browns. livestock and wildlife. White, G. C. 1996. NOREMARK: Population estimation surveys. Wildl. Soc. Bull. 24:50-52. White, G. C., and R. A. Garrott. 1990. Analysis of wildlife data. Academic Press, Inc., San Diego. 383pp. by Inc., for evaluating predation on Sta. Publ. B-1429. 42pp. Wade, Prepared tracking from mark-resighting radio-tracking �in 8 tra~zones. December 1993. 1994. 1995. andj996 Table 11 Elk ca~ture objectives and numbers of elk radiocollared Elk Collaredl Capture Calves Collared Hel ico~ter Corral Years Males Objectives TotallYear Females Years Trap 93 94 95 96 93 94 95 96 Total 93 94 95 96 93 94 95 96 Total 93 94 95 96 93 94 95 96 Zone Name A B C D E F G H Garfield Gibson. Uncle Bob West Divide Hightower Middle Manrn West Manrn Dry Hollow All 8 20 24 26 10 10 8 44 5 8 13 13 10 8 8 21 6 12 11 11 10 9 10 6 8 12 12 10 10 10 8 0 150 86 75 70 8 6 4 25 17 7 29 13 14 21111810 17 17 14 o 13 9 6 0 5 35 6b 0 6 7 13 11 17 o 13 6 0 o 0 9 7 14 8 19 29 21 17 12 4 14 0 9 7 14 10 12 4 14 0 100 67 68 70 141 83 71 70 • Helicopter = Helicopter net-gunning, Corral bAll 6 adult females captured 2 March 1995 4 4 14 18 14 9 5 0 = 0 6 0 0 0 0 0 35 0 10 0 0 0 0 0 6 0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 27 56 70 60 60 26 25 41 41 16 3 0 365 3 5 10 5 4 0 2 7 3 5 7 1 9 8 0 0 1 4 10 8 8 5 1 0 3 4 7 3 7 2 9 0 36 33 37 35 1 8 7 6 3 0 3 9 3 6 6 8 8 5 0 0 3 3 4 10 6 4 4 0 3 7 7 5 2 5 0 23 38 58 48 50 26 24 16 37 36 34 35 283 6 in Game Management Unit 42. Adult Females Collared 93 94 95 96 Total 4 12 12 10 10 0 1 19 0 6 0 2 0 0 0 6 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4 18 12 12 10 0 1 25 68 14 0 0 82 corral-trapping. Table 3. Survival rates of 6-11 month old calves with sexes pooled for the winter-spring time period 1 December - 14 June each year 1993-94 through 1996-97. Survival rates (S) calculated as a mean estimate of (alive)_j_lalive+ dead) and variance S (l-S)In collars. Elk Aqe and Time Period (dates) Calves Calves Calves Calves Calves 12/01/9312/01/9412/01/9512/01/96All Years 06/14/94 _OEiLl4L95 o~Ll~l96__ _ 06/14/97 Pooled Survival L 95% CI U 95% CI n collars Censored' Died Nonhunting Hunting 0.92 0.85 0.98 73 0 6 6 0 0.90 0.83 0.97 69b 0 7 7 0 0.88 0.81 0.96 69 2" 8 8 0 0.86 0.77 0.94 69 1d 10 10 0 0.89 0.85 0.93 280 3"·d 31 31 0 • Censored denotes collar failure andlor animallife/death status not known. • Includes (173.949194) male whose collar failed 12/94 but seen alive 1195, 1/96. Censored elk were (174.619/95) male for slipped collar and (174.800195) male for trap-related mortality. d Censored elk was (174.619/96) male for trap-related mortality. C ~ �60 Table 2. Causes of deaths in radiocollared elk between 1 December 1993 and 14 June 1997. Calves (M=ma1e, F=female) were 6-11 months old and collared at 6 months of age. Yearling males and females were 12-17 months old and juvenile males and females were 18-23 months old and all were collared as 6-month old calves. Adult males and females were> 24-months old at time of death. Elk Age/Sex Class at Death Cause of Death Calves Yearling Juvenile Adult Total and -Code M F M F M F M F M FAll Natural Causes o 2 2 4 o Malnutrition-6 2 2 o o o o Unknown-Suspect 1 4 o 3 Malnutrition-31 o o o 3 1 o o 7 1 5 12 7 4 Predation-Lion-3 o o o o o 1 o 1 o o 1 0 Predation-Bear-35 o o o o 1 1 o o Unknown Predator-5 o o o o 1 o o Unknown-Suspect 2 Predation-30 8 1 1 o 11 3 o 2 5 o 2 2 2 o Accident-Birthing-32 o o o o o 0 o Accident...,Fell-10 o 1 o 1 o 0 1 o o o o 2 2 Unknown Cause-11 4 6 o 2 1 o 1 o o Subtotals 42 19 23 17 14 Q Q Q 1. a ~ Legal Hunting Archery-33 Muzzleloading-34 Archery/Muzzle-27 Rifle-First-28 Rifle-Second-28 Rifle-Third-28 Rifle-Late-29 Subtotals Wounding Loss Archery/Muzzle-24 Rifle-Regular-25 Rifle-Late-26 Subtotals :Illegal Hunting Rifle-Regular-9 Out-of-Season-7 Subtotals Presumed Hunting Disappear-Rifle Seasons-22 Disappear-Archery/ Muzzle-21 Subtotals Totals o o o o o o o 2 2 o o o o 0 0 0 0 0 0 0 o o o o o o o o o o o o o o o o o Q Q Q .1 Q o 1 1 1 o 0 0 0 Q Q a o o o 0 0 9" o Q Q o 0 o o o o o o 8 2 8 4 4 1 o 4 1 o 3 1 13 7 7 4 7 4 13 7 4 7 17 14 7 o 6 o 4 6 Q 39 25 39 29 11 6 68 o o o o o o 1 1 2 2 4 2 6 3 6 o 3 o 9 3 1. Q Q 1 10 .2. 3 11 16 o o o o o 1 0 10 1 o 0 1 10 o o 2 2 2. Q Q 1. 1. 1. 10 ~ 12 o o 4 3 7 0 1 .§. o o 0 o o o o 1 0 1 Q Q 1. 1. Q Q .1 ~ .2. 1 6 2 3 17 14 12 47 45 78 68 12 7 1 146 • These elk were illegally shot as spike-antlered yearling males: (173.190/93), (173.232/93), (173.320193), (173.919/94), (174.059194) (174.140/95), (174.200195), (174.679/95), (174.729/95). • These elk disappeared during rifle bunting seasons and are presumably dead and legally harvested: (172.201193), (172.649193), (172.800/93), (173.309/93), (173.390/93), (173.439193), (174.001/94), (174.181194). One exception may be yearling male (173.309/93) that disappeared during the first rifle season in 1994 when spike-antlered bulls were not .Iegal .. �Table 4. Estimated causes of mortality and associated months old. 1 December 1993 - 14 June 1997. Frequency 1D / Trap Age" Bod Year Captured Zone Sex (months) wt. (kg) 86.0 F 9 C 172.899/93 102.0 F 10 G 173.000/93 108.0 M 9 C 173.262/93 111.0 M 9 C 173.289/93 82. O· 8 H M 173.461/93 77.0 M 10 H 173.469/93 117.0 12 F B 173.589/94 9 89.0 F C 173.640/94 100.0 F 9 E 173.789/94 86.0 F 8 D 173.870/94 140.0 M 9 F 174.119/94 .112.0 M 9 F 174.140/94 10 140.0 M B 174.170/94 127.0 M 10 F 174.220/95 120.0 M 10 F 174.230/95 115.0 M 10 E 174.240/95 79.0 E F 9 174.339/95 10 78.0 F C 174.500/95 10 120.0 B F 174.609/95 8 100.0 D M 174.689/95 11 M C 174.789/95 M 7 72.0 E 173.370/96 M 8 116.0 E 173.381/96 102.0 A F 9 173.640/96 F 10 89.5 A 174.520/96 11 M 103.0 174·.689/96 G E M 10 125.0 174.800/96 D F 8 105.5 174.910/96 91. 5 C F 8 175.059/96 A F 9 111. 0 175.130/96 M 77.5. 175.],.70/96 D 9 body condition Date Dead 03/18/94 04/25/94 03/18/94 03/22/94 02/07/94 04/25/94 06/01/95 03/30/95 03/20/95 02/21/95 03/01/95 03/14/95 04/25/95 . 04/08/96 04/24/96 04/17/96 03/27/96 04/16/96 04/15/96 02/05/96 OS/22/96 01/08/97 02/19/97 03/16/97 04/25/97 05/14/97 04/09/97 02/27/97 02/13/97 03/24/97 03/26/97 for 31 radiocollared Marrow % Fat 28.50d 8.03° 28.88d N/Af 0.21° 0.27° 51. 71° N/Af 14.71° N/Af 94.97° 64.44° 76.05° 41.08° 2.47° 86.28° 62.97° 34.56° 10.28° 91.90° 75.32· 79.89° 69.36° 30.50° 8.89° None' 51.89° 83.84° N/Af 8.03° 1.98° calf elk 6-11 Cause of Death & Code No. Lion Predation-3 Malnutrition-6 Unknown-11 Unknown-11 Malnutrition-6 Malnutrition-6 Unknown; suspect predation-30 Unknown; suspect predation-30 Unknown; suspect malnutrition-31 Lion Predation-3 Bear Predation-35 Lion Predation-3 Lion Predation-3 Unknown; suspect malnutrition-31 Lion Predation-3 Lion Predation-3 Unknown Predator-5 Unknown; suspect predation-30 Unknown; suspect predation-30 Lion Predation-3 Unknown; suspect predation-30 Lion Predation-3 Unknown; suspect predation-30 Malnutrition-6 Lion Predation-3 Unknown; suspect malnutrition-31 Lion Predation-3 Unknown; suspect predation-30 Unknown-11 Lion Predation-3 Unknown; suspect malnutrition-31 • Approximate age at death assuming 15 June birthdate. • Whole body weight at capture in December of 1993, 1994, 1995, or 1996. e Fat content as percent dry matter of bone marrow from either femur of carcass. • Fat content as percent dry matter of bone marrow from either mandible of carcass. • Fat content as percent dry matter of bone marrow from either humerus of carcass. r No suitable bones remaining with carcass. I No marrow observed in one long bone remaining with carcass. ~ �~ Table 5. Survival rates for winter-spring and summer-fall time periods from 1 December 1993-14 June 1997 for the cohort of 6-month old calves radiocollared in December 1993. Survival rates for male and female calves pooled when 6-11 months old were 0.92 (95% CI 0.86-0.98, n=73). Survival rates (S) calculated as a mean estimate of (alive)/~alive + dead) and variance S(1-S)/n collars. Elk Aqe (months) and Time Period (dates) 6 - 11 12 - 17 18 - 23 24 - 29 30 - 35 36 - 41 42 - 47 6 - 47 12/01/9306/15/9412/01/9406/15/9512/01/9506/15/9612/01/9612/01/9306/14/94 11/30194 06/14/95 11/30/95 06/14/96 11/30/96 06/14/97 06/14197 MALES Survival L 95% CI U 95% CI n collars Censored" Died Nonhunting Hunting 0.89 0.78 0.99 36 0 4 4 0 0.88 0.76 0.99 32 0 4b 0 4 FEMALES Survival L 95% CI U 95% CI n collars Censored" Died Nonhunting Hunting 0.95 0.87 1.00 37 0 2 2 0 0.97 0.92 1.00 35 0 1f 0 1 1.00 28 0 0 0 0 1.00 34 0 0 0 0 0.21 0.06 0.37 28c 0 22d 0 22 1.00 0.00 0.00 4 2· 0 0 0 0.50 0.00 1.00 4 0 2 0 2 0.97 0.91 1.00 34 0 1 0 1 0.97 0.91 1.00 33 0 1 0 1 0.84 0.71 0.97 32 0 5 0 5 • Censored denotes collar failure andlor animallifeldeath status not known. • Collar (173.309/93) "disappeared" during Fall 1994 hunting seasons and assumed dead. • Includes collar (173.241/93) that failed in August 1995 but bull seen alive Ianuary 1996. 4 Two collars (173.390193), (173.439/93) "disappeared" during Fall 1995 hunting seasons and assumed dead. • Two collars censored; (173.241193) failed, (173.381/93) slipped off 12/01195. r Collar (172.800/93) "disappeared" during Fall 1994 hunting seasons and assumed dead. I Includes (173.340/93) alive 5/17/97. 1.00 2g 0 0 0 0 1.00 27 0 0 0 0 0.06 0.00 0.14 34 2· 32 4 28 0.73 0.58 0.88 37 0 10 2 8 �Table 6. Survival 'rates for winter-spring and summer-fall time periods from 1 December 1994-14 June .1997 for the cohort of 6-month old calves radiocollared'in December 1994. Survival rates for male and female calves pooled when 6-11 months old were 0.90 (95% CI 0.83-0.97, n=69). Survival rates (S) calculated as a mean estimate of (alive) / (alive _+ dead>_and variance S (l-SLLn collars. Elk Aqe_(months)_ and Time Period (dates) 6 - 11 12 - 17 18 - 23 24 - 29 30 - 35 6 - 35 12/01/9406/15/9512/01/9506/15/9612/01/9612/01/9406/14/95 1l/_3QL95 06/14/96 11/30/96 06/14/97 06/14/97 MALES Survival L 95% CI U 95% CI n collars Censored" Died Nonhunting Hunting 0.91 0.81 1. 00 33b 0 3 3 0 0.90 0.79 1. 00 30b 0 3 0 3 0.96 0.88 1.00 26, 1c 1 1 0 0.08 0.00 0.19 25 0 23 0 23 2e 0 0 0 0 0.06 0.00 0.15 32 1c 30 4 26 FEMALES Survival L 95% CI U 95% CI n collars Censored" Died 'Nonhunting Hunting 0.89 0.78 0.99 36 0 4 4 0 0.97 0.91 1. 00 32 0 1 0 1 0.97 0.90 1. 00 30 1d 1 1 0 0.93 0.83 1. 00 29 0 2 0 2 0.96 0.89 1. 00 27 0 1 0 1 0.74 0.59 0.89 35 1d 9 5 4 • ~ • • • 1. 00 Censored denotes collar failure and/or animallife/death Status not known. Includes collar (173.949/94) that failed in December 1994 but male seen alive in January 1996 Censored elk was (173.949/94) failure, which was killed in first rifle season 1996, Censored elk was (173.719/94) slipped collar, Includes (174.030/94) alive 6/4/97. ~ �~ Table 7. Survival rates for winter-spring and summer-fall time periods from 1 December 1995-14 June 1997 for the cohort of 6-mbnth old calves radiocollared in December 1995 and survival rates for winter-spring for 6-11 month old elk calves radiocollared in December 1996. Survival rates for male and female calves pooled when 6-11 months old were 0.88 in 1995 (95% CI 0.81-0.96, n=69) and 0.86 in 1996 (95% CI 0.77-0.94, n=69). Survival rates (S) calculated as a mean estimate of (alive)/(alive+dead) and variance of S(1-S)/n collars. Elk Aqe (months) and Time Period (dates) 1995 Calf Cohort 1996 Calf Cohort 6 - 11 12 - 17 18 - 23 6 - 23 6 - 11 12/01/9506/15/9612/01/9612/01/9512/01/9606/14/96 _1_1130/96 06/14/97 06/14/97 06/14/97 MALES Survival L 95% CI U 95% CI n collars censored' Died Nonhunting Hunting 0.86 0.74 0.98 35 2b 5 5 0 0.83 '0.68 0.97 29 FEMALES Survival L 95% CI U 95% CI n collars Censored· Died Nonhunting Hunting 0.91 0.81 1.00 34 0 3 3 0 0.87 0.75 0.99 31 0 4 0 4 • Censored • Censored • Censored • Censored Ie 5 0 5 0.96 0.87 1.00 24 0 1 1 0 0.68 0.51 0.84 34 3b.e 0.85 0.73 0.98 34 11 6 5 5 5 0 0.9.3 0.82 1.00 27 0 0.74 0.58 0.89 34 0 9 4 5 0.86 0.98 0.74 35 0 5 5 0 2 1 1 denotes collar failure andlor animallife/death status not known. elk were (174.619195) slipped collar, (174.800195) capture mortality . elk was (174.660195) slipped collar July 1997. elk was (174.619/96) male trap-related mortality, Id �Table 8. Survival rates for winter-spring and summer-fall time periods from 1 December 1993 - 14 June 1997 for all radiocollared adult female elk ~12-months old. Survival rates (S) calculated as a mean estimate of (alive)/(alive + dead) and variance S(l-S)/n collars. Elk Aqe and Time Period (dates) Adult Adult Adult Adult Adult Adult Adult 12/01/9306/15/9412/01/9406/15/9512/01/9506/15/9612/01/9606/14/94 11/30/94 06/14/95 11/30/95 06/14/96 11/30/96 06/14/97 Survival 95t CI U 95t CI n collars Censored' Died Nonhunting Hunting L 0.87 0.80 0.94 100bf . 0 13c 1 12 0.96 0.91 1.00 68be 0 3 1 2 0.94 0.90 0.98 129bh 0 8d 0 8 0.96 0.92 1.00 95bg 0 4 1 3 • Censored denotes collar failure and/or animal life/death status unknown. c Collars (172.800/93,172.649/93) "disappeared" Fall 1994 hunting seasons,assumed dead. • Composition is 6 females 18+ and 62 females 30+ months old. • Composed of 34·18+, and 61-30+ months old females. I Composed of 30-18+ and 89-30+ months old females. k Composed of 31-12+.29-24+, and 83-36+ months old females. 0.94 0.90 0.98 119i· 21 7 4 3 0.89 0.84 0.94 143k 0 16 1 15 0.98 0.95 1.00 1271 0 3 1 2 Includes collar (172.011/93) that failed 4/1994 but seen alive in 1/1996 Collar (172.207/93) "disappeared" Fall 1995 hunting seasons, assumed dead. r Composed of 35-12+,6-24+, and 59-36+ months old females. h Composed of 32-12+,34-24+, and 63-36+ months old females. J Censored (172.011/93) failure and (173.719/94) slipped collar. I Composed of 27-18 + and 100-30+ month old females. b d Table 9. Survival rates for winter-spring and summer-fall time periods from 1 December 1993 - 14 June 1997 for the group of adult female elk that we~e ~12-months old when radiocollared in December 1993. Survival rates (S) calculated as a mean estimate of (alive)/(alive + dead) and variance S(l-S)/n collars. Elk Aqe and Time Period (dates) Adult Adult Adult .Adult Adult Adult Adult 12/01/93- 06/15/94- 12/01/94- 06/15/95- 12/01/95- 06/15/96- 12/01/9606/14/94 uU_L30nL 06/14/95 11/30/95 06/14/96 l_lj_3l>/9606/14/97 0.96 0.82 0.93 0.88 0.93 0;92 1.00 Survival 0.91 0.72 0.85 0.78 0.85 0.84 L 95t CI 1.00 0.91 0.99 0.97 1.00 1.00 U 95t CI 68be 65bf 53bf 49bf 42f 39f 36f n collars o 0 0 0 l' 0 o Censored' 3 12c 4 6d 3 3 o Died 1 1 103 0 o Nonhunting 2 11 3 6 0 3 Hunting o n __ • . Censored denotes collar failure and/or animal life/death status unknown c Collar (172.649/93) "disappeared" Fall 1994 hunting seasons, assumed dead. • Composed of 6-18+ and 62-30+ month old females. • Censored (172.011/93) failure of unknown status spring 1996. Includes collar (172.011/93) failed 4/1994 but seen alive 1/1996. Collar (172.207/93) "disappeared" Fall 1995 hunting seasons, assumed dead. r Composed of 100% females 24+ months old. b d ffi �Table 10. Annual survival rates from 1 December 1993-30 November 1996 for the group of adult female elk that were ~12-months old when radiocollared in December 1993. Survival rates (S) calculated as a mean estimate of (alive)/(alive+dead) and variance S(l-S)/n collars. Survival L 95% CI U 95% CI n collars Censored' Died Nonhunting Hunting • • • , Adult 12/01/93~11/30/94 0.78 0.68 0.88 68b,f o c Elk.Aae_and Time Periodldatesl Adult Adult 12/01/94 ••. J.1L30j9_5 __ 12J~l.j95-11/30/96 Adult 12/01/93-06/14/97 0.84 0.75 0.93 53b,f 0.86 0.75 0.97 42f 0.54 0.42 0.66 67f 0 1q 1q 31 d 15 10 6 2 1 3 6 13 9 3 25 Censored denotes collar failure and/or animallif~death status unknown. Collar (172.649/93) "disappeared" Fall 1994 hunting seasons, assumed dead. Composed of 6-18+ and 62-30+ month old females. Censored (172.011/93) failure 4/1996. b d I Includes collar (172.011/93) failed 4/1994 hut seen alive 1/1996. Collar (172.207/93) "disappeared" Fall 1995 hunting seasons, assumed dead. Composed of 100% females 24+ months old. Table 13. Data matrix for simple population model using spreadsheet software. We ~sed a post-hunt (December) population composition of SO calves:100 females (age ~ 12-months) and 4 adult males:16 yearling: 100 females and an estimated population of 3,600 elk. Annual population growth rate is about 7% when using this matrix. Survival Rate Fall Hunting Survival Rate Elk Sex/Age Class Summer-Fall Harvest Rate Elk Sex/Age Class Winter-Spring Adult Males 1.00 0.82 Adult Males 1.00 Age 24-29 months Age ~30 months Yearling Males Age 12-17 months 1.00 0.13 Yearling Males Age 18-23 months 0.97 Adult Females Age ~24 months 0.99 0.10 Adult Females Age ~30 months 0.98 Yearling Females Age 12-17 months 1.00 0.06 Yearling Females Age 18-23 months 0.97 Calves Age 0-5 months 1.00 0.02' .Calves Age 6-11 months 0.89 'Estimated from harvest surveys. s �Table 11. Management Frequency distribution of whole body weights for male and female elk calves trapped unit 42, December, 1993"';1996. Percentage 2er weight class shown in 2arentheses. Bod~ Weight Class {kg) Year Sex 60-§9 70-79 1993 1994 1995 1996 M M M M M 0(0.0) 0(0.0) 0(0.0) 0(0.0) 0(0.0) 1 (2.9) 1(3.0) 0(0.0) 3(8.6) 5(3.6) F F F F F 0(0.0) 0(0.0) 1(3.1) 0(0.0) 1(0.7) 1 (2.9) 2(5.7) 2(6.3) 0(0.0) 5 (3.7) ALL 1993 1994 1995 1996 ALL Table 12. Bod~ measurements 80-89 1( 1( O( O( 2( 90-99 100-109 6(17.1) 2.9) 3( 8.6) 2 ( 6.0) 6(18.2) 3.0) 3( 8.6) 0.0) 4(11.4) 6(17.1) 0.0) 3 ( 8.6) 1. 4) 12( 8.7) 21(15.2) 8(22.9) 4(11.4) 7(20.0) 4(11.4) 1 ( 3.1) 3 ( 9.4) 1 ( 2.9) 6(17.6) 10( 7.4) 24(17.6) 12(34.3) 10(28.6) 14(43.8) 8(23.5) 44(32.4) 110-119 120-129 130-139 12(34.3) 13(39.4) 8(22.9) 5(14.3) 38(27.50 10(28.6) 6(18.2) 9(25.7) 13(37.1) 38 (27.5) 1 ( 2.9) 2 ( 6.0) 10(28.6) 5(14.3) 18(13.0) 4(11.4) 10(28.6) 5(15.6) 10(29.4) 29(21.3) 6(17.1) 2 ( 5.7) 6(18.8) 8(23.5) 22(16.1) o( 0.0) 0.0) 0.0) 1 ( 2.9) 1 ( 0.7) o( o( in Game Total 140-149 1 (2.9) 2 (6.0) 1 (2.9) 0(0.0) 4(2.9) 35 (100) 33(100) 35(100) 35(100) 138 (100) 0(0.0) 0(0.0) 0(0.0) 0(0.0) 0(0.0) 35(100) 35(iOO) 32(100) 34(100) 136 (100) for elk calves tra22ed in Game Management Unit 42, December, 1993-1996. Male Calves Female Calves Mean SD Min Max n Mean SD Min Max n Measurement 112.7 Body Weight (kg) 191.2 Body Length, (em) Hindfoot Length (em) 56.3 0.59 Condition Index· 13.7 10.2 2.2 0.05 77.0 164.0 52.0 0.43 141. 0 210.0 61. 0 0.67 35 36 36 35 103.8 188.9 54.8 0.55 12.2 9.8 2.1 0.05 ·76.0 167.0 51. 0 0.46 1994 113.5 Body Weight (kg) 190.3 Body Length (em) Hindfoot Length (em) 56.9 0.59 Condition Index 14.2 9.2 1.8 0.05 71.0 168.0 50.0 0.42 140.0 204.0 59.0 0.69 33 32 32 32 103.0 186.3 55.1 0.55 13.3 8.8 2.1 0.06 70.0 128.0 166.0' 207.0 50.0 59.0 0.42 0.65 1995 119.1 Body Weight (kg) 194.0 Body Length (em) Hindfoot Length (em) 57.4 0.61 Condition Index 13.6 9.3 2.2 0.05 92.0 166.0 53.0 0.50 141. 0 206.0 61.0 0.71 35 36 36 34 105.7 189.4 54.9 0.56 15.2 11.4 2.3 0.06 60.0 158.0 47.0 0.38 129.0 .32 203.0 34 59.0 34 0.66 32 1996 114.3 Body Weight (kg) Body Length (em) 187.6 Hindfoot Length (em) 57.7 0.61 Condition Index 17.1 11.5 2.3 0.07 72.0 160.0 52.0 0.44 136.0 202.5 60.5 0.70 35 35 34 35 109.5 187.4 56.8 0.58 12.4 9.0 2.4 0.05 81. 5 170.5 52.5 0.47 136.0 210.0 62.0 0.69 34 35 34 34 '114.9 All· Body Weight (kg) 190.8 Years Body Length (em) Hindfoot Length (em) 57.1 0.60 Condition Index 14.8 10.3 2.2 0.06 71.0 160.0 50.0 0.42 141. 0 210.0 61. 0 0.71 138 139 138 136 105.5 188.0 55.4 0.56 13.4 9.8 2.3 0.06 60.0 158.0 47.0 0.38 136.0 210.0 62.0 0.69 136 140 138 135 Year 1993 • Condition index = (Weight/body length) . 35 123.0 207.0 35 59.0 34 0.64' 34 35 36 36 35 CJ) ...., �20 Table 14. Colorado. ~ Counts of elk on quadrats 11 and 12 February, 1997 in GMU 42 south of Rifle and Newcastle, Strata Size (mi2) Strata Garfield High Garfield Low E. Divide Low W. Div. East High W. Div. East Low W. Div. West High W. Div. West Low Hightower Low upper Alkali Ck. High Upper Alkali Ck. Low Low Alkal i Ck. Low upper DryHollow Low Lower Mamm Ck. Low W.Mamm-Grass Mesa Low W.Mamm-Grass Mesa High Totals Quadrats N n 10 17 16 10 17 16 10 6 6 11 11 5 5 6 4 4 9 5 9 9 4 5 10 10 13 10 10 13 5 7 5 7 137 137 9 4 Elk Counted 191 54 Elk/Quadrat Mean StdErr. 5 3 7 48 85 .58 86 14 244 19.10 6.75 2.67 34.17 12.25 22.25 9.00 0.00 22.00 16.00 17.00 14.50 17.20 4.67 34.86 72 1246 13.54 8 3 3 3 4 3 5 4 8 205 49 89 27 o 88 0.000 1.859 2.404 0.000 3.573 2.991 2.449 0.000 0.000 5.514 4.738 3.309 2.423 1.520 0.000 Population Size Total + 90% CI o 191 115 43 205 135 111 81 52 63 o 65 25 36 o o o 88 80 170 145 224· 23 45 78 54 52 13 244 o 1854 164 Table 15. Summary of helicopter flights used to estimate elk population size and density on 137 mi2 of winter ranqe .inGMU_42 south of Rifle and Newcastle, Colorado, during February, 1997. Proportion Unmarked Total Elk Flight Marked Elk Marked Elk Elk Proportion TYPe Dates Hours ._Available" Counted ID'db Counted Counted Counted Marked NonRandom1 2/10 Quadrats 2/11-12 NonRandom2 2/13 7.5 29.0 7.8 120 120 120 31 43 .33 29 40 29 0.2583 0.3583 0.2750 838 1203 1069 869 1246 1102 0.0357 0.0345 0.0299 ~umber of marked elk within the 132 mi2 sample area. "Number of marked elk individually identified during flights based on number/symbol identifier seen on collar. �Table 16.· Summary of flights Newcastle, Colorado. Method Estimator Flight Date to estimate elk population Area (mi2) FlownLHrs· size from 1994-1997 Minimum Elk CountedLFlight Population Estimate in GMU 42 south of Rifle Approximate 95% Conf. Int. and 95% C.I. %Precision Feb. 1997 Feb. 1997 Feb. 1997 Feb. 1997 Feb ..1997 ·All 1997 Quadrats-StRSb Quadrats-StRS~SBAc Quadrats-StRS-Lpd NonRandom Flight,REP1-LP NonRandom Flight,REP2-LP 3 Flights Pooled-JHEe 137/29.0 137/29.0 137/29.0 137/ 7.5 137/ 7.8 137/44.3 1246 1246 1246 869 1102 1246 1854 2105 3428 3289 3924 3610 1660-2048 1851-2359 2643-4213 2344-4233 2839-5010 3116-4251 11 12 23 29 28 18 Jan. Jan. Jan. Mar. Mar. Mar. Jan. Mar. All All Quadrats,REP1-StRS Quadrats,REP1-StRS-SBA Quadrats,REP1-StRS-LP Quadrats,REP2-StRS Quadrats,REP2-StRS-SBA Quadrats,REP2-StRS-LP NonRandom Flight,REP1-LP NonRandom Flight,REP2-LP 4 Flights Pooled-JHE 4 Flights Pooled-IEJHEe 132/19.6 132/19.6 132/19.6 132/19.1 132/19.1 132/19.1 132/ 8.5 132/ 8.0 132/55.2 132/55.2 885 885 885 1373 1373 1373 1354 1998 1998 1998 2165 2336 3463 3175 3387 3791 3177 3405 3472 3415 1265-3065 1294-3378 2477-4448 2098-4252 2189-4585 2975-4607 2560-3795 2941-3869 3168-3840 3129-3807. 42 45 28 34 3S 22 19 14 11 12 Quadrats-StRS Quadrats-StRS-SBA Quadrats-StRS-LP 177/17.5 177/17.5 177/17.5 361 361 361 1484 1698 3774 787-2181 928-2468 1993-5555 47 45 47 1996 1996 1996 1996 1996 1996 1996 1996 1996 1996 Feb. 1995· Feb. 1995 Feb. 1995 "Hours of helicopter time per flight. 'Stratified randoin sample of quadrats with counts of elk not adjusted for sighting bias. 'Stratified random sample of quadrats with counts of elk adjusted for sighting bias. "One sample Lincoln-Peterson mark-resight estimator. ·loint Hypergeometric mark-resight estimator for nonrandom and quadrat(unadjusted) flights pooled. 'lmmigrationlEmigration loint Hypergeometric mark-resight estimator that allows for movement of elk on/off intensive study area. iB �70 Table 17. Radiocollared elk captured as calves on winter range in GMU 42 during December 1995 that dispersed out of GMU 42 to a new winter range as of March 1997. Locations (GMUs) determined by aerial telemetry. Trap New Fregyency Age· Sex Zone Location DescriQtion GMU 173.899/95 F 21 F 42 West Battlement Creek 173.909/95LE F 21 F 421 Hawxhurst Creek 174.301/95 E 21 F 42 West Porucpine Creek 174.360/95 E 21 F Wallace Creek 42 West 174.370/95 21 E F 521 Drift Creek 174.391/95 D 21 F Paonia Reservoir 521 174.431/95LE D 21 F 521 Buck Mesa 174.451/95 D 21 F 43 Thompson Creek 174.491/95 C 21 F 42 West Holms Mesa 174.532/95 B 21 F 43 Thompson Creek 174.551/95 G 21 F 42 West Porcupine Creek 174.580/95 G 21 42 West Porcupine Creek F 174.589/95LE G 21 421 Hawxhurst Creek F 174.600/95 A 21 F 42 West Holms Mesa 174.119/95 21 B M 42 West Holms Mesa 174.629/95LE E 21 M 42 West Rulison 174.719/95 21 D M 521 Buck Mesa 174.770/95 C 21 421 Grassey Gulch M 174.780/95 C 21 °53 Anthracite Creek M 174.809/95 C 21 521 M Sommerset 174.851/95 21 C M 52 Oak Mesa • Age of elk in months as of March 1997. b LEO= Location elk routinely located. �71 30 25 0 w 0 !r w CD ~ 20 Z 10 ............................................................................................................ ··················································i; n = 61 !:g::::::::1 ADULT MALES _ ADULTFEMALES n=54 15 :J ......................................................................•.............................•....... ································1 5 0 . ~~ «.</}J #" ~Q;- .I ~~ ~0 ~'v MONlH Rg. 1. Timing of deaths for adlit elk >=12-months of age, 1993-94 - 1996-97. 12 10 0 w 0 !r w CD CALVES _ PREDATION E:m!::m!1 _ MALNUTRITION OTHER 8 6 ~ :J z 4 2 0 JAN FEB MAR MAY APR MONlH Rg.2. liming and estimated causes of deaths for calf elk 1993-94 -1996-97. months of age. 7 6 0 w 0 !r W CD ~ :J 5 4 - JUN Calves were of both sexes and 6-11 MALES n= 17 3 Z 2 1 0 JAN FEB MAR APR MAY JUN MONlH Rg.3. liming of deaths for male and female calf elk 1993-94 -1996-97. Calves were 6-11 months of age. �72 Appendix I. Mortalities approximate age in years Frequency 10/ Trap Year Captured Site Zone 172.030/93 BR A 172.039/93 GR B 172.070/93 BR A 172.080/93 GM A 172.090/93 GR B 172.101/93 SR B 172.128/93 GC B 172.139/93 BC C 172.160/93 CC C 172.181/93 MC C 172.201/93 GS C 172.207/93 SG D 172.258/93 HY C 172.277/93 GS C 172.290/93 SG' 0 172.290/94 SM 0 172.369/93 AC E 172.369/94 FM B 172.409/93 AC E 172.459/93 MH E 172.509/93 FS H 172.542/93 FS H 172.549/93 PG H 172.570/93 PG H 172.581/93 PG H 172.581/94 FM B 172.590/93 PG H 172.610/93 PG H 172.639/93 PG H 172.649/93 PG H 172.670/93 PG H 172.678/93 PG H 172.690/93 FM B 172.699/93 FM B 172.749/95 LS H 172.800/93 SR B 1,72.821/93 EG B 172.890/93 BC C 172.899/93 BC C 172.950/93 GH 0 173.000/93 WM G 173.041/93 GM A 173.060/93 MH E 173.081/93 FS H 173.120/93 GM A 173.140/93 PG H 173.160/93 FS H 173.190/93 BC C 173.201/93 GR B 173.210/93 GC B 173.219/93 BC C 173.232/93 BR AMY 173.262/93 BC C 173.262/94 HM B 173.269/93 MC C 173.279/93 MC C 173.289/93 MC C 173.289/94 PR A 173.300/93 GS C 173.309/93 HY C 173.320/93 HY C 173.320/94 HM B 173~332!93 GH 0 173.351193 SG 0 173.359/93 AC E 173.370/93 MH E 173.370/96 RG E, 173.381/96 RG E 173.390/93 MG 0 173.402/93 MG 0 173.410/93 WM G f73.4Z0/93 WM If of 146 radiocollared elk from 1 December 1993 through 14 June 1997. Age is of elk at death: C=calf 6-11 months old. Y=yearling 12-23 months old. Death Cause of Death & Code Nl.IIIber ' Sex Age Date Legal kill rifle season-28 F 5 27-0ct-95 Unknown-suspect predation-30 F 4 14-Jun-95 Unknown-11 F 11 12-Feb-96 Legal kill rifle season-28 F 6 05-Nov-94 Wounding loss late rifle season-26 F 11 16-Jan-94 Legal kill rifle season-28 F 8' 14-OCt-96 Wounding toss rifle season-25 F 8 31-0ct-96 Wounding loss rifle season-25 F 17 19-0ct-95 F 3 23-0ct-94 Legal kill rifle season-28 Wounding loss rifle season-25 F 3 03-Nov-94 F 3 15-Nov-94 Wounding loss rifle season-25 F 4 30-Nov-95 Disappear rifle season-22 , F 3 04-0ct-94 Legal kill archery/muzzle season-27 F 3 04-OCt-94 Wounding loss archery/muzzle-24 F 6 23-0ct-94 Legal kill rifle season-28 F 4 16-Sep-96 Legal kill mazzleloading season-34 F 3 18-Sep-94 Legal kill archery season-33 F 4 14-Jan-96 Legal kill late rifle season-29 F 8 29-Dec-94 Wounding loss late rifle season-26 Legal kill rifle season-28 F 8 24-0ct-96 F 3 21-0ct-95 Legal kill rifle season-28 F 6 01-Feb-94 Mountain lion predation-3 Legal kill late rifle season-29 F 7 28-Nov-95 'F 16 03-Nov-94 Wounding loss rifle season-25 Legal kill rifle season-28 F 3 17-OCt-94 Legal kilI late r.i fle season-29 F 3 08-Dec-95 F 5 24-Apr-96 Unlenown-11 Accident, birthing/calving-32 F 3 04-0ct-94 Accident, birthing/calving-32 F 16 14-Jun-96 F 2 30,Nov-94 Disappear rifle season-22 Legal leilllate rifle season-29 F 4 16-Jan-95 Wounding loss late'rifle season-26 F 9 22-Dec-94 IlLegal leill-7 F 8 24-Jan-94 Legal leiLlrifLe season-28 F 7 05-Nov-95 Unknown-suspect predation-30 F 11 07-Aug-96 Disappear rifle season-22 F Y 30-Nov-94 Legal leillarchery season-33 F 3 08-Sep-96 F 3 06-Nov-96 Legal kill rifle season-28 Mountain lion predation-3 F C 18-Mar-94 Legal kill rifle season-28 F 2 18-0ct-95 Malnutrition-6 F C 25-Apr-94 Legal leillrifle season-28 M 2 19-0ct-95 Legal leiII late rifle season-29 F 2 27-Dec-95 Legal leillrifle season-28 F 3 22-0ct-96 Legal leiLLrifle season-28 M 2 14-0ct-95 F 3 31-0ct-96 Wounding loss rifle season-25 Legal leillrifle season-28 F 3 13-Nov-96 Illegal leillrifle season-9 M Y 29-0ct-94 Wounding loss rifle season-25 M 2 30-Nov-95 Legal leillrifle season-28 M 2 19-0ct-95 Legal kill rifle season-28 M 2 06-Nov-95 ILLegaL leilLrifle season-9 15-Nov-94 Unknown-11 M C 18-Mar-94 Legal leiII rifle season-28 M 2 12-0ct-96 Legal k i ll rifle season-28 M 2 06-Nov-95 Legal kill rifle season-28 M 3 12-OCt-96 Unknown-11 M C 22-Mar-94 Legal leillmuzzleloading season-34 M 2 18-Sep-96 Legal leillrifLe season-28 M 2 24-0ct-95 Disappear rifLe season-22 M Y 30-Nov-94 Illegal leillrifle season-v M Y 20-0ct-94 Legal leillarchery season-33 M 2 14-Sep-96 Legal leillmuzzleloading season-34 M 2 12-Sep-95 Legal leillrifle season-28 M 2 05-Nov-95 Legal leillarchery season-33 M 2 06-Sep-95 Legal leillrifLe season-28 M 2 08-Nov-95 Mountain lion predation-3 M' C 08-Jan-97 Unknown-suspect predation-30 M C 19-Feb~97 Disappear rifLe season"22 M 2 30-Nov~95 'Legal leillrifle season-28 M 2 18-0ct-95 Wounding loss rifle season-25 M 2 02-Nov-95 Legal leillrifle season-28 M 2 24-0ct-95 ------------------------~--------------------------------------------------------~--~--------------- �73 Appendix I. (continued) Frequency ID/ Tra~ Year Ca~tured Site Zone 173.429/93 MH E 173.439/93 PG H 173.450/93 PG H 173.461/93 H PG 173.461/94 eM A 173.469/93 FS H 173.469/94 PR A 173.479/93 F.S H 173.492193 H FS 173.502193 FS H 173.510/93 FM B 173.521/93 FM B 173.540/94 OG B 173.549/94 HM B. 173.580/94 PR A 173.589/94 OG B 173.640/94 Me c 173.640/96 GM A 173.689/94 BG E 173.789/94 E MR 173.829/94 F MM 173.859/94 SM D 173.870/94 SM D 173.919/94 e Be 173.929/94 C Be 173.939/94 BC c 173.959/94 Be c 173.970/94 C MC 173.981194 MP D 173.990/94 C BC 174.001194 BG E 174.009/94 BG E 174.019/94 BG E 174.039/94 MR E 174.049/94 MM F 174.059/94 MR E 174.069/94 BG E 174.090/94 MR E 174.101194 F MM 174.109/94 MM F 174.119/94 F MM 174.129/94 F MM 174.140/94 F MM 174.140/95 GB B 174.150/94 MM F 174.160/94 MM F 174.170/94 FM B 174.181/94 FM B 174.200/95 MS F 174.220/95 MS F ;74.230/95 F MS 174.240/95 MD E 174.319/95 F MS 174.329/95 E MD 174.339/95 MD E 174.360/95 MD E 174.401/95 AP D 174.500/95 LB c 174.520/95 KR B 174.520/96 GM A 174.560/95 BL A 174.609/95 GB B 174.679/95 HG E 174.689/95 AP D 174.689/96 EW G 174.729/95 VM D 174.789/95 \II) C 174.800/96 BG E 174.809/95 \II) c 174.861195 B KR ,174.910/96 LM D 175.059/96 Be e 175.130/96 A GM 175.170/96 LM D Sex M M M M M M M M M M M M M F F F F F F F F F F M M M M M M M M M M M M M M M M M M M M M M M M M M M M M F F F F F F F F F F M M M M M M M M F F ·F M Ase z 2 2 .C Y C 2 2 2 3 2 2 2 Y Y C c e 2 ,C 2 2 e y 2 2 2 2 2 2 2 2 2 2 2 Y 2 2 Y 2 e 2 c Y 2 2 c 2 Y c c c Y Y C Y 2 c Y c Y C Y e e Y e e Y Y e e C e Death Date '4-Sep-9~ 30-Nov-95 15-0ct-95 07-Feb-94 19-5ep-95 25-Apr-94 20-Oct-96 10-Sep-95 03-Sep-95 13-Nov-96 28-Aug-95 17-0ct-95 13-0ct-96 07-Sep-95 21-Dec-95 01-Jun-95 30-Mar-95 16-Mar-97 31-0ct-96 20-Mar-95 10-Jan-97 23-Oct-96 . 21-Feb-95 02-Nov-95 16-Oct-96 18-Sep-96 02-Nov-96 20-Oct-96 02-Nov-96 20-Oct-96 30-Nov-96 12-0ct-96 13-Nov-96 05-Sep-96 14-Sep-96 17-0ct-95 26-Sep-96 20-Oct-96 24-May-96 19-Oct-96 01-Mar-95 14-oCt-96 14-Mar-95 30-Nov-96 14-0ct-96 15-Sep-96 25-Apr-95 30-Nov-96 18-0ct-96 08-Apr-96 24-Apr-96 17-Apr-96 02-Oct-96 03-Sep-96 27-Mar-96 22-Apr-97 22-Sep-96 16-Apr-96 21-Sep-96 25-Apr-97 14-May-97 15-Apr-96 13-Nov-96 05-Feb-96 14-May-97 17-0ct-96 22-May-96 09-Apr-97 22-Apr-97 31-Oct-96 27-Feb-97 --13-Feb~97 24-Mar-97 26-Mar-97 Cause of Death & Code Number Legai klii archery season:33 Disappear rifle season-22 Legal kill rifle season-28 Malnutrition-6 Wounding loss archery/muzzle-24 Malnutrition-6 Legal kill rifle season-28 Legal kill archery season-33 Legal kill archery season-33 Legal kill rifle season-28 Legal kill archery season~33 Legal kill rifle season-28 Legal kill rifle season-28 Legal kill archery season-33 Unknown-11 Unknown-suspect predation-30 Unknown-suspect predation-30 Malnutrition-6 Legal kill rifle season-28 Unknown-suspect malnutrition-31 legal kill late rifle season-29 Legal kill rifle season-28 Mountain lion predation-3 .Illegal kill rifle season-9 Legal kill rifle season-28 Legal kill archery season-33 Legal kill rifle season-28 Legal kill rifle season-28 Legal kill rifle season-28 Legal kill rifle season-28 Disappear archery/muzzle season-21 Legal kill rifle season-28. Illega l kill rifle season-9 . Legal kill archery season-33 Legal kill muzzleloading season-34 Illegal kill rifle season-9 Wounding loss archery/muzzle-24 Legal kill rifle season-28 Unknown-suspect predation-30 Legal kill rifle season-28 Bear predation-35 Legal kill rifle season-28 Mountain lion predation-3 Illegal kill rifle season-9 Legal kill rifle season-28 Legal k i II muzzleloading season-34 Mountain lion predation-3 Disappear rifle season-22 Illegal kill rifle season-9 Unknown-suspect malnutrition-31 Mountain lion predation-3 Mountain lion predation-3 Wounding loss archery/muzzle-24 Legal ki II archery season-33 . Unknown predator-5 Unknown-suspect predation-30 Legal kill muzzleloading season-34 Unknown-suspect predation-30 Legal kill muzzleloading season-34 Mountain lion predation-3 Illegal kill-7 Unknown-suspect predation-30 Illegal kill rifle season-9 Mountain lion predation-3 Unknown-suspect malnutrition-31 Illegal kill rifle season-9 Unknown-suspect predation-30 Mountain lion predation-3 Accident, fell-10 Wounding loss rifle season-25 Unknown-suspect predation-30 Unknown-11 Mountain· liOn predation-3 Unknown-suspect malnutrition-~ ��75 Colorado Division of Wildlife Wildlife Research Report July 1997 JOB FINAL REPORT state of Colorado Project No. ~W~-~1~S~3~-~R~-~1~0~ _ Work Plan No Moose Inyestigations Job No. Development of census methods and determination of movements, habitat selection, and degree of calf mortality of moose in North Central Colorado. Period Covered: Author: Mammals Research July 1, 1996 - June 30, 1997 R. C. Kufeld Personnel: D. Bowden, D. Younkin. ABSTRACT All results of this completed study have now been published. Six publications resulting from this study including 2 submitted and accepted for publication in ALCES during FY 96-97 are listed. ��77 DEVELOPMENT OF CENSUS METHODS AND DETERMINATION OF MOVEMENTS, HABITAT SELECTION, AND DEGREE OF CALF MORTALITY OF MOOSE IN NORTH CENTRAL COLORADO Roland P. N. C. Kufeld OBJECTIVES 1. To determine the proportion of moose counting moose in North Park. 2. To determine 3. To determine the degree of dispersal of young animals, and seasonal movements, home range size, and habitat selection of North Park moose. the extent of moose actually observed calf mortality when aerially in late winter. SEGMENT OBJECTIVES 1. To determine 2. To determine the degree of dispersal of young animals, and seasonal movements, home range size, and habitat selection of North Park moose. the extent of moose calf mortality in late winter. STUDY AREA The study area was described by Kufeld (1992). METHODS AND MATERIALS Data describing seasonal home range size, seasonal movements, selection, degree of dispersal of young animals, and survival radio-collared moose collected during the course of this study and 2 manuscripts were prepared and submitted for publication seasonal habitat and mortality of were analyzed in ALCES. RESULTS All results of this completed study have now been published. Six publications resulting from this study including the 2 submitted and accepted for publication in ALCES during FY 96-97 are as follows: Bowden, D. C., and R. C. Kufeld. 1995. size estimation applied to Colorado 851. Generalized mark-sight population moose. J. Wildl. Manage. 59:840- Kufeld, R. C. 1994. Neck circumference of Shiras calves during winter. Alces 30:63-64. Kufeld, R. C. 30:41-44. 1994. Status and management moose of moose (~ ~ in Colorado. shirasi) Alces �78 Kufeld, R. C. 1996. status and management of moose in the Colorado State Forest and adjacent area of North Park. 13 pages in Appendix of Colorado State Forest Ecosystem Planning Project strategic Plan. Colorado state Board of Land Commissioners, February, 1996. Kufeld, R. C., and D. C. Bowden. 1996. Shiras moose (~ ~ shirasi) Movements and habitat selection in Colorado. Alces 32:In Press. Kufeld, ~ Survival rates 32:In Press. Prepared R. C., and D. C. Bowden. shirasi) in Colorado. by Roland C. Kufeld LSS Res/Sci III. 1996. Alces of Shiras moose of (~ �79 Colorado Division Wildlife Research July 1997 of Wildlife Report JOB PROGRESS State of Project Work Colorado No. ~W~__ I&Sa3_-~R~-~1~0~ Plan No. _ Mammals SP1 Deer Job No. Period Author: REPORT Research Inyestigations Regulation of Mule Deer Population Growth by Fertility Control: Laboratory, Field, and Simulation Experiments Covered: July 1, 1996 - June 30, 1997 Dan L. Baker Personnel: T. M. Nett, M. W. Miller, and M. A. Wild ABSTRACT Wild ungulates can do serious and enduring harm to many plant communities, and preventing such damage requires controlling animal numbers. Despite the ecological and political importance of controlling ungulate populations, achieving such control by hunting may not always be feasible. In these situations, alternatives to hunting as a means of regulating ungulate numbers are needed. Fertility control offers a promising alternative. Here, we conduct research to develop and test a practical and acceptable method of contraception in mammalian wildlife. We propose to use conjugates of gonadotropin releasing hormone (GnRH) and cellular toxins to selectively destroy gonadotropin producing cells in the anterior pituitary, thereby preventing gamete production by the ovaries and testes. Using mule deer and elk as laboratory models, we determined the optimum dose of GnRH analog required for contraception during different phases of the reproductive cycle. Mule deer and elk were generally more responsive to GnRH analog during the breeding season than during anestrus. During the breeding season and anestrus, elk required three times more GnRH analog than mule deer to achieve maximum LH secretion (breeding season:3 vs 1 ~g GnRH analog/SO kg BW; anestrus: 3 vs. 10 ~g GnRH analog/SO kg BW). In contrast to these reproductive states, mule deer were less responsive than elk to all doses of GnRH analog during pregnancy (2 vs 4 ~g GnRH analog/SO kg BW). Maximum serum concentrations of LH in pregnant mule deer and elk declined exponentially during gestation and the pattern of decline was similar for both species. We conclude from these studies that the amount of GnRH analog required for maxim~ secretion of LH is species specific and depends on the reproductive status of the animal. ��81 REGULATION OF MULE DEER POPULATION GROWTH BY FERTILITY LABORATORY, FIELD, AND SIMULATION EXPERIMENTS CONTROL Dan L. Baker P. N. OBJECTIVES 1. To develop a practical and acceptable method for controlling populations using GnRH-toxin conjugates. 2. To demonstrate the feasibility the Rocky Mountain Arsenal. 3. To predict population simulation modeling. mule deer of such control in a field application impacts of alternative SEGMENT contraceptive at regimes using OBJECTIVES 1. To evaluate the effectiveness and duration of a single dose application of GnRH-toxin conjugate to prevent normal production of reproductive hormones in captive mule deer. 2. To prepare manuscripts summarizing results of GnRH dose-response experiments in captive mule deer and other wild ungulates. EVALUATION OF GnRB-TOXIN CONJUGATE IN MULE DEER Experiments to evaluate a single dose application of GnRH-toxin conjugate in mule deer were not conducted during this segment. Researchers at the Animal Reproduction and Biotechnology Laboratory, Colorado state University were unsuccessful in their attempts to conjugate the enzyme Rnase to gonadotropin releasing hormone analog and produce a stable hormonal cytotoxin that could be tested in mule deer and other captive wild ungulates. Additional studies are currently in progress to evaluate alternative methods of conjugation of these molecules. GnRB DOSE RESPONSE EXPERIMENTS IN CAPTIVE MULE DEER AND ELK INTRODUCTION controlling the growth of wildlife populations is fundamental to maintaining proper balance between animals and the habitats they use. Hunting has traditionally been used to maintain this balance, but there are an increasing number of circumstances where hunting animals to regulate there numbers is not feasible. In these cases, fertility control offers a promising alternative to hunting as a means of limiting population growth. Here, we conduct research to develop and test a practical and acceptable method of contraception in mammalian wildlife that overcomes many of the shortcomings of current �82 technology, safety. particularly problems of treatment duration and environmental We propose to use conjugates of gonadotropin releasing hormone(GnRH) and cellular toxins to selectively destroy gonadotropin producing cells in the anterior pituitary, thereby preventing gamete production by the ovaries and testes. In order to provide an estimate of the dose of GnRH-toxin conjugate required for sterilization, it is essential that the optimum dose of GnRH analog be first determined in the species of concern. Using mule deer(Odocoileus hemionus)and elk(Cervus elaphus)as laboratory models, we determine the most effective dose of GnRH analog for contraception during different phases of the reproductive cycle. MULE DEER METHODS AND MATERIALS Study 1: Breeding season/anestrus. We determined the pattern of LH secretion following i.v. administration of GnRH analog in two experiments with three captive adult mule deer (4 yr old; mean BW = 61 kg). The first experiment was conducted during the breeding season (9 Nov - 11 Dec 1992), the second during early anestrus (4 April - 4 May 1993). Breeding season was determined to begin in mid-November. Serum concentrations of progesterone were monitored in each animal at S-day intervals during this period to ensure that the ovaries were active (D.L. Baker unpublished results). Previous studies with white-tailed deer (Odocoileus virginianus), suggest a reduction in ovarian activity beginning after the end of March (Verme 1965). Mule deer does were considered anestrous when serum progesterone levels were below 1 ng/ml for two consecutive weekly samples (Plotka et ale 1977). During the breeding season, we measured the LH response of each female to each of four doses of GnRH analog (0.3, 1.0, 3.0, and 10.0 ~g/50 kg BW); during anestrus we challenged does with five doses of GnRH analog (0.3, 1.0, 3.0, 10.0, and 30.0 ~g/50 kg BW). Individual doses were made from one batch of analog, lyophilized, and stored at - 20° C. Before each experiment, doses were reconstituted in 10 ml sterile saline solution and refrigerated at 5°C until the experiment was completed. Doses were administered on the same day to all animals. Mule deer does were challenged with GnRH every S days until each animal received each dose. On the day before each trial, animals were weighed (± 0.5 kg), moved to individual isolation pens (5 x 10 m), sedated with xylazine hydrochloride, and fitted nonsurgically with indwelling jugular catheters. The sampling period began the next day at OSOO h and ended at 1600 h. We administered GnRH through the cannula and collected blood samples (5 ml) at 0, 30, 60, 90, 120, lS0, 240, 300, and 360 min postinjection. After collection, blood as held at 4° C for 24 h until serum was obtained by centrifugation. Serum was then stored at - 20°C until analyzed. study 2: Pregnancy We determined the pattern of LH secretion during the second trimester (26 Jan �83 - 2 Mar, 1994)in six captive adult mule deer (5 yr old; mean BW = 71 kg) challenged with each of six doses of GnRH analog (0.5, 1.0, 2.0, 4.0, 8.0, 16.0 ~g/50 kg BW) at 7-day intervals. Pregnancy was determined by real-time ultrasonography (Mulley et ale 1987) and measurement of serum concentrations of progesterone (Wood et ale 1986). Day of gestation was estimated by backcalculating from known parturition dates using a gestation period of 203 ± 5.4 (SE)days (Anderson 1966). Animal handling and blood sampling protocols were similar to those described in the first study with the following exceptions: l)two additional serum collections were conducted at 420 and 480 min postinjection to ensure that LH concentrations had returned to baseline by the end of the trial and 2)administration of GnRH analog and blood sampling were conducted on the same day that animals were sedated and catheterized. Sedative effects of xylazine lasted 2-4 h, making it possible to collect blood from the jugular cannula while animals remained recumbent or standing in isolation pens. This modification in protocol prevented overnight loss of catheters and appeared to be less stressful on experimental animals. Serum concentrations of LH were quantified by means of ovine (0) LH RIA (Niswender et ale 1969). Mule deer serum was demonstrated to inhibit binding of 125I_oLH to ovine LH antiserum in a parallel manner. Likewise, when varying quantities of oLH standard (NIH-OLH-S24) were added and samples were subjected to RIA, the values obtained were increased by the quantity of oLH added(r2 = 0.99, SEb = 0.22, P = 0.002). The limit of sensitivity of the assay was 0.4 ng/ml. These data indicate that the RIA provided a quantitative assessment of LH in mule deer serum. Serum concentrations of progesterone were determined by RIA (Niswender 1973). Sensitivity of the progesterone assay was 0.12 ng/ml. Intra-and interassay coefficients of variation for each of these assays were less than 10%. Responsiveness of the pituitary to GnRH analog challenge was assessed in three ways: l)maximum response (highest concentration of LH (ng/ml) achieved postinjection minus baseline), 2)time (min) required to reach maximal LH, and 3)total amount of LH secreted (ng/ml/min; estimated by calculating the area under the LH curve (Abramowitz 1968). Data were analyzed using least squares ANOVA for General Linear Models (Freund et ale 1986) and the SAS Interactive Matrix Language. Response to treatment was analyzed with a two-way factorial analysis of variance for a randomized complete block design with repeated measures structure. Levels of GnRH analog were treatments; individual animals were blocks. Factors in the analysis were dose and time. Treatment was tested using the animal-with in-treatment variance as the error term. Time was treated as a within-subject effect using a multivariate approach to repeated measures (Morrison 1976). We used a priori orthogonal contrast to test for differences among individual means (Miller 1966) • RESULTS study 1: AND DISCUSSION Breeding Season/Anestrus. Cycling and anestrus mule deer challenged with analogs of GnRH responded with a measurable increase in serum LH by the first post-treatment blood collection �84 (30 min), reached a peak about 2 h later, and then declined to pretreatment levels by 5-6 h postinjection. Mule deer does were generally more responsive to GnRH analogs during the breeding season than during anestrus. Pituitary responsiveness as measured by total amount of LH secreted (ng/ml/min) and by maximum LH response (ng/ml) was highly correlated (r2 = 0.998, slope = 0.976, P = 0.001) and is subsequently described in terms of the latter estimate. We observed nonlinear changes in response means with increasing doses of GnRH analog for both the breeding season and anestrus. During the breeding season, maximum LH response (mean = 23.1 ± 4.1 (SE) ng/ml) was induced with 1.0 ~g GnRH analog/50 kg BW. A statistically significant (p = 0.01) decrease in LH response was observed at higher concentrations of GnRH analog (Fig. 1). In contrast, anestrous mule deer required three times more GnRH analog (3 ~g/50 kg BW) to achieve maximum LH response (mean = 13.6 ± 2.8 (SE) ng/ml). Unlike estrous does, however, mean maximum LH response did not decline (p = 0.87) with increasing amounts of GnRH analog (Fig. 2). study 2: Pregnancy Similar to results in the first study, serum LH concentrations increased (p 0.011) in a nonlinear pattern in response to increasing doses of GnRH analog (Fig. 3). Mean maximal response was achieved with 4 ~g GnRH analog/50 kg BW; LH then declined but not significantly (p = 0.58) with higher doses of GnRH treatments. Time from injection of GnRH analog to maximum LH was similar (p = 0.55) for all treatments and all trials. Time to maximal LH did not change in response to day of gestation (p = 0.76)and no interaction between dose of GnRH analog and time of peak was detected (p = 0.32). Averaged across all levels of GnRH analog, the mean maximal LH response for pregnant mule deer occurred at 206.3 ± 1.03 (SE) min. We estimated mule deer does to be near the end of the first trimester of pregnancy (64 ± 5 (SE)days) when GnRH analog challenge trials began. Serum progesterone concentrations were 4.3 ng/ml or higher for each animal over the course of the experiment (36 days), and all animals delivered normal fawns in June. Pituitary secretion of LH in response to GnRH decreased exponentially advancing gestation (p = 0.001, Seb = 0.42, r2 = 0.98). Pituitary responsiveness to GnRH was lowest during. pregnancy, and this value progressively decreased during the period of gestation. LITERATURE with CITED Abramowitz, M., and I. A. stegun. 1968. Handbook of mathematical Dover Publications Inc., New York, NY, pp. 645- 652. Anderson, A. E. 1967. The breeding season in migratory Div. Game, Fish and Parks, Inf. Leafl., 60:1-4. functions. mule deer. Colorado Freund, R.J., R.C. Littell, and P.C. Spector. 1986. SAS system for linear models. SAS Institute, Cary, NC, 187-201. �85 30 I ,,-.. .§ Ol e: 20 I- :c ...J E ~ ::l E x r1 <IS ~ c 10 l- <IS CD ~ o I 0.1 1 I I t 10 100 GnRH Analog (ft.Q/50 kg BW) Fig. 1. Mean maximum serum LH concentration (ng/ml) for GnRH induced release of LH for female mule deer during the breeding season. 20 ,,-.. I• E Ol C :c ...J E ::l E 10 • 1 x <IS ::: c <IS CD ::: o 0.1 1 10 100 GnRH Analog (JA-g/50 kg BW) Fig. 2. Mean maximum serum LH concentration (ng/ml) for GnRH analog-induced release of LH for female mule deer during anestrus. �86 3 I - E Ol c 2 1 :I: _J E ::J E I ·xas :§: c 1 ! as CD :§: o 0.1 10 1 GnRH Analog (~g/50 100 kg BW) Fig. 3. Mean maximum serum LH concentration (ng/ml) for GnRH induced release of LH for female mule deer during pregnancy. 4 - E Ol c 3 J: .....I E ::J •... CD (/) 2 E ::J E x as :§: c 1 as CD :l: o 60 64 68 72 76 80 Gestation 84 88 92 96 100 (days) Fig. 4. Mean maximum serum LH concentration (ng/ml) in pregnant mule deer treated with GnRH analogs during pregnancy (r2 c 0.978, Seb c 0.~5). �87 Miller, R.G. 1966. Simultaneous statistical New York, NY, pp. 152-168. Morrison, D.F. 1976. Multivariate York, NY, pp. 145-194. statistical inference. McGraw-Hill Book Co, methods. McGraw-Hill Co, New Mulley, R.C., A.W. English, R.J.Rawlinson, and R.S. Chapple. 1987. Pregnancy diagnosis of fallow deer by ultrasonography. Aust. Vet. J. 64:257-258. Niswender, G.D., L.E. Reichert, Jr., A.R. Midgley, Jr., and A.V. Nalbandov. 1969. Radioimmunoassay for bovine and ovine lutenizing hormone. Endocrinology 84:1166-1173. ________ • 1973. Influence of site of conjugation on the specificity antibodies to progesterone. steroids 22:413-424. of Plptka, E.D., u.s. Seal, G.C. Schmoller, P.D. Karns, and K.D. Keenlyne. 1977. Reproductive steroids in the white-tailed deer (Qdocoileus yirg~n~anus borealis) 1. Seasonal changes in the female. Biol. of Reprod. 16:340 343. Verme, L.J. 1965. Reproductive Manage. 29:74-79. studies on penned Wood, A.K., R.E. Short, A.E. Darling, G.L.Dusek, 1986. Serum assays for detecting pregnancy deer. J. Wildl. Manage. 50:684-687. white-tailed deer. J. Wildl. R.G. Sasser, and C.A. Rudder. in mule deer and white-tailed ELK Results of GnRH challenge experiments with captive elk have been published. copy of this manuscript is presented in Appendix A. A �88 APPENDIX BIOLOGY OF REPRODUCfION 52, 1193-1197 A (1995) Gonadotropin-Releasing Hormone Analog-Induced Patterns of luteinizing Hormone Secretion in Female Wapiti (Cervus elaphus nelsoni) during the Breeding Season, Anestrus, and Pregnancy 1 D.L. BAKER/.3 M.W. MIlLER,3 and T.M. NElT Colorado Division of Wildlife,3 Research Center, Fort Collins, Colorado 80526 Animal Reproduction and Biotechnology Laboratory, ~ Department of Physiology, Colorado State University Fort Collins, Colorado 80523 ABSTRACT We conducted two studies to determine the pattern of GnRH analog·induced Uf secretion in adult female wapiti during the breeding season, anestrus, and pregnancy. In the first study, we measured Uf secretion during the breeding season and anestrus in five females challenged with each' of five doses of GnRH analog (0.3, I, 3, 10, and ,30 f.Lg/50 kg BW). In the second study, Uf response was determlaed in six pregnant females treated with six doses of GnRH analog (0.5, I, 2, 4, 8, and 16 f.L8/50 kg BW). Animals were fitted nonsurgI.qUy with indwelling jugular catheters, and blood samples were collected at 0, 30, 60, 90, 120, ,ISO, 2-ro, 300, 360, ~20, and ~ min postinjeCtion. Mean maximwil sawn concentrations of Uf were not different (p 0 ..(5) between females rreated during the breeding season (mean = 29.1 :I: 1.9 (SE] ng/ml) and anestrus (mean = 32.4 :I: 4.3 (SE] ngfml); however, anestrous females required 3.3 times more GnRH analog (3 vs. 10 f.Lg/50 kg BW) to produce the Same rc:sponsc:. Maximalindudble Uf conc:entratlons were lowest (mean = 7.7 :I: 2.5 (SE) ng/ml) for pregnant wapiti, and the mag. nitude of the response decreased exponentially (r 0.98, P 0.002, SF." 0.65) during gestation. Although the magnitude of the response declined over time, maximal release consistently occurred at the same dose (2 f.Lg/50 kg BW) (treatment X trial interaction, P 0.53). We conclude from these studies that the amount of Uf released in response to a given dose of GnRH analog depends on reproductive status. There is a marked difference in pituitary sensitivity to exogenous GnRH between the breeding season and seasonal anestrus, but the amount of releasable Uf is similar. In contrast, during pregnancy the amount of releasable Uf declines with advancing gestation, but there is not a detectable change in pituitary sensitivity. = = = = = INTRODUCTION domestic species may not be an appropriate model for wild ungulates [10,11). For example, in the domestic ewe, it is well established that the ability of the anterior pituitary to release lR in response to a challenge with GnRH analog is higher during the breeding season than during seasonal anestrus or pregnancy. Additionally, pituitary response to GnRH decreases during gestation [12-15]. To our knowledge; there have been no comparable studies of pituitary sensitivity to GnRH analog in female Wapiti. Thus, the objective of the present study was to assess the responses of pituitary gonadotrophs during the breeding season, anestrus, and pregnancy and to determine whether or not the relationship between response of LH to GnRH analogs is similar to that reported for the domestic ewe. In this investigation we tested the null hypothesis that the reproductive status of female wapiti has no effect on lli secretion in response to stimulation by exogenous GnRH administration. Wapiti (Cervus elapbus nelsoni), like other cervids, exhibit a' yearly breeding cycle that is influenced by photoperiod [1]. In temperate regions of North America, the majority of conceptions occur in late September during , decreasing day length, but recurrent estrous cycles of 21 days are possible through February if the initial heat is delayed or if females fail to conceive after breeding. In early spring, coincidental with increasing day length, reproductive cycles cease and females remain anestrous through August [2]. For pregnant females, parturition generally occurs in early June, after a gestanonpenod of about 255 days [2- 4]. The reproductive endocrinology of seasonal breeding' is not well characterized for female wapiti. Hormonal profiles of conspecific Eurasian red deer suggest that changes in the frequency of release of pulses of GnRH via the hypothalamus constitute the principal mechanism that dictates seasonal changes in gonadotropin secretion and thus reproductive aalvity [>-9~The pattern of lR secretion in response to exogenous GnRH, however, has not been studied extensively in wapiti or other wild ungulates, Studies that have been conducted suggest that the lR response described for ICI", MATERIALS AND METHODS Animals AcccpCed January 12, 1995. P.ec:IcIYedQaober 27, 1994. 'Supponed by Federal Aid to WIIdIlfe RestontIon Projea W.153·Ri. ~ Dr. Dan L Baker, Colorado DIvIsIon 01 Wildlife, Research Cen- ' 317 West Prospea Street, Fort CoIUns, CO 80526. FAX:(303) ~90-262J. Wapiti cows used in these studies were artifidally reared (16) and well conditioned to repeated handling and blood sampling. When not used in these experiments, they were maintained in 5~hectare paddocks and fed a diet consisting of CUbed alfalfa hay, long-stem grass hay, and high-energy supplement. These experiments were conducted with the 1193 �89 1194 BAKER ET AL. approval of the Colorado Division of Wildlife's Animal Care and Use Committee and in compliance with Federal Animal Welfare Regulations. . Experimental Protocol Animal handling and blood sampling protocols were Similar to those described in the first study with the following exceptions: 1) two additional serum oollections were conducted at 420 and 480 min postinjection to ensure that LH concentrations had returned to baseline by the end of the trial and 2) administration of GnRH analog and blood sampling were oonducted on the same' day that animals were sedated and catheterized. Sedative effects of xylazine lasted 2-4 h, making it possible to collect blood from the jugular cannula while animals remained recumbent or standing in isolation pens. This modification in protocol prevented overnight loss of catheters and appeared to be less stressful on experimental animals. Study 1: Breeding season/anestrus. We determined the pattern of ill secretion following Lv. administration of GnRH analog (o-AJa6-GnRH-Pr09-ethyIamide; Sigma Chemical Co., St. Louis, MO) in two experiments with five captive adult wapiti cows (5 yr old; mean BW = i77 kg). The first experiment was conducted during the breeding season (24 September-26 October 1991), the second during early anestrus (20 March-21 April 1992). Breeding season was estimated to begin in mid-September. Serum concentraRIA olU! tions of progesterone were monitored in each animal at Serum ooncentrations of LH were quantified by means 8-day intervals during this period to ensure that the ovaries of an ovine (0) ill RIA [18]. Wapiti serum was demon- , were active (Dl.. Baker, M. Miller, T. Nett, unpublished restrated to inhibit binding of 1Z5I-oLHto ovine LH antiserum suits). On the basis of data from seasonal hormone profiles in a parallel manner. likewise, when varying quantities of reported for red deer hinds [4] and measured serum prooIR standard (Nlli-Oill-524) were added arid samples 'were gesterone .concentrations of wapiti in this experiment, we subjected to RIA,the values obtained were increased by the estimated seasonal anestrus to begin in February and end quantity of oUl added (rz = 0.99, slope = 0.92, SEt, = 0.22, in early August. Wapiti OOVIIS were considered anestrous when P = 0.002). The limit of sensitivity of the assay was 0.4 ng/ serum progesterone levels were below: 1 ng/ml for two ml. These data indicate that the RIAprovided a quantitative consecutive weekly samples [8]. ,assessment of UI tn wapiti serum. Serum concentrations of In each experiment, we measured the ill response of progesterone were determined by RIA [19]. Sensitivity of each female to each of five doses of GnRH analog (0.3, 1, the progesterone assay was 0.12 ng/ml. Intra- and inter3, 10,' and 30 JLg/50 kg BW). Individual doses were made assay coeffidents of variation for each of these assays were from one batch of analog, lyophilized, and stored at - 20°e. less than 10%. Before each experiment, doses were reconstituted in 10 ml sterile saline solution and refrigerated at 5°C until the ex-. ' periment was oompleted. Doses were administered on the Statistical Analysis same day to all animals. Wapiti COVIIS were challenged' with Responsiveness of the pituitary to GnRH analog chalGnRH every 8 days until each animal received each dose, lenge was assessed in four ways: 1) maximum response On the day before each trial, animals were weighed (± (highest concentration of IR [ng/ml] achieved postinjec0.5 kg), moved to individual isolation pens (5 X 10 in), tion minus baseline), 2) time (min) required to reach maxsedated with xylazine hydrochloride (Rompun; Bayer AG, imal UI, 3) total amount of LH secreted (ng/ml/min; esLeverkusen, Germany; 25-30 mg/animal i.m.), and fitted timated by calculating the area under the LH curve) [20], nonsurgically with indwelling jugular catheters. The samand 4) the amount of GnRH analog required to induce haIfpling period began the next day at 0800 h and ended at maximal release of LH (ED50). We calculated the ED50 for 1600 h. We administered GnRH, through the cannula and each response curve Using a nonlinear least squares esticollected blood Samples (5 ml) at 0, 30, 60, 90, 120, 180, 240, 300, and 360 min postinjection. After oollection, blood mation program. Data were analyzed using least squares ANOYAfor Genwas held at 4°C for 24 h until serum was obtained by ceneral linear Models [21] and the SAS Interactive Matrix lan, trifugation. Serum was then stored at - 20°C until analyzed. guage. Response to treatment was analyzed with two-way Study 2: Pregnancy. We determined the pattern of IE factorial analysis of variance for a randomized complete block secretion during the fust trimester of pregnancy (2 Decemdesign with repeated measures structure. Levels of GnRH ber 1992-6 January 1993) in six captive adult wapiti cows analog were treatments, individual animals were blocks. (6 yr old; mean BW = 307 kg) challenged with each of six Factors in the analysis were dose and time Treatment was doses of GnRH analog (0.5, 'I, 2, 4, 8, and 16 JLg/SOkg BW) tested using the animaI-within-treatment variance as the er-' at 7-day intervals. Pregnancy was determined by real-time ror term. Tame was treated as a within-subject effect using ultrasonography [17] and measurement of serum concena multivariate approach to repeated measures [22]. We used trations of progesterone [5,8]. Day of gestation was estia priori orthogonal oontrast to test for differences among mated by back-calculating from known parturition dates using a gestation period of 255 ± 7 (SE) days [2, 3). individual means [23t �90 LH SECRETION RESULTS study 1: Breeding Season/Anestrus Individual patterns of GnRH:induced LH secretion were similar during the breeding season and seasonal anestrus. We observed a measurable increase in serum LH by the first post-treatment blood collection (30 min), a peak response about 2.5 h later, and then a decline to pretreatment levels by 5-6 h postinjection. With the exception of five individual LH response curves observed during the breeding season, concentrations of LH had approached or returned to baseline by 6 h postinjection. Failure of LH to return to baseline in these females was probably attributable to an increasing baseline secretion of this hormone that is characteristic of the estrous phase of the cycle. To correct for this changing baseline, the area under the curve for estrous animals was calculated first by using the preinjection baseline, Then, the area below a line drawn. from the preinjection value to the 6-h value was subtracted to obtain an area under the curve corrected for a gradually. Increasing baseline. Wapiti cows were generally more responsive to GnRH analogs during the breeding season than during anestrus. Pituitary responsiveness as measured by total amount of LH secreted (ng/ml/min) and by maximum LH response (ng/ ml) was highly correlated (r = 0.995, slope = 0.965, P = 0.001, SEt, = 0.14) and is subsequently described in terms of.the latter estimation. We observed nonlinear changes in response means with increasing doses of GnRH analog for both reproductive states (Fig. 1). During the breeding season, maximal LH response (mean = 29.1 ± 1.9 [SEj ng/ ml) was induced with 3 JJ.gGnRH analog/50 kg BW-.The EDso for this action was 0.78 JJ.g/50 kg BW. No statistically significant (p = 0.66) changes in LH response were observed at higher doses. In contrast, anestrous cows required more than three times as much GnRH analog (10 JJ.g GnRH analog/50 kg. BW) to achieve maximal LH response (mean = 32.4 ± 4.3 [SEj ng/ml, EDso = 3.5 JJ.g/50 kg BW). Unlike that in estrous cows, however, the mean maximum LH response declined (p = 0.04) at the highest dose (Fig. 1). Average time from injection of GnRH analog to maximal ui was similar for all doses and both reproductive states (p > 0.34). Mean time (min) to maximum LH concentration (ng/ml) for all animals during both reproductive states . and across all treatments was 158.1 ± 17.5 (SE) min. Study 2: Pregnancy Similar to results in the first study, serum LH concentrations Increased (p < 0.014) in a nonlinear pattern in response to increasing doses of GnRH analog (Fig. 1). Mean maximal response was achieved with 2 ..JJ.gGnRH analogi 50 kg BWj LH then declined (p < 0.03) and did not change (p = 0.52) with higher doses of GnRH treatments. The maximal lR concentration attained in pregnant females IN FEMALE WAPITI I ..J ~~---------------------------------, o Breeding season Anestrus E ~ I>. a Pregnancy 30 * EDso CI) ~ E::I~ 1195 20 .~.s :E c: 10 Sl :E 00.1 100 GnRH Analog (Jl.g1SOkgBW) FIG. 1. Mean maximum serum LH concentrations (ng/mll and ED •• for GnRH analog·induced release of LH for female wapiti during. the breeding season, anestrus, and pregnancy. (mean = 7.7 ± 2.5 [SED was 70% lower than that measured for estrous females, but the EDso required to induce maximal response was similar for both reproductive states (pregnancy EDso = 0.64 vs. breeding season EDso = 0.78 JJ.g/50 kg BW) (Fig. 1). . . Time from injection of GnRH analog to maximum ill was similar (p = 0.42) for all treatments and all trials. Time to maximal LH did not change in response to day of gestation (p = 0.58), and no interaction between dose of GnRH analog and time of peak was detected (p = 0.36). Averaged across all levels of GnRH analog, the mean maximal LH response for pregnant wapiti occurred at 154.4 ± 10.4 (SE) min. We estimated wapiti cows to be in the first trimester (53 ± 2.0 [SEj days) of pregnancy when the GnRH analog challenge trials began. Serum progesterone concentrations were 3.5ng/ml or higher for each animal over the course of the experiment (35 days), and all animals delivered normal calves in June. One cow was determined not to be pregnant and was removed from the study. Pituitary secretion of LH in ~~ponse to GnRH decreased exponentially with advancing gestation (p = 0.002, SEt, = 0.65, = 0.98) (Fig. 2). Although the magnitude of the LH response decreased over time, maximal release consistently occurred at the same dose (2 JJ.g/50 kg BW) for all r :J: ..J 15~--------------------------------~ ~ = 0.98 e = 0.002 sEt, 0 50 57 63 = 0.65 70 83 77 Gestation (days) AG. 2. Mean maximum serum LH concentrations wapiti treated with GnRH analogs during gestlltlon. (og/mll In pregnant �91 1196 BAKER ET AL. trials (treatment x day interaction, p > 0.51). Maximal LH concentration declined most sharply between 53 and 60 days of gestation with a more gradual decrease thereafter. By 88 days of gestation, serum LH concentrations had declined by 77% in relation to initial values. DISCUSSION Differences in GnRH analog-induced patterns of LH secretion were related to the reproductive status of female wapiti. There were marked differences in responsiveness of pituitary gonadotrophs to GnRH analogs. during the breeding season, seasonal anestrus, and pregnancy. Pituitary sensitivity of wapiti cows was greater during the breeding season than during anestrus; however, maximal LH response was similar. During the breeding season, the magnitude of GnRHstimulated LH release was more variable among individuals than during the other reproductive states. Estrous cycles were not synchronized, and as a consequence the treatments were administered randomly throughout the cycle (24). Therefore, the variation in maximum LH response may be largely attributable to the phase of the cycle during which cows were challenged with GnRH and the influence of fluctuating concentrations of estradiol and progesterone [25,26). Although wapiti cows were less responsive to GnRH analog stimulation during anestrus compared to the breeding season, this phase does not appear to represent a time of pituitary gonadotropic hormone depletion. Our results indicate that while pituitary sensitivity is diminished at this time in relation to the breeding season, the pituitary of seasonally anestrous females is capable of releasing LH when stimulated by exogenous GnRH. Stimulation of LH secretion by GnRH was concentration-dependent over the range of 0.3 to 10 ILg/50 kg BW. Maximum serum concentrations of LH attained during early anestrus were higher, although not Significantly higher (p > 0.34), than those values measured during the breeding season. Results from studies of other wild ungulates are generalIy Consistent with our Observations. Administration of GnRH analog to seasonally anestrous Pere David's deer (ElaphuIUS davidianus) resulted in increased LH concentrations comparable to those observed during the luteal phase of . the estrous cycle [10,11); these findings are similar to ours. Our observations of the GnRH-induced LH response in cycling and anestrous wapiti are broadly in accord with results for domestic sheep showing that the pattern and magnitude of the LH response are similar for the two reproductive states [12,26-28). However; in the ewe, maximum lli response during these two reproductive states Is achieved with the. same dose of GnRH, whereas wapiti cows required almost three times more GnRH analog. during anestrus to achieve the same response obtained during the breeding season. These· results lend support to the hypothesis that in early anestrus, the pituitary is less responsive to chal- lenge by GnRH in wild cervids compared to domestic sheep [lO,l1J. Pituitary responsiveness to GnRH was. lowest during pregnancy, and this value progressively decreased during the period of gestation in which the experiment was conducted (Days 53-88). Although LH secretion declined over time, maximum release consistently occurred at the same dose of GnRH (2 ILg/50 kg BW), suggesting that in wapiti, pituitary sensitivity to GnRH remains constant. To our knowledge, pituitary responsiveness to GnRH during pregnancy has not been previously reported for wild ungulates. In the domestic ewe, however, several studies have documented a similar decline in pituitary secretion of LH during gestation [12,15). In pregnant sheep, GnRH-induced release of LH has been shown to be highly correlated to the amount of LH contained in the pituitary [12,15). Furthermore, it has been reported that in sheep a constant percentage of LH is released in response to a maximally stimulatory dose of GnRH. That is, more LH is released early in gestation because more LH is contained in the pituitary, not because of a change in pituitary sensitivity to GnRH [12,15). Our observations suggest that a similar mechanism operates in wapiti. In summary, we conclude that the GnRH-induced patterns of LH secretions observed in this study are consistent with those described for the domestic ewe with one notable exception: during anestrus there is a marked reduction in pituitary sensitivity to GnRH in female wapiti, but not in sheep. Further studies with wild cervids are needed to elucidate the mechanism responsible for reduced sensitivity of the hypothalamic-pituitary axis during this reproductive state. ACKNOWLEDGMENTS The aWlors appreciate and ad<nowIedge AI- Cze for :I5SisWlce throughout Ihese . experiments, including blood sampling, RIA, and overall execution of the study. We are also Indebted to J.e. Ritchie for assrst2nce In animal handling and sample colleaIon. A special thanks to Dr. MA. Wild ror the care, maintenance, and veterinary attention required to condition experimental wapiti for these trialS and for assistance whenever needed to complete the stUdy. We thank Dr. D.e. Bowden for statistidll consultation and analysis. OJ. Freddy and Dr. MA. Wild offered panicularly helpful comments on early dr2fts. REFERENCES , 1. Uncoln GA Seasonal breeding in deer. In: Fennessy PF, Drew KR (eds.), Biology of Deer Produaion. Wellington, New Zealand The Royal Sociely. of New Zealand; 1985: 165-179. . 2. Momson jA. Olar2cteristics or estrus In captive elk. Behavior 1960; 16:84-92. 3. Morrison.JA, Trainer CE, Wright PL Breeding season In elk as determined from known-age embryos. J Wildl Manage 1959; 23:27-6. ". Gulness F, Uncoln GIl., Short RV. The reproduatve cyde or the remale red deer (Cetvus ~). J Reprod Fertil 1971; 2N27-eB. 5. Kelly RW, MaI2t1y KP, Moofe GH, Ross 0, Gibb M. Plasma concentrationS or IJ{, prolactin, oestradiol, and progesterone In Canale red deer (C6t1UUIapbUS) duro Ing pregnancy. J Reprod Fertll 1982; 604:-475-483. 6. Loudon ASI, Mllne.JA, Curlewls}D, McNeIlly AS. A comparison or the seasonal honnone changes and patterns or growth. voluntary food inW<e and reproduction In juYenlle and adult red deer (Cetvus eIapbus) and ~ DaYkfs deer (EIa· pburus davtdJanus) hInds. J EndocrInoI 1989; 122:733-7"5. �92 LH SECRETION 7. Argo CM, loudon AS!. Effect of age and time of day on the timing of the surge In luteinizing hormone, behavioral oestrus and mating in red deer hinds (Cervus elapbus). J Reprod Fenil 1992; 96:667-672. 8. Adam a.. Moir CE, Alkinson T. Plasma concentrations of progesterone in female red deer (Cervus elapbus) during the breeding season, pregnancy and anoestrus. J Reprod Fenil 1985; 74:631-636. 9. Asher CW, Fisher MW,Jabbour HN, SmithJF, Mulley ac, Morrow q, Veldhuizen FA, Langridge M. Relationship between the onset of oestrus, the preovulatory surge in luteinizing hormone and ovulation follOWing oestrus synchronization and superovulation of f:umed red deer (Cervus e/apbus). J Reprod Fenil 1992; 96:261-273. 10. Curlewis)O, Md.eod BJ, Loudon AS!. Uf secn:tion and response to GnRH during seasonal anestrus of the Pe.re David's deer hind (EIap:mrus davidianus).") Reprod Fenil 1991; 91:131-138. 11. Md.eod B), Brinklow BR, Curlweis )0, loudon AS!. EffiC2CYof intermittent or continuous administration of GnRH in Inducing ovulation in early and late seasonal anoestrouS in Pere David's deer hind (EJapburus davidianus). ) Reprod Fertll 1991; 229-238. 12. )enldn G, Heap RB, Symons DBA. Pituitary responsiveness 10 synthetic Uf-RH and pituitary Uf content at various reprod~ stages In sheep.) Reprod Fertll 19n; 49:207-211. 13_ Crighton DB, Foster )p, Hareslgn W, Scott SA. Plasma Uf and progesterone levels after single or multiple Injeaions of syr:tthetic Uf-RH In. anoestrouS ewes and comparison with ~ during the oestrous cyde.) Reprod FertlI 1975; «:121124. 1"- 0wnIcy WA, fIndIay]K, Cumming IA, Budanaster JM, GodingJR Effea of pregnancy on the Uf response 10 syn!hcUc gooadoIropln-celeasing hormone In the ewe. Endocrinology 1974; 9-4:291-293. 15. Crowder ME, GUles PA, Tam2IlInI C, Moss os, Nett, TM. Pituitary content of g0nadotropins and GnRH-ceceptors In pregnan!, pastpanum and steroi<kreared OYX ewes. J Anlm Sci 1982; 54:1235-1242. .:16. Wild MA, Miller MW. BoaIe-f2islng wild ruminants In captivi!},. Colo DiY of Wild! Outdoor Facts 1991; llH--6. IN FEMALE WAPITI 1197 17. Wilson PRoBingham CM. Accuracy of pregnancy diagnosis and prediction of calving date in red deer using real-time ultrasound scanning. Vet Rec 1990; 126:133135. 18. Niswender GO, Reichen LE)r, Midgley AR)r, Nalbandov AV. Radioimmunoassay for bovine and ovine luteinizing hormone. Endocrinology 1969; 8(1166-1173. 19. Niswender, GO. Influence of the site of conjugation on the specifici!}, of antibodies to progesterone. Steroids 1973; 22:413-424. ZO. Abramowitz M, Stegun IA. Handbook of Mathematical Functions. New Yot1c:Dover PubliC2tions Inc.; 1968: 645-652. 21. Freund RJ, Uttell RC, Spector PC. SAS system for linear models. Cary, NC: SAS Institute; 1986: 187-201. 22. Mocrison OF. Multivarl2te Statistical Methods. New York: McGraw·HiII Book Co.; 1976: 145'-194. 23. Miller RG. Simultaneous Statistical Inference. New York: McGraw-Hili Book Co.; 1966: 152-168. 24. Reeves JJ, Arimura A, SchaIly AV. Pituitary responsiveness to purified luteinizing hormone-releasing hormone (Uf·RH) at various Slages of the estrous cycle in sheep.) Anim Sci; 32:123-126. 25. Crowder ME, Nett TM. Pituitary content of gonadotropms and receptors for gonadottopln-releasing hormone (GnRH) and hypothalamic content of GnRH during the preovulatory period In the ewe. Endocrinology 1984; 114:234-239. 26. Goodman RL, K2rsch. 1:1. Pulsatile .seereuon of lutelnizln8 hormone: differential suppression by ovarian steroids. Endocrinology 1980; 107:1286-:-1290. 27. Foster)p, Crighton DB. Lutdnlzlng hormone (Uf) release after single Injeaions . of a syiuheIic Uf-releasing hormone (Uf-RH) In the ewe at dirce dlIferenl reprodualYe stages and 00mparts0n with naruraJ Uf release at oestrus. Theriogcnology 1974; 2:87-100. 28. Symons AM,.CunningIwn NF, Saba N. The gonadotroplc.hormone response of anoestrouS and cyclic ewes to synthetic lutelnizln8 hormone-releasing hormone. ) Reprod Fertll 197"; 39:11-21. . 29. Ownley WA, Fmdlay)K,)onas H, CurnmIng lA, GOdingJR Effea of pregnancy on the FSH response to synthetic gonadotrophln·releasing hormone in ewes. ) Reprod FertlI 1974; 37:109-112. �93 Colorado Division Wildlife Research July 1997 of Wildlife Report JOB FINAL REPORT HEART RATE AS A POTENTIAL INDICATOR OF STRESS: .APPLICATION TO BIGHORN SHEEP EXPOSED TO HUMAN DISTURBANCE Period Covered: Authors: Personnel: July 1, 1996 - June 30, 1997. M. A. Wild, D. L. Baker, and D. Bowden. L. D. Bowser, A. L. Case, J. Ritchie, J. L. Schaefer, E. Wheeler. To better understand the effects of disturbances on bighorn sheep, a reliable, longterm, discrete, quantitative indicator of stress is required. We evaluated remote monitoring of heart rate to meet this need and performed experiments to investigate the correlation between heart rate and serum cortisol in captive bighorn sheep. We reported a new technique for remote monitoring of heart rate in a draft manuscript submitted for publication to the Journal of Wildlife Diseases. The manuscript is titled KSurgical Implantation and Evaluation of Heart Rate Transmitters in Bighorn Sheep" by Margaret A. Wild, Donald L. Piermattei, R. Bruce Heath, and Dan L. Baker. We investigated the correlation between heart rate and serum cortisol levels by analyzing data collected in 1996 from 15 captive adult non-pregnant ewes. We found a positive, linear relationship between heart rate and serum cortisol. Further, we used linear regression to determine the correlation between heart rate response and change in cortisol (difference between -5 min value and 20 min post-stressor value) (r2 = 0.43) and also heart rate response and peak cortisol response for a trimmed data set including those animals whose initial cortisol level was at baseline (r2 = 0.57). We used multiple regression to find a model for the stressor induced change in cortisol. The model y = -10.2 + 0.07(Bl) + 0.11(B4) + 0.08(B5), where B1, B4, and B5 are the harmonic mean heart rate 1,4, and 5.min post-stressor initiation, respectively, was acceptable (r2 = 0.605); however, the model was markedly improved by addition of the initial (-5 min) cortisol value (r2 = 0.756). These analyses suggest that for the best predictive value of heart rate, the bighorn sheep should initially be at baseline cortisol levels. In a more general sense, we found that increases in heart rate >125 bpm were associated with cortisol values >20 ng/ml. A conservative interpretation of these data would suggest that disturbances resulting in heart rate responses of >125 bpm be avoided if the manager wishes to minimize distress to bighorn sheep. Additionally, we performed experiments to investigate the effect of pregnancy on the correlation between heart rate and serum cortisol and to determine the circadian and seasonal cycles of heart rate and serum cortisol in bighorn sheep. These data have not yet been analyzed. Although this project has been terminated due to budgetary cons~raints, we expect to complete analysis and publication of our data as time permits in the future. Hopefully, in the future we, or others, will have the opportunity to evaluate the feasibility and practicality in a field application. Given results from experiments with captive bighorn sheep, we believe that heart rate is a promising technique that could be used to monitor stress in free-ranging bighorn sheep. ��95 HEART APPLICATION RATE AS A POTENTIAL INDICATOR OF STRESS: TO BIGHORN SHEEP EXPOSED TO HUMAN DISTURBANCE Margaret A. Wild, Dan L. Baker, and David Bowden P. N. OBJECTIVES 1. To develop a safe, reliable, and unobtrusive heart rate in bighorn sheep over an extended system period to remotely (~ 1 year). 2. To determine the correlation between heart rate and serum cortisol levels in bighorn sheep and to understand the effects of some other physiologic parameters on this correlation. 3. To determine the impact of human disturbance on free-ranging ~~heep using heart rate telemetry and the heart rate-cortisol . correlation. SEGMENT bighorn OBJECTIVES 1. To evaluate function of heart 15 captive bighorn sheep. 2. To determine the correlation between heart level in adult, female bighorn sheep under graded stressors. 3. To perform experiments to determine the influence circadian cycles and reproductive status on heart cortisol levels. METHODS monitor rate transmitters surgically implanted rate and serum cortisol controlled situations with of seasonal and rate and serum AND MATERIALS In performing this study, we generally followed methods described in the Program Narrative (Wild and Baker 1995) and modifications reported in Wild Baker (1996). Modifications to the protocol are described below. Evaluation of Heart A draft manuscript Diseases. Graded Stressor Rate Transmitters was prepared Experiment in in Bighorn and submitted - Non-pregnant and Sheep to the Journal of Wildlife Ewes We performed preliminary analysis on data collected from captive bighorn sheep during January-March 1996. We used two methods to investigate the relationship between heart rate and serum cortisol data. First, we plotted the harmonic mean heart rate of the first 5 min after stressor initiation vs. either the 20 min post-stressor cortisol value or the change in cortisol associated with the stressor (difference between the -5 min sample and 20 min sample value). The second method that we used was multiple regression to find a model to explain the change in cortisol that occurred as a result of the. stressor. We determined this change in cortisol level by finding the �96 difference between the ~5 min cortisol sample (presumably a baseline) and the 20 min cortisol sample (the approximate time of peak cortisol response). We then tested various models to explain the change in cortisol level using independent variables that included the harmonic mean heart rate over 1 min periods from 1-5 min post stressor and baseline (-5 min) cortisol value. Graded c- stressor Experiment - Pregnant Ewes Graded stressor trials were performed using pregnant ewes during early (6 January - 7 February 1997; early second trimester) and late gestation (17 March - 11 April 1997; third trimester). These trials were similar to those performed using open ewes in winter 1996; however, based on data from the 1996 trials, we made the following modifications, a) the acclimation period prior to application of the stressor was increased to about 105 min, b) heart rate data were collected for 30 min prior to application of the stressor, and c) collection of heart rate and serum cortisol data was limited to 80 min postst~f!e.-~. Circadian and Seasonal Cycles We continued to study the circadian and seasonal cycles of heart rate and serum cortisol levels. We collected continuous heart rate data and blood samples every 2 hr for a 24 hr period on 19-20 September and 21-22 December 1996. RESULTS AND DISCUSSION Evaluation of Heart Rate Transmitters in Bighorn Sheep A draft manuscript was submitted for publication to the Journal of Wildlife Diseases. Following is the abstract of that manuscript "Surgical Implantation and Evaluation of Heart Rate Transmitters in Bighorn Sheep" by Margaret A. Wild, Donald L. Piermattei, R. Bruce Heath, and Dan L. Baker. ABSTRACT: A safe, reliable, and unobtrusive telemetry system to monitor heart rate of bighorn sheep (Qyia canadensis) over relatively long ranges was developed. We surgically implanted Telonics model HR400 transmitters on the dorsolateral thorax of 15 captive adult bighorn sheep ewes in April .•. May and October-November 1995. No complications or marked impairment of function were associated with the surgery; however, a transmitter was passively expelled from one ewe 19.5 mo post-implantation. Twelve of 15 transmitters remained functional ~1 yr, while three failed 3.5 to 4.5 mo following implantation. Heart rate data collected from the transmitters using a Lotek SRX_400 telemetry receiver/datalogger equipped with W9 EVENT_LOG accurately reflected "true" heart rate. Line of sight signal range was at least 800 m in 95% (37/39) of collections made from standing ewes, while data could be collected reliably (74%; 29/39) to 600 m from bedded ewes. We recommend this approach to remotely monitor heart rate in ungulates >50 kg body mass when a long lasting unobtrusive system is required. Graded Stressor Experiment - Non-pregnant Ewes The trend of change in serum cortisol level associated with change in heart rate can be appreciated by plotting the harmonic mean heart rate of the first 5 min after stressor initiation vs. the 20 min post-stressor cortisol value �97 (Fig. 1); however, due to marked variance in the initial cortisol levels of the bighorn sheep, further analysis was required. We first analyzed these data using the change in cortisol after the stressor (difference between the 5 min sample and 20 min sample value; Fig. 2). We also analyzed the data using a trimmed data set (Fig. 3). The trimmed data set included only those response data from animals whose initial (-5 min) cortisol value was ~10 ng/ml (i.e., "baseline"). These findings suggest that in order to best predict response of serum cortisol after a stressor, the animal would need to be at baseline initially. However, in the field we would not likely know the initial cortisol value (although an estimate based on heart rate and behavior might be possible). In a more general sense, we found that increases in heart rate >125 bpm were associated with cortisol values >20 ng/ml (Fig. 1). In this study, we condidered values <10 ng cortisol/ml to be baseline. A conservative interpretation of these data would suggest that disturbances resulting in heart rate responses of >125 bpm be avoided if the manager wishes to minimize distress to bighorn sheep. ~, . The overall best multiple regression model to explain the change in cortisol included the intercept, the baseline cortisol value (cortO), and the harmonic mean heart rate 1 min (B1) and 5 min (B5) post-stressor initiation. Model 1: y = -7.7 + 0.8(cortO) + 0.07(B1) + 0.09(B5) This model nicely explained the change in cortisol with an R-square of 0.756; however, as discussed above, the baseline cortisol value would likely not be known in free-ranging animals. A model using the intercept and the harmonic mean of min 1, 4 (B4), and 5 post-stressor explained less, although a significant amount (P<0.0001), of the variability (r2 0.605). = Model 2: y = -10.2 + 0.07(B1) + 0.11(B4) + 0.08(B5) These findings suggest that an increase in heart rate is strongly correlated with an increase in cortisol; however, the specific amount of change is somewhat difficult to predict if the baseline cortisol value is not known. Graded Stressor Experiment - Pregnant Ewes Data were collected from six ewes during early gestation and from five during late gestation. Based on back-calculations from lambing dates, ewes ranged from 36-67 days gestation at study initiation and 131-162 days gestation at study termination. In general, bighorn sheep performed well and experimental techniques were successful. Serum cortisol levels were determined and heart rate data compiled. Although due of lack of funding this is the Job Final Report for this project, we will complete data analysis and publication as time permits in the coming year. We will perform similar analyses as described above to determine the correlation between heart rate and serum cortisol level. We will compare results to those obtained using open ewes in winter 1996 to determine the effect of reproductive status on the predictive value of heart rate as an indicator of stress in bighorn sheep. Circadian and Seasonal Cycles Heart rate data and serum cortisol values were collected from six bighorn sheep at the autumn equinox and from five bighorn sheep at the winter solstice. Although due to lack of funding this is the Job Final Report for �98 this project, as time permits in the future we will analyze these data in addition to those collected at the spring equinox and summer solstice 1996. These analyses will provide information on the circadian and seasonal cycles of heart rate and serum cortisol levels in bighorn'sheep. Effects of other physiologic factors, such as feeding and exercise, on the correlation between heart rate and serum cortisol levels were not performed due to project termination. Field study Although in the Program Narrative (Wild and Baker 1995) we described an experiment to test the system's performance under field conditions, due to lack of funding this project has been terminated. Hopefully, in the future we, or others, will have the opportunity to evaluate the feasibility and practicality in a field application. Given results from experiments with caR~, bighorn sheep, we believe that heart rate is a promising technique . that could be used to monitor stress in free-ranging bighorn sheep. An ideal application would be to evaluate disturbance of bighorn sheep in a wildlife viewing area in response to various practices for managing human activity. This information would give managers a quantitative tool to assist in formulating management practices. The technique could likely be successfully adapted to monitor disturbance in other species of wildlife as well. LITERATURE CITED Wild, M. A. and D. L. Baker. 1995. Heart rate as a potential indicator stress: application to bighorn sheep exposed to human disturbance. Colorado Div. Wildl. Res. Rep., Jul 1994 - Jun 1995, Fort Collins. of Wild, M. A. and D. L. Baker. 1996. Heart rate as a potential indicator stress: application to bighorn sheep exposed to human disturbance. Colorado Div. Wildl. Res. Rep., Jul 1995 - Jun 1996, Fort Collins. of �99 • • • • • • • •• • • •• ... ~.:.. . ..', . ... • •••••••• • • • • • ., • • . • ~ ~ ••• • 25 50 75 100 125 150 175 200 225 250 Mean Heart Rate (bpni) Fig. 1. Relationship between the harmonic mean heart rate for the first 5 min post-stressor initiation and serum cortisol 20 min post-stressor initiation for 15 captive bighorn sheep exposed experimentally to graded stressors. -.e C) 40 30 •• c '-" • • • •• • •• • -10~--~--~--~--~~~~~--~--~----~~ o 25 50 75 100 ·125 150 175 200 225 250. Mean Heart Rate (bpm) Fig. 2. COrrelation between the harmonic mean heart rate for the first 5 min post-stressor initiation and the change in serum cortisol associated with the stressor for 15 captive.bighorn sheep exposed experimentally to graded stressors. �100 -.ECl 40 c •...•.. 15 en ~ 820 E ::s ••• Q) 25 50 75 100 125 150 175 200 225 250 Mean' Heart Rate (bpm) Fig. 3. Correlation between the harmonic mean heart rate for 1 min poststressor initiation and serum cortisol 20 min post-stressor initiation for captive bighorn sheep exposed experimentally to graded stressors. Response data are reported only for bighorn sheep whose initial cortisol values were at baseline. �101 Colorado Division Wildlife Research July 1997 of Wildlife Report JOB FINAL REPORT state of Colorado Proj ect No. --llW~...• 1...•• 5!..:13t....--,R;)....... 1k.lOIL_ Mammals Research Work Plan Hultispecies No.__~~ _ Job No. Inyestigations Animal and Pen Support Facilities for Mammals Research Period Covered: July 1, 1996 - June 30, 1997. ~, Author: M. A~ Wild Personnel: P. E. Bleicher, D. Leinart, Wallick, E. Wheeler. J. Tollefson, D. Stallnecht, L. A. ABSTRACT The Colorado Division of Wildlife's Foothills Wildlife Research Facility (FWRF) maintained captive animals (up to 128 wild ungulates of 5 species) and facilities supporting five major research projects. Six weaned bighorn sheep lambs from Wyoming Game and Fish Department, Sybille Wildlife Research Unit were added to the herd. Eleven bighorn sheep lambs and 14 viable mule deer fawns were born to captive females at FWRF in spring 1997. During the year, reductions in animal numbers were due to natural mortalities (11 adults and 1 neonate) and euthanasia of six white-tailed deer fawns and two pronghorn bucks as part of facility management programs. Chronic wasting disease (CWO) has become a significant source of mortality in captive mule deer. Between July 1995 and June 1997, half of the 18 mortalities occurring in mule deer were due to CWO. Thirty-three percent of male and 14% of female mule deer housed at FWRF in the past 2 years (excluding fawns born in 1997) have died of CWO. However, endemic CWO has provided an opportunity to study epizootiology and antemortem diagnosis of the disease. Over the past 6 years, FWRF operation has emphasized economy and utility in function in addition to improvements in animal welfare. The high quality of animal care is in part reflected in the results of animal welfare inspections. Economy and utility of FWRF function have increased in at least four areas. First, the use of volunteer workers has increased markedly. Well trained volunteers contributed 443.5 man hours in FY1997. Second, a conservation oriented approach to the use of utilities and services for FWRF has lead to a reduction in resource use relative to the size of the facility over the past 6 years. Third, implementation of standard operating procedures for animal caretaking and feeding has improved overall animal health and feed efficiency. And finally, as older portions of the facility are repaired and replaced, the need for unscheduled daily repairs has decreased; however, numerous maintenance and improvement projects were performed again in FY1997 to increase usefulness, efficiency, quality, and/or safety of facilities at FWRF. Although availability of research animals and facilities has significantly benefited the research program in the past, �102 budgetary constraints dictate that future support of FWRF be dependent solely on the budgets of specific research projects. In the future, descriptions of animal and facility maintenance will be reported in Job Progress reports of those research projects. �103 ANIMAL AND SUPPORT FACILITIES MAMMALS RESEARCH Margaret FOR A. Wild P. N. OBJECTIVES 1. To provide and maintain captive wildlife populations and facilities supporting CDOW's Terrestrial Wildlife Research Program, as well as programs of CDOW cooperators. 2. To develop improved animal and facility management practices provide maximum research opportunities at minimum cost. 3. To enhance facilities to serve a growing diversity that will of anticipated research -4~?s. SEGMENT OBJECTIVES 1. Maintain and modify animal research 2. Coordinate all animal rearing, training, activities. 3. Provide or oversee maintenance for up to 20 elk, 40 mountain sheep, 25 pronghorn, 45 mule deer, and 11 white-tailed deer under applicable federal and institutional animal welfare regulations and in suitable health for use in research experiments. 4. Conduct management experiments to increase efficiency and efficacy feeding, husbandry, and maintenance activities related to research facility operations. 5. Follow a conservation-oriented approach for providing services to operate research facilities. 6. Follow Standard Operating facility records. Procedures and holding facilities. maintenance, in maintaining and research utilities detailed of and animal and METHODS AND MATERIALS Over the past 6 years, FWRF operation has been generally guided by the objectives outlined in the 1990 Program Narrative (Miller 1990). Emphasis has been placed on economy and utility in FWRF function. These considerations have been concurrent with increased emphasis on quality of animal care and animal welfare. Animal care and facility maintenance standard operating procedures (SOP's) were developed during FY1993 and have been followed for routine procedures since that time, and modified as needed. Given this general guidance, and the direction required to meet upcoming terrestrial research needs, we performed the following tasks: �104 Animal Maintenance General: Again this year, routine feeding and caretaking of research animals, including health observations, training, weighing, and clean-up, was performed primarily by well trained work-study and temporary employees, as well as volunteers. FWRF was inspected by USDA APHIS for compliance with federal animal welfare regulations on 18 March 1997. To maintain optimal population size of the captive herd, 12 bighorn sheep ewes and 10 mule deer does were bred in fall 1996. Additional bighorn sheep lambs obtained from Wyoming Game and Fish Sybille Wildlife Research Unit (SWRU) were also added to the herd. Six white-tailed deer fawns born unexpectedly in late September and october were euthanized. Nutritional Maintenance ~,,,,",,,-,,,,_~"'..9.i~,protocols: Feeding protocols were as previously described (Miller . 1990, Wild et al. 1992, Wild 1993, Wild and Graffam 1994; Wild and Schaefer 1996). For mule deer, we continued partial supplementation with a pelleted browser maintenance ration (1 kg/head/day; PMI Feeds, St. Louis, MO 63144). Health Maintenance General: We continued to monitor animal health using FWRF sOP's. Animal health care was provided as required and as mandated by the preventive medicine program and chronic wasting disease protocols. Chronic wasting disease: We followed protocols for the preventive medicine program (Wild 1995) and management of chronic wasting disease (CWO) (Wild and Graffam 1994). All animals at FWRF were monitored closely for clinical signs of CWO. Tissues from all mortalities occurring at FWRF were examined for evidence of infection with CWO. We initiated a study to investigate the utility of several techniques for the antemortem diagnosis of CWO. Samples were collected in summer and winter from all mule deer and elk at FWRF. Details of the methods and results are reported in the Program Narrative and Progress Report for WP2J17. Epizootic hemorrhagic disease in bighorn sheep: We determined serologic titers to epizootic hemorrhagic disease (EHD) in bighorn sheep in the FWRF herd following mortality of an adult captive ewe in fall 1995. Paired serum samples were collected from 34 bighorn sheep and 7 mule deer and submitted to the Southeastern Cooperative Wildlife Disease Study. EHD and bluetongue virus (BTV) titers were determined using serum neutralization. Facility/Maintenance/Repairs/Improvements A variety of scheduled and unscheduled maintenance and repair activities were necessary to support facility operation and ongoing research programs. We worked toward a conservation-oriented approach for facility care by undertaking preventive maintenance projects, and performing high-quality new construction and repairs to existing facilities. Facility repair and construction projects were prioritized based on animal welfare concerns and anticipated research needs. �105 Research Projects Facility operations offered support for pilot studies and for research projects conducted by CDOW personnel and other collaborators that were initiated, conducted, or continued using FWRF animals and facilities throughout the year. Educational Contributions Facility tours and educational lectures were provided to school, university, and professional groups visiting FWRF. We emphasized the importance of maintaining captive wildlife for performing controlled experiments and the contributions made by research projects performed at FWRF. FWRF animals and facilities were also used occasionally for hands-on training for professional groups. RESULTS AND DISCUSSION Animal Maintenance General: In FY1997, temporary, work-study, YCC employees and volunteers performed the majority of tasks of animal and facility maintenance at FWRF. Nine volunteers contributed 443.5 hr work to FWRF. These volunteers performed primarily caretaker tasks and also assisted in weighing and collecting samples from animals. contributions by volunteers represented a savings to FWRF of about 0.25 TFTE and $4124 (vs. cost of temporary employees). The animal welfare inspection by USDA APHIS revealed only one minor infraction from compliance. The error was corrected immediately. This finding highlights the high standards of animal care provided by FWRF employees and volunteers. At the close of FY1997, FWRF maintained 19 elk, 37 bighorn sheep, 11 whitetailed deer, 39 mule deer, and 16 pronghorn. During the year 11 bighorn sheep lambs and 14 viable mule deer fawns (one set of triplets died within 24 hr of birth) were born at FWRF. An additional six bighorn sheep were obtained from SWRU. Twelve natural moralities occurred (Table 1). Additionally, six whitetailed deer born unexpectedly in late September and October and two aggressive pronghorn bucks were euthanized as part of facility management. The USDA Animal Damage Control provided financial support for 11 white-tailed deer at FWRF. OVer the last 6 years, the number of elk, pronghorn, and bighorn sheep at FWRF has remained relatively stable. The number of mule deer has increased significantly and white-tailed deer were added to the captive herd. As a result of a decline in need for birds held in captivity for research purposes, waterfowl and upland birds are not currently maintained at FWRF. Although availability of research animals and facilities has significantly benefited the research program in the past, budgetary constraints dictate that future support of research animals at FWRF be dependent solely on the budgets of specific research projects. Therefore, in the future, descriptions of animal maintenance will be included in those projects' progress reports. �106 Nutritional Maintenance Feeding protocols: Individuals in all species maintained reasonable body condition on available diets. Subjective evaluation of clinical health and body condition suggested that supplementation of mule deer diet with browser maintenance pellets may be beneficial; however, confounding effects of age, sex, and occurrence of disease (chronic wasting disease and failure to thrive syndrome) preclude analysis. Although feed costs have not been reduced over the last 6 years, the quality and appropriateness of diets have improved (Wild 1995). Waste associated with feeding alfalfa hay has been reduced markedly by feeding cubed alfalfa to elk and grass hay to bighorn sheep. Vitamin and mineral supplementation in pelleted feed has been altered to better accommodate FWRF captive ungulates (Wild and Graffam 1994). Health Maintenance ~, General: OVerall, captive wildlife maintained at FWRF remained healthy throughout the year. Chronic wasting disease (CWO) has emerged as a significant. source of mortality in captive mule deer (see below). General animal health has improved (Wild and Schaefer 1996) and unexpected deaths have declined over the last 6 years. Unexpected fawning occurred in four white-tailed deer between 29 September and 11 October 1996. Due to limited availability of pens, white-tailed deer bucks were housed with does except during the rut (late August-early March). As in previous years, bucks were returned to the doe pen about 10 days after their antlers were shed (antlers shed 26 February). This management decision was based on the belief that testosterone levels are extremely low when antlers are shed and therefore, males should be inactive sexually (especially these bucks that are inexperienced breeders). However, based on a 200 day gestation period, the first does were bred on 14 March and sexual activity likely continued through the month. The fawns were euthanized at about 24 hr of age because of their poor prognosis for survival and normal function and to avoid undue negative impact of a autumn/winter lactation on the does. In the future, white-tailed deer bucks will be houaed throughout the year with mule deer bucks to avoid a repeat of this situation. Interesting, however, this event does show that even after antlers are shed, viable sperm remain present and sexual drive continues even in inexperience breeders. Further, whitetailed deer does continue to exhibit estrous cycles into March. Chronic wasting disease: Despite the strict protocols, CWO was diagnosed in four mule deer (three bucks, one doe) during FY1997. Since the depopulation in 1985, 10 mule deer have died or been euthanized due to chronic wasting disease. Seven of 10 of these CWO deaths have occurred in April. Overall, 18 moralities occurred in the 43 mule deer housed at FWRF between July 1995 and June 1997 (excluding fawns born in June 1997). Nine of these moralities were due to CWO. Of 15 males, five died of CWO (33%), while four died of other causes. Of 28 females, four died of CWO (14%), while five died of other causes. The higher incidence of CWO in captive male mule deer may indicate a predilection for disease in males or increased transmission of CWO in one pasture where 12 of 15 male mule deer (and all CWO positive males) have been housed. �107 Epizootic hemorrhagic disease in bighorn sheep: Of the 34 bighorn sheep tested, four adult ewes had positive titers to EHD (1:320). One lamb had a positive titer to BTV-13. This suggests that two viruses may have been present at FWRF. None of the samples collected from FWRF mule deer were positive. Although uncommon, these results indicate that bighorn sheep may become infected with EHD virus. Facility Maintenance/Repairs/Improvements A conservation oriented approach to the use of utilities and services for FWRF has likely lead to a reduction in resource use relative to the size of the facility over the past 6 years. Analysis of changes in resource use are not possible because of confounding effects of facility size, intensity of use, and because prior to 1993, CDOW did not receive bills for electricity service which is the most significant utility consumed at FWRF (CSU unknowingly paid all FWRF electric bills). However, insulation of buildings, installation of automatic waterers in all west side pastures, revegetation of east side a~~s with drought resistant grasses, and employee awareness of conservation techniques has likely helped minimize use of electricity and water. Trash production has been minimized through more efficient feeding programs which produce less waste. As older portions of the facility are repaired and replaced, the need for unscheduled daily repairs appears to be decreasing. Although numerous specific facility modifications were budgeted for and performed, the costs of routine repair and maintenance have been reduced about 47% over the last 6 years. Maintenance projects continue to be important for animal safety and facility function. In the last 6 years FWRF has been improved in many ways, but the most significant inciude the addition of a surgical suite, renovation of the office space, expansion of animal pens, and improvement of animal handling facilities. Significant maintenance/repair/improvement projects completed at FWRF this year included: - Paint and repair gates and animal shelters on east side Replace visual barrier on pen fences as needed Installation of insulation in the shop and east side lab Seal roofs on office/shop and labs Pour concrete pads for waterers on east side Repair and replacement of plumbing in east side lab Removal of composted hay stems and other wastes from pen cleanings Construction of wing fence in pen E4 Removal of y-maze structure in pen E2 Construction of four creep-feed areas and four covered feeders in mule deer pens Replace east side scale Repairs to old roofs after damage from extreme winds Although a centralized management approach for FWRF is likely the most effective, budgetary constraints dictate that in the future facility maintenance, repair, and improvements be performed and funded by specific research projects. Therefore, in the future, descriptions of facility maintenance will be included in those projects' progress reports. �108 Research Projects During the last 6 years, FWRF has provided animals and support in conducting numerous research projects involving elk, bighorn sheep, pronghorn, mule and white-tailed deer, and waterfowl. In addition to ongoing facility management experiments and improvements described above, the following pilot studies and research experiments were initiated, conducted, or continued using FWRF animals and facilities this year: Feasibility of using liposomes as deer immunocontraceptive carriers--L. Miller and B. Johns (ADC). - Evaluation atipamezol of medetomidine and ketamine immobilization in elk--M. Miller, W. Lance. - Heart rate as a potential and D. Baker. ~~ Educational and reversal with indicator of stress in bighorn sheep--M. - Evaluation of antemortem tests to diagnose cervids--M. Wild, M. Miller, T. Spraker. - Etorphine immobilization W. Lance. oral vaccine with naltrexone chronic wasting disease reversal Wild in in elk--M. Miller and Contributions FWRF provided formal educational instruction for special interest grade school through university groups. Numerous other informal tours were provided individually to visiting professionals. Youth in Natural Resources - CSU Veterinary Students in Summer Research Program CSU Pre-vet Symposium - CSU Wildlife Techniques class - Loveland High School zoology class Project WILD teacher training class Front Range Community College forestry and wildlife class - Blevins Junior High School zoology class-mentoring program - Several special interest grade school groups LITERATURE CITED Baker, D. L. and N. T. Hobbs. 1985. Emergency feeding of mule deer during winter: tests of a supplemental ration. J. Wildl. Manage. 49:934-942. Miller, M. W. 1990. Animal and pen support facilities for mammals research. Colorado Div. Wildl. Res. Rep., WP1a, J1, Jul 1989 - Jun 1990, Fort Collins. Wild, M. A. Colorado Collins. 1993. Animal and pen support facilities for mammals research. Div. Wildl. Res. Rep., WP1a, J1, Jul 1992 - Jun 1993, Fort �109 Wild, M. A. Colorado Collins. 1995. Anima.l and pen support facilities for manunals research. Div. Wildl. Res. Rep., WP1a, Jl, Jul 1994 - Jun 1995, Fort Wild, M. A, and W. S. Graffam. manunals research. Colorado Jun 1994, Fort Collins. 1994. Animal and pen support facilities for Div. Wildl. Res. Rep., WPla, J1, Jul 1993 - Wild, M. A, M. W. Miller, B. J. Maynard, and D. R. Magnuson. 1992. Animal and pen support facilities for manunals research. Colorado Div. Wildl. Res. Rep., WPla, Jl, Jul 1991 - Jun 1992, Fort Collins. Wild, M. A., and J. L. Schaefer. 1996. Animal and pen support facilities for manunals research. Colorado Div. Wildl. Res. Rep., WPla, Jl, Jul 1995 - Jun 1996, Fort Collins. Table 1. Summary Species Bighorn of mortalities A82 L97 14 0 Neoplasia Trauma Ea93 R693 Rb91 Rc91 X86 Y92 3 3 5 5 10 4 Chronic wasting Capture myopathy Chronic wasting Chronic wasting Pneumonia Chronic wasting 236 Da95 Db95 Jo88 095 Sb95 12 1 1 8 1 1 Old age changes Euthanized - aggressive Euthanized - aggressive Lumpy jaw; brain abscess Trauma Trauma Bb96 K96 L96 Ta96 Tb96 Euthanized 0 Euthanized 0 Euthanized 0 Euthanized 0 Euthanized 0 Euthanized Pronghorn Ba96 FY 1997 Cause Mule deer White-tail at FWRF during Age (yrs) Animal sheep in hoof stock 0 ID of Death - pleochromocytoma disease disease disease disease ��111 Colordo Division of Wildlife Wildlife Research Report July 1997 Job Progress state of Project Report Colorado No. ~W~-~1~5~3~-~R~-=-1~0~ _ Mammals Research Work Plan No. Hultispecies Job No. Mammals Inyestigations 2 Research Administration ~, Period Covered: Author: July 1, 1996 - June 30, 1997 R. Bruce Gill Personnel: R. Bruce Gill and Diane K. Haerter ABSTRACT Plans were developed, research on mammalian Research research Mammals issues. approved, and budgeted for 9 projects relating species other than deer, elk, and moose. and administrative staff members. Research support staff provided was provided technical input to Mammals into several to 1 and Mammals ongoing 2 policy Project personnel contributed to news releases and public information programs to inform the public on various wildlife management and policy issues. ��113 Mammals 2 Research Administration R. Bruce Gill P.B. Objectiye Administer research studies within the Mammals productivity and the lowest cost. Segment 1. 2 Unit for the highest Objectives Lead and administer research on mammalian species in the Mammals Research Program other than deer, elk, and moose. RESULTS Research and technical activities of six full-time employees were supervised during the segment. In addition contract services were obtained and supervised for 2 additional research projects. Results included: • Plans were developed, approved, and budgeted for 9 projects relating to research on monitoring and managing wildlife health in Colorado; managing infectious diseases in mountain sheep populations; experiments with the efficacy of lungworm treatment in mountain sheep populations; pronghorn population performance; pronghorn population estimation; mountain goat dispersal dynamics; experimental black bear inventory; kit fox status; and swift fox status. Work was satisfactorily funded projects. completed and reported for all federally • Research and administrative support was provided to. Mammals 1 and Mammals 2 research staff members. captive mule deer, white-tailed deer, elk, mountain sheep, and pronghorn and experimental holding facilities were maintained to augment various research projects requiring handling and close observation of experimental animals. Expenditures were encumbered and budget status was tracked using the Colorado Financial Reporting System. Support staff assisted in the recruitment and administration of temporary employees and volunteer staff. • Mammals Research staff provided technical input into several ongoing policy issues including the management of wildlife predation on domestic livestock, swift fox and kit fox recovery; black bear damage prevention/mitigation; pronghorn hunting regulations; diagnosis and management of chronic wasting disease; alternative livestock regulations; coyote season regulations; and trapping regulations. • Project personnel contributed to news releases and public information programs to inform the public on various wildlife �114 management and policy issues. In addition, project personnel participated as invited speakers in over two dozen professional and non-technical meetings, symposia, and public forums. Prepared by: R. Bruce Gill Mammals Program Leader �115 Colorado Division Wildlife Research July 1997 of Wildlife Report JOB FINAL REPORT State of Project Work Colorado No. W-153-R-IO Mammals Plan No. Multispecies Job No. e~Covered: Research Inyestigations Monitoring and Managing Wildlife Health in Colorado July 1, 1996 - June 30, 1997 Authors: M. W. Miller Personnel: W. J. Adrian, J. Bredehoft, G. Byrne, A. L. Case, D. Clarkson, M. Cousins, B. Davies, H. Dietrick, R. Forde, D. Freddy, T. Fulk, D.M. Getzy, K. Green, J. Jackson, K. Kinney, S. Kolus, M. Lamb, M. Leslie, C. Leonard, R. Mason, K. Madriaga, C.W. McCarty, C.A. Mehaffy, B. Olmstead, J. Ritchie, G. Schoonveld, H. Spear, M.L. Stevens, R. Spowart, T. R. spraker, S. Tracy, W. Travnicek, C. Wagner, E. S. Williams, and E. Zimmerman ABSTRACT Wildlife populations throughout Colorado were monitored for occurrence of disease using a combination of extensive and intensive approaches. We continued to develop and modify a statewide surveillance program for acquiring, examining, reporting on, and summarizing sporadic wildlife disease cases occurring throughout Colorado. At least 82 carcasses and/or tissue samples representing 73 wildlife cases were submitted for diagnostic examination during July 1996-June 1997. Trauma, brain abscesses, locoism, bronchopneumonia, nonsuppurative meningitis/encephalitis, contagious ecthyma, keratoconjunctivitis, and neoplasia were detected among ruminant submissions (chronic wasting disease cases are now reported separately under Work Plan 2, .Job 17). Trauma was the cause of death in all carnivore cases. Tularemia was diagnosed in a beaver. About 250 snow geese were lost during an apparent epornitic of colibacillosis in southeastern Colorado, and sporadic salmonellosis cases in passerine birds continued through August 1996. Other mammalian and avian cases appeared to represent isolated incidents or unusual maladies. For 26 cases, cause of death could not be determined from samples submitted. Aside from locoism in South Park elk, salmonellosis in songbirds, colibacillosis in geese, and enzootic occurrences of pasteurellosis and contagious ecthyma in bighorn sheep described previously, all cases completed to date appeared to represent isolated cases of trauma, intoxication, or disease. We continued developing a generalized, stochastic, individual-based simulation model of infectious disease in wild ungulate populations. We constructed �116 models for 2 wildlife disease problems of current interest: chronic wasting disease in a free-ranging mule deer population and bovine tuberculosis in a Michigan white-tailed deer population. Preliminary results of chronic wasting disease modeling suggest lateral transmission is probably an important component of maintaining this disease in a free-ranging population and that predicted impacts on poulation performance are substantially reduced if observed differences in prevalence between sexes (males » females) prove correct. Preliminary results of bovine tuberculosis modeling suggest assumptions for scenarios where Mycobacterium bovis first infected the Michigan wild white-tailed deer population >30 yrs ago are more plausible than assumptions required for scenarios where tuberculosis was introduced <10 yrs ago, and that management strategies directed at reducing deer densities alone are unlikely to eliminate tuberculosis from the affected population within the next 25 yrs. Although this document is designated as a "Job Final Report", activities previously undertaken in Work Plan la, Job 6 will continue next segment under .orit""P'!ckage7610 • �117 MONITORING AND MANAGING WILDLIFE HEALTH IN COLORADO M.W. Miller P. N. OBJECTIVES Develop and implement a program for enhancing statewide efforts manage health of Colorado's terrestrial wildlife populations. to monitor and AGREEMENT OBJECTIVES 1. Modify and improve systems for submitting, diagnosing and reporting sporadic disease cases in wild animals throughout Colorado. on 2:...#, v,elop and use databases for assimilating and analyzing data on and/or guiding management of wildlife disease problems identified through surveillance and surveys. 3. Provide- assistance in investigating outbreaks in Colorado. and managing wildlife disease Maintaining healthy wildlife populations is a fundamental component of sound wildlife management practices. Habitat degradation, high animal density, extreme weather, and disease can act singly or in combination to compromise the overall health of a wildlife population. As Colorado's wildlife managers, we have developed a variety of tools for monitoring and assessing the effects of habitat loss, animal numbers, and weather on wildlife populations. We have also invested considerably in developing tools to manage these factors to optimize performance of the wildlife populations in our stewardship. In contrast, monitoring and managing the effects of disease on wildlife population performance have received relatively little attention (with a few notable exceptions). This lack of attention may be rooted to some extent in a widely-held belief that wildlife diseases are symptoms of larger underlying population problems that will be resolved if those larger problems are managed properly. Despite this belief, disease can be a significant obstacle to effective and efficient wildlife management in Colorado. Disease outbreaks account for substantial mortality in some wildlife populations. Introduced pathogens have potential to decimate local wildlife populations. Some diseases depress wildlife population performance to levels below resource-based carrying capacity. Many wildlife diseases are shared with domestic animals and/or humans, and in some cases wildlife populations serve as reservoirs for these agents. Disease also detracts from the aesthetic value of wild animals, and may convey a perception of mismanagement to uninformed publics. For these reasons, diseases should be regarded as an integral part of wildlife population dynamics and wildlife management. Select wildlife health problems have been monitored in Colorado for more than 30 years. These longstanding efforts have provided useful information on the diseases studied. However, because these efforts have not always been coordinated on a statewide basis, and because some findings have not been widely available to managers and policy makers, applications to overall �118 management programs have been limited. In order to improve our collective ability to manage wildlife health in Colorado, we need a more coordinated and systematic approach for monitoring, investigating, and reporting on health problems in free-ranging wildlife. A more complete understanding of wildlife diseases and their effects on population performance is fundamental to comprehensive wildlife management. Enhanced surveillance efforts will provide a mechanism for detecting health problems throughout the state before serious impacts to wildlife populations occur. Assimilating diagnostic data will aid in assessing trends suggestive of population-level disease problems. Programs for conducting extensive and intensive surveys for potential and realized wildlife diseases will provide reliable prevalence and distribution data for managers and administrators to use in decision making. Expertise in investigating and managing epizootics and epornitics will ameliorate efficacy and efficiency of efforts to control outbreaks. Improved techniques for diagnosing and studying wildlife diseases wi~~.?vide a firm foundation for health management programs designed to enhance the quality of Colorado's wildlife populations. MATERIALS AND METHODS Disease Surveillance We monitored wildlife populations throughout Colorado for occurrence of disease using a combination of extensive and intensive approaches. These were organized and conducted as follows: statewide Surveillance: We continued to develop and modify a program for acquiring, examining, reporting on, and summarizing sporadic wildlife disease cases occurring throughout Colorado. All carcass submissions were subjected to necropsy. Ancillary diagnostics, including histopathology, bacteriology, virology, serology, parasitology, and toxicology were performed at the discretion of CDOW personnel and/or the attending pathologist. preliminary examination and/or test results were telephoned or faxed to CDOW's Wildlife Research Center Laboratory, usually within 3-5 days of completion, and a final report were usually provided within 15 business days of submission. Copies of reports were filed and sent to appropriate field personnel. Pertinent data from preliminary and final reports, including species, age, sex, location, number affected, diagnosis, and other information (as available) were entered into a permanent computerized database. This database was used to generate quarterly and annual wildlife morbidity and mortality reports. In addition, data are available for analysis of long-term trends in select wildlife disease problems. Surveys: No surveys were conducted in this segment. Disease Investigations Assistance was provided in investigating a contagious Loveland during December 1996-January 1997. ecthyma epizootic near Experimental Approaches Epizootic modeling (C.W. McCarty and M.W. Miller): We continued developing a generalized, stochastic, individual-based simulation model of infectious disease in wild ungulate populations. We plan to use this model in predicting consequences of disease introductions, improving understanding of the epizootiology of select disease problems, and evaluating potential disease �119 management strategies. In this model, populations display density-dependent sigmoid growth in the absence of disease or other limiting processes. We employed a novel mathematical approach for estimating pathogen transmission within simulated populations, and assumed transmission propabilities are a function of prevalence. We constructed models for 2 wildlife disease problems of current interest: chronic wasting disease in a free-ranging mule deer population and bovine tuberculosis in a Michigan white-tailed deer population. A manuscript describing our novel mathematical treatment transmission as applied to epizootiology of tuberculosis was submitted to the Journal of Wildife Diseases. RESULTS .~, 1sease of disease in white-tailed deer AND DISCUSSIQH . Surve111ance Statewide Suryeillance:Wildlife populations throughout Colorado were monitored for occurrence of disease using a combination of extensive and intensive approaches •. We continued to develop and modify a statewide surveillance program for acquiring, examining, reporting on, and summarizing sporadic wildlife disease cases occurring throughout Colorado. At least 82 carcasses and/or tissue samples representing 73 wildlife cases were submitted for diagnostic examination during July 1996-June 1997. Trauma, brain abscesses, locoism, bronchopneumonia, nonsuppurative meningitis/encephalitis, contagious ecthyma, keratoconjunctivitis, and neoplasia were detected among ruminant submissions (chronic wasting disease cases are now reported separately under Work Plan 2, Job 17). Trauma was the cause of death in all carnivore cases. Tularemia was diagnosed in a beaver. About 250 snow geese were lost during an apparent epornitic of colibacillosis in southeastern Colorado, and sporadic salmonellosis cases in passerine birds continued through August 1996. other mammalian and avian cases appeared to represent isolated incidents or unusual maladies. For 26 cases, cause of death could not be determined from samples submitted. Aside from locoism in South Park elk, salmonellosis in songbirds, colibacillosis in geese, and enzootic occurrences of pasteurellosis and contagious ecthyma in bighorn sheep described previously, all cases completed to date appeared to represent isolated cases of trauma, intoxication, or disease. We will continue adding new accessions throughout the coming fiscal year to our computerized database for diagnostic case information, as well as data from archived reports as they become available. Surveys: Disease No surveys were conducted in this segment. Investigations Bighorn sheep inhabiting the lower Bigh Thompson Canyon west of Loveland showed signs and lesions of contagious ecthyma during December-January. Although most animals observed appeared affected to varying degrees, no associated mortality was reported. Affected animals recovered spontaneously. Attempts to demonstrate Parapox virus via electron microscopy were unsuccessful. Consequently, these presumptive diagnoses were based on clinical, gross, and histological evaluations. �120 Experimenta1 Approaches Epizootic modeling: Preliminary results of chronic wasting disease modeling suggest lateral transmission is probably an important component of maintaining this disease in a free-ranging population and that predicted impacts on poulation performance are substantially reduced if observed differences in prevalence between sexes (males » females) prove correct.. Preliminary results of bovine tuberculosis modeling suggest assumptions for scenarios where Mycobacterium bovis first infected the Michigan wild white~tailed deer population >30 yrs ago are more plausible than assumptions required for scenarios where tuberculosis was introduced <10 yrs ago, and that management strategies directed at reducing deer densities alone are unlikely to eliminate tuberculosis from the affected population within the next 25 yrs. Although this document is designated as a ~ob Final Report~ activities previ:ously undertaken in Work Plan la, Job 6 will continue next segment Work Package 7610. kr~ under dgments The statewide wildlife health monitoring and surveillance program described above relies heavily on efforts of dedicated field personnel throughout the Colorado Division of Wildlife, and truly represents a division-wide effort to improve our'understanding and management of wildlife disease problems. In addition to those specifically listed, we collectively thank all of those regional and area biologists, district and area wildlife managers, and others who assisted by submitting diagnostic cases throughout the year. Prepared by ~ Wildlife Research Veterinarian .Table 1- At least 82 carcasses and/or tissue samples representing 73 wildlife cases were submitted for diagnostic examination during July 1996-June 1997. FY1996-1997 DATE 06-14-96 06-19-96 06-24-96 07-11-96 07-12-96 08-02-96 08.:05-96 08-08-96 08-09-96 08-29-96 09-06-96 09-09-96 09-11-96 SPECIES mdeer mdeer geagle rabbit ovine suffolk elk red crossbill bat mdeer mdeer bighorn sheep e~ mdeer DIAGNOSTIC AGE SEX REGION 7+ f f f na nw nw sw nw na na na f m na m m se se ne se w ne ne ne f a 2mo 1.5 na na na na a na 1-2 a SUBMISSIONS CAUSE OF DEATH suggestive of toxoplasma canine bite wounds blunt trauma to heart unknown mtn. lion or coyote bites locoism salmonellosis neg, rabies . unknown bronchopneumonia . shot loco poisoning? trauma-wasting disease ACC# 956-31747 956-32167 956-32735 967-01272 967-01852 967-03630 967-03912 967-04216 967-04286 967w1094 967-07093 967-07219 967-07605 --------------------------------------------------------------------------------------------- �121 Table DATE 09-12-96 09-12-96 09-12-96 09-17-96 09-20-96 09-26-96 09-30-96 10-04-96 10-08-96 10-11-96 10-15-96 10-18-96 10-28-96 l. Continued. SPECIES 12-31-96 deer deer deer mdeer goshawk mdeer mt.lion mdeer mdeer kestrel mdeer deer mdeer elk elk mdeer bh sheep mdeer mdeer moose magpie llama mdeer beaver elk bh sheep canine bhsheep mdeer raccoon elk elk mdeer elk 01-01-97 01-06-97 01-07-97 01-08-97 mdeer bald eagle mdeer mdeer 01-08-97 01-08-97 01-08-97 snow goose 8 mallard ducks elk ~, 11-04-96 11-07-96 11-22-96 11-24-96 11-26-96 12-03-96 12-03-96 12-06-96 12-10-96 12-13-96 12-13-96 12-13-96 12-17-96 12-18-96 12-19-96 12;.30-96 12-30-96 01-24-97 02-03-97 02- -97 02-02-97 02-02-97 02-10-97 02-14-97 02-17-97 snow goose elk mdeer elk elk . 2 avian duck bhsheep bhsheep AGE a y f f SEX REGION CAUSE OF DEATH ACC# f f w w w ne ne ne trauma starvation suppurative nephritis unknown trauma no medical cause peritonitis \ wasting disease gunshot wasting disease unknown not wd results ? neg. wasting disease wasting disease unknown wasting disease unknown lab results bact report unknown. undetermined fibromuscular intimal proliferation unknown lion kill abcess on the lung tularemia peritonitis pasteurella shot after attacked man shotunknown wasting disease killed by animal lymph nodes normal wasting disease wasting disease leptomeningitis, cerebral thromosis assoc. w/mycotic infection trawnacar toxinDDE septic abscess of jaw chronic conjunctivitis and chronic necrosuppurative keratitis e coli (?) unknown bilateral retrobulbar malignant lymphoma acute necrotizing enteritis wyo. lab report wasting disease nonsuppurative meningitis wasting disease(head) unknown e.coli? unknown HE stain pregnant 967w1119 967wl120 967w1116 967-08184 967-08560 967-08987 967-09342 967-09873 967-10156 967-10553 967-10831 967-11127 697-11938 967-11940 967-12545 967-11938 967W1209 96wl0650 967-14562 967-15125 967-15124 967-15372 967-15754 967-16042 967-16046 967-15959 967-16274 967-16353 967-16416 967-16895 967-16894 967-15513 967-15489 2.5 5 a m f f a f 4/5 a 5mo m f m ne w ne ne ne ne se w a a a f m m f f m -5 4-5 yng m m f a. a a f m f a ynga a a m m f m a 3m, 5f f ne .w ne ne ne ne w se ne ne ne w ne ne w ne ne w se w w se a 4+ 15+ 10+ m f f 10+ a f f ne ne ne ne w 967-17134 967-17171 967-17643 967-17782 967-17916 967-17914 967w1259 967-17915 967-19346 W833 967-16908 967-20265 967-20266 967-21219 967-21626 97W479 --------------------------------------------------------------------------------------------- �122 Table DATE 1. Continued. SPECIES 02-17-97 02-25-97 03- -97 bhsheep elk mdeer 03-03-97 03-03-97 03-04-97 03-05-97 elk deer goose can. house finch 03-00-97 wtdeer 03-00-97 mdeer 3~7._., AGE -9mo a SEX m f REGION W ne -1 f f ne ne ne elk 03-16-97 03-17-97 elk .bh sheep 2+ f f ne 03-19-97 03-21-97 03-21-97 03-22-97 03-24-97 03-25-97 10+ a 1~ a yng 4 f m f f ne ne ne ne ne ne 04-5-97 04-7-97 04-10-97 04-11-97 04-13-97 04-16-97 04-16-97 04-16-97 04-16-97 04-21-97 04-23-97 04-26~97 04-28-97 04-30-97 04-30-97 04- -97 05-05-97 elk elk elk mdeer red-thawk mdeer mtn. lion elk g-howl mdeer mdeer mdeer wtdeer pocket gopher elk barn owl mdeer bhsheep deer raccoon elk elk elk elk 5+ a 2 8-10 3-4 f m f m m 05-08-97 05-09-97 05-16-97 bhsheep deerm deerwt 05-17-97 05-21-97 deer deerm 05-27-97 06-19-97 elk raccoon a 10mo f ·f f m f 1.5 ne ne ne se w ne ne w ne ne ne f 10+ f 13+ 2 m 1 f f ne f f ne m CAUSE OF DEATH bronchopneumonia neg. brain stains leg from arsenal chonic dermatophyte infection of the hair . modert autolysis 2 brains neg. brain staining presumptive lead poisoning proliferative conjunctivitis avian poxvirus infection head~191031097 neg. for wasting disease head#mdm02031097 neg. for wasting disease head#em009031297 pos. for wasting disease peritonitis fibrinour bronchopneumonia verminous pneumonia old age peritonitis and emaciation wasting disease wasting disease trauma wasting disease head neg to wasting disease wasting disease trauma head wasting disease severe parasitism wasting disease unknown unknown encephalitis\bronchopneumonia emaciation trauma. not wd chronic bronchopneumonia severe parasitism euthanasia. severe burns head unknown neg. wd head unknown neg. wd head unknown not wd head ef020-050497 neg. for wasting disease unknown MDF020050997 diagnosis inconclusive 3WfFOO9051697 unknown. no wasting dis. blunt trauma MDF029052197 no wasting disease may be a bacteroal septicemia neg wasting disease cruelty complaint acid death? ACC# 967-21844 967-22796 967-24002 967-23439 967-23506 967-23440 97w1239 967-24913 %7-24914 967-24915 967-24755 967-1360 967-25151 967-25485 %7-25486 967-25563 967-25689 967-25717 %7-25859 967-27061 967-27082 967-27659 967-27779 967-27851 967-28277 967-28279 967-28241 967-28278 967-28678 967W1458 967-29306 967-29502 967-28010 %7-30280 %7-30776 %7-30992 %7-31771 967-31809 %7-3Z235 %7-32708 %7-35335 . �123 Colorado Division Wildlife Research July 1997 of Wildlife Report JOB state of PROGRESS REPOR~ Colorado Project No. W-153-R-I0 Mammals Research Work Plan No. 2A Mountain Job No. Sheep Inyestigations strategies for Managing Infectious Disease in Mountain Sheep Populations ~, Period Covered: July 1, 1996 - June 30, 1997 Authors: M. W. Miller Personnel: B. J. Kraabel, and H. J. McNeil A. L. G. Lotto, N. DuTeau, and M. D. Salman ABSTRAC~ We used ribosomal RNA fingerprinting and in vitro measures of leukotoxin production to compare Pasteurella haemolytica isolates from eight indigenous Rocky Mountain bighorn sheep herds in Colorado. Using ribosomal RNA gene restriction patterns, at least 26 distinct strains of P. haemolytica were identified among isolates (n = 59) from these herds; we identified one to seven distinguishable ribotypes within individual herds. Of the 26 ribotypes identified, 21 appeared unique to individual herds, four others (E, N, T, BB) were shared by only two herds, and one (A) was common to 3 herds. In vitro evaluation of leukotoxin production by genotypically-distinct P. haemolytica isolates revealed further differences among strains: 4 ribotypes (AA, B, E, 0) showed marked leukotoxin production -- bighorn neutrophil death rates @ 150 ~g culture supernatant were 4-9 times those of an Enterobacter sp. control (7.1±0.4% neutrophil death @ 150 ~g supernatant); cytotoxicity of the other 22 strains examined approximated control levels. These findings support hypotheses that strains of P. haemolytica carried by healthy bighorn sheep may vary within and among wild populations. These isolates are being further compared and characterized via PCR. We continued investigations of multivalent Pasteurella haemolytica vaccines in captive bighorn sheep (Ovis canadensis). Both strain selection and blending procedures may have influenced antigenicity of our modified Pasteurella haemolytica supernatant vaccine. A vaccine lot that combined bighorn and domestic strains appears to contain antigen copmarable to that included in the original multivalent vaccine. Data from ~ilot studies in rabbits suggest this most recent combination of domestic and bighorn strains may approximate antigenic stimulation provided by the original vaccine. However, further testing in bighorn sheep will be necessary before the modified vaccine can be reliably incorporated into delivery system and field evaluations. ��125 EXPERIMENTS TO IDENTIFY AND MANAGE STRESS IN MOUNTAIN SHEEP POPULATIONS M. W. Miller and B. J. McNeil P. N. OBJECTIVE To develop strategies for managing population performance. infectious diseases affecting bighorn sheep SEGMENT OBJECTIVES 1. Analyze and report on data comparing genotypic and phenotypic characteristics of Pasteurella spp. isolates among different indigenous ~ighorn populations. 2. Analyze and report on data evaluating efficacy of a multivalent haemolytica vaccine in protecting captive bighorn sheep from challenge with pathogenic Pasteurella haemolytica. Pasteurella 3. 4. Refine experimental Pasteurella haemolytica vaccine to incorporate cytotoxic field isolates identified through ongoing epizootiological investigations. Design and begin conducting experiments evaluating delivery of modified haemolytica vaccine to captive bighorn sheep via biobullet implant and/or oral poly (methylacrylic acid) hydrogel. Pasteurella [Objectives 5 and 6 were included in the original were not funded under Project W-153-R-IO.] draft Segment Narrative, but MANAGEMENT OF BACTERIAL AND VIRAL DISEASES IN MOUNTAIN SHEEP POPULATIONS Inability to control infectious disease outbreaks and subsequent mortality in mountain sheep populations represents a significant obstacle to long-term success in their management. Although the "bighorn pneumonia complex" has been studied intensively for over 3 decades, little is known about many aspects of its etiology and epizootiology. Moreover, management interventions recommended for preventing or controlling this problem remain untested. Although viral, bacterial, and parasitic agents have all been incriminated in these outbreaks, Pasteurella spp. are perhaps the most common pathogens associated with bronchopneumonia in bighorns. Two' species, P. haemolytica and P. multocida, and several biotypes and/or serotypes within those species, have been isolated from bighorns during epizootics. Unfortunately, despite extensive diagnostic and experimental investigation, the epizootiology of pasteurellosis in wild bighorn populations is poorly understood. In the absence of knowledge about the epizootiology of pasteurellosis, effective strategies for managing pneumonia in bighorn populations have not emerged. Here, we report on a series of ongoing research stUdies designed to improve knowledge about various aspects of pasteurellosis epizootiology and management in bighorn sheep. �126 METHODS AND MATERIALS Management of Bacterial and Viral Diseases in Mountain Sheep Populations In conjunction with numerous cooperators, we continued developing and improving tools available for use in studying etiology, epizootiology, and prevention or control of disease outbreaks in bighorn populations: Epizootiology of pasteurellosis in indigenous bighorn populations (Miller, Spraker, Mills, Snipes, and Kraabel): We used ribosomal RNA fingerprinting and in vitro measures of cytotoxin production to compare Pasteurella haemolytica isolates from eight indigenous Rocky .Mountain bighorn sheep herds (Almont/Taylor River, Avalanche Creek, Chalk Creek, Cottonwood Creek, Grant, Tarryall Mountains, Texas Creek, Waterton Canyon) in Colorado. Genomic fingerprinting (Snipes et al. 1992) of remaining untyped isolates (n ~ 50) was completed. We also continued evaluating potency of cytotoxins derived from genotypically-distinct P. haemolytica isolates in vitro using methods described by Silflow et al. (1993). ~~, Differentiation of potentially pathogenic from nonpathogenic Pasteurella isolates (Lotto, DuTeau, Miller, and Salman): We continued examining the ability of 8 primer pairs chosen from the leukotoxin A region of P. haemolytica to differentiate between cytotoxic and noncytotoxic strains of P. haemolytica. We performed polymerase chain reaction (PCR) procedures on 9 primers in 8 different pairing combinations: 2 upper and 2 lower primers amplified areas of the leukotoxin promoter region; 2 upper and 3 lower primers amplified parts of the leukotoxin A coding region. haemolytica Efficacy of a multivalent Pasteurella haemolytica toxoid-bacterin in protecting captive bighorn sheep (Ovis canadensis) from challenge with pathogenic Pasteurella haemolytica (Kraabel, Miller, Conlon, McNeil, and Bulgin): We prepared a draft manuscript describing results of last segment's challenge trial; that manuscript is currently in review for publication in the Journal of Wildlife Diseases. Refinement of experimental Pasteurella haemolytica vaccine (McNeil, Miller, and Shewen): Three strains of Pasteurella haemolytica obtained from previous field investigations were cultured according to standard methodology in order to assess the production of toxin in a cytotoxicity assay. Although all three strains resulted in poor cytotoxicity results relative to the standard domestic A1 strain, we proceeded with vaccine synthesis to determine protein profiles of individual strains. All strains were grown in a 2 litre fermenter until the desired optical density was reached. The supernatant was then filtered, dialyzed and resuspended 20 times concentrated in sterile water for injection. This material was kept frozen until the vaccine was blended. Concentrated material was then used to compare the protein profile with that of the original multivalent P. haemolytica vaccine used successfully in bighorns. Electrophoretic gels were run and then transferred to run Western blots. The material was first blotted with standard bovine anti-Po haemolytica sera. The resulting profiles were encouraging enough to proceed with blending vaccine. Blended vaccine was then used in a rabbit study in order to confirm antigenicity before use in bighorn sheep. Three rabbits were given the original vaccine used previously in bighorns (lot 1940902) and three others received the newly blended vaccine. On day 0, all rabbits were bled and �127 and vaccine was administered (O.Sml) intramuscularly in the left haunch. On day 14 of the study, the rabbits received a second dose of vaccine. On day 21, blood was taken from all rabbits in order to establish if the sera contained neutralizing antibodies against P. haemoly~ica toxin. All rabbits had neutralizing titres of 0 for the prebleed. The three rabbits receiving the newly blended vaccine failed to respond, while the three receiving 940902 had titres of 3,4 and 6. Consequently, a third dose of vaccine was administered and rabbits were exsanguinated 14 days later. During the rabbit study, further Western blots were performed using a rabbit anti-sera raised in the laboratory against the LktA portion of P. haemoly~ica toxin. Monoclonal antibody (601B) generously provided by Rhone Merieux was also used to blot the same samples. One of the differences between 940902 and the new vaccine (other than the choice of isolates included) was the method of concentration. The original va~, had been concentrated using a tangential flow membrane. The . concentrated salts and dyes in the new vaccine may have had an effect on antigenicity. In an attempt to reconcile apparent discrepancies between original and modified vaccine performances, we modified our approach. A new lot of vaccine was blended that included two of the three CDOW strains (100, D2) as well as two other strains that were in the original vaccine (A2, T10). The supernatants were to be concentrated using a Millipore Minitan System w~th SOOOMWC membranes. Each isolate of Pas~eurella haemoly~ica was grown on a blood agar plate overnight in a 37°C incubator. The overnight culture was used to inoculate a small volume of Brain Heart Infusion Broth (BHIB) that was incubated on a shaking platform at 37°C for approximately 18 hours. This primary culture was used to inoculate 2 litre of RPMI in a 2L fermenter (to achieve a previously established optical density reading). The fermenter was run at 200rpm @ 37° C until the desired optical density was reached. The material was then centrifuged to remove the bacteria. The remaining supernatant was run through a Millipore Minitan System using SOOOMWC membranes until the material was approximately 20X concentrated. Concentrated material from each of the four fermenter runs was syringe-top filtered and stored at -20° C for further testing and vaccine blending. Samples of the membrane concentrated material were run on a 12% precast (PAGE) gel along with samples of the original components of the lot 940902 vaccine. Two identical gels were run. These gels were then transferred onto nitrocellulose for Western blots. One was blotted with a standard anti-Po haemoly~ica bovine serum and the other was blotted against a rabbit serum that had been vaccinated three times with vaccine lot # 940902. These protein profiles showed great similarity between the newly prepared concentrated samples and those used in the original vaccine (lot#940902). Based on these preliminary findings, the new vaccine was blended. Vaccine was blended according to the original vaccine (lot #940902) formulation. The vaccine is comprised of 71% total antigen (A2, 100, T10, D2), 23% Al(OH)3' 0.5% Thimerosol (2% solution), 1.5% NaHC03 (7.5% solution), 4% RPMI, and 0.3% QuilA (1.5% solution). Vaccine was sterility checked for bacterial contamination, then bottled in 30 ml bottles and labelled �128 "Pasteurella EXPERIMENTAL haemolytica USE ONLY". supernatant vaccine lot # 970520 2ml dose FOR Nineteen (19) full bottles were stored at 4°C. A second rabbit study was implemented in order to test the safety and antigenicity of vaccine lot # 970520. Three SPF rabbits were purchased and were housed in the Central Animal Care Facility. The animals were sedated and bled to obtain prevaccine sera and then were vaccinated with 0.5 ml of the vaccine in the right haunch. The rabbits received a booster dose of vaccine at day 14 of the study. The rabbits were test bled at day 21 and their sera was run in a leukotoxin ~eutralizing assay as well as in direct agglutination assays using P. haemolytica A2 and T10 as antigens. There was only a marginal neutralizing titre achieved in one of the three rabbits so a third booster was administered. At this time, two more PAGE gels were run using the same samples as the previously run gels. These were also transferred for use in Western blots. Once again, one was blotted with serum from a rabbit that had received 3 doses of~~940902 from the initial rabbit study. The other was blotted with the serum from the one rabbit that seemed to have responded to the lot #979520 vaccine. The protein bands that were recognized by both rabbit sera were very similar and· thus encouraging. The rabbits were bled again 10 days after receLvLng the 3rd dose of 970520 vaccine. These sera were run in the leukotoxin neutralizing assay, and this time all three rabbits were responding to vaccination. The rabbits were then bled out and the sera stored at -20 C. Deliyery of Pasteurella haemolytica vaccine to bighorn sheep (McNeil and Miller): We prepared a draft study plan describing an experiment to evaluate vaccine delivery systems in captive bighorn sheep. Initiation of that experiment is planned for next segment, pending outcome of a pilot study evaluating antigenicity of our modified Pasteurella haemolytica supernatant vaccine. RESULTS AND DISCUSSION Epizootiology of pasteurellosis in indigenous bighorn populations: Using ribosomal RNA gene restriction patterns, at least 26 distinct strains of P. haemolytica were identified among isolates (n = 59) from these herds; we identified one to seven distinguishable ribotypes within individual herds. Of the 26 ribotypes identified, 21 appeared unique to individual herds, four others (E, N, T, BB) were shared by only two herds, and one (A) was common to 3 herds. In vitro evaluation of cytotoxin production by genotypically-distinct P. haemolytica isolates revealed further differences among strains: Four ribotypes (AA, B, E, 0) showed marked cytotoxin production -- bighorn neutrophil death rate @ 150 ~g culture supernatant was 4-9 times that of an Enterobacter sp. control (7.1±0.4% neutrophil death @ 150 ~g supernatant) (Fig. 1); cytotoxicity of the other 22 strains examined approximated control levels. All three indigenous bighorn herds that yielded markedly cytotoxic P. haemolytica strains have recent histories of pneumonia epizootics: pasteurellosis outbreaks occurred in the Taylor River herd in 1979 and again in 1991, in the Waterton canyon herd in 1980, and in the Chalk Creek herd in 1981. One of these strains (E) was also recovered from dead bighorns during recent epizootics in the Alamosa Canyon (1989) and Rock Creek (1990) herds -- �129 these latter herds can be linked to Taylor River by bighorn translocation activities during the last decade. Our findings support hypotheses that strains of P. haemolytica carried by healthy bighorn sheep may vary within and among wild populations. Our preliminary results also suggest the combination of genomic fingerprinting and cytotoxicity determination may offer a useful approach for studying the epizootiology of pasteurellosis within and among bighorn herds and may provide insights into strategies for effectively preventing or managing pneumonia epizootics. We are currently preparing a manuscript summarizing the findings reported here. Differentiation of potentially pathogenic from nonpathogenic Pasteurella isolates (Lotto, DuTeau, Miller, and Salman): Among the 8 PCR primer paring combinations examined to date, results of the most successful primer pairs are described here. One primer pair (AMU/AEIL) that has yielded encouraging results amplifies a sequence near the middle to the end of the Lkt ~g region. Using this primer pairing, we performed several different thermal cycling techniques, including variations of both a "hot Fang" method (low initial annealing temperature that increases after a number of cycles) and a "modified stepdown" method (high initial annealing temperature that decreases after a number of cycles). haemolytica Promising results in differentiating cytotoxic from noncytotoxic P. isolates were obtained with both thermal cycling techniques. The hot Fang method consistently distinguished 2 cytotoxic controls from 2 noncytotoxic controls. In further testing with 5 other randomly selected isolates, results with 4 of 5 isolates were consistent with in vitro cytotoxin assay data gathered previously (M.W. Miller, unpubl. data). The modified stepdown method was also consistent in correctly distinguishing cytotoxic from noncytotoxic controls. Moreover, modified stepdown results were consistent with in vitro cytotoxicity data for all 5 randomly selected isolates. Another promising primer pair (AMU/AE2L) replicated a slightly smaller sequence than AMU/AE1L in an overlapping genomic region. Hot Fang thermal cycling methods employing AMU/AE2L yielded results consistent with cytotoxin assay data in classification of positive and negative controls as well as 5 out of 5 blindly chosen isolates. A third primer pair (PMU/PEL) that may also prove useful after further evaluation amplifies a region in the middle to the end of the leukotoxin promoter region. Using modified stepdown methods, results with this primer pair were consistent with cytotoxin assay data in classigying cytotoxic and noncytotoxic controls, as well as 3 out of 5 other isolates. haemolytica Based on these data, we believe that PCR can be used to distinguish cytotoxic from noncytotoxic strains of P. haemolytica isolated from free-ranging bighorn sheep. Further evaluation of these techniques on additional isolates will be completed next quarter. Efficacy of a multivalent Pasteurella haemolytica vaccine in bighorn sheep: protection from challenge with pathogenic Pasteurella haemolytica: We examined the efficacy of the multivalent Pasteurella haemolytica toxoid-bacterin (A1, A2, T10) in reducing morbidity and mortality in captive bighorn sheep (Ovis canadensis) associated with exposure to a pathogenic strain of P. haemolytica (biotype T, serotype 10, ribotype Eco; "Alamosa Canyon" strain). Fifteen captive bighorns were divided equally into 3 treatment groups based on vaccination status: control (no vaccination), 1 dose 10 days prior to challenge, or 1 or 2 doses 57 wk prior to challenge. At challenge, each �130 bighorn received about 5 ml (6.2 X 107 CFUs) of P. haemolytica suspension sprayed into the proximal trachea. Vaccination reduced mortality rates (P = 0.1) in bighorns vaccinated 10 days prior to challenge, as compared to controls; although mortality rates in bighorns vaccinated 57 weeks prior to challenge did not differ from controls (P ~ 0.2), a trend in reduced mortality was apparent. Necropsy scores were not significantly different between vaccinated animals and controls (P 5 0.22). Leukotoxin neutralizing antibody titers to P. haemolytica were elevated at challenge in bighorns vaccinated 10 days previously (P = 0.0034), and titers in bighorns from both vaccinated groups were elevated at postmortem 5 7 days after challenge (P 5 0.0044). In contrast, titers of agglutinating antibody to P. haemolytica serotype Al or T10 surface antigens did not differ between vaccinated and unvaccinated bighorns (P ~ 0.19). Based on these data, we believe that this experimental P. haemolytica vaccine is safe and can stimulate protective immunity from pneumonic pasteurellosis in bighorn sheep. Further evaluation of this vaccine as a tool in preventing and managing pasteurellosis in wild bighorn sheep ap~q" warranted. Refinement of experimental Pasteurella haemolytica vaccine (McNeil, Miller, and Shewen): Both strain selection and blending procedures may have influenced antigenicity of our modified Pasteurella haemolytica supernatant vaccine. Data from pilot studies in rabbits suggest the most recent combination of domestic and bighorn strains may approximate antigenic stimulation provided by the original multivalent vaccine. However, further testing in bighorn sheep will be necessary before the modified vaccine can be reliably incorporated into delivery system and field evaluations. Delivery of Pasteurella haemolytica vaccine to bighorn sheep (McNeil and Miller): A draft study plan is appended. This experiment will be initiated next segment, pending outcome of vaccine testing described above. Wildlife Research veterinarian �131 Appendix. A STUDY PLAN State of Project Work Colorado No. Package Task No. Cost Center W-153-R-ll No. Mammals 3004 3 3430 Program Management of Other Ungulates Strategies for Managing Pasteurellosis in Mountain populations Sheep EXPERIMENTAL EVALUATION OF ALTERNATIVES FOR DELIVERING MULTIVALENT Pasteurella haemolytica SUPERNATANT VACCINE TO BIGHORN SHEEP (OVis canadensis) Successful bighorn sheep (OVis canadensis) management appears dependent on eliminating pneumonia epizootics in otherwise thriving herds. Periodic pneumonia· outbreaks cause extensive mortality in bighorn sheep populations throughout North America (Rush 1927, Potts 1937, Marsh 1938, Buechner 1960, Forrester 1971, Feuerstein et al. 1980, Wishart et al. 1980, Foreyt and Jessup 1982, Onderka and Wishart 1984, Spraker et al. 1984, Schwantje 1986, Coggins 1988, Festa-Bianchet 1988, Miller et al. 1991a, CDOW unpublished data) . Although viral, bacterial, and parasitic agents have all been incriminated in these outbreaks, Pasteurella spp. are perhaps the most common pathogens associated with bronchopneumonia in bighorns. Two species (P. haemolytica and P. multocida), and several biotypes and/or serotypes within these species, have been isolated from bighorns during epizootics (Potts 1937, Marsh 1938, Post 1962, Feuerstein et al. 1980, Wishart et al. 1980, Foreyt and Jessup 1982, .onde rka and Wishart 1984, Spraker et al. 1984, Schwantje 1986·, Coggins 1988, Festa-Bianchet 1988, Onderka and Wishart 1988, Onderka et al. 1988, Foreyt 1989, Miller and Hobbs 1989, Miller et al. 1991a,b, M.W. Miller, unpublished data) . Despite extensive diagnostic and experimental investigation, the epizootiology of pasteurellosis in wild bighorn populations is poorly understood. Hypotheses regarding the ecology and epizootiology of pneumonia outbreaks in bighorn sheep have not been rigorously tested, and the relative importance of endogenous and introduced strains of P. haemolytica as factors limiting bighorn abundance remains particularly unclear (Miller et al. 1991a,b, Wild and Miller 1991, Hobbs and Miller 1992). In the absence of knowledge about the epizootiology of pasteurellosis, effective strategies for managing pneumonia in bighorn populations have not emerged. Wildlife managers can neither predict nor prevent outbreaks in bighorns. Consequently, long-term attempts to manage bighorns often fail. Difficulties in understanding and managing pasteurellosis also plague the domestic sheep industry world-wide. Differences in overall species susceptibility and strain-specific pathogenesis notwithstanding (Onderka and Wishart 1988, Onderka et al. 1988, Foreyt 1989, Silflow et al. 1991, 1993), the epizootiology of pasteurellosis in domestic sheep (reviewed by Gilmour and Gilmour 1989) is strikingly similar to that observed in bighorn sheep (Foreyt 1990, Miller et al. 1991a,b, Hobbs and Miller 1992). Because of its wide distribution and sporadic nature, recent attempts to manage pasteurellosis in. domestic sheep have focused on prevention through vaccination. The efficacies of vaccines developed for domestic sheep have varied widely (reviewed by Gilmour and Gilmour 1989), and many either fail to prevent or exacerbate disease. However, experimental vaccines containing cytotoxin and soluble cell �132 surface antigens from P. haemolytica offered ~86% protection against experimental challenge (Sutherland et al. 1989, J.A. Conlon, unpublished data); moreover, humoral immune responses stimulated by one of these vaccines (Sutherland et al. 1989) simulated those observed in lambs allowed to recover from experimental P. haemolytica infections (Donachie et al. 1986). The effects of a multivalent P. haemolytica supernatant vaccine (A1, A2, T10) on humoral and cellular immune responses of bighorn sheep has also been examined (Miller et al. 1995). Bighorns receiving the vaccine demonstrated marked elevations in both P. haemolytica cytotoxin neutralizing antibody titers and agglutinating antibody titers to serotype A1 surface antigens. Administration of this same multivalent vaccine also proved protective against intratracheal challenge with a pathogenic strain of P. haemolytica (Kraabel et al., unpublished data). Because captive bighorn sheep that survived pneumonic pasteurellosis have shown resistance during subsequent pneumonia epizootics (Miller et al. 1991b), it follows that vaccines stimulating immunity to P. haemolytica cytotoxin and soluble cell surface antigens might afford protection against naturally occurring pasteurellosis in bighorns as well. It is believed that an effective vaccine against P. haemolytica would represent a significant and needed advance 'n~~ife management by protecting herds from massisve die-offs caused by Past'eurella species, and the subsequent low lamb recruitment following the initial mortality (Foreyt 1992). Determining an acceptable way to administer such a vaccine to wild free-roaming populations of bighorn sheep provides the focus of this study. ' Conventional hand injection methods, although effective, are not desirable due to the stress on the animals and the man-power costs involved.' Bighorn sheep have effectively been treated orally with anthelmintics and antibiotics for many years (Coggins 1988, Feuerstein et al. 1980, Bailey 1990). New technology using poly(methacrylic acid) hydrogels for oral administration of vaccines to ruminants has been shown to have potential (Bowersock 1994a,b). The use of biobullets to administer drugs and vaccines is becoming more widely used due to its cost-effectiveness and seemingly minimal affect on the treated animals (DeNicola et al. 1996, Jessup et al. 1992, Herriges et al. 1991). It has been shown that a similar vaccine to the one described above (Presponse *, LangfordCyanamid, Canada) can be safely lyophilized and resuspended (McNeil, unpubiished data). Resuspended vaccine was administered to domestic lambs and shown to increase both neutralizing and agglutinating titres. We propose an experiment to evaluate and compare the serological responses of captive bighorn sheep to a multivalent P. haemolytica supernatant vaccine using conventional hand injection, biobullet delivery, and oral vaccination methods. B. OBJECTIVES The objective of this study is to compare and evaluate serological responses captive bighorn sheep receiving an experimental P. haemolytica supernatant vaccine via different delivery systems (hand injection, biobullets, oral vaccination) . C. in EXPECTED RESULTS AND BENEFITS Managing pneumonia is essential to improving success of management programs for Rocky Mountain bighorn sheep. Preventing pasteurellosis outbreaks would dramatically diminish the impacts of this disease on bighorn population dynamics. The potential efficacy of a yearly vaccination program as well as the efficacy of vaccinating in the face of an outbreak have been investigated (Kraabel et al., unpublished data). If biobullet or oral vaccination of this multivalent P. haemolytica supernatant vaccine stimulates serological responses equal or greater to those already known to provide protection against challenge with pathogenic P. haemolytica, then it may be feasible to vaccinate wild freeroaming populations of bighorn sheep as a tool for preventing or managing pneumonia epizootics. If proven effective, such a tool could ultimately become �133 an integral component of comprehensive programs for bighorn sheep in Colorado D. herd health monitoring and elsewhere. and management APPROACH 1. Hypotheses serum neutralizing leUkotoxin; and serum antibody antibody titres titres to P. haemolytica to P. haemolytica will not differ among groups of bighorn sheep injection, biobullet implantation, or orally, unvaccinated control animals. surface antigens receiving vaccine via hand and will not differ from 2. Methods ~~udy animals and their care - Captive Rocky Mountain bighorn sheep (0. canadensis canadensis) (n =41) will be used in this experiment. All bighorns will be housed at the CDOW's Foothills Wildlife Research Facility (FWRF) throughout the study. Groups of animals will reside in 37 ha .pastures throughout the study. Grass/alfalfa hay mix and a pelleted high-energy supplement will be provided as prescribed under FWRF feeding protocols for bighorn sheep; fresh water and mineralized salt blocks will be provided ad libitum. b. Experimental design - Experimental animals will be divided into groups receiving hand injected vaccination (treatment group 1), biobullet vaccination (treatment group 2), oral vaccination (treatment group 3) or placebo vaccination (treatment group 4). Animals will be allotted into groups according to baseline neutralizing titres to P. haemolytica leukotoxin. Sex and age differences will also be considered to keep the groups as homogeneous as possible. Potential efficacy of the various delivery systems will be evaluated through a comparison of humoral immune responses; specifically, leukotoxin neutralizing ability and direct agglutination titres. Having a sample size of 10 animals per group will be adequate to show differences if they exist. c. Experimental vaccine - The experimental P. haemolytica supernatant vaccine will be an inactivated bacterial cell-free biologic extracted from culture supernatants of three serotypes (A2, T10, A6,8,9,11,12) of P. haemolytica; it will contain leukotoxoid and serotype-specific surface antigens, and will be adjuvanted with a combination of Quil A and Al(OH)3 (Vanselow et al. 1985, Lofthouse et al. 1995, Langeveld et al. 1994) To modify the vaccine used previously (Miller et al., 1995) it is necessary to examine these serotypes recovered from bighorn sheep to evaluate the production of cytotoxic material. Each of the three serotypes of P. haemolytica will be grown separately in 1L of Brain Heart Infusion Broth in a 2 L flask for 4.5 hours at 37° C and agitated at 100 rpm. The culture will be centrifuged at 8000 rpm for 15 minutes and the pellet resuspended in 2L warm (37° C) RPMI containing 7 % fetal calf serum (FCS) in a 4 or 6 L flask. The culture will be grown at 37° C, 100 rpm for 1hour. The culture will then be centrifuged as noted above and the supernatant will be filtered through a .2 !-lmfilter using positive pressure. The supernatant will be dialysed (6000-8000 mwc) against distilled water for 48 hours at 4° C. The dialysed supernatant will then be shell-frozen in a methanol bath and lyophilized. Lyophilized material will be tested for cyotoxic activity using a modified in vitro assay that measures the ability of the supernatant kill BL-3 cells (Greer and Shewen 1986). A sub-sample will be heat to �134 inactivated at 56°C for 1 hour to confirm lability. Bovine and bighorn serum with previously established cytotoxin neutralizing titres may be re-evaluated usihg the three lyophilized supernatants to compare behaviour of these supernatants in a neutralization assay. Each of the three serotypes will be prepared separately, using the same procedure: An overnight (O/N) culture of bacteria on a blood agar plate will be used to inoculate a small volume of RPMI media containing 0.5% BHIB. This culture will incubate O/N at 37° C, 100 rpm. This primary culture (200mls) will be used to inoculate 1800 mls of RPMI in a 2L fermenter. Fermenter conditions will be set at 37°C and 200 rpm. The start optical density (OD) will be between .1 and .2 at 525nm. The culture will be fermented to a stop OD of .58-.70 at 525nm (approximately 2-4 hours). The culture will then be centrifuged at 8000 rpm and filtered through a .2 ~m filter. A sample of each culture will be removed and 7% FCS added in order to test for cytotoxicity. This supernatant will be concentrated 20X .. A sub-sample from each culture will be aliquotted for use in gel analysis. Gel profiles and Western blots will be analyzed in order to project the immunogenicity of the ~accine. The vaccine will be blended by according to the following: 90 mls each of the ·20X concentrated supernatants with 88.6 mls of 3% Al (OH)3 , 1. 9 mls of 2% thimerosal, 6.2 mls of 7.5% NaHC03 and 16.8 mls of RPMI. This mixture will stir DIN at 4°C. Quil A will then be added to the mixture (1.5 mls). This blended vaccine will then be aliquotted into vials. This vaccine will be used in all experiments involved in this study plan. The above vaccine will be lyophilized and put into biobullets as supplied by the CDOW. Each biobullet will contain the dry weight equivalent of a 2 ml dose of vaccine. The use of hydrogels for oral delivery of a vaccine against p. haemolytica has been previously investigated (Bowersock et al. 1994a,b). Hydrogels will be loaded with the above vaccine by soaking dried hydrogels in vaccine for 48 hours. These fully swollen hydrogels will then be placed in a 37° C incubator for 48 hours to dry completely. These dried hydrogels will be mixed with apple pulp (Feuerstein et al. 1980) for oral vaccination. Bighorn sheep in treatment group 1 will be aseptically injected with 2 ml of experimental supernatant vaccine intramuscularly (1M) ; biobullets will be implanted using the appropriate device to bighorns in treatment group 2; hydrogels will be orally administered to bighorns in treatment group 3; treatment group 4 will receive 2 ml of sterile PBS administered 1M. d. Sample collection be collected from at 1, 2, 4, 8, 12 for serology will serum collected. and preparation - For serology, blood (10-12 ml) will each bighorn immediately prior to vaccination and again weeks post vaccination. All blood samples collected' be held for 1-4 hr at about 22 C, centrifuged, and Serum will be stored at -20 C until analyzed. If necessary, adult bighorns may be sedated with xylazine HCI (5-20 mg IV or 25-100 mg 1M) or immobilized with a cocktail of carfentanil HCI (1.5 mg), ketamine HCI (100 mg), and xylazine HCI (20 mg), delivered 1M by projectile syringe, to facilitate collections; if immobilization is required, narcotic effects will be reversed with naltrexone HCl (150 mg SC + 50 mg IV). e. Serology measured - Levels of cytotoxin neutralizing antibodies in sera will be using a modified in vitro leukotoxin neutralization assay �135 (Greer and Shewen 1986, Shewen and Wilkie 1988). Serial two-fold dilutions of test sera will be preincubated with leukotoxic P. haemolytica culture supernatant for 30 minutes at 22°C. Supernatant concentration will be adjusted to produce a standardized titre with positive control bovine serum. A 50 ~l volume of test serum/toxin mixture will be transferred to a microtitre plate containing bovine leukemia-derived B cells (BL-3) and 50~1 RPMI. Plates will be incubated at 37° C for 1 hour, and cell viability will be measured by uptake of neutral red dye as previously described (Greer and Shewen 1986). Titres will be expressed as the highest 10g2 dilution that yields ~50% neutralization of toxicity. Levels of serum antibody against serotypespecific surface antigens will be measured using a direct microagglutination assay (Shewen and Wilkie 1982) that incorporates washed formalinized P. haemolytica as antigen. Response to each of the three serotypes used in the vaccine will be investigated. Agglutination titres will be expressed as the reciprocal 10g2 of endpoint dilutions. f. Record keeping - In addition to experimental data gathered, we will ~intain a log of experimental vaccine received and administered; , recorded information will include volumes, lot and vial numbers, and dates of receipt or administration. ' E. ANALYSI:S, Serum neutralizing antibody titres to P. haemolytica leukotoxin and serum antibody titres to P. haemolytica surface antigens will be compared between groups. Serology data will be analyzed using least squares ANOVA for General Linear Models (Freund et ale 1986). All statistical comparisons will use 0:=0.1. F. (No dates SCHEDULE August-September, 1 November, 1997 1997 confirmed) All bighorn sheep placed in pastures at CDOW's FWRF; collect blood for baseline titres. Collect blood from all animals according to group. and vaccinate November, 1997 February, 1998 Collect blood at 1-4 wk intervals. March-May, Analyze data, prepare G. 1998 PERSONNEL Heather McNeil Principal Investigator Michael W. Miller Co-Principal Investigator Jennifer A. Conlon Co-Principal Investigator Patricia E. Shewen Co-Principal Investigator Margaret A. Wild Attending veterinarian report & draft manuscript. �136 H. I. BUDGET Personal Services Operating Supplies and Services Animal Care Serology Miscellaneous supplies Total Operating $ 28, 000 Travel Expenses Capital Expenditures $ TOTAL $ 46,800 LITERATURE $ 8,600 $ 7,200 S 2.000 $ 17,800 ~ 1,000 Q CITED Bailey, J.A. 1990. Management of Rocky Mountain bighorn sheep herds in Colorado. ~piv. of Wild., Spec. Rep. No. 66. 24 pp. Bowersock, T., Shalaby, W., Levy, M., Blevins, W., White, M., Borie, D., and K. Park. 1994a. The potential use of poly(methacrylic acid) hydrogels for oral administration of drugs and vaccines to ruminants. J. Contr. Releases 31:245-254. Bowersock, T., Shalaby, W., Levy, M., Samuels, M., Lallone, R., White, M., Borie, D., Lehmeyer, J., Park, K. 1994b. Evaluation of an orally administered vaccine, using hydrogels containing bacterial exotoxins of Pasteurella haemolytica, in cattle. Am. J. Vet. Res. 55:502-509. Buechner, H.K. 1960. The bighorn sheep in the United States, its past, present, and future. Wildl. Mono. 4:74. Coggins, V.L. 1988. The Lostine Rocky Mountain bighorn sheep die-off and domestic sheep. Bienn. Symp. Northern Wild Sheep and Goat Counc. 6:57-64. DeNicola, A. J., D. J. Desler, and R. K. Swihart. 1996. delivery system. Wildlife. Soc. Bull. 24:301-305. Ballistics of a biobullet Donachie, W., C. Burrells, A.D. Sutherland, J.S. Gilmour, and N.J.L. Gilmour. 1986. Immunity of specific pathogen free lambs to challenge with an aerosol of Pasteurella haemolytica biotype A serotype 2. Pulmonary antibody and cell responses to primary and secondary infections. Vet. Immunol. Immunopathol. 11:265-279. Festa-Bianchet, M. 1988. A pneumonia epizootic in bighorn sheep, with comments on preventive management. Bienn. Symp. Northern Wild Sheep and Goat Counc. 6:66-76. Feuerstein, V., R.L. Schmidt, C.P. Hibler, and W.H. Rutherford. 1980. Bighorn sheep mortality in the Taylor River-Almont Triangle area, 1978-1979: a case study. Colo. Div. of Wild., Spec. Rep. No. 48. 19 pp. Foreyt, W.J. 1989. Fatal Pasteurella haemolytica pneumonia in bighorn sheep after direct contact with clinically normal domestic sheep. Am. J. Vet. Res. 50:341-343. Foreyt, W.J. 1990. Pneumonia in bighorn sheep: effects of Pasteurella haemolytica from domestic sheep and effects on survival and long-term reproduction. Proc. Biennial Symp. Northern Wild sheep and Goat Counc. 7:92-101. Foreyt, W.J. and D.A. Jessup. 1982. Fatal pneumonia of bighorn sheep following association with domestic sheep. J. Wildl. Dis. 18:163-169. Foreyt, W.J., K.P. Snipes, and R.W. Kasten. 1994. Fatal pneumonia following innoculation of healthy bighorn sheep with Pasteurella haemolytica from healthy domestic sheep. J. Wildl. Dis. 30:137-145. �137 Foreyt, W.J. 1992. Failure of an experimental Pasteurella haemolytica vaccine to prevent respiratory disease and death in bighorn sheep after exposure to domestic sheep. Bienn. Symp. North. Wild Shaeep and Goat Counc. 8:155-163. Forrester, D.J. 1971. Bighorn sheep lungworm-pneumonia complex. pp. 158-173 in J.W. Davis, and R.C. Anderson, eds. Parasitic Diseases of Wildlife. Iowa State Univ. Press, Ames. Freund, R.J., R.C. Littell, and P.C. Spector. SAS Institute, Cary, NC, .187-201. Gilmour, N.J.L. and J.S. Gilmour. Pasteurella and Pasteurellosis. Ltd., San Diego, CA, 341 pp. 1986. SAS system for linear models. 1989. Pasteurellosis of sheep. pp. 223-262 in C. Adlam and J.M. Rutter, eds., Academic Press Greer, C.N. and P.E. Shewen. 1986. Automated colorimetric assay for the detection of Pasteurella haemolytica leucotoxin. Vet. Micro. 12:33-42. Herriges, J.D., Jr., E.T. Thorne, and S.L. Anderson. 1991. Vaccination to control b~~osis in free-ranging elk on western Wyoming feed grounds. pp. 107-112 in R. D. Brown, ed ..The biology of deer. Springer-Verlag, New York, N.Y. Hobbs, N.T. and M.W. Miller. 1992. Interactions between pathogens and hosts: simulation.of pasteurellosis epizootics in bighorn sheep populations. pp. 997-1007 in Wildlife 2001: Populations. D.R. McCullough and R.H. Barrett, eds., Elsevier Science Publishers, Ltd., London, England, 1163 pp. Jessup, D.A., J.R. DeForge, and S. Sandberg. 1992. Biobullet vaccination of captive and free-ranging bighorn sheep. Proc. International game ranching symposium 2:429434. Langeveld, J.P.M., J.I. Casal, E. Cortes, G. van de Wetering, R.S. Boshuizen, W. M.M. Schaaper, K. Dalsgaard, and R.H. Meloen. 1994. Effective induction of neutralizing antibodies with the amino terminus of VP2 of canine parvovirus as a synthetic peptide. Vaccine. 12:1473-1480. Lofthouse, S.A., A.E. Andrews, A.D. Nash, and V.M. Bowles. 1995. Humoral and cellular responses induced by intradermally administered cytokine and conventional adjuvants. Vaccine. 13:1131-1137. Marsh, H. 1938. Pneumonia in Rocky Mountain bighorn sheep. J. Mammal. 19:214-219. Miller, M.W. and N.T. Hobbs. 1989. Epidemiology of pneumonia outbreaks in captive bighorn lambs. Pages 95-104 in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-2, WP2a, J4. Colorado Division of Wildlife, Fort Collins, Colorado, USA. Miller, M.W., N.T. Hobbs, and M.A. Wild. 1991a. Management of bacterial and viral diseases in mountain sheep populations. Pages 101-122 in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-4, WP2a, J4. Colorado Division of Wildlife, Fort Collins, Colorado, USA. Miller, M.W., N.T. Hobbs, and E.S. Williams. 1991b. Spontaneous pasteurellosis in captive Rocky Mountain bighorn sheep (avis canadensis canadensis): clinical, laboratory, and epizootiological observations. J. wildl. Dis. 27:534-542. Miller, M.W., B.J. Kraabel, J. A. Conlon, H. J. McNeil, and J. M. Bulgin. 1995. Strategies for managing infectious diseases in mountain sheep populations. Pages 151-161 in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-8, WP2a, J4. Colorado Division of Wildlife, Fort Collins, Colorado, USA. (in press) Onderka, D.K. and W.D. Wishart. 1984. A major bighorn sheep dieoff from pneumonia in southern Alberta. Bienn. Symp. Northern Wild Sheep and Goat Counc. 4:356-363. �138 Onderka, D.K. and W.D. Wishart. 1988. Experimental contact transmission of Pasteurella haemolytica from clinically normal domestic sheep causing pneumonia in Rocky Mountain bighorn sheep. J. Wildl. Dis. 24: 663-667. Onderka, D.K., S.A. Rawluk, and W.D. Wishart. 1988. Susceptibility of Rocky Mountain bighorn sheep and domestic sheep to pneumonia induced by bighorn and domestic livestock strains of Pasteurella haemolytica. Can. J. Vet. Res. 52: 439-444. Post, G. 1962. Pasteurellosis of Rocky Mountain bighorn sheep (Ovis canadensis). Wild1. Dis. 23:1-14. Potts, M.K. 1937. Hemorrhagic septicemia in the bighorn of Rocky Mountain National Park. J. Mammal. 18:105-106. Rush, W.M. 1927. Notes on disease in wild game animals. J. Mammal. 8:163-165. Schwantje, H.M. 1986. A comparative study of bighorn sheep herds in southeastern British Columbia. Bienn. Symp. Northern Wild Sheep and Goat Counc. 5: 231-252. e~. E. and B. N. Wilkie. 1988. Vaccination of calves with leukotoxic culture supernatant from Pasteurella haemolytica. Can.J. of Vet. Res. 52:30-36. Shewen, P.E. and B.N Wilkie. 1982. Antibody titers to Pasteurella Ontario Beef Cattle. Can. J. Compo Med. 46:354-356. haemolytica Al in Silflow, R.M., W.J. Foreyt, S.M. Taylor, W.W. Laegreid, H.D. Liggitt, and R.W. Leid. 1991. Comparison of arachidonate metabolism by alveolar macrophages from bighorn and domestic sheep. Inflammation 15:43-54. Si1flow, R.M., Foreyt, W.J., and R.W. Leid. 1993. Pasteurella haemolytica cytotoxin dependant killing of neutrophi1s from bighorn and domestic sheep. J. Wildl. Dis. 29:30-35. Spraker, T.R., C.P. Hibler, G.G. Schoonve1d, and W.S. Adney. 1984. Pathologic changes and microorganisms found in bighorn sheep during a stress-related die-off. J. Wildl. Dis. 20:319-327. Sutherland, A.D., W. Donachie, G.E. Jones, and M. Quirie. 1989. A crude cytotoxin protects sheep against experimental Pasteurella haemolytica serotype A2 infection. Vet. Microbiol. 19:175-181. Vanselow, B.A., I. Abetz, and K. Trenfie1d. 1985. A bovine ephemeral fever vaccine incorporating adjuvant Quil A: A comparative study using adjuvants Quil A, aluminium hydroxide gel and dextran sulphate. Vet. Record. 117:37-43. Wild, M.A. and M.W. Miller. 1991. Detecting nonhemolytic Pasteurella haemolytica infections in healthy Rocky Mountain bighorn sheep (Ovis canadensis canadensis): Influences of sample site and handling. J. Wi1dl. Dis. 27:53-60. Wishart, W.D., J. Jorgenson, and M. Hinton. 1980. A minor die-off of bighorns from pneumonia in southern Alberta. Bienn. Symp. Northern Wild Sheep and Goat Counc. 2: 229-247. �139 Colorado Division Wildlife Research July 1997 of Wildlife Report JOB FINAL state of Colorado Project No. W-153-R-I0 Work Plan No. REPORT Mammals Research Mountain 2A Job No. Sheep Inyestigations Experimental Evaluation of Mountain Sheep Transplanting and Disease Treatment Period Covered: July 1, 1996 - June 30, 1997 Authors: M. W. Miller, Jurgens J. Vayhinger, S. Roush, T. Verry, A. Torres, and V. Personnel: R. Dobson, J. Duran, B. Elkins, J. George, D. Getzy, R. Green, R. Hancock, M. Lamb, R. Myers, S. Ogilvie, G. Roberts, B. Thornton, R. Zaccagnini. ABSTRACT We conducted a 4-year management experiment to examine the effects of alternative lungworm treatment strategies on lamb survival and population performance in Rocky Mountain bighorn sheep (Ovis canadensis canadensis) herds in southcentral Colorado. Beginning in December 1991, 2 bighorn herds in the Tarryall Mountains and 2 herds in the Collegiate Peaks were managed under 1 of 4 alternative lungworm treatment regimes: baiting with alfalfa hay and apple pulp treated with fenbendazole (about 3 g/adu1t ewe) (B/T), baiting with alfalfa hay and apple pulp without fenbendazole (B), placing fenbendazo1etreated salt blocks (1.65 g fenbendazole/kg) on winter ranges (T), and withholding bait and fenbendazole (control) (C). Treatments were rotated annually under a predetermined, randomly-selected schedule. We monitored lamb production and survival among radiocollared ewes in each herd from May through October 1991-1995. Both production and survival varied widely and affected recruitment (lambs/marked ewes) through October. Annual recruitment rates through October ranged from 0.13 to 0.88, but mean recruitment rates did not differ among herds (P = 0.51) or between years· (P = 0.80). Among the 4 study herds, sick and coughing lambs were observed every summer in 1 herd and 1 summer (1995) in a second. Mean lamb production (estimated by observations of marked ewes with ~ 2 wk old lambs at heel) ranged from 0.83 (B/T) to 0.95 (B), but did not differ among treatments (P ~ 0.23). Mean lamb survival (lambs in October/lambs born to marked ewes) through October ranged from 0.59 (B/T) to 0.81 (T), and was also unaffected by management treatment (P ~ 0.15); moreover, baiting combined with fenbendazole administration failed to prevent catastrophic (86%) lamb mortality in 1 herd. overall, neither baiting (P = 0.45) nor fenbendazole treatment (P = 0.62) enhanced lamb recruitment among �140 . these 4 apparently healthy bighorn populations during the 4 years of our study. Our results demonstrate that annual parasite treatment is not prerequisite for lamb survival among southcentral Colorado's wild bighorn populations. Moreover, annual parasite treatment may not prevent catastrophic losses of lamb cohorts among treated herds. Based on these data, we question the need for annual baiting and parasite treatment in the herds studied here and believe such practices need to be reevaluated elsewhere as well. In making these reevaluations, we encourage managers to weigh the potential benefits of baiting and treatment against the "costs" of such interventions those costs may include locally increased bighorn densities (that could lead. to infectious disease outbreaks), alteration or loss of movement and distribution patterns, increased vulnerability to predators (including man), and dependence on artificial feed sources. In light of the potential detrimental effects of baiting and treating wild sheep herds annually, we recommend that anthelmintic therapy be reserved for specific situations where verminous pneumonia in lambs is not only documented, but is occurring at rates sufficient to depress long-term performance of individual bighorn populations • ~, [No~e: Addi~ional objec~ives originally lis~ed for ~he 1996-1997 segment were no~ funded wi~h Federal Aid monies; consequen~ly, progress ~oward ~heir comple~ion is no~ repor~ed here.] �141 EXPERIMENTAL EVALUATION OF MOUNTAIN SHEEP TRANSPLANTING AND DISEASE TREATMENT M. W. Miller, J. Vayhinger, S. Roush, T. Verry, A. Torres, and V. Jurgens P. N. OBJECTIVE Design, conduct, and report on management experiments to evaluate efficacy transplanting and disease treatment practices for managing mountain sheep pulations. of ~, AGREEMENT Report on a management level experiment parasite control program. OBJECTIVE evaluating Colorado's mountain sheep [Note: Additional objectives originally listed for the 1996-1997 segment were not funded with Federal Aid monies; consequently, progress toward their completion is not reported here.] Rocky Mountain bighorn sheep (Ovis canadensis canadensis) populations throughout North America are plagued by pneumonia outbreaks caused by a complex of bacterial, parasitic, and/or viral agents. These epi~ootics and subsequent population declines represent a significant obstacle to long-term success in bighorn sheep management. Treating with anthelmintics to reduce parasitic lungworm (Protostrongylus spp.) burdens has become an integral part an aggressive management program to reduce disease outbreaks and enhance productivity of mountain sheep herds in Colorado. Although some combination of treating, trapping and other management activities has increased sheep numbers statewide over the last 2 decades, it is unclear which strategies were most influential in this achievement. In light of the growing number of significant mountain sheep herds throughout Colorado, the apparent intensity of management required to perpetuate individual populations, and the costs associated with applying intensive management, it has becoming increasingly important to explore alternatives for efficient but efficacious management practices. Here, we describe a management experiment conducted to examine effects of alternative lungworm treatment strategies on bighorn lamb survival and population performance. Our experiment tested the hypothesis that bighorn lamb production and survival rates for herds would not differ in response to various management treatments. �142 MATERIALS AND METHODS Beginning in December 1991, we began managing each of 4 study herds [Tarryall Mountains: Twin Eagles (TE) and Sugarloaf (SL); Collegiate Mountains: Chalk Creek (CH) and Cottonwood Creek (CW)] under 1 of 4 alternative lungworm treatment regimes: Control Treat OnlyBait Only Bait/Treat- no treatment -- bait and fenbendazole withheld (C); fenbendazole-treated salt blocks placed on bait stations (T); baited with alfalfa hay and apple pulp but not treated with fenbendazole (B); baited with alfalfa hay and apple pulp and treated with fenbendazole (B/T). Treatments were assigned to study herds as prescribed rotating schedule (Table 1.; Year 1 = 1992). .' by a randomly selected, ~, We attempted to apply experimental treatments uniformly across the 4 study herds. We baited with third cutting alfalfa hay (about 2 kg/head/day) and fermented apple pulp (about 1 kg/head/day) at both Band B/T sites for about 8-10 weeks beginning in mid-December. In addition, B/T sites were also treated with fenbendazole (about 3 g/adult ewe) added to apple pulp on 2 occasions 1 to 2 wks apart. Four or five treated salt blocks (1.65 g fenbendazole/kg; 15 kg/block) were available to sheep at T sites during January to May; 1 additional block held in a wire cage during that same period was used as an environmental control. Blocks were replaced if they disappeared before 1 May. For Band B/T herds, we recorded baitsite attendance and receipt of fenbendazole treatments for each radiocollared ewe. For T sites, we weighed medicated blocks to estimate consumption but were otherwise unable to document exposure of marked ewes to the treatment. Control herds were monitored periodically but received no other management intervention. Adult bighorn ewes from each herd were initially captured using drop nets in February and March 1991 and marked with individually identifiable radiocollars; collar symbols, color combinations, and radio frequencies were identifiable to individuals and herds of origin. Additional adult ewes were captured opportunistically in some herds each of the following winters to achieve target sample sizes (15 to 20 ewes per herd) or replace mortalities. We used net gunning, aerial darting, or darting from the ground in supplemental captures. No anthelmintics or antibiotics were used during supplemental captures. Each year, we assessed effects of winter treatments on lamb production and survival by observing radiocollared ewes (n = 11 to 18) from all 4 herds about once every 2 weeks from May through October to determine whether they produced lambs, and whether their lambs were still alive and healthy. We defined annual lamb production as the proportion of marked ewes seen with a lamb nursing or at heel at least once between May and October. Annual lamb survival was the proportion of lambs seen with marked ewes that were still alive at the end of October. Annual lamb recruitment, the product of these two processes, was the proportion of marked ewes with lambs alive at the end of October. We recognize that the foregoing approach may have slightly underestimated production and overestimated survival by misclassifying periparturient mortality as failed production. However, because verminous �143 pneumonia primarily affects survival of lambs >6 wks of age, we believed our approach more sensitively measured the treatment responses of primary interest in this study. In addition to lamb survival data, we recorded approximate UTM coordinates, habitat type, and group size and composition for each radiocollared ewe observed. All field data were transcribed into a computerized database to aid in mapping seasonal range movements and determining annual lamb production and survival rates. Radiocollared ewes were also monitored every 2-4 weeks to detect mortality and movements during November through April in conjunction with a USFS/CDOW cooperative project to identify critical winter and transitional ranges of these 4 herds. These data will be reported separately elsewhere. RESULTS AND DISCUSSION l~ked ewes maintained fidelity to .their respective herds of origin throughout the study. Consequently, we believe treatments were applied to ewes within each herd only as prescribed by the foregoing experimental schedule.· Treatment Rates As controls, herds received no treatments or intervention (aside from supplemental captures) and were essentially undisturbed by management activities other than monitoring or inventory during December-May. Use of treated salt blocks varied considerably among herds. Adjusting for estimated environmental losses, block consumption equated to about 13 ewe treatments (2 X 3 g/ewe) at CH, 26.5 at CW, 6 at SL, and 6.5 at TE; under a moderate dosing regime (3 X 0.75 g/ewe) (Foreyt and Coggins 1990), block consumption equated to about 35 ewe treatments at CH, 70 at CW, 16 at SL, and 17 at TE. We observed marked and unmarked sheep using blocks on several occasions in late January-April, but mule deer or elk also may have used treated blocks at some sites. Apparent differences in salt block consumption between study herds in the Collegiate Peaks and Tarryall Mountains are supported by field observations suggesting marked differences in affinity for natural mineral licks between ewes in these 2 discontinuous mountain ranges. Whether these observed differences reflect natural mineral deficiencies in some ranges and/or greater abundance of salt associated with domestic stock grazing remains undetermined. However, our data suggest such differences, combined with consumption by other species, could influence the potential use and efficacy of fenbendazole-treated salt blocks among diverse bighorn sheep ranges in Colorado and elsewhere. Responses to bait, with or without fenbendazole, were generally more consistent than to treated blocks in 3 of the 4 study herds. This may be in part because all of the herds we used had been baited an~ treated annually for a decade or more preceding initiation of this experiment. In years when bait alone was applied, marked ewes averaged (± sd) 42 (± 13.7) days on bait at CH, 26 (± 15.1) days at CW, 38 (± 1.9) days at SL (6 days missing data), and 44 (± 1.0) days at TE. In contrast to the other 3 herds, most marked ewes at CW visited the bait site infrequently during December-January. Many ewes in this herd stayed on alpine winter ranges until heavy snows apparently forced them �144 to lower elevations markedly. in February, when CW baitsite attendance increased In years when baiting was combined with fenbendazole treatment, marked ewes averaged (± sd) 51 (± 10.8) days on bait at CH, 23 (± 6.3) days at CW, 45 (± 1.1) days at SL, and 47 (± 4.5) days at TE. In addition to bait, 14 (93%) of 15 marked ewes at CW, 8 (50%) of 16 marked ewes at CW, and all 17 marked ewes at both SL and TE also were present to presumably receive at least 1 fenbendazole treatment during their scheduled treatment periods.. The relatively low treatment rate at CW was attributed to failure of marked ewes to remain on low elevation winter ranges in 1995 despite daily baiting. Lamb Production and Survival Both production and survival varied widely and affected recruitment (lambs/marked ewes) through October. Annual recruitment rates through October ranged from 0.13 to 0.88; however, mean recruitment rates did not differ among e~p = 0.51) or between years (P = 0.80). Among the 4 study herds, sick and coughing lambs were observed every summer in the Chalk Creek herd and during the summer of 1995 in the Cottonwood Creek herd; no sick lambs were seen in e'ither of the Tarryall Mountain herds during the 4-year study period. Mean lamb production ranged from 0.83 (B/T) to 0.95 (B), but did not differ among treatments (P ~ 0.23). Mean lamb survival (lambs in October/lambs born to marked ewes) through October ranged from 0.59 (B/T) to 0.81 (T), and was also unaffected by management treatment (P ~ 0.15); moreover, baiting combined with fenbendazole administration failed to prevent catastrophic (86%) lamb mortality in the Cottonwood Creek herd in 1995. Although recruitment varied widely, neither baiting (P = 0.45) nor fenbendazole treatment (P = 0.62) enhanced lamb recruitment among these 4 apparently healthy bighorn populations during the 4 years of our study. Range Use and Movement Patterns Relatively consistent and predictable range use and movement patterns for each of the 4 study herds have emerged since monitoring began in 1991. Plots of May 1991-March 1994 location data (UTM coordinates) for radiocollared ewes revealed apparent differences in distribution and movement patterns among herds (Fig. 2). Ewes from the CW herd have consistently shown widest distribution and greatest movements; CH ewes were the most limited in their range use and movement. Although ranges of the SL and TE herds appeared to overlap considerably, to date we have observed no exchange of radiocollared ewes between these 2 herds. Disturbances by hikers (CW) and hunters (CH, SL) appeared to influence movements of ewe/lamb groups on occasion. Location data gathered since March 1994 will be added to further define key ranges and migration corridors for these 4 herds. Population Parameters and Performance OVerall, noncapture mortality rates of adult ewes in these 4 herds averaged about 0.08 (se = 0.01) annually over the 56 months covered by our study, but causes and annual rates (ranging from 0 to 0.28 at CW in 1994) of ewe mortality appeared to vary among herds. No consistent health problems were detected among marked ewes. In total, 20 radiocollared ewes (2 at CH, 5 at TE, 5 at SL, and 8 at CW) died of noncapture causes between February 1991 and October 1995. Of these, lion predation appeared to have caused 6 losses in �145 the Tarryalls (3 at TE and 3 at SL), injuries from falls may have killed 2 ewes (at CW), pneumonic pasteurellosis killed 1 ewe (at CW), and lightning claimed 1 ewe (at CW); causes of death or disappearance for 10 other ewes (2 at CH, 2 at TE, 2 at SL, and 4 at CW) could not be determined, although we speculated lightning strikes also may have been involved in 2 deaths and 2 disappearances at CW and 1 death at CH during late June-early July. Six of 10 ewe mortalities in the Collegiate Peaks herds since 1991 occurred in apparently healthy ewes during mid June-mid August and may have been lightning-caused; whether telemetry collars are a predisposing factor in these losses was uncertain. Despite observed variation in recruitment and adult mortality rates, winter range counts during 1991-1995 suggest all 4 bighorn herds under study remained stable or grew during the course of this exper iment. CONCLUSIONS AND MANAGEMENT RECOMMENDATIONS ~ults demonstrate that annual parasite treatment is not prerequisite for lamb survival among southcentral Colorado's wild bighorn populations. Moreover, annual parasite treatment may not prevent catastrophic losses of lamb cohorts among treated herds. These findings should not be altogether surprising -- there are clearly a multitude of factors besides disease that can influence lamb production and survival in bighorn populations. And even in herds where signs of respiratory disease are observed among lambs, anthelmintic therapy in ineffective in treating bacterial pneumonia (e.g., pasteurellosis) that appears to be far more prevalent than verminous pneumonia among bighorn populations in Colorado and elsewhere. Based on these data, we question the need for annual baiting and parasite treatment in the herds studied here and believe such practices need to be reevaluated elsewhere as well. In making these reevaluations, we encourage managers to weigh the potential benefits of baiting and treatment against the "costs" of such interventions -- those costs may include locally increased bighorn densities (that could lead to infectious disease outbreaks), alteration or loss of movement and distribution patterns, increased vulnerability to predators (including man), and dependence on artificial feed sources. In light of the potential detrimental effects of baiting and treating wild sheep herds annually, we recommend that anthelmintic therapy be reserved for specific situations where verminous pneumonia in lambs is not only documented, but is occurring at rates sufficient to depress long-term performance of individual bighorn populations. Wildlife Research Veterinarian �146 Table 1. Treatment assignments for 4 bighorn herds included in a 4-year management experiment to examine effects of alternative lungworm treatment strategies on bighorn lamb survival and population performance. HERD CQLLEGIAIEMQU~IAI~S IABBYALL MQU~IAI~S YEAR CHALK CREEK COTTONWOOD CREEK SUGARLOAF MOUNTAIN TWIN EAGLES 1992 8' C T 8IT 1993 8IT T 8 C 1994 C 8 8IT T 1995 T 8IT C 8 Treatment assignments: 8IT = bait with alfalfa hay and apple pulp treated with fenbendazole; 8 = bait with alfalfa hay and apple pulp without fenbendazole; T = fenbendazole-treated salt blocks on bait stations; and C = withhold all bait and fenbendazole (control). �147 Colorado Division Wildlife Research July 1997 of Wildlife Report JOB FINAL REPORT state of Project Colorado No. W-153-R-10 Work Plan No. 3A Job No. Period Author: Mammals Research Pronghorn Inyestigations Habitat Selection and population Performance of a Pioneering Pronghorn Population Covered: July 1, 1996 - June 30, 1997 T.M. Pojar ABSTRACT The rate of increase (ROI) for the Middle Park pronghorn population continues the downward trend as the population size increases and exhibits a strong density dependence (~ = 0.0141). The density dependence feedback is linked to some factor other than the fawn to doe ratio because there was no apparent influence of density on this parameter (~ = 0.258). The summer distribution area remained very similar throughout the tracking period with animals moving from the Kremmling area north and west to Muddy Pass/Diamond Creek summer range and east to Granby. There were, however, consistent reports of a small group (4 ± animals) summering in the Fraser valley; none of these were radioed. The maximum count of pronghorn south of the Colordo river was 27 in August, 1996 which included fawns of the year. This represents less than 5% of the population; none of these animals remain south of the river during winter. The distribution of yearlings radioed for the mortality investigation is essentially the same as the population as a whole. Sixty 6-month-old fawns (30 males and 30 females) were radioed for the mortality aspect of this study during 1993-1995; the mortality rates were nearly even for males and females. Twenty-four males and 25 females were still alive at the end of 1996. A Program Narrative was written to describe the data analysis and reporting for this project (Appendix I). ��149 HABITAT SELECTION AND POPULATION PERFORMANCE PRONGHORN POPULATION Thomas OF A PIONEERING M. Pojar P.N. OBJECTIVE Describe population dynamics pronghorn population. and habitat SEGMENT 1. 2. Describe seasonal population. and annual Monitor natural mortality ~ales and females. of habitation use of a pioneering, OBJECTIVES distribution and movement using expanding of the Middle patterns Park pronghorn of radioed yearling 3. Map areas the GIS format. 4. Monitor population dynamics of Middle Park pronghorn with: a. Ground counts to describe changes in population size. b. Ground counts to quantify population sex and age composition. STUDy AREA The study area is described (1993) in Pojar METHODS SEASONAL AND ANNUAL (1988) and a map of the area is in Pojar AND MATERIALS DISTRIBUTION Tracking was done mostly from the ground; fixedwing aircraft was used if an animal could not be located after a reasonable effort from the ground. Legal descriptions of animal locations were recorded to the nearest quarter mile then converted to UTM (U.S. Army 1973) coordinates for computer processing. All radioed animals have been located biweekly (with very few exceptions) since January 1, 1987. POPULATION SIZE AND STRUCTURE Herd structure estimates were obtained by classifying all animals that accompanied the animals that are radioed. The herd structure estimate used in population projections is the one with the largest sample size obtained in August or September. Total counts are made during winter by counting all animals associated with radioed animals. With the increased population size, it is not always possible to get an accurate count of total mature bucks (1.5 yrs and older) in the population. However, it is still possible to get very accurate counts of bucks in 60-80% of the population. The proportion of bucks in this portion of the population is then extrapolated to the total population to estimate totai mature bucks. Total population count during winter, estimated number of mature bucks from the winter count, and recruitment based on fawn to doe ratios from late summer are used for the population projection. �150 Population projections 1. are based on the following assumptions: Winter counts represent the total population and the estimated number of mature bucks in Middle Park. Late summer age ratio estimates represent "recruitment" into the population. Annual survival of mature bucks and does and female fawns is 92.5%. Annual survival of males in their first year (after weaning) is 50%. (This severe mortality on male fawns is arbitrary, however, it allows the number of mature males in subsequent years to match fairly well with winter counts.) 2. 3. 4. After several year's data collection on the natural mortality of yearling males, it is concluded assumptions 3 and 4 above are in error. Future projections should assume mortality of animals from fawns (6 months) to 3 years old is approximately equal for males and females. Therefore, for the uck to doe ratios to match observed ratios, it is necessary to shift the i~ntial natural mortality to older age class bucks. NATURAL MORTALITY OF MALES AND FEMALES The methods for estimating differential natural mortality between female pronghorn are outlined in Appendix I of Pojar (1994). male and RESULTS SEASONAL AND ANNUAL DISTRIBUTION The winter distribution during 1996-g7 was similar to the previous winter distribution although the Wolford Mountain area was not used as it was in 1995-96. Groups of 50+ spent the early part of the winter in Sulphur Gulch and Antelope Creek vicinities. As the winter progressed, they resumed occupation of their traditional wintering areas on Red Mountain and within a 5 square mile area to the north and east. They occupied this area through spring thaw and co-existed there with several hundred deer and elk. Although access to GIS methods for mapping areas of habitation was not available for this report, efforts will continue to obtain such access. POPULATION SIZE AND STRUCTURE Total population size estimates are obtained during winter and herd structure estimates are obtained in late summer (Table 1). The annual changes in population size are used to calculate the rate of increase which is regressed on population size to project the K-value for the population. The ROI is calculated as ROI = p +P. "_2__ 1 P1 where P1 is the population size at time 1 and P2 is the population at time 2 (Table 2). The rate of increase for 1996-97 is 0.07 (Table 2). Based on the linear relationship of population size and ROI, the projected K-value for this population is 621. The projected winter 1997-98 population size is 622, �151 Table 1. Herd structure of. Middle Park pronghorn based on a sample obtained by locating radioed animals in late summer. The population size is from the subsequent winter counts with harvest added back into the population to get the pre-hunt population size, e.g. 1996 pre-hunt population was 594, 579 winter count plus 15 harvest. YEAR POP. SIZE 19861 1987 1988 1989 1990 1991 9~' 1993 1994 1995 1996 1997 80 122 160 223 261 308 347 425 466 .5.35 594 6372 NO. RADIO % RADIO B:I00D RATIO F:100D RATIO SAMPLE % OF POP. 7 24 22 17 13 39 31 58 52 56 44 5.7 15.0 10.2 6.5 4.2 11.2 7.3 12.4 9.7 9.4 6.9 36 54 40 56 22 23. 26 10 29 32 37 42 77 77 32 50 47 65 48 66 46 42 42 39 47 63 108 161 148 148 286 266 332 437 353 414 59 52 68 72 66 48 82 63 71 82 59 65 This year's data based on the sample of the population trapped 16 December 1986. 2 Population size projected from previous year's wintering population and the late summer fawn to doe ratio estimate (see Table 3). 1 Table 2. Population size of the Middle Park pronghorn herd during winter and the calculated rate of increase. Population size reflects the removal of 1315 animals per year by harvest beginning in 1990, .i.e. the 1996-97 winter population was 594 before harvest and 579 after harvest. YEAR 1986-87 1987-88 1988-89 1989-90 1990-91 1991-92 1992-93 1993-94 1994-95 1995-96 1996-97 1997-98 POP. SIZE (projected) 80 122 160 223 246 292 332 410 453 520 579 622 RATE OF INCREASE .52 .31 .39 .10 .19 .14 .23 .10 .15 .11 .07 �152 therefore, given "normal" climatic conditions, the ROI for 1997-98 should be near zero and the population should cease to grow. The winter of 1996-97 was relatively mild through January with low snow fall and accumulation, and no extended periods of sub-zero (Fo) temperatures. About mid-January, heavy snows and deep accumulation in higher elevations forced several hundred deer and elk into the general vicinity of the pronghorn wintering area. Table 3. Population projection text for the assumptions. I POPULATION I BUCKS ~TER '96-97 ~ 1 for the Middle Park pronghorn I DOES I FAWNS population. I TOTAL 111 321 137 569' WINTER MORTALITY 111X .075 = 8 MORT 321 X.075 = 24 MORT 68X.5=34B 68X.075=5D 71 PRE ... FAWNING 1997 111- 8 = 103MATURES + 34 YRLS TOTAL =137 321 - 24= 297 MATURES + 63 YRLS TOTAL = 360 LATE SUMMER 1997 MATURE 103 YRLS 34 TOTAL 137 MATURE 297 YRLS 63 TOTAL 360 This total represents NATURAL MORTALITY I 497 @ 39F:100D 360 X .39 = 140 FAWNS the actual number counted on December OF YEARLING See 637 19, 1996. MALES AND FEMALES For the third year, the net-gun technique (Helicopter Wildlife Management, Salt Lake City, Utah) was used to capture target animals for the mortality study. Fourteen males were captured in December 1996 and fitted with radios. Thus far, 62 pronghorn have been captured using this technique with 1 known capture related mortality. To avoid the problem of accommodating neck growth and neck swelling during the rut for males, solar power transmitters mounted on ear tags were used in 1995. As of this writing, the dependability of these radios is disappointing. Signal transmission is sporadic depending on the angle of the solar panels to direct sunlight. subsequent discussions with the manufacturer has revealed that there was an error in construction of the circuitry resulting in the need for excess power requirements to trigger the radio signal emission. Tracking of these radios will be sporadic at best. Sixty radios were deployed during 1993-95 for estimation of differential natural mortality between males and females. Of these, 30 were put on males and 30 on females. Twenty-four of the males are still alive (as of January 1, �153 1997) and 25 of the females are alive. Natural mortality of young (age 6 months to 3 years) pronghorn is equal for males and females. Therefore, the reason for the observed buck to doe ratio digressing from a one-to-one ratio must be due to either biased buck to doe ratio estimation or higher mortality on males in the older age classes. LITERATURE Pojar, CITED T.M. 1988. Habitat selection and population performance of a pioneering pronghorn population. colo. Div. Wildl. Res. Rep. July, 181-192. 1993. Habitat selection and population pronghorn population. Colo. Div. Wildl. 1994. ~~ronghorn u.s. Habitat selection and population population. Colo. Div. Wildl. Army. 1973. Technical Manual: Headquarters, Dep. of the Army, pp performance of a pioneering Res. Rep. July, pp 199-207. performance of a pioneering Res. Rep. July, pp 125-136. Universal Transverse Mercator Grid. Washington D.C. TM No. 5-241-8, 64 pp. ��155 APPENDIX I PROGRAM NARRATIVE State of Colorado Project No. W-153-R Work Package No. --=3"""0=04......_ Task No. _.LI :!oOo&<....:!:2<-A. _ _ Cost Center 3430 Mammals Program Management of Other Ungulates Pronghorn Data Analysis and R~orting NEED According to the Colorado Wildlife Commission's Long range Plan (March II, 1994) it is the Commission's ol~ "The Division will manage it's Wildlife-related recreation programs based upon sound iolo'gicaJ principles and up-to-date demographic information on public expectations and value." To accomplish this, the Long Range Plan states a goal for the Division of Wildlife to "Use hunting as the primary tool to manage big game populations to maintain a balance between these species and their habitats (Goal 10. I). State-of-the-art wildlife population inventory methods are an essential component in maintaining the balance between wildlife species and their habitats. Pronghorn are no exception. Over the past one and a half decades the Division has invested considerable effort into improving methods for estimating pronghorn numbers and sex and age composition; this expanded our knowledge of pronghorn population performance under a variety of environmental circumstances and habitat constraints. This project proposes to analyze, sununarize, and report on the most recent data sets which address these issues. Two primary tasks have been the focus of this work; the first monitored performance of a pioneering pronghorn population in Middle Park Colorado and the second compared several methods for estimating population parameters. Historically, pronghorn were abundant in Middle Park but were extirpated at the turn of the century largely as a consequent of over hunting. In the late 1970s, pronghorn began to pioneer into Middle Park from North Park on a seasonal basis and by the early 1980s, they were once again yearlong residents. This pioneering population provided a unique opportunity to study the growth and distribution of an unhunted population that was re-establishing itself into historic range. Monitoring population characteristics, movements, and habitat occupation as it expanded provided critical information for modeling of this species and insights into environmental resistance characteristics that are useful in establishing a balance between pronghorn and habitats in sagebrush steppe ecosystems. Traditional pronghorn inventory methodology has consisted of trend counts or total area counts. Trend counts assumed (but did not establish) that trend areas were characteristic of the entire population and that annual variation in trend counts reflected variation in population trajectory. Total area counts assumed that pronghorn could be counted over large areas without error, or at a minimum, with consistent error. Neither of these premises were verified. Work over the past several years have incorporated principles of sampling into pronghorn inventory efforts and have tested for various counting biases. Field activities for both tasks have been curtailed and the existing data awaits analysis, sununary, and reporting. B. OBJECTNE .ThslU Test for density dependence in Middle Park population parameters and sununarize the fmdings in manuscript format suitable for submission to a professional wildlife management or ecological journal. �156 Task 2 Analyze and summarize the results of experiments in pronghorn inventory methods and report the fmdings in manuscript format suitable for submission to a professional wildlife management journal. C. EXPECTED RESULTS OR BENEFITS Produce manuscript(s) and submit them to the appropriate peer reviewed publications. D. APPROACH In.sk.l Summarize population size and herd structure data and animal movement data in appropriate form to describe the effects of increasing density as the population expanded over the years of the study. Some specific topics to be addressed are: ~ a. Population dynamics - growth and structure Seasonal and annual area of habitation by sex and age class c. Dispersal and mortality of males and females in relation to density d. Summer home range fidelity and size by individual animals related to density e. Seasonal habitat use f. Investigate models to describe density dependence Iask.1 Summarize and compare pronghorn density estimates and precision of fixedwing line transect surveys and helicopter quadrat surveys. E. LOCATION Data analysis and reporting will be done at the Colorado Division of Wildlife Research Center, 317 W. Prospect Rd., Ft. Collins, CO 80626. F. ESTIMATED COST Fiscal Year 1997-98 Estimated Costs $1,000 PFTEs 1.00 �157 Colorado Division Wildlife Research July 1997 of Wildlife Report JOB FINAL REPORT state of Project Colorado No. W-153-R-I0 Work Plan No. 3A Job No. Period Author: Mammals Research Pronghorn Inyestigations Experimental Pronghorn Surveys Using Fixedwing Line Transects and Helicopter Ouadrats Covered: July 1, 1996 - June 30, 1997 T.M. Pojar ABSTRACT The first year of a 3 year study was completed to compare pronghorn population density estimates obtained by fixedwing line transect and helicopter quadrat surveys in the North Park Colorado and Big Creek Wyoming area. According to area biologists, these areas constitute a discrete population and should be surveyed and managed as such. Line transects were run on 1.609 km (1 mile) intervals and 10% of the area was surveyed by quadrat. The estimate from the quadrat survey was 2,607 pronghorn with a 90% confidence interval of ± 26%. In addition to a population estimate all animals seen were classified. The B:100D ratio was 49B:100D based on a sample of 263; the 90% confidence interval was large as is usually the case with B:100D ratio estimates - ± 48%. A reasonable fit to the line transect data was obtained using the half-normal function with 1 cosine adjustment term. This resulted in a population estimate of 1,850 with approximate 90% confidence limits of ± 27%. The group size used in this estimate was 2.0145 which is the mean for all groups seen in the first 2 bands. A manuscript authored by Pojar, Bowden, and Madison will be prepared and submitted to the Wildlife Society Bulletin; it will include data from this effort as well as the line/quad comparison in the Craig area from 1993-1995. The original Program Narrative describing the study is included in Appendix I; although the original PN called for the first data collection to begin in 1997-98, it was begun in 1997 because cost center administrators were able to fund it early. Since this project was terminated after only 1 year of an approved 3-year study, a Program Narrative was written to describe the data analysis and reporting for this project (Appendix II). �158 MANAGEMENT RECOMMENDATIONS The relationship between quadrat and line transect surveys should be established for each area that line transect sampling is contemplated. The relationship could vary with terrain, vegetative type, and pronghorn density. The minimum data set to estimate this relationship is 3 years of data. It is then preferable to make this same comparison every third or forth year thereafter. Line transect methodology must be followed meticulously and observers must be trained in the principles and techniques of line transect methods to get the best results. Sampling intensity should be 10% for quadrats and line samples should be at 1.609 km (1 mile) intervals for these comparisons. However, sampling at every third or fourth 1.609 km may be sufficient in some areas of Colorado if an adequate number of groups can be observed. �159 APPENDIX I PROGRAM NARRATIVE PRONGHORN INVESTIGATIONS State: Colorado T. M. Pojar Project Title: Project Number: Experimental pronghorn surveys using transects and helicopter quadrats. Work Plan 3Ar Job 7 October 31, 1996 fixedwing line Transect surveys are popular because they make efficient use of air time in terms of surveying geographic area. Fixedwing aircraft are most commonly used for transect surveys because they are usually available to biologists and they are more economical to operate than helicopters. Conventional strip r~t surveys attempt to count all pronghorn (Antilocapra americana) on contiguous strip transects that cover 100% of the area (Springer 1950, Hailey 1979, Gill et al. 1983, and Allen and Samuelson 1987). The total number observed -is then taken as the total for tnat population with no adjustments for detection errors, despite abundant evidence that subjects of aerial surveys are nearly always undercounted (Bergerud 1963, Graham and Bell 1969, pennycuick and Western 1972, caughley 1974, Parker 1975, Norton-Griffiths 1976, Melton 1978, Bayliss and Giles 1985, Packard et al. 1985, Pollock and Kendall 1987, Bayliss and Yeomans 1989, Firchowet al. 1990, Johnson et al. 1991, Lefebvre and Kochman 1991). Line transect surveys are attractive because, to be effective, they do not require that all subjects within the bounds of the transect are seen (Burnham et al. 1980). Other attractive features of line transects are that they require minimal effort to establish (White et al. 1989), they can be done from fixedwing, and they make efficient use of air time. The most stringent requirement, however, is that all subjects in the first interval are accurately enumerated. Accurate delineation of all other distance interval boundaries is important to ensure that the detection function is fit to the real distribution of observations. Line transect methodology was used to estimate mule deer (Odocoileus hemionus) density using a helicopter as the observation platform (White et al. 1989). Pojar et al. (1995) used helicopter line transects to estimate pronghorn density in 2 areas of Colorado. A specially equipped fixedwing aircraft was used by Johnson et al. (1991) to survey pronghorn herds in Wyoming and they obtained adequate line transect samples in ~ 50% of the flight time required for conventional trend counts. Although fixedwing line transect surveys are attractive for economic reasons, estimates of large ungulate densities based on this technique have not been compared with known densities or other methods where bias has been estimated. Bias was examined for heiicopter quadrat surveys by Pojar et al. -(1995). Overcount bias was not detected and undercount bias would have been proportional to the number of animals that did not flush and were not detected during a search of the quadrat by 2 helicopter search crews. These authors believe that helicopter surveys of quadrats is the least biased of any method they tested - helicopter line transect, wide strip transect, and narrow strip transect. Relative to helicopter quadrat surveys, line transects are perceived to be easier to execute and operation costs of fixedwing aircraft are about onethird that of helicopter costs. Therefore, if fixedwing line transect surveys produce estimates of comparable density and precision to helicopter quadrat �160 surveys at less cost, then they can be considered for more extensive application as a management tool in Colorado. Line transect data analysis is burdened with subjective decisions on grouping, truncation, and model selection, any of which can dramatically affect density and variance estimates. In contrast, quadrats feature a finite population sampling approach and the analysis procedure is direct with no subjectivity in the calculation of density and variance. From an analysis perspective, quadrats offer a standardized procedure which does not add to the variability of the density and variance estimates. However, in the past it has been expensive to locate and mark quadrat corners which was one of the principle obstacles to more widespread use of quadrat surveys. Recent technology known as the Global Positioning System (GPS) provides sufficiently accurate navigation to eliminate the need to physically mark quadrats. Although the typical error for a GPS receiver is claimed to be 20-35 m (Hurn 1989), errors may be somewhat larger but still within acceptable limits for aerial quadrat surveys. Compare pronghorn density estimates and estimated precision of fixedwing line transect surveys with those of helicopter quadrat '.surveys. 2. Evaluate consistency of line transect data analysis. 3. Compare costs of fixedwing line transect and helicopter quadrat surveys Expected Results and Benefits: Good management of any species is anchored to reliable estimates of density. Fixedwing aircraft cost about one-third as much to operate as a helicopter. Therefore, if a survey method using fixedwing produces comparable estimates of density and variance to a helicopter quadrat survey, then the' management agency will realize a significant savings. The conventional fixedwing strip transect method of estimating pronghorn density in North Park, Colorado, has been used exclusively in the past. This method has serious shortcomings (Pojar et al. 1995). A Management Plan has recently been drafted for this pronghorn herd management area (PH-3) (Steinert a~d Snyder 1996) and a more defensible, statistically sound method of estimating pronghorn density is necessary. Use of the fixedwing line transect method is desirable because it is more economical than helicopter quadrat surveys. A sound basis for both the Management Plan and use of fixedwing line transects is possible by establishing the relationship between quadrat and line surveys for this area. Approach: The following hypotheses will be tested: 1. Ho: There is no difference in the expected value of the density estimator between fixedwing line transect and helicopter quadrat surveys. 2. Ho: Variance component associated with differences in data set analysis of the same data set among scientists is small compared to other variance components. The following criteria will also be used to evaluate the 2 survey methods: 1. Precision as measured by the 90% confidence interval. 2. Cost, in terms of time and money, of the surveys. The fixedwing line transect will be conducted as outlined by Johnson et al. (1991) and analyzed according to the guidelines provided by Buckland et al. (1993) and Laake et al. (1993) using the program DISTANCE •. Navigation �161 will be with GPS equipment. Quadrats will be surveyed as described by Pojar et al. (1995) with the exception that quadrats will be located with GPS navigational equipment. Latitude and longitude coordinates will be estimated for each quadrat corner from USGS topographic maps. A GPS receiver mounted in the helicopter, using an external antenna, will be used to locate the quadrat corners. A 3 person crew (pilot, navigator, and primary observer) will conduct the quadrat survey. The t-test will be used to compare estimated pronghorn densities from line transect and quadrat surveys. For purposes of this experiment, quadrat results will be the standard. Location: There are approximately 410 square miles of yearlong pronghorn habitat in North Park (Steinert and Snyder 1996). This will be the area surveyed. Forty randomly drawn quadrats (a 10% sample) will be used in this experiment for the quadrat survey. A line transect will be run on the center line of each i~de strip through the area of investigation. The line transect survey will be done within 1 week of the quadrat survey. Schedule: 1997-98 . 1997-98 1997-98 1998-99 1998-99 1999-01 1999-01 April - estimate quadrat corner coordinates from USGS maps. May - conduct both fixedwing line and helicopter quadrat surveys. June-July - data analysis and preliminary report. May - conduct both surveys. June-July - data analysis and interim report. May - conduct both surveys. June-December - data analysis and final report. Personnel: Principal Investigator: Tom Pojar Consultants: Dave Bowden, Bruce Gill, Gary white, Dave Anderson, and Ken Burnham Cooperators: Jeff Madison, Jim Olterman, Kirk Snyder, Steve Steinert, Chuck Wagner, J. Wenum, and Susan Werner of the CDOW; and Biff Burton, Rich Guenze1, and Tim Thomas of the Wyoming Game and Fish. Literature Allen, Cited: S. H. and J. M. Samuelson. 1987. Precision and bias of a summer aerial transect census of pronghorn antelope. Prairie Nat. 19:19-24. Bayliss, P., and J. Giles. 1985. Factors affecting the visibility of kangaroos counted during aerial surveys. J. Wildl. Manage. 49:686-692. Bayliss, P., and K. M. Yeomans. 1989. Correcting bias in aerial survey population estimates of feral livestock in northern Australia using the double-count technique. J. Appl. Ecol. 26:925-933. Bergerud, A. T. 1963. Aerial winter census of caribou. J. Wildl. Manage. 27:438-449. Buckland, S. T., D. R. Anderson, K. P. Burnham, and J. L. Laake. 1993. Distance Sampling: Estimating abundance of biological populations. Chapman and Hall, NY. 471pp. �162 Burnham, K. P., D. R. Anderson, and J. L. Laake. 1980. Estimation Of density from line transect sampling of biological populations. Wildl. Monogr. 72. 202pp. Caughley, G. 1974. Bias in aerial survey. J. Wildl. Manage. 38:921-933. Firchow, K. M., M. R. Vaughan, W. R. Mytton. 1990. Comparison of aerial survey techniques for pronghorn. Wildl. Soc. Bull. 18:18-23. Gill, R. B., L. H. Carpenter, and D. C. Bowden. 1983. Monitoring large animal populations: The Colorado experience. N. Am. Wildl. Conf. 48:330-341. Graham, A., and R. Bell. 1969 (68) • Factors influencing the countability of animals. E. Afr. Agric. For. J. Special Issue 34:38-43. Hailey, T. L. 1979. A handbook for pronghorn antelope management in texas. Texas Parks and Wildl. Dep. No. 20. 66pp. Hurn, J. 1989. GPS A Guide to the Next Utility. Trimble Navigation, P.O. Box 3642, Sunnyvale, California. 76pp. Johnson, B. K., F. G. Lindzey, and R. J. Guenzel. 1991. Use of aerial line transect surveys to estimate pronghorn populations in Wyoming. Wildl. ·~~~-bC. Bull. 19:315-321. Laake, J. L., S •.T. Buckland, D. R. Anderson, and K. P. Burnham. 1993. DISTANCE User's Guide V2.0. Colorado Coop. Fish and Wildl. Res. Unit, Colorado State Univ., Fort Collins, CO. 72pp. Lefebvre, L. w., and H. I. Kochman. 1991. An evaluation of aerial survey replicate counting methodology to determine trends in manatee abundance. Wildl. Soc. Bull. 19:298-309. Melton, D. A. 1978. Undercounting bias of helicopter censuses in Umfolozi Game Reserve. Lammgergeyer 26:1-6. Norton-Griffiths. M. 1976. Further aspects of bias in aerial census of large mammals. J. Wildl. Manage. 40:368-371. Packard, J. M., R. C. Summers, and L. B. Barnes. 1985. Variation of visibility bias during aerial surveys of manatees. J. Wildl. Manage. 49:34735l. Parker, G. R. 1975. A review of aerial surveys used for estimating the numbers of barren-ground caribou in northern Canada. Polar Record 17(111):627-638. Pennycuick, C. J., and D. Western. 1972. An investigation of some sources of bias in aerial transect sampling of large mammal populations. E. Afr. Wildl. J. 10:175-191. Pojar, T. M., D. C. Bowden, and R. B. Gill. 1995. Aerial counting experiments to estimate pronghorn density and herd structure. J. Wildl. Manage. 59:117-128. Pollock, K. H., and W. L. Kendall. 1987. Visibility bias in aerial surveys: A review of estimation procedures. J. Wildl. Manage. 51:502-509. Springer, L. M. 1950. Aerial census of interstate antelope herds of California, Idaho, Nevada, and Oregon. J. Wildl. Manage. 14:295-298. Steinert, S. F. and K. F. Snyder. 1996. Antelope management plan: Data analysis unit PH-3. Colorado Div. Wildl. 16 pp xerox. White, G. C., R. M. Bartmann, L. H. Carpenter, and R. A. Garrott. 1989. Evaluation of aerial line transects for estimating mule deer densities. J. Wildl. Manage. 53:625-635. �163 APPENDIX II PROGRAM NARRATIVE state of Project No. Work Package Task No. A. Colorado W-1S3-R No. 3004 1 & 2 Cost Center 3430 Mammals Program Management of Other Ungulates Pronghorn Data Analysis and Reporting NEED According to the Colorado Wildlife Commission's Long range Plan (March 11, 1994) it is the Commission's policy that "The Division will manage it's wildlife-related recreation programs based upon sound biological principles and up-to-date demographic information on public expectations and value." To aC~lish this, the Long Range Plan states a goal for the Division of ildlife to "Use hunting as the primary tool to manage big game populations to maintain a balance between these species and their habitats (Goal 10.1). state-of-the-art wildlife population inventory methods are an essential .component in maintaining the balance between wildlife species and their habitats. Pronghorn are no exception. Over the past one and a half decades the Division has invested considerable effort into improving methods for estimating pronghorn numbers and sex and age composition; this expanded our knowledge of pronghorn population performance under a variety of environmental circumstances and habitat constraints. This project proposes to analyze, summarize, and report on the most recent data sets which address these issues. Two primary tasks have been the focus of this work; the first monitored performance of a pioneering pronghorn population in Middle Park Colorado and the second compared several methods for estimating population parameters. Historically, pronghorn were abundant in Middle Park but were extirpated at the turn of the century largely as a consequent of overhunting. In the late 1970s, pronghorn began to pioneer into Middle Park from North Park on a seasonal basis and by the early 1980s, they were once again yearlong residents. This pioneering population provided a unique opportunity to study the growth and distribution of an unhunted population that was re-establishing itself into historic range. Monitoring population characteristics, movements, and habitat occupation as it expanded provided critical information for modeling of this species and insights into environmental resistance characteristics that are useful in establishing a balance between pronghorn and habitats in sagebrush steppe ecosystems. Traditional pronghorn inventory methodology has consisted of trend counts or total area counts. Trend counts assumed (but did not establish) that trend areas were characteristic of the entire population and that annual variation in trend counts reflected variation in population trajectory. Total area counts assumed that pronghorn could be counted over large areas without error, or at a minimum, with consistent error. Neither of these premises were verified. Work over the past several years have incorporated principles of sampling into pronghorn inventory efforts and have tested for various counting biases. Field activities for both tasks have been curtailed and the existing data awaits analysis, summary, and reporting. �164 B. OBJECTIVE Task 1 Test for density dependence in Middle Park population parameters and summarize the findings in manuscript format suitable for submission to a professional wildlife management or ecological journal. Task 2 Analyze and summarize the results of experiments in pronghorn inventory methods and report the findings in manuscript format suitable for submission to a professional wildlife management journal. C. EXPECTED RESULTS Produce manuscript(s) publications. D. OR BENEFITS and submit them to the appropriate peer reviewed APPROACH ~,,,,, ~"":'Tasl{ 1 Summarize population size and herd structure data and animal movement data in appropriate form to describe the effects of increasing density as the population expanded over the years of the study. Some specific topics to be addressed are: a. b. c. d. Population dynamics - growth and structure Seasonal and annual area of habitation by sex and age class Dispersal and mortality of males and females in relation to density Summer home range fidelity and size by individual animals related to density e. Seasonal habitat use f. Investigate models to describe density dependence Task 2 Summarize and compare pronghorn density fixedwing line transect surveys and helicopter E. estimates and precision quadrat surveys. LOCATION Data analysis and reporting will be done at the Colorado Division Research center, 317 W. Prospect Rd., Ft. Collins, CO 80626. F. of ESTIMATED Fiscal COST Year 1997-98 of Wildlife Estimated $1,000 Costs 1.00 �165 Colorado Division Wildlife Research July 1997 of Wildlife Report JOB PROGRESS REPORT state of project Work Colorado No. ~W~-~1~5~3~-~R~-~1~0L- Plan No. __~4~A~ _ Mammals _ Mountain Research Goat Inyestigations Mountain goat numbers distribution, and dispersal in the northern Collegiate range. Job No. ~, Period Author: Covered: July 1, 1996 - June 30, 1997 D. F. Reed ABSTRACT Mark-resight population estimates, distribution and habitat use, and the pioneering of mountain goats in the northern Collegiate range has been investigated since 1994. During three winters and three hunting seasons, 8 collared animals have died and 8 have been harvested. Seven additional telemetry collars were attached in 1996 to replace some of those lost. However, one of the 7 additional collars was put on an animal out of the markresight study area northwest of Mount Elbert. This individual, in a group of 20, has apparently remained in this "dispersal" area throughout the year. Helicopter counts were conducted 25 August 1994, 30 August and 1 September 1995, 18 July, 2 August, and 7 October 1996, and 25 August 1997. The results of these counts ranged from a low number of the marks (radio-collars) being observed and poor identification of distinguishing marks (numbered or color coded collars) (9 of 39 marks; sightability = 0.23) to a high number of marks being observed and good identification of marks (29 of 36; sightability 0.81). The best estimate for the study area population (north of Clear Creek) appears to be from the count on 30 August 1995 (N = 173 [CI 147-203J). Movements of up to 17 km have been detected for both males and females. The group of 20 goats found northwest of Mount Elbert (in which Black 74 radiocollar was placed) was 13 km north of Lake Creek and Highway 82. This group plus another group of up to 44 animals south of Independence Pass appear to be significant dispersal movements. Planned manuscripts for both the "dispersal" and "mark-resight" questions addressed in this study are titled "Dispersal in introduced mountain goats" and "Mark-resight population estimates of mountain goats in the Quail Mountain - La Plata Peak area of Colorado." ��167 * MOUNTAIN GOAT NUMBERS, DISTRIBUTION, AND DISPERSAL COLLEGIATE RANGE IN THE NORTHERN Dale F. Reed P •N. OBJECTIVE To improve estimates of mountain goat populations by mark-resight methodology, to determine distribution, and to estimate dispersal rates in an increasing mountain goat population. SEGMENT OBJECTIVES 1. Periodically locate radio-collared mountain goats in the northern Collegiate range (G-3N) to track movements, habitat use, and dispersal. ~- 2. Estimate mountain goat numbers using mark-resight methodo'logy. STUDy AREA The study area is described in the Program METHODS Narrative (Reed 1995). AND MATERIALS The methods are outlined in the Program (Appendix A in Reed 1995, Reed 1996). Narrative and 1996 progress report RESULTS Results are reported in the 1996 progress report (Reed 1996) or the manuscripts in preparation titled "Dispersal in introduced mountain goats" "Mark-resight population estimates of mountain goats in the Quail Mountain La Plata Peak area of Colorado". LITERATURE and - CITED Reed, D. F. 1995. Mountain goat numbers, distribution, and dispersal in the northern collegiate range. Colo. Div. Wildl. Res. Rep. July, 232-234pp. Reed, D. F. 1996. Mountain goat numbers, distribution, and dispersal northern Collegiate range. Colo. Div. Wildl. Res. Rep. July, Prepared by _ Dale F. Reed in the pp. ��Colorado Division Wildlife Research July 1997 of Wildlife Report JOB FINAL REPORT state of Project Work Colorado No. W-153-R-10 Plan No. Black Research Bear Research Development of black Inventory Techniques Job No. Period Mammals Covered: Author: Personnel: Thomas July bear 1, 1996 - June 30, 1997 D. I. Beck J. Aragon, T. Beck, S. Birch, J. Eussen, D. Finley, R. Firth, J. Frothingham, B. Holder, L. Willmarth, D. Younkin, CDOW; D. Harper, D. Marlow, Colo. state Patrol; D. Schmidt, USFS; D. Bowden, G. White, Colo. State Univ. ABSTRACT A black bear (~ americanus) resighting system utilizing 35-mm cameras activated by active infra-red sensors was employed throughout a 404 kro2 area. Thirty-nine cameras were distributed, one each per 10.4 kro2 quadrant. During 6 resighting sessions of 14 days each, 108 pictures of bears were obtained. Seventy-seven were tagged, of which 70 were discretely identified. A new, collar-based marking system proved reliable. Data from both the Uncompahgre and Middle Park study areas were reanalyzed based on 4 resighting sessions because of large degrees of population shuffling during the latter two sessions; which coincide with onset of berry ripening. The estimated population size in Middle Park study area was 33, (95% CI 25-43), for a density estimate of 8.1 bears/100 kro2• The estimated population size for the Uncompahgre study area was 182, for a density estimate of 39.0/100 kro2• Total capture of bears was a poor index of relative black bear density which would over-estimate black bear presence in the lower-density habitats. Little difference was observed between areas in distribution of visits to camera sites by day or time; with over 85% of visits in the first 10 days and peak activity occurring in the last 3 hrs of light. Nocturnal pictures represent less than 16% of total resightings. The estimation procedure still provides wide confidence intervals for the Middle Park study, primarily because of the small population size and the marginally acceptable resighting rate of 28%. The described procedure can result in adequate captures (>50% of N) and minimally acceptable resighting rates (30%) for a cost of approximately $40,000 per 450 kro2 to obtain black bear density estimates. It is unknown if a higher density of cameras would increase resighting rate but costs would escalate rapidly. ��171 DEVELOPMENT OF BLACK BEAR IHVEHTORY Thomas P.R. bias D. I. Beck OBJECTIVE 1. Evaluate a capture-sight program estimating black bear density. 2. Document age and gender hunting seasons. TECHNIQUES utilizing cameras in vulnerability set on bait of black 3. Obtain density estimates of black bears in 3 heavily markedly different vegetation communities. bears hunted stations during areas for autumn of SEGMENT OBJECTIVES 1. Evaluate the use of infra-red triggered bears in the Middle Park study·area. 2. Evaluate NOREMARK for estimating 3. Describe fall habitats Colorado. of black 4. Capture bears and mark black cameras black bears AND marked black bear density. in the northern on a study METHODS to resight montane area in south-central forests of Colorado. MATERIALS study Area Description The Middle Park study area was located in GMU 18 in an area of near-continuous montane conifer forest. Mountain areas vegetated with spruce-fir (~ ~) and lodgepole pine (~ contorta) forests account for 30% of the statewide black bear habitat (Beck 1991) and the Middle Park area is representative of these forest communities. The 404 km2 area was divided into 2 39 quadrants, each 10.4 km • The area is roughly 22 km north-south and 19 km east-west. The west boundary is the E. Fork Troublesome Creek, the east is a north-south line over Little Gravel Mtn., the north boundary is just south of the Continental Divide, while the south boundary parallels the Colorado River. The south boundary is the only one with a distinctive vegetation change with the lower boundary being the upper limits of extensive sagebrush (Artemisia spp.) hills. Elevations range from 2380 m to 3540 m. The average frost-free period is approximately 45 days. Three quadrants in the northwest corner which were used for trapping were not used for resighting because of severe access problems. The southern row and western 2 columns of quadrants exhibit a mosaic of vegetation with substantial stands of aspen (Populus tremuloides). The remainder of the quadrants are characterized by relatively homogenous conifer stands of lodgepole pine or spruce-fir interspersed with meadows. �172 Resighting Black Bears The resighting system was comprised of active infra-red triggers, a recorder, and a 35-mm point-and-shoot camera which was commercially available from TRAILMASTER (Lenexa, KS). set-up of the units basically duplicates the system used on the Uncompahgre Plateau, as described by Beck (1995), with the following modifications. In an attempt to reduce the number of dark exposures taken in nocturnal periods, a newer Olympus camera (Infinity Mini DLX) was purchased for 25 of the units. The newer camera had a different electronic configuration which was expected to insure that all initial pictures had an operational flash. However, this model did not have a CONTINUOUS SHOOT mode, as did the older models (AF-1 Twin). Thus, to obtain multiple pictures of each bear in order to maximize chances of identify~ng unique markers, the recorder was programmed to have a 0.2 minute delay after each beam break. This contrasts with the 15minute delay used with the older units, which normally took 3-4 pictures at the initial beam break. For analysis purposes, both camera systems were treated the same. Each IS-minute period was treated as a separate sighting and all pictures in that period were counted as one sighting. The possibility of some animals triggering an entire roll of film was balanced against the concern that the dark exposures were missing nocturnal visits of bears. Thus, the split system would allow analysis of the trade-offs. Six resighting sessions were conducted in 1996 at biweekly intervals beginning 12 June and ending 1 September. Three different sites were used in each quadrant, with 2 consecutive sessions occurring at each site. Attractant baits varied between sessions, in the following order: rotting fish, rotting beaver, Very Berry synthetic lure, rotting chicken, Shellfish synthetic lure, and rotting fish. Camera height was standardized at 2 m and distance from camera to infra-red transmitter was selected to be 4 m, or as close as allowable given terrain. Because of cold evening temperatures, camera batteries (AA) were changed at each session. Batteries (C) to operate the infra-red units were changed after 2 sessions. Fuji print film, ASA 400, in 36-exposure rolls was used. All film was developed to negative rolls, examined and data recorded. Negatives of black bears were then developed into prints for examination for unique collar marks. All negative and photo examination was done on a light table with the aid of an 12X magnifier. Photographed black bears were recorded as: unknown classification, tagged but unable to discern tag combination, unique tag identified, untagged, and cub. Also, within the untagged group, subjective evaluations of the yearling cohort was attempted. Other wild and domestic animals were recorded by species. When no animals were observed in photographs the category SITE was tallied. Completely dark exposures were tallied as DARK SITE. Data back units on the camera were programmed to print each photograph. These were then compared to recorded bern as recorded on the TRAILMASTER unit. date and time of day on breaks in the infra-red A total of 42 black bears were collared and residing iIi or near the study area during 1996. Aerial radio-tracking was conducted every Tuesday (with 2 exceptions) during the months of June, July, August, and early September. Locations were plotted by UTM's and also recorded as IN or OUT of the 39- �173 quadrant study area or as NOT FOUND. If a black bear was recorded as IN during any part of the session, it was considered as available for resighting during that session. Not all bears were located each flight and occasionally subjective decisions on IN or OUT status were made based on locations during previous and subsequent flights, location data from the previous year, patterns of movements, and age and sex class of the bear. Judgment decisions were noted separately from unambiguous aerial locations. Density Estimation The population estimation technique utilized was the Bowden Estimator available through PROGRAM NOREMARK (White 1996) and described by Bowden and Kufeld (1995). This is a mark-resight procedure which utilizes the frequency of multiple sightings of marked individuals to estimate the number of unmarked bears observed in the resighting procedure. Input data required for each session were: number of marked bears in area, number of marked bears observed, the number of times each marked bear was observed, and the number of unmarked bears observed. The resultant population estimate includes all bears yearling and older. It does not include first-year cubs. This is an important point when comparing to other studies. Description of fall·habitat use Limited radio-tracking in 1995 left more questions than answers as to the important fall feeding areas and the important fall foods. Radio-tracking in 1996 was to be ground-based in September to identify the types of foods being utilized and the general habitat conditions where such foods would be found. Limited ground-based tracking would be conducted in August as time permitted while operating the camera systems. The point of primary interest was the species of bear food present at late-summer, early-fall black bear locations. Identify a study area in South-central Colorado and mark bears A 6-day reconnaissance was conducted in the areas both east and west of Trinidad, CO in May, 1996. General habitat conditions were examined, extensive walking surveys for black bear si9n were conducted, and initial meetings with potentially cooperative landowners were conducted. The land tenure in this area is primarily large holdings in private ownership. A typical 450 km2 study site would involve 2-4 owners. RESULTS Resighting Black AND DISCUSSIQN Bears The modified camera-recorder system was effective at obtaining resightings of black bears (Table 1). The total number of pictures was substantially less in Middle Park than on the Uncompahgre (294 vs. 1735) The combination of greater user experience in Middle Park and the newer designed cameras resulted in a lower proportion of site pictures and higher proportions of bear pictures. Black bear pictures accounted for 30% and 37% of all pictures on the Uncompahgre and Middle Park study areas, respectively. Similar percentages for site pictures were 34 and 22 and for dark site pictures 6 and 1. Noticeably fewer cattle pictures were obtained in Middle Park because of both differing grazing intensities and a concerted effort to avoid cattle in Middle Park. �174 The use of colored dowels attached to the radio collar was an effective unique marker. Loss or destruction of dowels has not been observed and 90% of photographed tagged bears were accurately identified based on dowel color combinations and/or ear tag numbers. The dowel system was far superior to the ear tag/streamer combinations used in the Uncompahgre study. Of 94 black bear photographs which we recorded date of visit, 52% were in the first 5 days, 39% in the second five days, and 9% in the last 5 days. Comparable splits for the Uncompahgre study had been 51%, 30%, and 19%. Thus it seems that a 15-day sighting period is reasonable and that the acent; baits remain efficacious throughout this time. For the effort involved, periods of less than 10 days will result in substantial data loss. The proportion of black bear photographs taken during daylight hours in Middle Park was 84%; very similar to the 88.5% on the Uncompahgre. Thirty-two percent of all Middle Park activity was in the 3-hr period preceding darkness (dusk; 1800-2100). The need for ASA 1000 film is not warranted, and in fact would not likely provide the clarity needed during daylight hours. The use of a 0.2 minute delay on the newer cameras was not a problem. The maximum number of sequential pictures taken was 5 and that only 1 time. The modal number of pictures per bear was 1. This was anticipated given that over Table 1. Photographic CATEGORY summary 1 for Middle 2 5 6 23 13 4 2 2 2 17 13 12 7 2 2 2 3 22 12 1 6 2 1 108 70 7 13 14 4 11 10 8 1 66 3 1 2 2 4 1 10 5 3 7 33 2 8 1 11 22 17 1 1 3 12 8 1 Site-no animal Site-dark 17 11 2 9 2 2 1 Cattle Dog 1996. SESSION 3 4 BEARS: Tagged Tagged, unk untagged 2+ untagged Yrl Unknown Cubs 2 1 Park, People 2 1 1 Elk Moose Deer Puma Chipmunk Red squirrel Snowshoe hare Marten Coyote Striped skunk Moth porcupine Marmot 7 1 2 5 1 6 6 9 3 2 1 2 2 2 5 1 1 1 1 7 1 2 1 1 2 1 4 1 2 2 2 1 2 3 2 1 1 1 4 1 1 TOTAL 9 5 20 2 1 3 3 1 �175 60% of Uncompahgre bear activity at the infra-red min. We observed time splits on 32 sequences of quadrant. Thirty of these were separated by more minutes, and 1 by 1 minute. There appears to be bears to leave the immediate site as soon as the Density beam was for periods < 1 bear photos at the same than 60 minutes, 1 by 15 a strong tendency for black cameras activate. Estimation Estimates of the population size were made based on the first 4 sessions and all 6 sessions. The estimate for 4 sessions was 33 (95% CI = 25-43). The estimate for 6 sessions was 37 (95% CI = 26-51) (Table 2). During anyone session, the number of coilared bears available for sighting ranged from 25 to 30, averaging 27, although 38 different collared black bears were in the study area over the course of the summer. The resighting rate for the sessions varied from a high of 38% (Session 1) to a low of 19% (Session 5) and averaged 28% (Table 2:Col 2 divided by Col 1). The proportion of unmarked bears which were yearlings, and thus not available for tagging in 1995 since we did not handle cubs, was 64% (9 of 14) for the first' 4 sessions and 52% (14 of 27) for all 6 sessions. To calculate the trapping efficiency (proportion of population marked in 1995), use the 4session population estimate of 33 resident bears, of which 27 were marked, and of which 4 of the 6 unmarked are yearlings. Thus the high estimate for trapping efficiency would be 93%. The low end of trapping efficiency can be represented from the 6-session numbers: population estimate of 37, average marked of 27, 5 of 10 unmarked are yearlings. This would result in a trapping efficiency of 84%. Either calculation indicate the trapping program resulted in a very large majority of resident bears being marked in one season. Table 2. Bowden Estimator (PROGRAM NOREMARK) population, Middle Park study area, 1996. Sess. 29 28 25 25 27 30 1 2 3 4 5 6 a Marked Bears Number No. Mark Observed No. Time Seen 11 7 6 8 5 9 17 8 13 13 7 12 of yearlings ?? Ident in (); unavailable 1 1 4 0 0 1 estimates of black No. Unmark 4 2 4 4 5 8 Popln Est. (3) a (2) (2) (2) (3 ) (2) for tagging 35 34 30 32 43 47 bear 95% CI 28-44 26-43 22-41 23-43 26-69 33-67 in 1995. Simulation work on the sensitivity of the population estimation procedure was conducted prior to either field study by Dr. Gary White, colo. State Univ. Based on these simulations, the minimal targets for field work were to capture and mark 50% of the resident population, obtain a resighting frequency of at least 30%, and conduct at least 4 resighting sessions (G. White, pers comm). Increases in precision would be aided most by increasing resighting rate, followed by increasing proportion marked. While resighting rate cannot be calculated for the Uncompahgre study because of pulled ear-tags, the capture efficiency there was nearly 59% (based on capture of 89 bears, 32% estimate of yearlings, and population estimate of 182). Therefore, the trapping intensity �176 was adequate to capture sufficient bears in both high and low density ,study areas. However, the resighting rate is at the low margin; even with a camera site every 10.4 km2• While data are presented for all 6 sessions, there are reasons to believe the analyses from only the first 4 sessions are most appropriate. On the Uncompahgre study area, a significant shuffling, both in-migration and out~ migration, occurred between Session 4 and 5. This was triggered by the seasonal movement of black bears to low-elevation berry and acorn supplies. For Session 6, 36% of collared bears were not present in the study area. On a given flight, as many as 48% were outside the study area. In the Middle Park area, even though much higher in elevation and with different mast species, the late summer shuffle also began in early August. Although few collared bears left the area entirely, they made dramatic shifts in location and a high proportion were along boundaries. It was during Session 6 that the most nonyearling unmarked bears were observed in photographs. Although increasing the number of sessions is expected to reduce variance of the estimate, in the Middle Park area it did the opposite; presumably because of the shuffling of bears into the study area. The lay-out o~ the study area had the misfortune to include a major late-summer black bear concentration area along the east boundary (Little and Big Gravel Mountains, Trail Mountain). For these reasons, further comparisons of data will use the estimates derived from Sessions 1-4 for both study areas. The entire data sets were presented for those that do not agree with this rationale. Attenuation to bait sites without reward (no food, only scent) was a serious concern which was addressed by changing scents each session. There is no evidence that a significant attenuation occurred through 6 sessions in Middle Park (Table 1). While there was a decline in pictures per session for the last 2 sessions on the Uncompahgre (95 for 1-4, 72 for 5-6) it is not possible to attribute cause since there was so much seasonal out-migration occurring then as well. The Uncompahgre Plateau is subjectively considered to be among the best black bear habitats in Colorado by the principal investigator. This contention is supported by 18 years of hunter killed black bear records. Conversely, the coniferous forests of the high-elevation parks is subjectively considered to be among the poorest black bear habitats in Colorado; also supported by the kill data. Thus it may be of interest to look at the comparative data. The density estimate for the Uncompahgre area was 39.0 bears/100 km2 and for Middle Park 8.1 bears/100 km2; a ratio of 4.8:1. Is trapping with near-equal intensity a good indicator of relative density? On the Uncompahgre area, 89 bears were captured in 45 quadrants (1.98 bears/quad) while 49 bears were captured in 39 quadrants in Middle Park (1.26 bears/quad); a ratio of 1.6:1. In the Middle Park area, it appears that even with a relatively large study area, the number of bears captured included a few transient bears and several border area bears along with a large majority of the resident bears. Thus using capture rate to compare areas of different habitat does not appear warranted. Somewhat surprisingly, the sighting frequency closely followed the estimated densities. The sighting rate on the Uncompahgre was 2.11 pictures/quad/session while on Middle Park area it was 0.47 pictures/quad/session; a ratio of 4.5:1. Description of fall habitat use The berry crop in 1995 was quite poor in the study area. Collared bears moved extensively during the August and early-September period but no concentrations near any berries were detected. Examination of bear scats and actual �observation strongly indicated that nearly all feeding during this period was on mushrooms. There was one area of concentrated use near Little Gravel Mtn. and again, there were no berries found but mushrooms were abundant. At about the same time period in 1996, collared black bears began changing their areas of use. The general shift was to go to stands of monotypic lodgepole pine with buffalo berry (Sheperdia canadensis) understory. Most of the destination areas were the same as in 1995. However, 1996 was .a vastly different year and buffalo berries were very abundant in the understory of specific lodgepole pine stands. Many of the lodgepole pine stands have near-monotypic ground cover of Vaccinium spp.; none of which were observed to produce berries in either 1995 or 1996. Unfortunately, by the time field crews were free of camera duties in September, nearly all the buffalo berries had been consumed so more detailed inquiries were not warranted. Based on aerial radio-tracking in both years, black bears appear to return to lodgepole-sheperdia stands each summer in late-July. The buffalo berries appear to be the critical mast crop for black bears in Middle Park. Detailed studies of the understory relationships in lodgepole stands are warranted based on the berry production value of Sheperdia canadensis. Clearly this is a valuable feeding regime for both bears and numerous species of birds in the area. Prior to this study, it had been suggested by the principal investigator that a proportion of the bears would migrate northwest to more productive berry (Prunis, Amelanchier) and acorn (Quercus) production areas approximately 45 km away. However, no such movements were detected from the collared bears in either 1995 or 1996. The bears tricked the PI once again. Identify study area in south-central Colorado and mark bears Extensive hiking and vehicle-based surveys were conducted in the area from Culebra Peaks east to Trinidad, mostly south of the Purgatoire River. Additional surveys east of Trinidad, on Raton Mesa, were also conducted. The general impression was that the areas of highest quality (Raton Mesa and Culebra Peaks) were connected by many miles of generally good, but not outstanding, black bear habitat between. From initial conversations with landowners (facilitated by DWM B. Holder) it appeared that a reasonable study area could be found and landowner cooperation obtained. However, more black bear studies did not survive the extensive reallocation program that the Terrestrial section, CDQW, conducted in Spring/Summer 1996. Therefore, there was no trapping effort conducted. CONCLUSIONS 1. The Bowden estimator method of precision, provides a reasonable estimate, for black bear density studies. 2. Study areas need to be at least 420 km2, preferably physiographic or vegetative boundaries. with with a quantified Borne natural 3. Trapping, based on a network of 10.4 km2 quadrants, at an intensity of 3040 trap-nights over a 60-day period with the newly developed cage traps, will provide for sufficient marked black bears (> 55% of residents). 4. The newly developed cage trap for bears appears to have a high capture efficiency in a variety of habitats (Colorado, Utah, New Mexico studies). It allows for individual workers to safely handle bears alone. Most �178 importantly, it eliminates major injuries to black minor injuries in less than 5% of captures. 5. The active infra-red triggered effective tools for resighting camera systems black bears. bears marketed and results by TRAILMASTER in are 6. A system of 2 colored dowels (2.5 X 10 cm) affixed to the radio collar provides an effective unique identifier for marked bears in photographs. Ear-tag/streamer combinations were not effective, with low rates of retention. 7. A resighting intensity of 4 sessions each 14 days in length with 1 camera/10.4 km2 will provide a minimally acceptable level of resighting (30%). However, at this resighting level, 95% confidence intervals will still be wide. 8. The cost of conducting a density estimate on a single study area is approximately $40,000; much of which is manpower costs. secondary benefits include data on age and sex composition of trapped sample and seasonal movement patterns. 9. In order to project study area densities to larger areas of the state, a series of randomly selected study areas in each vegetation complex would need to be sampled. See Miller et ale 1997 (pgs. 36-39 for more detailed discussion). The current estimates do provide some direction as to the best and worst to be expected in Colorado; from 8 to 39 black bears/100 km2~ LITERATURE Beck, T.D.I. 1991. Black bears Tech. Publ. 39. 86 pp. CITED of west-central Colorado. Colo. Div. Wildlife _____ • 1995. Development of black bear inventory techniques. Colo. Wildlife Fed. Aid Prog. Rep. W-153-R-8, WP 5A, J2. 11 pp. Div. Bowden, D. C. and R. C. Kufeld. 1995. Generalized mark-resight population size estimation applied to Colorado moose. J. Wildl. Manage. 59(4):840-851. White, G. C. 1996. NOREMARK: Population estimation surveys. Wildl. Soc. Bull. 24(1):50-52. from mark-resighting �179 Colorado Division Wildlife Research July 1997 of Wildlife Report JOB PROGRESS state of project Work Colorado No. W-1S3-R-I0 Plan No. Mammals Research Kit Fox Studies lOA Job No. Period REPORT Kit fox (Yulpes macrotis) Status in Colorado Covered: Author: Personnel: July 1, 1996 - June 30, 1997 T. D. I. Beck T. Beck, UNC R. B. Gill, CDOWi J. Eussen, D. Finley, J. P. Fitzgerald, ABSTRACT All survey work conducted in 1992-1996 was synthesized into a summary document on status of kit fox in western Colorado by the primary contractor. This document will provide the guidance and support for detailed conservation strategies. Program planning documents were prepared for continued recovery work. Continued monitoring of kit fox in Montrose East further documented the poor survival of both adults and juveniles. A 69 km dispersal of a juvenile male was documented. ��181 KIT FOX (VULPES MACROTIS) THOMAS P. Document the geographic western Colorado. IN' COLORADO D. I. BECK N'. distribution STATUS OBJECTIVE and relative abundance of kit fox in SEGMENT OBJECTIVES 1. Prepare state recovery 2. Prepare plan. PROGRAM 3. Monitor radio-collared plan NARRATIVE for kit fox. for programs necessary kit fox in Delta, Montrose, to implement and Mesa recovery counties. METHODS AND MATERIALS All of the general survey work, trapping results, monitoring by radiotelemetry, survival, and reproduction data were summarized for the period 1992-1996. Detailed maps of all areas covered by either trapping or spotlight surveys were prepared. Basically, this was an effort to synthesize all we presently know about kit fox status in Western Colorado and present in a single document. This effort was lead by the contractor, J. P. Fitzgerald, of Univ. of Northern Cqlorado. Monitoring of radio-collared kit foxes was done at approximately monthly intervals with greater effort in Feb.-Mar. to document pairing and den location. Nearly all radio-tracking was ground-based as only 4 collared were transmitting and all were in the same general area. foxes Trapping efforts to mark the juveniles of a collared pair were conducted in July, 1996, using the techniques described in detail in Appendix A. Trapping efforts were also made in spring, 1997 to an effort to replace radio-collars on 2 adults. Several on-the-ground searches were made to follow up on reportings of kit foxes in areas where earlier efforts had produced no evidence of foxes. RESULTS AND DISCUSSION' All of the work for the period 1992-1996 under the direction of J. P. Fitzgerald, UNC, was synthesized into a final report to the Colorado Division of Wildlife, and is appended to this report (Appendix A). Following distribution of this report, several discussions were held as to the most urgent needs for kit fox recovery. A number of options were discussed, involving a wide array of both financial and manpower resources. Development of a detailed recovery plan was delayed pending the outcome of a major reallocation of resources within the Terrestrial section of CDOW. This reallocation process was a Division-wide activity dictated by an outside �182 management plan until review. It seemed prudent to refrain from formalizing the level of future support could be ascertained. a recovery A modified PROGRAM NARRATIVE was developed as part of the 5-year renewal of project documents. The new PN focused on recovery activities designed to increase the geographic distribution of kit fox into all appropriate habitats in the Uncompahgre, Gunnison, and Colorado River valleys. Work will initially focus in the Uncompahgre and Gunnison valleys; which are now isolated from the Colorado River valley by altered land uses (irrigated farming, subdivisions). Two radio-collared kit fox (#23, #307) paired in the Montrose East area and raised a litter of pups successfully, even thought the female (#23) died during June, 1996 of unknown causes. She was located buried under 1 m of fresh mud in a steep arroyo < 0.5 km from the den. Three pups were trapped at the den; all were males and each was ear-tagged. One male pup (#316) was subsequently trapped and collared but slipped the collar during the winter. He had stayed within 5 km of the den site during this time. Another of the male pups (#315) was picked up along Highway 50 on 04 Feb 1997. He had been killed by a vehicle collision the previous night. The location of death was at the far north end of the Gunnison valley habitat, a distance of 69 km straight-line from his capture site at the extreme south end of available habitat in the Uncompahgre valley. Clearly he had to disperse through the Peach Valley area, cross the Gunnison River, and continue north through a long expanse of what appears to be good fox habitat but an area where we have yet to document resident kit fox. The only resident kit foxes located north of Peach Valley (Uncompahgre unit) are at the extreme north end of the Gunnison valley; where a family group has a den complex about 2 km from the location of #315's death. We have documented limited dispersal into the Gunnison valley and this was the third kit fox killed along Hwy. 50. The Gunnison valley will be the focus of the next phase of recovery work as it appears to have large expanses of habitat devoid of foxes. Photographic surveys will be conducted to cross-check the trapping surveys. Prey-base studies will be conducted as well. A working hypothesis for this area is that all aspects of kit fox habitat are present except other kit fox. Thus, when dispersing animals move into the area, they keep moving in search. of other kit fox. Relocation of kit fox families from the Uncompahgre valley to the Gunnison valley will be explored carefully. The adult male (#307) in Montrose East paired up with another collared adult female (#51) in spring 1997 and had a litter of pups within the Montrose landfill. On 18 May.1997 only the adult male could be located at the den and one dead pup was discarded at the mouth of the den. Subsequent searches resulted in the location of the dead adult female 2 km to the north. Although the carcass was quite desiccated, it appeared to be a probable case of canid predation. The adult male abandoned the den by 7 June 1997 and all pups were presumed dead. The Montrose East area, although the location of most trapped kit foxes in the past. 3 years, is an area of highly fragmented habitat with high levels of human activity, including wandering domestic pets. While reproduction has been documented, survival of young appears to be quite poor. Detailed mapping of this portion of the kit fox range will be conducted as spatial arrangements of irrigated land and other human activities may be pinching this group of foxes into a narrow finger of habitat. �Spotlight searches have documented the continued presence of some kit fox in the Peach Valley area; but none were trapped in 12 trap-nights. In the Colorado River valley, a citizen observation resulted in the confirmation of kit fox activity adjacent to irrigated land just east of the town of Fruita. Most of the earlier trapping effort in this area was conducted farther north of the irrigation canal as red fox (yulpes yulpes) and coyote (~ latrans) are believed to commonly hunt along the ditch banks and irrigated fields. This kit fox location was approximately 14 km from the nearest known kit fox den sites. No effort was made to capture these kit fox. APPENDIX A. Status and Distribution of the Kit Fox (vulpes macrotis) in western Colorado. Report to Mammals Research, Colorado Div. of Wildlife, pp. Prepared by J. P. Fitzgerald, Univ. Northern Colorado. 78 ��185 AppeodixA Contractor's Final Report Status and Distribution of the Kit Fox (Vulpes macrotis) in Western Colorado Report Submitted to the Colorado Division of Wildlife Report Submitted by: Dr. James P. Fitzgerald Biological Sciences University of Northern Colorado Period Covered: 1992-1996. �186 CONTENTS ACKN"OWLEDGMENTS ABSTRACT INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. METHODS AND MATERIALS Selection of Study Areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Unifying Characteristics of Trapping Areas Climatic Conditions Field Protocol Live Trapping Foxes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Other Search Methods Handling Captured Foxes , Habitat Descriptions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. RESULTS AND DISCUSSION . '. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Trapping Effort. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Foxes Observed But Not Captured Non-target Species Capture Baits Habitat Characteristics of Sites With Foxes Den Site Characteristics Condition of Captured Foxes Fox Populations Kit Foxes in the Lower Colorado Drainage Kit Foxes in the Lower Gunnison Drainage, Foxes North and West of Delta Peach Valley-Montrose East Fox Population ' Population Dynamics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Sex and Age Structure Reproductive Success and Pairings Survival and Mortality Movements of Peach Valley-Montrose East Foxes Use of Dens Movements and Home Range Estimates Disturbance Factors Spotlighting OUTLINE OF MANAGEMENT ASSUMPTIONS AND NEEDS General Assumptions Colorado Population Assumptions MANAGEMENT RECOMMENDATIONS RESEARCH RECOMMENDATIONS LITERATURE CITED . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. APPENDICES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Appendix A - Tables of trap nights by area Appendix B - Tables oflocations and dates of capture of all radio collared or ear tagged animals . Appendix C - Descriptions of Areas Trapped 188 189 190 192 192 193 193 194 194 195 196 196 197 197 199 200 200 201 203 206 207 208 210 210 211 211 212 213 215 218 220 222 225 225 226 226 226 227 228 228 231 231 233 237 �187 FIGURES 1. Distributional map of the kit fox (Yulpes macrotis) in North America and Colorado (modified from Fitzgerald et al.1994) 2. Kit fox in live trap, Peach Valley, Colorado 3. Location of male (open circle) and female (black circle) kit foxes, and one fox of unknown sex (half-shaded circle) trapped in western Colorado, 1992-96. Base map modified from Anderson et al. 1992 187 4. Habitats in which kit foxes were captured. Clockwise from upperleft: Rabbit Valley, Corcoran Point, Peach Valley, Montrose East 5. Movements of selected radio-collared female kit foxes from the Peach Valley-Montrose East complex, 1992-96 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6. Movements of selected radio-collared male kit foxes from the Peach Valley-Montrose East complex, 1992-96 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 7. Den locations in Peach Valley; Southern (upper) Peach Valley on left, northern (lower) Peach Valley on right. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 8. Location of dens used by foxes in Montrose East, 1994-95 Numbered dens were used by family groups May-August. Lettered dens were used by individuals located by Boyle August-December . . 9. Kit fox dens, clockwise from upper left: den in a deep wash, Peach Valley; den on rim of a slope, Peach Valley; den in eroded clay-loam soil, Montrose East; den by road in Peach Valley . . . . . . . .. Page 191 195 202 219 219 221 222 224 TABLES 1. Estimated harvest of kit fox based on trapper questionnaire returns to the Colorado Division of Wildlife, 1975-92 (Fitzgerald 1994). Kit fox were reported in 9 of the 17 years ... . . . . . . . . . . . .. 2. Elevation (m) and weather station normals for precipitation (em), temperature (C), and frost free days, averaged 1951-80, for areas surveyed for kit fox. Values are from the closest weather stations (Hadeen 1990) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 3. Sex, age class, and month and year of first capture of kit foxes, western Colorado, 1992-95. No new captures were made in 1996 4. Trapping success by sex and month of capture 1992-96, for all kit foxes captured or recaptured in western Colorado 5. Non-target species captured in kit fox traps, 1992-96, western Colorado 6. Summary of baits used to capture or recapture kit fox, 1992-96 7. Habitat characteristics of sites where kit fox were captured in western Colorado 8. Livestock grazing statistics for areas with kit fox, western Colorado. Specific allotment boundaries are shown on maps appended to the report. (Data from rangeland program summary updates from Montrose and Grand Junction offices USDI, BLM.) . . . . . . . . . . . . . . . . . . . . . . . . . .. 9. Aspect, distance from roads, slope, and number of entrances, for kit fox dens in Peach Valley. A and B were whelping dens (Link 1995) 10. Percent frequency of the commonest vegetation along four (NESW) transects at 10 kit fox dens in Peach Valley (Link, 1995) ,....................................... 11. Percent frequency of cover, average vegetation height,and slope at 12 kit fox dens in western Colorado 196 12. Weight (kg) and body measurements (em) of adult male and female kit foxes, western Colorado, compared to adult male and female swift foxes from eastern Colorado . . . . . . . . . . . . . . . . . . . . . .. 13. Number of kit foxes captured in the Peach Valley-Montrose East complex compared with data from Egoscue (1975) in western Utah, and Berry et al. (1987) in California 14. Capture dates, location, estimated age, and status of kit foxes trapped in the lower Colorado River Drainage, Mesa and Garfield Counties 1994-96 191 193 199 199 200 201 203 204 205 205 208 208 209 �188 15. Capture dates, location, estimated age, and status of kit foxes trapped north of the Gunnison River in Mesa and Delta Counties, 1992-96 16. Estimated age, and status of male kit foxes trapped in the Peach Valley-Montrose East complex, 1992-96 202 17. Estimated age, and status of female kit foxes trapped in the Peach Valley-Montrose East complex, 1992-1996 203 18. Radio-collared males and females at Peach Valley and Montrose East during the annual breeding season (January-May) 1992-96 19. Pairs or trios of kit foxes by dates observed, western Colorado 1992-96 20. Known mortality of 17 marked and 5 unmarked foxes from Mesa, Delta, and Montrose Counties, 1992-96 207 21. Estimated age at death, or age at last radio contact, for 42 kit foxes in the Colorado and Gunnison River Drainages, 1992-96 , . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 22. Number of foxes using dens and numbers of dens used, Peach Valley, August-December 1993. Use days compared to total days by fox reflects the number of days a den was shared 23. Number of foxes using dens and numbers of dens used, Montrose East, August-December, 1994.. 24. Estimated home range size for 9 kit foxes, Peach Valley and Montrose East complex, October- . January 1994-95. (Boyle 1995) 25. Estimated size of home range based on distances between denning sites for 8 kit foxes radiocollared for at least 7 months in the Peach Valley and Montrose East populations 26. Spotlighting time and distance in road km surveyed for kit fox by study area, 1992-93 (Link 1995) 211 215 216 217 220 221 223 223 225 ACKNOWLEDGMENTS The Colorado Division of Wildlife funded the project. T. Beck and B. Gill of Mammals Research handled project coordination and administration. The University of Northern Colorado Research Corporation handled paperwork for me and the Biology Department contributed workspace. Many CDOW people at the Grand Junction and Montrose Regional Offices helped on everything from where to park a trailer to showing field crews around their districts. Thanks to B. Pech, B. Divergie, P. Creeden, V. Graham, 1. Leslie, 1. Morris, G. Bock, D. Coven, M. McLain, 1. Wolfe, T. Antonelli, R. Arant, K. Stransky, M. Zgainer, and R. Olterman. Special thanks to 1. Olterman for his flights and coordinating things at Montrose. Thanks to 1. Ellenberger and 1. Gray for helping us at Grand Junction. Success of any field project depends on field crews. Thanks to M. Link, C. Parmeter, T. Verbeck, A. Anderson, 1. Eussen, L. Dent, M. Reddy, D. Watson, 1. Prather, S. Boyle, R. Hays, C. Hatch, S. Lechman, and R. Basagoitia .: I needto emphasize thanks to M. (Shelly) Link for working two hard, lonely field seasons by herself to make lots of contacts, talk to lots of ranchers, landowners, and other locals to finally find some kit foxes in western Colorado. Thanks also to all of those ranchers that helped by providing information, company, and even showers for field crews. Shelly, Jim Eussen and Lonnie Dent deserve added thanks for helping with annual and final reports. We also thank the trappers, taxidermists (especially D. McCauley), and ADC people that tried to steer us to areas they thought had foxes. Twice we received assistance of veterinarians at the Redlands Clinic and Nancy Limbach of the Pauline Schneegas Wildlife Foundation for treatment and rehabilitation of injured foxes. BLM biologists especially R. Lambeth were helpful all along on the project. Thanks to the NPS people at Colorado National Monument and U.S. Fish and Wildlife personnel at Browns Park Wildlife Refuge for use of facilities or providing trapping permits. It was gratifying to see several local residents take particular interest in the fox project. Special thanks to Jim (Melvin) Baker for helping us keep track of the Montrose East foxes and for his quality video of a family of foxes. �189 ABSTRACf From March 1992 to July 1996, 8,518 trap nights of effort were expended in attempting to capture kit foxes in 8 counties in western Colorado. The area trapped was estimated to be 533 mi2 (1,380 knr'). In 6,099 trap nights of effort in 4 counties (Garfield, Mesa, Delta, Montrose) 47 individual foxes were captured including 23 mares, 23 females, and one animal of undetermined sex. Kit foxes were trapped at 7 different locations in those counties. Capture of foxes in Delta and Montrose counties extends the eastern border of the reported range of the species in Colorado by 60 mi (96 Ion). No foxes were captured in 2,419 trap nights of effort (28% of trap effort) in Moffat, Rio Blanco, San Miguel and Montezuma Counties although kit or swift foxes were observed in Moffat County. All kit fox captures were in the lower Colorado and lower Gunnison River drainages at elevations below 6,000 feet (1,829 m) in cold desert shrublands. All sites where foxes were captured were grazed by domestic sheep or cattle and received considerable use by offroad recreational vehicles. Foxes were captured or recaptured in all months of the year with the highest numbers of captures in May, July, August, and September. Capture success averaged 1.25 foxes per 100 trap nights in areas known to be occupied by foxes. Kit fox dens were in a variety of situations ranging from the bottom of deep, steep-sided washes to under rock outcrops or on rims of ridges. Dens averaged 2 entrances. Bare ground at den sites usually exceeded 60%, grass cover averaged 20%. Average vegetative height near dens was 9" (23 em), Most foxes captured appeared in good condition. Two females suffered broken jaws from trap injury. One died in rehabilitation while the other was released to the study area after her recovery. Ear lengths of male and female kit foxes were significantly longer than ear length measurements from swift foxes captured in eastern Colorado. Tail lengths of male kit foxes were significantly longer than tail lengths of male swift foxes. Female kit fox tail lengths were not significantly longer than those for swift fox females. Fewer than 100 kit foxes are thought to exist in the Colorado and Gunnison River Drainages. There is no evidence populations are selfsustaining. Sex ratios for adults and pups approximated 1:1. Pups averaged 5 months (range 2-8) at their time of capture and 13 months (2-24) at their death or loss of radio contact. The oldest male was estimated to be 3.2 years at his time of death, the oldest female 4.3 years. The maximum average length of time radio-collared adult kit foxes were located was 10 months for males and 15 months for females in the Peach Valley-Montrose East complex. Kit foxes in western Colorado probably mate in mid-February based on emergence of pups in mid-May. Litter size averaged 2.9 (range 24) based on counts from 7 litters. Eighty-nine percent of marked pups were dead or missing by the end of their yearling year. No pups survived beyond 3 years of age. Twenty-two kit foxes, including 17 marked animals, were reported dead. Seven deaths were attributed to coyotes, 3 to motor vehicles, 2 to drowning, 2 to trapping, and 1 to trap injury. Cause of death was not determined for the other foxes. Only 4 of the 47 foxes captured are known to still be alive. Lack of reproductive aged males and failure of yearling females to breed may be limiting fox populations in western Colorado. One adult female moved over 25 mi (40 Ion) from her normal range, an adult male moved about 20 mi (32 Ion) from his natal den. Radiocollared foxes used an average of 3.6 dens (range 2-6). Home range size averaged 2.0 mi2 (5.2 knr') based on a combination of radio tracking and mapping of dens used by individual animals. Late fall and early winter single night movements were estimated to average 3.8 mi (6.1 Ion) but may have been influenced by the observers presence. �190 INTRODUCTION The kit fox (Yulpes macrotis) occupies arid portions of California, Nevada, Arizona, Utah, New Mexico, western Colorado, southern Oregon, and southern Idaho to central Mexico (Fig. 1) (McGrew 1979, Hall 1981). The kit fox is considered by some authors to be a sub-species of the swift fox (Yulpes yelox) (Dragoo et al. 1990). Their taxonomic relationship is unclear (Armstrong 1972, Rohwer and Kilgore 1973, Hall 1981, O'Farrell 1987, Scott-Brown et al. 1987, Samuel and Nelson 1982). The swift and kit fox both have a diploid chromosome number of2n=50 (Wayne et at. 1987). Kit fox have longer ears (average 80 nun vs. 64 nun) with ear bases close to the midline of the skull, a narrower rostrum, and larger (deeper) bullae than the swift fox (Thorton and Creel 1975, Dragoo et al.1987, Dragoo et al. 1990). The eyes of the kit fox are more slanted. Their tail length is about 62% of their total body length compared to 52% for the swift fox (Thorton & Creel 1975, O'Farrell 1987). The species hybridize in zones of contact (Rohwer & Kilgore 1973). Dragoo et at. (1990) provide electrophoretic data which suggest that they should be synonymized. Using similar techniques, Mercure et at. (1993) concluded the kit and swift fox could be considered different species. For management purposes in Colorado it makes sense to recognize the swift fox as the species on the plains east of the Rocky Mountains and the kit fox on the western slope (Fitzgerald et at. 1994). Comparative measurements of the two species in Colorado presented later in this report support this distinction. Literature on the kit fox has been reviewed by McGrew (1977) and O'Farrell (1987). Species biology has been studied in California (MorreI1972, O'Farrell 1983, Cypher and Scrivener 1992, White and Ralls, 1993), Arizona (Zoellick et at. 1989, Zoellick and Smith 1992) and Utah (Egoscue 1962, McGrew 1977, Smith 1978, Daneke and Sunquist 1984, O'Neal et at. 1986). Little literature exists on the species distribution in Colorado, northern New Mexico, northeastern Arizona, or eastern Utah. Findley et at. (1975) reported 4 specimens of kit fox from near Farmington and Broomfield, San Juan County, New Mexico about 20 miles (32 km) from the Colorado border. Halloran and Taber (1965) recorded a kit fox near Wide Ruins, on the Navajo Reservation in Arizona. Armstrong (1982) reported kit fox in Canyonlands National Park, Utah, 42 miles (67 km) from the Colorado border. McGrew (1977) reported kit foxes in Carbon and Emery Counties within 82 miles (132 km) of the Colorado border, and possibly in Grand County within 52 miles (83 km) of the border. There are 3 published records of kit fox in western Colorado. Miller (1964) observed a live icit fox in Colorado National Monument, 1.2 miles (2 km) west of Red Canyon View on Rim Rock Drive, Mesa County. Miller and McCoy (1965) reported 2 young kit foxes killed 12 miles (19 km) south of Mack, Mesa County. Egoscue (1964) reported 2 kit fox skulls from McElmo Canyon, Montezuma County, Colorado. There have been few reports of kit fox in the state since those publications. Black-footed ferret survey crews reported spotlighting 3 kit foxes in the Bitter Creek area, and 2 in the Mack-Lorna areas in Mesa County in 1983 (Grieb 1983). Colorado Division of Wildlife harvest statistics have reported kit fox to be taken on an irregular basis from several western counties (Table 1). However, no specimens were verified to species and harvest numbers are prone to considerable error (Fitzgerald 1994). In 1985, Colorado mammalogists raised concerns over lack of knowledge of kit fox while harvest was allowed (Crumpacker and Winternitz 1986). As a result of increasing interest among some mammal researchers within the CDOW and mounting public inquiry regarding the status of some harvested furbearers the Division in 1992 contracted with the University of Northern Colorado for a study to clarify distribution and status of the kit fox. Preliminary results from the study led the Colorado Wildlife Commission on 10 March 1994 to close seasons on kit fox and to impose trap restrictions in portions of the Gunnison River Drainage. On 22 March 1994 the Director of the Division of Wildlife designated the kit fox a species of special concern. In September 1994 the area with trapping restrictions was increased to include the lower Colorado River Drainage. �191 . ..... . '.' .' .. ....•... ,.: :.~ :'. " :~ o0, ._ Figure 1. Distributional map of the kit fox from Fitzgerald et aL 1994). (Yulpes •••••••••••.••. _,4_ macrotis) in North America, and Colorado (modified Table 1. Estimated harvest of kit fox based on trapper questionnaire returns tothe Colorado Division of Wildlife, 1975-92 (Fitzgerald 1994). Kit fox were reported in 9 of the 17 years. Year County 75 77 Delta 2 Garfield. Gunnison La Plata Mesa 3 1 Moffat Montezuma Montrose 2 Rio Blanco 1 Totals 7 2 * Nwnbers probably are a statistical error. 78 79 81 86 87 89· 2 3 5 91 6 2 13 5 33 2 3 192* 4 26* 10 5 5 6 10 41 13 198 4 33 41 Prior to 1975, separate statistics were not kept on this species. 2 This paper summarizes results of our investigations to determine the current range and ecological distribution of kit fox in western Colorado. �192 METHODS AND MATERIALS SELECTION OF STUDY AREAS Areas surveyed for kit fox were selected on the basis of: 1) Elevational, topographical and ecological similarity to habitats in Utah that had kit fox; 2) Records of kit fox from Colorado including trapper reports; 3) Interviews with area individuals; 4) Areas recommended by T. Beck, Mammals Research, Colorado Division of Wildlife. McGrew (1977) characterizing kit fox distribution across Utah found them at mean elevations ranging from 4,839-4,924 feet (1,475-1,501 m) with extreme elevations of2,401 and 6,102 feet (732 and 1,860 m). He reported foxes from shadscale (Atriplex canescens), mountain sagebrush (Artemisia tridentata), pinyonjuniper (pinus edulis-Juniperus sp.), and creosote bush (Larrea tridentata) communities. Egoscue (1962) in western Utah found foxes at elevations of 4,311-4,402 feet (1,314-1,342 m). Vegetation included shadscale, grey molly (Kochia yestita), seepweed (Suaeda torreyana), shadscale, rabbitbrush (Cluysothamnus sp.), greasewood, horsebrush (Tetrndymia canescens), shrubby buckwheat (Eriogonum sp.), and Indian Ricegrass (Oryzopsis hymenoides). In Washington County, Utah, Daneke and Sunquist (1984) found foxes in areas with halogeton (Halogeton glomeratus), rabbitbrush, black sagebrush (Artemisia nova), and winterfat (Krascheninnikovia lanat!!). In Millard County, in western Utah, O'Neal et al. (1986) reported kit foxes at elevations between 5,2495,372 feet (1,600-1,700 m) with vegetation including winterfat, budsage (Artemisia spinescens), black sagebrush, shadscale, saltbush, rabbitbrush, and Indian Ricegrass. Smith (1978) in Utah and Toole Counties, Utah found foxes where winterfat, cheat grass (Hromus tectorum), greasewood, sagebrush, shadscale, and pinyon-juniper dominated. Based on those reports we focused trapping efforts in cold desert shrub lands and on the margins of pinyonjuniper woodland. Virtually all valley floors in western Colorado are bordered by pinyonjuniper or juniper woodland where Pinus edulis intermingles with Juniperus scopulorum, J. monosperma, or L utahensis at elevations ranging from 5,500-8,500 feet (1,680-2,597 m). Stands are modified locally by intrusion of saltbush, rabbitbrush, or other shrubs (Costello 1954). In western Colorado cold desert shrub lands occur typically below 6,900 feet (2,100 m) on dry hillsides or valley floors associated with eroded shales or sandstones (Galatowitsch 1988). Three plant communities make up most of these shrub lands: greasewood, saltbush, and sagebrush or sage-grassland. Greasewood communities are found along washes usually in areas of alkaline soil (Armstrong 1972. Common plants in addition to greasewood are cheat grass, saltgrass (Distichlis sp.), summer cypress (Kochia sp.), peppergrass (l&pidium sp.), saltbush, and sagebrush. Saltbush communities are dominated by Atriplex sp., with associates including winterfat, sagebrush, rabbitbrush, ricegrasses, summer cypress, and buckwheat species. Herbaceous cover is often sparse. Soils are slightly less alkaline, and usually better drained than those of greasewood communities (Armstrong 1972, Galatowitsch 1988). Mat saltbush (A. corrugaW) often covers large areas with other Atriplex species, wild rye (Elymus saline), and ricegrass. In mat saltbush areas, cover rarely exceeds 10% (Galatowitsch 1988). Alkali sinks or marshes may be present in both greasewood and saltbush communities. Sagebrush communities support a variety of grasses or mixed forte understory. Species of Artemisia vary with sites and elevations in western Colorado (Galatowitsch 1988). Primarily on the basis of vegetative type and elevation, thirteen trapping areas were selected in eight counties, listed from North to South: 1. Browns Park, Moffat County. 2. Blue Mountain and Mellen Hill, Rio Blanco County. 3. Lower Colorado River Valley (Grand Valley) including the Colorado National Monument, Mesa County. �193 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. Gunnison River Valley, Mesa and Delta Counties. Peach Valley, Delta and Montrose Counties. Gateway along the Dolores River, Mesa County. Sinbad Valley, Montrose County. Paradox Valley, Montrose County. Big Gypsum Valley, San Miguel County. Disappointment Valley, San Miguel County. McIntyre Canyon, San Miguel County. McElmo Canyon, Montezuma County. Colorado River Valley, DeBeque-Parachute, Garfield County Descriptions of these sites are provided in Link (1995) and sununarized in the Appendix of this paper. Areas from which foxes were trapped are described in detail in the results section. Kit fox are believed to occupy portions of the Ute Mountain Indian Reservation in Montezuma County but permission to trap tribal lands was not obtained. UNIFYING CHARACTERISTICS OF TRAPPING AREAS All trapping areas met the habitat and elevational conditions reported in Utah kit fox studies. Most sites were in linear valleys draining to Utah, theoretically allowing corridors for immigration of kit foxes from any existing populations in eastern Utah. Most sites were isolated, except by linear riparian corridors, from each other and from the Utah border by wooded uplifts that would hinder kit fox movements into the areas. All areas included a mix of land uses and land ownership patterns. Public lands were primarily managed by the U.S. Bureau of Land Management, with lesser amounts of U.S. Forest Service, U.S. Park Service, and U.S. Fish and Wildlife Service holdings. Private lands ranged from urban areas and intensively irrigated crop lands to pastureland: Most trapping effort was confmed to public lands. CLIMATIC CONDITIONS Weather data were derived from records at the closest weather stations (Table 2). The average annual precipitation in western Colorado valleys ranges from 7.5-13" (19-33 em), Average annual temperature ranges from 44-52 F (6.7-1l.1 C) (Hadeen 1990). Extremes in temperature ranged from 100F (38 C) in Grand Junction to - 38F (-27 C) in Rangely. The frost free season ranges from 126-176 days (Hadeen 1990). Weather for Cortez 6,217 feet (1,895 m) is probably harsher than McElmo canyon 5,098 feet (1,554 m) because of its higher elevation. Table 2. Elevation (m), and weather station normals for precipitation (em), temperature (C), and frostfree days, averaged 1951-80, for areas surveyed for kit fox. Values are from the closest weather stations (Hadeen 1990. Extreme Average Frostfree Area Elev. Temp. Temp. Days Precip. Colo. Nat. Mmt. Cortez Delta Dinosaur Fruita Gateway Grand. Junct. Montrose Rangely 1763 1895 1504 1806 1366 1388 1468 1768 1613 3l.0 28.5 19.3 23.4 25.0 24.5 23.4 19.2 27.9 38.3/-16.1 35.0/-23.8 37.2/-23.3 37.2/-2l.1 37.2/-25.0 37.7/-16.7 38.3/-2l.7 33.9/-20.0 36.7/-27.2 9.7 9.3 9.2 8.2 9.1 11.1 10.7 8.3 7.7 176 146 152 133 145 158 149 153 153 �194 FIELD PROTOCOL Scientific collecting permits to trap foxes were obtained annually from the Colorado Division of Wildlife. A federal permit was obtained to trap in the Colorado National Monument and in Browns Park on U.S. F. & W. S. refuge lands. The University of Northern Colorado, Institutional Animal Care and Use Committee, approved protocols used during the study. In each trapping area the following procedures were used: Contact the local CDOW personnel; work with them to identify potential trapping sites; determine land ownership; identify and contact owners, or other residents; obtain permission to trespass or trap, and trap. Over 130 individuals were contacted by telephone or site visits. Those contacted included ranchers and other private land owners, trappers, taxidermists, veterinarians, local residents, and federal agency personnel with the Forest Service, Bureau of Land Management, Natural Resources Conservation Service, U.S. Fish and Wildlife Service, and National Park Service. A number of individuals contacted us with information on kit fox locations as a result of CDOW news releases about the project. LIVE TRAPPING FOXES Sampling was primarily by live-trapping. Because of the large area surveyed each area was censused once unless foxes were captured, fox sign observed, or foxes were reported to field workers. Areas with foxes were trapped over multiple years. Our approach, using live traps as the primary search method, varies from approaches by most other researchers who have relied on foot searches (O'Neal et al. 1986, Beedyet al. 1987, Orloff 1987), scent-station surveys (Beedy et al. 1987, O'Farrell 1987, Orloff 1987, Orloff et al. 1993) or spotlighting surveys (Beedy et al. 1987; Orloff 1987, O'Farrell 1987) to initially locate populations. We believed it was as manpower efficient and economical to place live traps as it was to construct and maintain scent-stations even using smoked plates (Orloff et al. 1993). Field crews ran 10-25 traps in a trapline daily. The major added expense being the traps ($35-40/each) and extra time to process captured animals. During the first half of the field work only 1 person was assigned to the project; in subsequent years 2-person crews were used to speed up setting traps and handling captured animals without having to use immobilizing agents. Commercial live traps measuring 11 x 12 x 32 in (28 x 30 x 82 em), with either 1" (2.5 em) or 1" x 2" (2.5 x 5.1 em) mesh were the standard trap (Fig. 2). O'Farrell (1987) and others reported kit foxes readily enter live traps of that dimension. Because of jaw injuries to two animals the smaller mesh traps were used exclusively after 1994. In a few selected areas, Woodstream soft-catch 1.5 leg-hold traps were used in dirt hole sets. When initially trapping an area, traps were placed subjectively in areas that appeared suitable habitat and at sites likely to attract foxes while hunting. Trap locations included bottoms or rims of washes, alongjeep trails, roads or livestock trails, near stock tanks, or adjacent to culverts (Berry et al. 1987, Orloff 1987). Traps were set for three consecutive nights and checked daily, beginning at sunrise. No literature recommends trap spacing for kit fox. Home ranges of 0.3-4.2 mi2 (0.8-11.0 knr') have been reported (O'Neal et al. 1986, White and Ralls 1993). Wood (1959) reported traps spaced 0.2 miles (0.32 km) apart were most effective for censusing gray foxes (Urocyon cinereoargenteus) and red foxes (Yulpes vulpes). We decided to place traps 0.2-1.0 mile (0.32 - 1.6 km) apart. Numbers of traps per trap line varied from 7 to 23. If no foxes were trapped after three nights of trapping effort, it was assumed there were few to no kit fox in the area. If a kit fox was captured or observed, or fox sign was present, a more intensive trapping effort was made using traps in lines or on a grid system no more than 0.6 miles (1 km) apart. That trapping lasted at least three days. We trapped all months of the year. Traps were checked beginning at sunrise every morning to minimize time in the trap. Workers in Canada and Colorado have trapped swift fox in winter and summer weather with no harm to animals as a result of spending the night under exposed conditions. Captured non-target animals were recorded and released if alive. Although black-tailed jackrabbits (Le.pus californicus) are known to be effective baits (Egoscue 1962, O'Neal et al. 1986) we decided at the onset of the study that we would not killiagomorphs or prairie dogs (Cynomys �195 Figure 2. Kit fox in live trap, Peach Valley, Colorado. leucurus) for bait. A field crew violated that condition in 1994 and was not rehired in 1995. Most live traps were baited with turkey chicks culled by Longmont Foods hatchery, Longmont, Colorado. Chicks had been used successfully in trapping swift fox in eastern Colorado, were easy to obtain, and kept well when frozen. As available, road-killed lagomorphs (Sylyilagus audubonii rimarily), prairie dogs, elk (Cervus elaphus), and deer (Odocoileus sp.), were also used. In 1995 we used a commercial lure (chicken and fish) from Erickson's On Target ADC, Inc. in combination with all other baits. We used dirt-hole sets at all leg-hold trap sites baited with small quantities of commercial lure. All trap locations and sites of fox capture were recorded and mapped by township, range, and section. Maps are attached to the report. OTHER SEARCRMETHODS No studies have been made on the relative effectiveness of trapping vs other survey techniques for either kit or swift fox but there are limitations with other methods. Live-trapping provided positive identification of the species and data on sex and condition of the animal. Field workers did not have to tty to identify smudged, . faint, windblown, or partial tracks of animals at scent station. A number of other medium sized mammals in western Colorado have front feet similar in shape to small foxes including cottontails.jack-rabbits, domestic cats and small domestic dogs. Over much of the study area grey foxes occupy similar habitat and unless tracks are exceptionally clear they can easily be confused with kit fox, although Orloff et al. (1993) discounted this as a problem when using clear tracks on aluminum plates. Spotlighting had limitations as a survey technique in western Colorado because of large expanses of shrub lands where vegetative height limits visual field. Trying to spotlight while driving on rough, unmaintained roads or dirt tracks is also dangerous especially if the driver is working one of the lights. Such roads were primacy access to most potential kit fox habitat in western Colorado. Spotlighting was done during the first 2 years in all areas trapped. The vehicle was driven at 5-14 mph, while spotlighting with a hand-held Brinkman Q-Beam 200,000 candlepower spotlight. Spotlighting periods lasted �196 from 1-4.5 hours. Some spotlighting was done during the early morning hours before daylight although most was conducted during the late evening and night time hours. Studies in California and Utah suggested lagomorphs were the main prey of kit fox and that dynamics of kit fox populations depended on numbers and distribution oflagomorphs (Egoscue 1962, O'Farrell 1987). During the first 2 years of the project Link (1995) recorded numbers oflagomorph sightings per km driven. However, Peach Valley, the only area where she found foxes, had the lowest lagomorph sightings. We discontinued most spotlighting efforts in subsequent years. Four other methods were tried briefly. In 1992, 1994, and 1995 scent stations were set in areas known to have kit fox present. Procedures followed Linhart and Knowlton (1975). In 1995, road killed deer carcasses were placed at scattered sites in portions of the Gunnison River valley and visited to look for fox scats or tracks. Two all-terrain 4-wheel units were used in 1995 to try to locate tracks following snows. Three, remotely triggered, 35 rom camera systems (Olympus, CAM- TRAKKER 9000XG) were used for 5 nights during the summer of 1992 to try to photograph pups and adults at whelping dens. Cameras were placed 3 meters from the point of focus. . Scats or food debris deposited at or near dens or voided by captured animals were collected. Specific location and dates were recorded at the time of collection, but for most scats the exact time of deposition was unknown. Scats obtained from traps were labeled by sex. Scats will eventually be analyzed by Jim Eussen as a M.A. thesis project at UNC. HANDLING CAPTURED FOXES When approaching a fox the handler noted condition including any visible injuries such as bloody jaws or other wounds. A handling bag of darkcolored heavy cloth with a drawstring closure was used to remove captured kit fox from the traps (O'Farrell 1987). Most foxes entered the bag voluntarily. A few had to be driven into the bag. Foxes were weighed in the bag using a hand-held spring scale. By working the head of the fox out of the bag they could be radio collared and ear tagged. All foxes large enough to collar were fltted with 52 g units (HLPM 2180 LD - Wildlife enough to collar were fitted with 52 g units (HLPM 2180 LDWildlife Materials, Inc.) with a 7" (18.5 em) flexible cable antennae. Radio collars had frequencies of 150151 Hz. Collars did not have mortality sensors. All foxes were tagged in both ears with style 893, Jiffy Wing Bands, National Band and Tag Company, Newport, Kentucky. Standard measurements of ear length, tail length, and total length were taken on most adults. During the project ketamine hydrochloride was carried in case particular foxes needed immobilization during tagging and collaring. At dosages of 15 mg/kg body weight it is a reliable immobilant for kit fox with recovery taking 30 minutes (Scott-Brown et al. 1987). Pentobarbital was also carried in case an animal with serious injury had to be euthanized. The first field worker (Link) was given pre-exposure rabies vaccination. After 1992 post-exposure rabies vaccination was given in the event a worker was bitten (1 case - Hays). HABIT AT DESCRIPTIONS Field workers categorized trap sites in terms of the predominant vegetation and estimated percent cover. Habitat type names followed those used by Costello (1954), Galatowitsch (1988), or Armstrong (1972). At dens used by kit foxes, 100' (30.5 m) long transects were run north, east, south, and west from the main entrance of the den. A modified Parker-loop system was used to determine cover at every 1 foot (30.5 em) interval along the transects. The categories used were: rock, bare ground, litter, fortes, shrubs, annual grass, or perennial grass. All plant species names follow Harrington (1954) or Weber and Wittman (1992). �197 RESULTS AND DISCUSSION TRAPPING EFFORT From March 1992 to July 1996, 8,518 trap nights of effort were expended including 8,414 trap nights using live traps, and 104 nights using leg-hold traps. The area trapped was estimated at 533 mF (1,380 km'). January-July 1996 trap effort was miniinal (50 trap nights) concentrating on the capture of specific foxes in the Peach Valley-Montrose East complex. In 6,099 trap nights of effort foxes were captured at 7 sites in the Colorado River-Gunnison River Drainages in 4 counties (Fig 3., Appendix A - Table 1). Twelve areas in 7 counties were trapped 2,419 trap nights (28% of trap effort) with no captures (Appendix A - Table 2). Figure 3. Locations of male (open circle) and female (black circle) kit foxes, and one fox of unknown sex (half-shaded circle) trapped in western Colorado, 1992-96. Base map is modified from Anderson et al. 1992. �198 We captured and marked 46 individuals; 22 males, 23 females, and 1 of unknown sex (Table 3). An adult, unmarked male is included in the Table although it may have been later recaptured and marked. Two recaptured foxes (ADF 21 and JM 184) were the only animals captured in 1996. At least 1 fox was first captured in every month except January, March, and October. Capture success measured in catch per 100 trap nights varied annually: 1992 - 0.5; 1993 - 0.7; 1994 -1.2; 1995 - 0.3; 19961.0. Most trapping effort in 1992 was conducted in areas where we did not capture foxes or find sign of their presence. Conversely in 1995 and 1996 almost all effort was centered in the Gunnison and Colorado River Drainages, much of it in areas where foxes were present yet our catch per unit effort was very low. That may reflect increasing wariness in previously captured foxes and lower numbers of naive foxes available for trapping. Success in Peach Valley, where 20 (43%) of the 46 marked foxes were captured tends to support this observation. In 2,163 trap nights of effort in Peach Valley, annual success rates in new fox captures per 100 trap nights of effort were: 1992 - 1.4; 1993 - 1.1; 1994 - 0.3; 1995 - 0.8. No effort was made to trap in Peach Valley in 1996. . When capture and recapture data are combined we averaged 1.25 animals per 100 trap nights in areas where fox were present in at least one year of the project. These figures are below those reported over 11 years by Cypher and Scrivener (1992) in California where their highest success was 20 foxes per 100 trap nights and their lowest was slightly less than 1 fox per 100 nights. Their average capture rate was around 5 animals per 100 trap nights. Foxes were captured or recaptured in every month (Table 4). The greatest numbers were trapped during the months of May, July, August, and September. We had good success in capturing foxes close to natal dens in May and early June when pups were above ground and adults seemed less wary then when pups were still below ground. The late summer captures reflect increasing opportunities for capture of juvenile animals or adults that may have trouble fmding abundant food during the hottest time of the year. O'Neal et al. (1986) discussed the problem of kit foxes becoming trap shy following their initial capture. We observed similar learning patterns, however this seldom seems to be considered when estimating population size (Harris et al. 1987). Of 40 radio-collared foxes (20 mare, 19 female, and 1 of unknown sex) we recaptured and changed radio-collars on 13 (6 females, 7 males) of them once and on 5 (4 females, 1 male) twice. We recaptured some individuals as many as 7 times but 22 (55%) of the animals were never recaptured. We made a total of 61 recaptures of marked animals with several animals captured on successive or alternate nights within the same trapping bout. We could not verify presence of kit foxes in Rio Blanco, or eastern Garfield Counties. We did not trap in Gunnison County from which 2 kit foxes were reported trapped in 1989. Some of our trapped foxes were about 20 miles from the Gunnison County line and a few may venture that far east but it is unlikely since higher elevations and dense oakbrush (Ouercus gambelii) or pinyonjuniper woodlands predominate along the border. We do not believe kit foxes exist in the isolated valleys that we trapped: Gateway, Sinbad, Big Gypsum, Paradox, Disappointment, McIntyre. That opinion is based on the small, fragmented numbers of foxes present in the lower Gunnison and Colorado River basins, our lack of trap success, and residents lack of knowledge of the species in those areas. The LaSal Mountains and extensive pinyon-juniper woodland would appear to be efficient barriers to kit fox range expansion into most of those valleys. The kit fox captures made in the lower Colorado basin in Rabbit Valley, and Prairie Canyon were expected based on Miller (1964), Miller and McCoy (1965), reports of kit foxes from CDOW personnel (DWM P. Creeden and black-footed ferret survey crews (Grieb 1983) or from harvest reports for Mesa County (Fitzgerald 1994). The kit foxes captured in the Peach Valley and Montrose East areas represent an eastern �199 Table 3. Sex, age class, and month and year of first capture of kit foxes, western Colorado, 1992-95. No new caEtures were made in 1996. Month Year Sex Age Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Totals 1992 Female Male 1993 Female Male 1994 Female Male 1995 Female Male AD AD Juv AD AD Juv AD Juv AD Juv Unk. AD Juv AD Juv Totals 1 1 2 1 1 1 1 I 4 2 2 0 1 1 1 1 9 1 1 1 3 3 1 3 3 1 2 1 1 15 2 3 2 1 1 1 9 5 0 2 1 1 3 2 1 1 4 1 9 6 5 6 1 2 2 1 3 47 Table 4. Trapping success by sex and month of capture 1992-96, for all kit foxes captured or recaptured in western Colorado. Jan Feb Mar Apr Mav Jun Jul Aug Sep Oct Nov Dec Total Male Capture Recap Female Capture Recap SexUnk. Totals 1 2 1 2 1 1 3 2 2 2 2 1 6 5 6 8 3 4 1 2 11 1 28 2 5 1 14 2 3 1 7 2 1 2 9 5 5 2 13 6 25 1 1 4 23 29 1 23 32 6 108 1 2 range extension of the kit fox in Colorado by about 60 miles (96 km). We expected to capture fox in the McElmo Canyon area based on the report of Egoscue (1964). However, much of the area along the Colorado-Utah border seems marginal habitat. The Utah side of the border does have extensive cold desert shrub lands suitable for kit fox and the foxes reported by Egoscue (1964) for Colorado may have come from Utah or represented animals straying into our state. FOXES OBSERVED BUT NOT CAPTURED A CDOW employee (R Basagoitia) observed two "pups" dead on Highway 50 on 3 Sept 1994, 1.2 miles (2 km) northwest of the Adobe Flats Reservoir by the junction of the Alkali Flats road. He also observed an adult kit fox crossing Highway 50 due south of Cheney Reservoir on 5 Sept 1994. Basagoitia photographed �200 2 collared kit foxes and one unmarked fox at two different guzzlers near Corcoran point, Mesa County and reported kit fox sign in Wells Gulch that same year. Dent reported a second fox with the animal he captured in Prairie Canyon, Garfield County in late summer of 1994. Two pups observed at a den at Corcoran Point in late summer 1994 were not captured because of their small size. A recreational trapper from Delta reported to Verbeck taking 2 kit foxes of unknown sexfrom "west of the Gunnison Gorge in fall of 1993. In May and June of 1996, two litters of kit fox pups were observed but not captured at Montrose East, 4 animals were in one litter, an unknown number in the other. A small population of kit or swift foxes inhabits extreme northern Moffat County. However, in 636 trap nights of effort from 1993-1995 we were unable to capture any individuals. M. Link spotlighted an animal close to the National Wildlife Refuge Boundary in the summer of 1993. BLM and U.S. Fish and Wildlife biologists Mike Albee and Bob Leachman observed a fox in the Vermillion Creek area in the summer of 1993 while working on black-footed ferret surveys. CDOW biologist Clait Braun saw a swift or kit fox in August of 1994 on Colorado Road 318 near West Boone Draw close to where it joins Vermillion Creek. Scat believed to be kit or swift fox was reported by S. Lechman and 1. Prather in 1995 from near a road killed antelope in Powder Wash. We speculate the Moffat County animals are swift foxes (Y. yelox) that have occupied the area from habitats in eastern Wyoming rather than kit foxes moving across more formidable habitat from Utah to occupy the area. Studies of "swift fox" distribution in Wyoming tend to support this thesis but Wyoming investigations are based on track surveys with individual animals not verified to species (Woolley et a1. 1995). NON-TARGET SPECIES CAPTURE During the trapping effort a number of non-target species were captured (Table 5). All individuals, except 7 rock squirrels (SpermQphilus yariegatus) that were dead in traps, were released. Many of the areas we trapped were in or close to colonies of rock squirrels and white-tailed prairie dogs. Table 5. Non-target species captured in kit fox traps, 1992-96, western Colorado. Number Name Rock squirrel Striped skunk Domestic cat White-tailed prairie dog Raccoon Badger Magpie Coyote (pup) SpermQphilus variegatus Mephitis mephitis Felis domesticus Cynomys leucurus Procyon rotor Taxidea taxus Pica pica Canis latrans 32 17 7 6 4 2 2 1 BAITS During the study there was some concern, especially among field crews working the summer of 1994, that turkey chicks were not effective baits and were contributing to the low capture rate for kit foxes. Opinions for that included a belief that there was rapid desiccation of the chicks with little scent produced and that they were novel to the area. Baits used by other researchers include: black-tailed jackrabbits (O'Neal et al. 1986, Zoellick et al. 1989, Zoellick and Smith 1992), cottontail rabbit (Sylvilagus audubonii) (Berry et al. 1987), canned mackerel (Zoellick et a1. 1987), smoked salmon (O'Neal et al. 1986), raw chicken (Briden et al. 1987), and live deer mice (peromyscus maniculatus) (O'Neal et a1.1986). �201 There is no clear indication that one bait works better than another based on our captures in western Colorado (Table 6). Many variables interact to influence trapping success including season of the year, trap placement in relation to home ranges of foxes, and placement of the trap, as well as the bait and or scents used. In eastern plains studies on swift fox we have captured over 200 animals using the turkey chick and scent lure method (Fitzgerald unpublished). Table 6. Summary of baits used to capture or recapture kit fox, 1992-96. Bait Cottontail Cottontailffurkey Prairie Dog/Turkey Prairie Dog/Cottontail Prairie Dog/Cottontailffurkey Prairie DoglBeef Prairie Dog Turkey Turkey/Lure Rock Squirrel Rock SquirrellBeef Scraps Pheasant Chicken/Cottontail Beef Scraps Not Recorded Total Foxes Caught % of Capture 18 7 7 3 9 17 6 6 3 8 2 17 6 16 2 1 1 1 9 5 100 2 18 7 17 2 1 1 1 10 5 108 We had 2 situations where we captured large numbers of foxes in short periods of time when using bait other than turkey, or turkey combined with other baits. In late May and early June 1994 we tried to trap as many animals as possible from family groups in Montrose East. We mixed turkey chicks with road-killed whitetailed prairie dogs and cottontail rabbits. That combination resulted in capture of 9 kit foxes in 8 days. Boyle (1995) captured or recaptured 16 kit foxes at that same location over a 4-day period in late August 1994 using baits ranging from beef scraps to prairie dog and rock squirrels. We believe that any meat bait used with an attractant scent is a workable combination for trapping kit foxes. Several field workers experimented with various commercial lures and scents placed at either scent stations (Link, Eussen, Dent) or at trap sites, including leg-hold sets (Hatch, Parmeter, Eussen, Lechman, Prather) with no captures or reports of fox visiting the sets. HABITAT CHARACTERISTICS OF SITES WITH FOXES All 7 areas in which foxes were captured were within the ColoradoGunnison Drainages and are similar in parameters (Table 7) to those outlined by McGrew (1977). Sites ranged in elevation from 4,800 feet ( 1,463 m) at Rabbit Valley to close to G,OOOfeet (1,829 m) at Cheney Reservoir. Most captures were at elevations of 5,000-5,600 feet (1525-1708 m). Precipitation for the areas variedfrom an estimated low of 7.5" (20 em) for Peach Valley to close to 10" (25 em) for Rabbit Valley and the southern edge of the Book Cliffs. . Vegetation ranged from sage-saltbush grasslands in Montrose East, to mat saltbush in much of Peach Valley to greasewood-saltbush stands in Prairie Canyon, and shrub-grassland intermingling with pinyon-juniper woodlands at Rabbit Valley (Figs. 4-5). Soils over most of the area represent weathered sandstone or shale parent materials often with high clay to clay-loam content. �2 Figure 4. Habitats in which kit foxes were captured. Clockwise from upper left: Rabbit Valley, Corcoran Point, Peach Valley, Montrose East. �203 All areas have livestock grazing as the predominant use (Table 8). The BLM Grand Junction office provided information on general range condition, the Montrose office indicated they had no recent data on range condition. The majority of kit fox captures were in Peach Valley grazing units which received high levels of stocking of sheep with most grazing during the winter and spring. This suggests the potential for coyote . control programs conflicting with kit fox recovery goals unless carefully conducted and monitored. The CDOW and Department of Agriculture will have to shape their respective management plans accordingly in this area. . Table 7. Characteristics of sites where kit fox were captured in western Colorado. Hlbltat Ellwml elaol Q2DlDlUO~ Blbblt ¥lIIIX mixed grass shrublands eml[11 QlOJ20 greasewood sagebrush FealUms ~ 121111All]lQtl saltbush adobe ell~b~lllel saltbush adobe MQOImII Ell! sage grassland >24 >15 >12 >12 >14 sandstone shale, canyons sandstone shale cliffs scattered surface rocks. mesa rolling hills flats adobe hills flats rolling hills swales All 7 areas have soils derived from Chipeta. Persayo. Fruita, Billings. or Badlands associtations except for Cheney Reservoir with Shavano-Lazear soils. Soils are loams ranging from stony and gravelly loams to fine sandy or fine silty clay loams. All soils are derived from shales and sedimentary rocks. EImIII2D al) emgpbal120 Suda~ sandstone . rock outcrops ~blolxBes. grassland saltbush >24 Veg!!lalKm H!!lgbl (10.) >18 SUrfa~ Corm;zmo eQlot mixed grass shrublands (10.) WilI!![ Land MaOag!!Dl!!DI QISlao~1Q 4800-5300 5040-5200 5360-5680 5680-6000 5100-5167 5412-5740 5412-5904 9-10 10-11 10-11 9-10 8-9 8-9 9-10 stock tanks none guzzlers reservoir ditches none ditches seeps ditches seeps grazing oil I gas grazing oil I gas. ORV grazing ORVrec. grazing ORVrec. grazing ORVtec. Grand Junction 35 miles Grand Junction II miles Delta 17 miles Delta 4 miles Delta 9 miles . grazing ORVrec. Grand Junction. 27 miles Montrose 3 miles All sites where we captured foxes receive off-road vehicle use by recreationists. The Grand Junction BLM office had no estimates for recreational visits. The Montrose office estimated 10,000 on-road visits annually in Peach Valley and 15,000 off-road visits. They did not estimate distribution of use by time of year or by location. The largest numbers of kit foxes were found within 10 miles of Delta or Montrose, and locations of family groups were known to several private citizens in the area. Two foxes used rock outcrops for denning sites including adult M194 in Rabbit Valley and a rehabilitated female (F 1) in Peach Valley. None of the areas being used by kit foxes in western Colorado offer habitat characters that appear to be uniquely different from hundreds of square miles of the Colorado-Gunnison River drainage. DEN SITE CHARACTERISTICS Link (1995) presented information about 26 dens used by kit foxes in Peach Valley. Site characteristics at 17 of them are presented in Table 9. The only data for the other 9 dens was location and number of entrances (5 had 2 entrances, 4 had 1 entrance). Most dens had a southerly aspect, all were within 1 mile (1.6 km) of' irrigation canals. Occupied dens were identified by the presence of fresh scat, animal remains, freshly excavated dirt and fresh tracks. Seven dens were within 164 feet (50 m) of dirt roads, 3 were less than 13 �204 Table 8. Livestock grazing statistics for areas with kit fox, western Colorado. Specific allotment boundaries are shown on maps appended to this report. (Data from rangeland program summary updates from Montrose and Grand Junction Offces, USDI, BLM). Allotment Grand Junction District. Rabbit Valley Rabbit Valley Baker Bitter Cr. Prairie Canyon Corcoran Point Mt Garfield Hunter Wash Cheney Res. Montrose District Wells Gulch & Alkali Flats Peach Valley Selig Canal UpperPV LowerPV Green Mountain Brush Point Adobe South Allot # AUM's Acres Livestock Use Season 6613 6612 6616 1,347 1,026 668 15,848 15,947 25,645 Cattle Sheep Cattle F,W, Sp F,W,Sp Sp,Su,F 6509 6504 6202 1,900 1,411 3,698 27,249 16,032 29,710 Cattle Cattle/Sheep Cattle F,W, Sp W,Sp Su,W,F 4016 4017 1,078 35,439 Sheep. W,Sp 5003 5007 5004 4017 50085030 700-800 1,000 800-1,200 580 1,153 95 1,935 3,727 2,420 21,170 18,205 1,611 Sheep Sheep Sheep Sheep Sheep Cattle Sp,F W Sp,F, W All Year Sp,W Sp,Su,F feet (4 m) from the road. Two dens were on the vertical sides of washes. One den had its main entrance at the bottom of a 16 foot (5 m) deep vertical wash (see Figure 9). This wash had surface water within 13 feet (4 m) of the den and the den would probably have flooded during severe rainstorms. Link (1995) measured ground cover along transects in 4 compass directions (N,E,S,W) at 10 dens in Peach Valley (Table 10). Point samples of cover were taken at 1 foot (30.4 em) intervals. Plants with highest percent frequency of occurrence were shadscale, clasping peppergrass Q.&pidiumperfoliatum), and skeleton 'mustard (Schoenocrambe linifolia). The most common grasses included galletagrass (Hilariajamesii), cheatgrass, and foxtail barley (Hordeum jubatum). Other plants common near den sites included horsebrush, Indian pipeweed (Eriogonum inflatum), poison hemlock (Conium maculatum), winterfat, Esteve's pincushion (Chaenactis steyioides), crane's bill (E. cicutarium), Russian thistle (Salsola iberica), yellow stonecrop (Amerosedum lanceolatum), ~ and Opuntia sp. Eleven of 13 dens had less than 50 percent vegetative cover around them. Bare ground surrounding dens averaged 69% (range 37-100%). Dens G, H, and L had no vegetation along transect lines. ,- Percent cover was estimated at 12 other dens since Link's work (Table 11). Den sites had large amounts of bare ground and little vegetative cover. Slope and aspect varied widely depending on location. Many dens in western Colorado differ from those described in the literature by often being on steep slopes or in gullies and rarely having more than 3 openings. Egoscue (1956, 1962) and O'Neal et al. (1986) in Utah, and Morrel (1972) in California, reported dens grouped on areas with well drained loose textured soils with little or no relief. Silty clay soils were favored because of easier digging (Egoscue 1956, Hoffmeister 1986). Egoscue (1962) and O'Farrell (1987) reported �205 Table 9. Aspect, distance from roads, slope, and number of entrances, for kit fox dens in Peach Valley. A and B were whelEing dens !Link 1995}. Den A B C D E F G H I J K L M N 0 P V Aspect West Southwest Southwest Southwest Southeast Southwest Southeast South Southwest South Southeast South East South Southeast Southeast West Distance from Road (m) Degree of Slope within 25 m Number of Entrances 15 40 10 35 40 40 45 45 30 20 20 20 30 4 2 100 2 800 200 20 50 400 50 50 300 100 3 2 2 2 1 3 1 1 1 1 1 2 2 4 3 3 2 Table 10. Percent frequency of the commonest vegetation along four (NESW) transects at 10 kit fox dens in Peach Vall~ {Link 1995}. Cover Stonecrop Cactus Den Peppergrass Skeleton Mustard Schadscale Grass * A B C D E F 21 25 22 40 25 0 40 67 78 14 I J K M * Primarily 31 16 26 37 25 0 60 33 22 71 0 20 0 18 0 0 0 0 0 0 19 14 5 3 0 0 0 0 0 14 15 5 36 0 0 0 0 0 0 0 0 0 0 0 0 100 0 0 0 0 Hilaria jamesii, Bromus tectorum. and Hordeum jubatwn. entrances to often be key-hole shaped and about 8-10" (20-25 em) high and less than 8" (20 em) wide. They attributed such shape as preventing easy entry by either badgers (Taxidea taxus) or coyotes. We saw few dens we would describe as key-hole shaped. Kit fox dens have been reported to have 2-25 entrances (O'Farrell 1987, O'Neal et al. 1986). Egoscue (1956) reported 3 as average, O'Neal et ala (1986) reported 6 natal dens had 3-10 entrances. Egoscue (1975) speculated that multi-entranced Whelping dens represented sites used by generations of kit foxes and that such areas were critical to successful breeding. Most dens in Colorado have 2 entrances. The low number of den entrances including natal dens (discussed later) may indicate that kit foxes in the Peach Valley and Montrose East areas of Colorado are relatively new colonizers. �206 Table 11. Percent frequency of cover, average vegetation height, and slope at 12 kit fox dens in western Colorado. % Freguency Location Bare Ground Grass Forb Shrub Litter Total % Veg Ht. (in.) Degree Slope Cheney Den A Corcoran Den A Montrose East 1 Montrose East 2 Montrose East 3 Peach Valley 1 Peach Valley 2 Peach Valley 3 Peach Valley 4 Peach Valley 5 Peach Valley 6 Peach Valley 7 Average 79 54 42 50 72 57 49 50 37 71 48 80 57 4 2 11 10 14 12 13 10 7 7 21 3 10 1 1 7 1 1 1 0 20 27 9 11 1 7 14 8 22 21 8 8 12 15 19 8 20 12 14 2 35 19 18 5 22 25 5 10 5 0 4 12 100 100 100 100 100 100 100 100 100 100 100 100 100 4 9 10 13 10 10 6 6 8 10 14 6 9 flat 40ENE 15SE flat 54SW 40NE 40NE 15NE 16NE 85S 85SW 26SW Ramp-shaped berms are often reported to be at the entrances to occupied dens. O'Farrell (1987) reported the average length of underground burrows as 7-16' (2-5 m) with maternity chambers 12-24" (30-60 em) wide and 12" (30 em) high. We did not excavate any dens. Height of vegetation along transects measured at 12 dens in western Colorado averaged 9" (23 em) (Table 11). Link (1995) estimated percent cover and height of vegetation in all major sampling areas in the study (Appendix C - summary in this report) and results are similar to the literature. About 80% of kit fox dens in western Utah were found in sparsely vegetated shadscale flats where the vegetation averaged 8-10" (20-25 em) in height (Egoscue 1956). In Pine Valley, Utah foxes were located in flat, shrub-grassland areas with a vegetation height between 12-35" (30-90 em) (Daneke and Sunquist 1984). O'Neal et al. (1986) reported average shrub height of from 2-15" (5.0-37.7 em) at den sites in Utah. Low vegetation is believed to be favored at den sites to reduce chance for ambush predation . . CONDITION OF CAPTURED FOXES With the exception of 2 adult females that broke lower jaws in traps and a female with a severe axillary laceration, all captured animals were in good to excellent condition based on weights, pelage condition and . absence of significant wounds or injuries. Both females with broken jaws were treated by local veterinarians who pinned their jaws. Both were housed with Ms Nancy Limbach of the Pauline Schneegas Wildlife Foundation, Silt, Colorado for rehabilitation. One female injured on 31 May 1992 in Peach Valley was released back to the wild in mid-September and found dead on 23 November 1992 about 8 miles (13 km) from her release site. Necropsy revealed bite marks on the thorax, right front shoulder, left foreleg, and neck attributed to a coyote attack. She had 5 placental scars, 2 in the right and 3 in the left uterine hom. The second female captured at Cheney Reservoir 13 July 1994 died in rehabilitation from unknown causes and was disposed of by the rehabilitator without providing us the carcass. Miscellaneous injuries included tom ears on 1 male and 1 female, tom out ear tags (1 ear on 2 females and 1 male; both ears on 2 females and 4 males), and superficial cuts and wounds. An adult female recaptured on 14 April 1995 favored her back right leg and would not put weight on it. She was located a number of times after that and seemed to be fine. An adult male had a right shoulder laceration of about 1.5" (4 em). An adult �207 male judged an older animal had a tom eyelid, and badly stained and worn teeth when captured 4 December 1995. Several other animals had trap injuries including a female with a scrape on her lower jaw, a male with a cut on his lower right lip, and a juvenile male with a broken upper left canine. One adult female recaptured 5 September 1994 had some hair loss and minor abrasions associated with her radio-collar. During the study period ketamine was used on 2 foxes, both captured by Verbeck in 1993 while working alone. He reported animals totally recovered from the effects of the drug about 45 minutes after injection. The 2 females with broken jaws were not given any treatment in the field. They were immediately taken to veterinary clinics for medical treatment. No reliable techniques exist for aging kit or swift foxes past 4-5 months of age (O'Farrell 1987, Scott-Brown et al. 1987). Carbyn and Klausz (1995) report aging swift foxes in the field and then having keepers at captive breeding facilities age the same animals. Keepers estimated all animals to be juveniles « 13 months) while field crews estimated 60% to be adults. Since 1983 the Canadian swift fox recovery program has released over 706 animals to the wild and bred hundreds of animals in captivity (Brechtel et al. 1994) yet have not arrived at a suitable aging technique for live animals. It is likely some animals we have shown as adults in the present study were juveniles. In estimating ages in later parts of this paper we have been conservative for animals over 5-6 months of age. However, we have assumed that any males and females known to have bred or paired during the breeding season were at least 22 months of age since breeding in yearlings is rare in the species (O'Farrell 1987). Standard measurements and weights of adult kit foxes from western Colorado are similar to those reported in the literature (O'Farrell 1987). Colorado foxes had little difference in mean weights between males and females (Table 12). Kit foxes of both sexes from western Colorado show Significantly greater earlength (P < .0001, t values = 13.9 females, 10.5 males) than swift foxes captured in eastern Colorado. Male kit foxes had significantly longer tails (P < .0002, t = 4.2) than male swift foxes. Female kit foxes had longer tails than female swift foxes but the differences were not significant (P < .075, t = 1.8). Comparative mean tail length to mean body length percentages were 60% for female and 64% for male kit foxes. In swift foxes the percentages were 48% for females and 47% for males. These are similar to the 62% and 52% reported in the literature (O'Farrell 1987). Iffoxes are captured in Moffat County external measurements should allow for identification to species. FOX POPULATIONS The low numbers of animals captured and recaptured, lapse time between captures, and distances between captured foxes make it impossible to estimate numbers in the lower Gunnison and Colorado River drainages. We speculate fewer than 100 animals. There is no evidence a self-sustaining population of kit foxes exists in the region. The Peach Valley/Montrose East fox complex is the only-group with documented reproduction for more than 2 years. Minimal known numbers of kit foxes in the Peach Valley/Montrose East group are compared with numbers from Egoscue (1975) in Utah and Berry et al. (1987) in California (Table 13). Egoscue (1975) used capture and eartagging to estimate numbers of animals on his study area. Data from Berry et al. (1987) are numbers of animals radio-collared in each year of the study so their total population was larger than reflected in the Table. Our minimum numbers of marked foxes in the Peach Valley-Montrose East complex in 1994 and 1995 were not much below numbers reported by those 2 studies. �208 Table 12. Weight (kg) and body measurements (em) of adult male and female kit foxes, western Colorado, compared to adult swift foxes from northeastern Colorado. N Mean Std. Dev. Minimum Maximum Kit Foxes Males Weight 2.5 11 0.3 2.2 3.1 Tot Length 78.5 8 6.9 71.0 89.0 Tail Length 8 30.6 1.8 28.0 33.0 Ear Length 8.2 4 0.5 8.0 9.0 Females Weight 1.7 14 2.2 0.3 2.6 Tot Length 12 79.0 6.1 69.0 88.0 Tail Length 12 29.4 2.0 26.0 33.0 Ear Length 10 8.0 0.5 7.5 9.0 Swift Foxes Males Weight Tot Length Tail Length Ear Length Females Weight Tot Length Tail Length Ear Length 49 28 31 27 2.5 84.4 27.1 6.7 0.3 3.9 2.0 0.3 2.0 77.0 23.0 5.0 3.0 92.0 32.0 7.0 43 24 31 25 2.4 84.3 27.2 6.0 0.3 3.4 2.0 0.2 1.9 78.0 24.0 5.4 2.9 90.0 33.0 6.3 Table 13. Number of kit foxes captured in the Peach Valley-Montrose East complex compared with data from Egoscue (1975) in western Utah, and Berry et al. (1987) in:California. California (74 mi2)** PV-ME (45 mi2)* Utah (40 mi2)* Year FoxlMi FoxlMi Year FoxlMi Year 1992 1993 1994 1995 6 7 21 21 0.13 0.15 0.46 0.46 1966 1967 1968 1969 22 21 21 17 0.55 0.52 0.52 0.42 1980 1981 1982 1983 1984 1985 1986 57 28 56 52 48 10 19 0.77 0.38 0.76 0.70 0.65 0.13 0.26 * Total count of all foxes known to be alive that year. ** Numbers of animals radio-collared by year. Kit Foxes in the Lower Colorado Drainage In four years of trapping at 922 trap sites with 1,930 trap nights of effort, we trapped 208 mf (539 knr') of approximately 348 mi? (901 km') of potential fox habitat in the lower Grand Valley. Potential habitat was considered to be areas with natural vegetation at less than 6,000 feet (1830 m) in elevation, and primarily on �209 public lands. Trapping covered from the Utah border eastward to Mount Garfield at the mouth of DeBeque Canyon and south into Colorado National Monument. Most sites were trapped more than once. (See map of trapping sites filed with this report). Only 10 individual foxes have been captured in the area (Table 14). Four,2 females, 1 male, and 10f unknown sex, were captured on the extreme western border of the Grand Valley in the Prairie Canyon and Rabbit Valley areas covering about 119 mf (300 km") In 156 trap nights of effort in and around Prairie Canyon one animal of unknown sex was captured and a second animal observed. In 303 trap nights of effort in Rabbit Valley, 2 females and 1 male were captured. Neither female was paired with the male. The male was about 4 miles (6.4 Ian) west of the closest female. Females were captured about 1 mile (1.6 Ian) apart. The minimum straight line distance between the Rabbit Valley and Prairie Canyon animals was 14-15 miles (22.5 Ian). All foxes from Rabbit Valley and Prairie Canyon were within 6 miles (9.6 Ian) of the Utah border, with the latter less than 0.5 mile (0.8 Ian) from the border. Table 14. Capture dates, location, estimated age, and status of kit foxes trapped in the lower Colorado River Drainagez Mesa and Garfield Counties 1994-96. Status as of Site and Estimated Ag~ (mQ) Last Date Located 7/1/96 Fox CaEture Date CaEture· Rabbit Valley 25 Unknown ADM 194 8/02/94 16 37 Unknown ADF 196 8/02/94 28 ADF198 16 Unknown 8/10/94 16 Prairie Canyon Unknown 16 UNK 153 8/21/94 16 Corcoran Point 25 Unknown ADM 130 16 7/17/94 Unknown JM 128 5 7/15/94 5 13 Unknown JM310 8/12/95 5 ADF 132 37 Unknown 7/19/94 28 ADF311 26 Dead 11/03/95 20 Unknown JF 126 7/15/94 5 12 Estimated Average Age 21 16 All Rabbit Valley and Prairie Canyon kit foxes captured were radiocollared; none were recaptured. Radio signals were not picked up from ADF 198 and UNK 153 after their capture dates. Male (ADM 194) and female (ADF 196) in Rabbit Valley was located periodically for over 8 months but remained alone throughout that time. The Rabbit Valley foxes may represent animals that periodically invade Colorado from habitat in Utah but for unknown reasons fail to establish a population. We captured foxes in Rabbit Valley only once during our 4 years of trapping effort. All animals were estimated to be over 1 year old at time of capture yet we did not capture any pups and neither female was in close proximity to the male. Female 196 did have heavily pigmented, hard nipples suggesting she had pups in the spring. The Rabbit Valley grazing allotment has a long history of livestock grazing by sheep and cattle. In recent years cattle have been grazed rather than sheep. We do not have data on the history of predator control for the area but it was likely intensive when sheep pastured the valley. Miller and McCoy (1965) report the surprise of a rancher when he killed 2 kit foxes near Mack in the early 1960's. The rancher was unfamiliar with kit foxes from the area and it may be that they have been uncommon in and near Rabbit Valley for decades. �210 We captured 1 kit fox and observed another in Prairie Canyon. Diggings, scats and tracks observed since that date suggest a small group offoxes has inhabited the area since at least August 1994. The location is not farfrom a reported sighting of2 kit fox pups made by CDOW, DWM Paul Creeden in 1992. However, years of black-footed ferret surveys over much of the western edge of Mesa county have not resulted in other sightings of kit foxes (Link 1995) and Link did not capture any or observe signs of their presence in 1992. Use of night photography systems developed by Beck on the bear project should be considered for this and the Rabbit Valley location. In 1994 a small, probably family group, of foxes was located near Corcoran Point north of Grand Junction. Link (1995) summarized trapping efforts in that area in 1992-1993. Although she and Verbeck put in over 500 trap nights of effort in that part of the Valley (Appendix A - this report) no animals were captured. However, since mid-1994 we have located dens, and captured 6 kit foxes (1 adult mare, 2 adult females, 2 male pups, 1 female pup) from that area. All were radio-collared (Table 14). The fate of the 4 animals radiocollared in 1994 are unknown. The radio signal from adult female (F311), radio-collared in early November 1995, was tracked to the porch of a land-owner east of the Grand Junction airport in late June 1996. The land-owner indicated that the animal had drowned in one of his cattle tanks. He removed the radio-collar but did not know who to contact regarding the animals death and had left the collar on his porch. The other Corcoran Point fox (JM310) was last located alive in April 1996. Fox scat is still present around the Corcoran Point dens and it may be that several animals are still present in that area. The radioed animals at Corcoran Point have foraged and denned primarily along the base of the Book Cliffs but hunt southward onto extensive saltbush flats. They are 23-28 miles (37-45 km) from Rabbit Valley or Prairie Canyon. They probably occupied the area by moving along the base of the Book Cliffs rather than across the open, partially irrigated valley floor southwest of their site. It is unlikely these foxes will persist over time unless supplemented by immigrants from other areas. In summary, in the lower Grand Valley, over 200 mi" (518 knr') of sagebrush and saltbush rangelands, clay barren areas, and shrubgrasslands appear to be suitable habitat for kit foxes but are either unoccupied or occupied at very low levels. Research in areas with relatively stable populations reported 1 fox per 2-4 mf (5-10 knr') in Utah (Egoscue 1975) or 3 adults/mi? (1.2 adults/krrr') in California (Morrell 1975). Kit fox numbers in the lower Colorado basin in Mesa and Garfield counties are far below that. The Grand Junction area is one of the fastest growing regions in Colorado. That growth is forming an urban corridor that may block movement of kit foxes from the Colorado to the lower Gunnison River drainage. More people will produce increased recreational pressure on public lands used by foxes. Kit Fox in the Lower Qwmison Drainage Foxes North and West of Delta We placed 400 traps over an estimated 114 mf of 190 mi2 (295 km2 of 492 knr') (60%) of potential kit fox habitat from the town of Whitewater in Mesa County to Delta in Delta County including a few areas south of the Gunnison River. In 2,040 trap nights of effort we captured 5 foxes, 4 of them for the first time, 1 a recapture. Foxes came from around Cheney Reservoir and near the Delta Airport (Table 15). The Cheney Reservoir foxes appear to represent a small, probably related, group similar to those at Corcoran Point and at Prairie Canyon. We spent 1,036 trap nights of effort around Cheney Reservoir. Three hundred trap nights of effort in 1992 resulted in no captures. Since June 1994 the remaining trap nights yielded 2 males and 1 female. The status of juvenile male (MI0l) captured in July 1994 is unknown. Adult male (M312) captured in December 1995 was recovered dead in Wells Gulch in June 1996. The uncollared adult female broke her jaw on the trap on her day of capture and died in rehabilitation. She may have been the mother of JM 10 1 captured at the same time. �211 In addition to the trapped animals, 2 road killed pups, neither collected, were reported (R Basagoitia) on Highway 50 in August 1994 just south of Cheney Reservoir. Basagoitia reported kit fox tracks and scat at a guzzler in Wells Gulch in early summer 1994. Project trapping crews have also found fox scats in Wells . Gulch, including fresh scat on 19 June 1996 the day the carcass ofM312 was recovered. It appears that a few foxes persist in the Cheney Reservoir-Wells Gulch area. This areas is about 35 miles (56km) from the Corcoran Point animals and 15-20 miles (24-32 km) from the Peach Valley-Montrose East complex. Two males have been captured in an area northeast of Delta close to the airport. A juvenile male (M17) radio-collared in late 1993 was never located again (Table 15). Male 33, captured as ajuvenile in Montrose East 27 August 1994, was recaptured near the airport late in 1995 at about the same location as M17's capture site. On a flight on 4 December 1995, Jim Olterman located 2 other foxes radio-collared in Peach Valley (M26 and F5) in Alkali Gulch west and north of Delta This is about 20 miles (32 km) (Figure 5) from their capture sites, 8 miles (13 km) from Wells Gulch, and 5 miles from the Delta Airport site. We have no evidence that a resident group of foxes inhabit the Airport or Alkali Gulch areas on a consistent basis but it appears that these areas are used by migrants. Table 15. Capture dates, location, estimated age, and status of kit foxes trapped north of the Gunnison River in Mesa and Delta Counties, 1992-96. Site and Estimated Age (mo) Status as of Fox 7/1/96 Capture Date Capture Last Date Located Cheney Reservoir MJ 101 7/11/94 7/11/94 ADF Unmarked ADM 312 12/4/95 Delta Airport 11/22/93 MJ 17 M33* 11/17/95 * Recaptured animal from Montrose East. 5 28 21 5 29 26 Unknown Dead Dead 8 20 8 20 Unknown Unknown Peach Valley-Montrose East Fox Population Population Dynamics South of Delta is the. Peach Valley-Montrose East complex an area comprising about 56 mf (145 km") of BLM lands below 6,500 feet (1981 m) in elevation. About 45 mi? (117 km') of the area was trapped. The north-east side of Peach Valley ill the vicinity of the Smith Mountain BLM grazing allotment and the area north and east of Flattop Mesa in the Lower Peach Valley BLM allotment were not extensively trapped. Both areas contain fox habitat but private landowners blocked on-road access to the area northeast of Flat Top Mesa. Access is possible across BLM lands using off-road vehicles. Summer crews in 1992-95 did not have ATV's for their use. Boyle (1995) using an ATV found kit fox tracks after a snowstorm. Eussen, using an ATV in July 1996 found fox scats and signs in the same area. It is likely that kit foxes use some of the area and it may be the source for most unmarked animals that have shown up on the study area each year. However, Eussen reported coyote sign also abundant in the area. In the complex we captured 33 kit fox (10 adult males, 6 male pups, 10 adult females, 7 female pups) in 2,677 trap nights of effort (1.25 foxes/100 trap nights). The foxes are distributed from south of the Gunnison River to south of Flat Top Mesa in Delta and Montrose counties, east and north of Highway 50 (Fig 3). A few individuals have made excursions north of the river, and one female moved south of the area �212 across Highway 50, east of Montrose. This is the only group of foxes found in 4 years of trapping large enough to be considered a population and the only area where foxes were taken in all years of the study. Our limited knowledge about kit fox biology and reproductive success in Colorado comes largely from this group. Sex and Age Structure Sex ratios approximate 1:1 for pups and adults. For marked animals, we estimated average age at capture as 16 months for 10 adult males and 20 months for 10 adult females (Tables 16, 17). However, age estimates for foxes over 5-6 months are inaccurate as discussed in the methods section. Average age at the time of last radio-contact or death was 26 months (2.2 years) for adult males (range 14-39 months) and 35 months (2.9 years) for adult females (range 22-51 months). Juvenile animals averaged 5 months of age at their time of capture for both sexes. Juvenile males averaged 13 months at the time of their last contact or death, juvenile females 8 months. We estimated the longest lived male to be 39 months (3.2 years) when he died and the longest lived female to be age 51 months (4.3 years) at her death. The longest life of any marked pup was 24 months. Egoscue (1975) estimated average annual age of his fox population at 1.8 to 2.2 years with one animal living to age 7. Table 16. Estimated age, and status of male kit foxes trapped in the PeachValley-Montrose East Complex, 1992-96. Site and Status as of E:!tim~ted Age (mQ) Fox Last Date Located 7/1/96 CaEture Date CaEture Adults Peach Valley ADM 0 5/14/92 14 14 Unknown ADM 4 6/25/92 15 15 Unknown ADM 2 18 9/28/92 18 Unknown ADM 8 2/23/93 11 39 Dead 13 ADM; 12 4/21/93 24 Dead ADM 13 34 9/20/93 18 Unknown 20 ADM 16 9/30/93 18 Unknown Montrose East 38 Alive ADM 184 6/3/94 15 17 37 Dead ADM 30 8/26/94 20 Dead ADM 48 8/27/94 17 26 Average Age 16 Juveniles Peach Valley JM 192 JM305 JM307 Montrose East JM24 JM26 JM33 Average Age 7/18/94 7/26/95 7/27/95 5 5 5 8 9 15 Dead Dead Alive 5/29/94 8/24/94 8/27/94 2+ 5 5 5 2+ 24 20 13 Unknown Dead Unknown �213 Reproductive Success and Pairings Kit foxes are monestrus. Most vixens breed at 22 months with only small numbers breeding at 10 months (Morrell 1972, O'Farrell 1987). Breeding females in the San Joaquin Valley return to whelping dens as early as September and October with pairing occurring in October or November (MorreI1972). In Utah, kit foxes copulate in early January with 3-5 pups born in late February or early March after an estimated 49-55 day gestation period (Egoscue 1956, 1962, O'Neal et al. 1986). Pups emerge from the den at about one month of age (Egoscue 1962). Table 17. Estimated age, and status offemale kit foxes trapped in the Peach-Valley-Montrose East Complex, 1992-96. Site and Status as of Estimmed A~ (mQ) Fox Last Date Located 7/1/96 Ca~ture Date Ca~ture . Peach Valley ADF 1 5/31192 26 31 Dead ADF5 51 Unknown 9/28/92 30 ADF 14 18 36 Dead 9/28/92 Dead ADF7 2/23/93 11 37 Unknown ADF 190 16 22 7/14/94 ADF 309 28 37 Alive 7/28/95 Montrose East . 5/29/94 37 Alive ADF21 14 ADF22 14 34 Unknown 5/29/94 Dead ADF23 5/29/94 14 37 ADF 180 Dead 5/31194 26 28 Average Age 20 35 Juveniles Peach Valley JF 186 JF 188 JF306 JF 308 Montrose East JF 176 JF 178 JF 182 Average Age 7/13/94 7/13/94 7/26/95 7/26/95 5 5 5 5 5 5 12 12 Unknown Unknown Dead Dead 5/30/94 5/30/94 5/30/94 2+ 2+ 2+ 4 10 2+ 8 8 Unknown Unknown Dead Using those data and our field observations we estimated breeding seasons for foxes in western Colorado. Link (1995) found pups above ground in Peach Valley on 13 May 1992 .. In 1994, pups were reported to us in Montrose East at about the same date (J. Baker to A. Anderson pers. commun.). We estimated some Montrose East pups observed in late May were 2-3 weeks out of the den while others looked like a later litter. If we assume pups in western Colorado emerge around 10 May they were born during the first 7-10 days in April and copulation would have occurred in mid February. Colorado litters are probably born almost I month later than those in southern Utah. �214 In mid-May and early June 1992, Link (1995) found 2 whelping dens in Peach Valley each with 2 pups. The pups and females at both dens were not captured, an adult male (ADM 1) was captured at 1 of the dens. Of 16 adult females captured over the study period, 2 (F7, F311) were taken at times of the year when field crews could not determine if they had whelped. Two females, F 14 and F 198 did not show darkened nipples or other evidence they had bred. In Montrose East in 1994,3 females, probably yearlings, (F21, F22, F23) were nulliparous when captured in May and early June. In 1995, F21 and F23 were recaptured in April and May and showed no sign of whelping. Both of them had pups in 1996. We necropsied only 1 adult female (Fl). She had 5 clearly visible uterine scars at the time of her death in November 1992. Eight females showed evidence oflactation: F198 from Rabbit Valley, F132 from Corcoran Point; F5, F190, and F309 (in 1995) from Peach Valley, and F180, F21, and F23 from Montrose East. Female 309 has not been recaptured in 1996 but was radio-tracked to a den which appeared to be occupied by pups based on small tracks and accumulated food debris. Female F5 had pups in 1994 but none in 1995. The unmarked female from Cheney Reservoir that died in rehabilitation may have had pups since a male pup was taken close to her site of capture at the sametime she was captured. However, her reproductive condition was not noted after taking her to the veterinarian. Since 1992 we know of 6 litters of pups in Peach Valley. Three litters had 2 pups, 1 litter had 3 pups, 1 litter had 4 pups. The number of pups F309 had this spring is unknown. Two litters from Montrose East had an estimated 7-8 pups using a single den in 1994. One litter (F23's) in Montrose East in 1996 had 4 pups, the size of F21's litter for 1996 is not known. Female 23 was killed by a coyote in early June and it is not known if her mate (M307), a young male, will be able to rear the pups. A litter at Corcoran Point in 1994 had at least 3 pups. A litter at that site in 1995 was of unknown size. Two litters of unknown size were present at Cheney Reservoir based on captured pups, or pups reported dead on the road near that point. Lack of reproductive aged males, and failure of yearling females to breed may limit fox populations in western Colorado (Table 18). Females 21, 22, and 23 in Montrose East (Table 18) were healthy animals yet none of them bred in 1994 as yearlings and two did not breed until about 33 months of age. Morrell (1972) and Egoscue (1975) did not believe females bred until 22 months of age. O'Farrell (1987) reported breeding in some females at 10 months of age. Only 1 male (M307) and no females of 10 male and 8 female pups (Tables 14, 15, 16, 17) we have radio-collared is believed to have bred at less than 12 months of age. Some pups occasionally remain with vixens (Egoscue 1956, 1962, Morrell 1972), and if female, may become helpers at a den (O'Neal et al. 1986). This appears to be the situation with the 3 non-reproductive females in Montrose East in 1994. We have videotapes taken by J. Baker of Montrose that clearly show several adult foxes freely mixing with pups at a whelping den. Unfortunately none of the animals were marked at the time the films were made and we cannot determine sexes of adult animals visiting with the pups. Of 10 male pups marked during the study only two (M310 at Corcoran Point, M307 at Montrose East) are still known to be alive. We do not believe male pups JM33 and JM26 ever mated although they both survived more than 20 months. We speculate that the large area in the Colorado and Gunnison river drainages and low fox numbers may force young males to emigrate to find unmated females. Their risk from coyote predation or other mortality would increase and some females may be left at reproductive age with no mates. However, O'Farrell (1984) reported most pups of both sexes stayed within 7 miles (11 km) of their natal dens and usually stayed within denning ranges of their parents. If so, that would seem to not favor outbreeding in the species. The instability in kit fox populations in western Colorado, and probable lack of mates is further demonstrated by lack of strong pair bonding (Table 19). None of the foxes we have observed have demonstrated long term fidelity with mates. Morrell (1972), and Egoscue (1956, 1962) reported some kit foxes paired for life while �215 Table 18.. Radio-collared males and females at Peach Valley and Montrose East during the annual breeding season pan-May) 1992-96. Year & 92 93 94 95 96 Area PV ME PV PV ME ME PV ME PV ME Female F5 nl x x P F6 x x x D F7 x D x F 190 P U F309 p p F 21 xn xn P F22 xn U F23 xn xn P F 180 D xl Male Ml x M8 x x U M 12 x D M13 x U M30 -xD M26 -xD M 184 x x x M307 x x = could have mated; U = location unknown; D = dead; xn = nonlactating; xl = lactating; P = pups observed; -x- = moved from Montrose to Peach Valley and assumed not to have bred. others changed mates. Both parents generally provide food and care for pups until they are 4-5 months old (Morrell 1972, O'Farrell 1987) with pups dispersing in late summer (Morrell 1975) or by October (Sheldon 1992). Two of the Montrose East juvenile males (JM26, JM33) did not disperse until mid-winter. Survival and Mortality We obtained mortality information on 22 foxes including 8 marked males and 9 marked females (Table 20). The average estimated minimum age at death for females was 27 months (range 12-37 months) and 24 months (range 8-39 months) for males. Seven of the recovered foxes were in advanced stages of decomposition making it impossible to determine their cause of death. Seven deaths were attributed to coyotes, 3 to motor vehicles, 2 to drowning, 2 to recreational trapping, and 1 while rehabilitating with a broken jaw. One death listed as drowning may represent an illegal human kill with the radio-collar (or carcass) thrown into a canal near Delta. We have been unable to recover the collar because of high water flows. Two foxes (M12 and an unmarked fox) located by Anderson were within 100 yards (33 m) of each other in Peach Valley and within 100 yards (33 m) of a well traveled road; they may have been shot. It seems unlikely both animals would have succumbed to coyote predation as they were within several hundred yards of dens. A CDOW, DWM (Covin) is presently investigating a report of a kit fox being shot in the Peach Valley-Montrose East area. We know of only 2 animals taken by recreational trapping during the study. Both were taken by the same trapper in fall of 1993 from "west of the Gunnison Gorge." Only 4 (9%) of the 47 foxes captured are known to be alive on 1 July 1996: AM 184, JM 307, AF 21, AF 309. Seventeen (36%) are known dead, the remainder (55%) cannot be located. Using estimated age at first capture statistics (Table 14-17), 86% of marked foxes died or disappeared before reaching 36 months of age (Table 21). Forty-five percent of dead or missing animals were classified as pups, with 89% of them �216 Table 19. Pairs or trios of kit foxes by dates observed, western Colorado 1992-96. Location MonthNr Pairing Observed Peach Valley May Jun Dec Jan Jun Jul Sep Sep Oct Oct Oct Apr Oct May Dec Dec Jan Apr Apr Oct May Jun Jul Jul Dec Unmarked Male, Female 2 pups Unmarked Male, no Female 2 pups. M12, F14 M12,F14 M13, Unmarked female M 192, F190 M8,F7 M12, F6 M8,F7 M16, F5 and F7 M12, M16 and FS M30, F5 M26, M30 and F23 Male, 2 Females unmarked 8-11 pups M(J)26, F21 M184, F(J)176 M184, F(J)176 M184,F21 M130, F132 M26, F23 M(J)307, FA23 FA21, Unknown male M130 and F132 Unmarked Female, no Male M26,FA5 Montrose East Corcoran Point Cheney Res Alkali Flats 92 92 93 94 94 94 94 94 94 94 94 95 95 94 94 94 94 95 95 95 96 96 94 94 95 Offspring 0 0 4 pups 2 pups 0 0 0 0 0 0 0 0 0 0 0 0 0 4 pups not counted 3 pups 1 pup 0 disappearing or dead by the end of their yearling year. Forty percent of the marked animals were estimated to be yearlings (13-24 months old) at capture with 62% of them disappearing or dying in that same year. Only 6 animals were classified as 2 year olds at capture. Five (83%) of them were dead or missing at the end of their third year of life. During the study we only trapped and marked one complete litter of pups (M305, M307, F306, F308) those of female 309's captured in late July 1995. Male 307 is the only pup still alive. Berry et al. (1987) constructed a survivorship curve for kit foxes on the Naval Reserve in California based on 144 animals marked as pups of the year. They estimated 74% mortality in pups during their first year, with 9% surviving past their second year. Berry et al. (1987) estimated 63% of their 211 collared foxes to be dead before age 3 and 86% dead by age 4. Juvenile mortality accounted for 50% of deaths. Our losses approximate those numbers if most missing animals are dead. Kit fox mortality has been attributed to predators, road kills, cyanide guns, other predator controls, construction, trappers, and hunters (Egoscue 1956, Smith 1978). Egoscue (1975), in Utah, reported adult mortality (or emigration) ranged from 10-58% annually with pup mortality close to 75% annually. Coyotes have been implicated as major mortality sources in several studies and relatively unimportant in others. Our losses from coyotes (7 of 17, 41 % ) are lower than most reports from the literature. O'Neal et al. (1986) reported 17% mortality from coyotes in 23 investigated kills in Utah. In California, O'Farrell ( 1984) reported that of 44 radiocollared San Joaquin kit foxes, 5096 were killed by predators, usually coyotes, 14% were killed by vehicles, and 36% were too badly decomposed to determine cause of death. At the Naval Petroleum Reserves in California, coyote predation was the primary cause of mortality for kit foxes (Cypher �217 and Scrivener 1992). On the Carrizo Plain of central California, 13 of 18 (72%) kit foxes were killed by coyotes (Martin 1993). Another California study (Disney and Spiegel 1992) found predation by bobcats (Felis rufus), coyotes, and domestic dogs (Canis domesticus) accounted for 64% of mortality. Table 20. Known mortality of 17 marked and 5 unmarked foxes, Mesa, Delta, and Montrose counties, 199296. Minimwn EstiMinimwn Fox & Date Carcass Sex Cause of Death Located CaJ2ture Date ABe at CaJ2ture mated ABe Death Fl 5/31192 F14 9128192 F7 2123193 F180 5/31194 UF 7/11194 F306 7126/95 F308 7126195 F23 5129194 F3II 11/3/95 Average Estimated Minimwn Age Males M8 2/23/93 M12 4121193 M192 7/18/94 M26 8/24/94 M30 8/26/94 M48 9/27/94 M305 7/27/95 M312 12/4195 Average Estimated Minimwn Age 11119/92 3/15/95 4/14/95 7/28/94 3/15/96 3/15/96 6/18/96 6119/96 26 18 II 26 27 5 5 14 20 31 36 37 28 29 12 12 37 25 27 Coyote Undetermined Coyote Undetermined In Rehab Coyote Coyote Coyote Drowned II 13 5 5 17 17 5 21 39 24 8 24 37 20 9 27 24 Undetermined Undetermined Undetermined Undetermined Drowned? Coyote Automobile Coyote at Death 6128195 3126/94 11118/94 3/15196 6/18/96 11122/94 12/12/95 6119/96 at Death Reported Mortality of Unmarked Animals Fall 93 U Fall 93 U 4/14/94 U U 9/3/94 9/3/94 U Trapper* Trapper* Undetermined** Road Kill*** Road Kill*** Unk Unk 13 6+ 6+ • Reported to Verbeck by a Delta Recreational Trapper . •• Found by A. Anderson in Peach Valley close to M12 . ••• Reported by R. Basagoitia. Table 21. Estimated age at death, or age at last radio contact, for 42 kit fox in the Colorado and Gunnison River Drainages, 1992-96. Estimated Age at Marking (years) 0 1 2 Totals Nwnber of Animals 19 17 6 42 0 D 4 4 ABe at Death/Loss of Radio Contact b~ Year of ABe 4 2 3 1 D L D L D L D L L II II I 2 I 8 3 9 1 3 4 5 2 1· 5 3 2 2 0 1 1 �218 The effects of predator control programs on kit foxes is unclear. :Kitfoxes (Robinson 1953) and swift foxes (Bunker 1940) have been poisoned and trapped during campaigns directed against coyotes and wolves. Robinson (1953) reported 1080 poison stations in 7 areas in Wyoming, Colorado and New Mexico did not reduce bobcats, skunks (Me.phitis me.phitis), badgers, and raccoons (Procyon rotor) over 1 decade. Only 1 kit fox was recorded trapped in 1940-41. In Utah, where poison and coyote-getters were used for several years, 36 kit foxes were taken compared to 13 coyotes (Robinson 1953). Robinson (1953) assumed control methods had a negative effect on sedentary kit fox populations. Kit foxes reportedly tolerate eating kangaroo rats killed with zinc phosphate (Schitoskey 1975). Mortality to kit foxes by diseases or parasites is unknown (O'Farrell 1987). O'Neal et al. (1986) speculated that external and internal parasites might influence mortality without directly causing death. Canine distemper and plague antibodies are reported for swift foxes in Wyoming but mortality from such diseases is unknown (Dr. Elizabeth Williams, Univ. Wyoming Veterinary Clinic - pers. commun.). Movements of Peach ValievlMontrose East Foxes Foxes monitored in Peach Valley during 1992-1993 made few shifts in home ranges (Link 1995). Most movements were less than 3 miles and involved shifts to new den sites. The farthest distance moved was about 8 miles (13 km). That animal, adult Fl, covered that distance over a 2-month period following her release from rehabilitation after recovering from a broken jaw. From 1994-96 several foxes made long distance movements from the Peach Valley and Montrose East sites (Figures 5, 6). Female F14, estimated to be 3 years old, was a stable resident in Peach Valley from 19921994. She was found dead on 15 March 1995, 12 miles (19 km) south of her home range. To get to that area she had to cross irrigated agricultural or urban-urbanizing lands and cross U.S. 50. The area in which her carcass was located was trapped on two occasions in 1994 and 1995 with no evidence of other foxes in the area. Female F5 (estimated to be about 4.2 years old) was a Peach Valley resident since 1992. On 4 December 1995, Olterman located her in Alkali gulch about 25 miles (40 km) northwest of her normal range. She has not been located since then. At her last known location she was close to JM26 and may have been traveling with him. Four females (F308, F309 collared in Peach Valley in summer 1995; and F21 and F23 both collared in Montrose East in summer 1994) were located on 14 February 1996 in Montrose East, 7 miles south of 308 and 309's original capture locations. On 9 April the carcass of F308 was located in the northern portion of Peach Valley about 9 miles north of the Montrose East site. On 18 June, female 309 was located alive at a den believed to have pups about halfway between Peach Valley and Montrose East. F21 is still alive in Montrose East while F23 was recovered dead from coyote injuries on 18 June 1996 in Montrose East where she was known to be rearing pups. Several males made considerable movements. Two (ADM30, JM26) were in Montrose East on 10 January 1995 and moved about 6 miles (10 km) to Peach Valley by 27 January (M30) and 13 April (M26). In October 1995 Olterman located M33 north of the Delta Airport near where it was subsequently caught in mid-November by field crews. M33 was captured as a pup on 27 August 1994 in Montrose East. The move to its November location covered over 20 miles (32 km) and required crossing the Gunnison River. Unfortunately, the field crew failed to replace M33's radio-collar in November and its batteries are probably dead. Male 26, reported with F5 in Alkali Gulch in December 1995, was back in Peach Valley on 14 February. His carcass was recovered on 9 April in Peach Valley close to a domestic sheep carcass and within a few miles of his February location. Male 184 captured in June 1994 in Montrose East was located on 4 December, 7 miles north in Peach Valley close to F306 and M305 by June 18 he had moved back to Montrose East. Male 305, a Peach Valley pup in 1995, was found dead on Highway 50 on 23 December close to Sweitzer lake about 7 miles northwest of his capture site. The reasons for substantial movements by foxes in Peach Valley and Montrose East are unknown but it has probably resulted in increased mortality as we have had 6 foxes (MJ305, M26, M30, FJ306, FJ308, FA23) �Del~ 65~ ;> 65~ ~ .. OSlO Figure 5. Movements of selected radio-collared female kit foxes from the Peach Valley-Montrose East complex, 1992-96. . miles ? o S JOmiles Figure 6. Movements of radio-collared male kit foxes from the Peach Valley-Montrose East complex, 1992-96 I\) ..•. (0 �220 die since early December 1995. We also lost F311 at Corcoran Point and M312 from Cheney Reservoir during this same time span; both recovered considerable distances from their capture sites. Instability and mortality translates into low reproductive success. O'Neal et al. (1986) reported an adult female to move 40 miles (64 km). O'Farrell (1987) reported pup dispersal distances averaged 7 miles (11 km) with most dying before establishing home ranges. Use of Dens Kit fox use of dens was monitored in Peach Valley in 1992-93 and in 1994-95 in Montrose East (Tables 22, 23, Figs. 7, 8). Twenty-six active dens were located on BLM land in Peach Valley. Den N was a whelping den pointed out to Link by a local resident. Most dens were located by following radio-collared animals. Only 2 of the observed dens were used in both 1992 and 1993. In mid-December 1993, a mated pair (M8, F18) centered their activity around den Z. During the fall and early winter of 1993, several foxes (F14, M12, M13, F5) showed considerable movement between den sites located 0.6-1.5 miles (1-2.5 km) apart. Those movements were interpreted as pre-breeding behaviors that would result in pair bonding. Dens R and V were used by 4 unpaired foxes. In May and June of 1994, at least 3 non-lactating females, 1 lactating female, and 1 adult male occupied a linear cluster of dens in Montrose East where pups were being reared (Fig. 8, dens 1-7). The dens were located along a wash and occupied an area of around 200 acres (80 ha). In late fall and early winter of 1994, Boyle (1995) tracked foxes in the Montrose East area. Nine foxes used 15 of21 dens in the valley from late August to late December 1994 (Table 23, Fig. 8 -lettered dens). They did not use any of the dens used in spring of 1994 for whelping and pup rearing. Foxes used often shared dens. Examples of kit fox dens are illustrated in Figure 9. Foxes in both areas used from 2-6 dens (average 3.6 dens/fox). Foxes at other sites were not monitored closely enough to establish den use patterns although the Corcoran Point foxes primarily used 4 dens during the summer and fall of 1994. The adult male in Rabbit Valley spent almost all of his time at a den located under a sandstone overhang. Egoscue (1956, 1962) reported 9 adult kit fox in a 12 mf (29 knr') area with as many as 8-10 dens located within 2-5 acres (1-2 ha). Dens were occupied exclusively by mated pairs or family groups (Egoscue 1962). In California, up to 39 dens occurred in a 320-480 acre (130-195 ha) area (MorreI1972) also reported to be their denning range (O'Farrell and Gilbertson 1986). The distribution of dens used in the Peach Valley and Montrose East sites in Colorado were not as closely clustered. Table 22. Number offoxes using dens and numbers of dens used, Peach Valley, August-December, 1993. Use days compared to total days by fox reflects the number of days a den was shared. n Fox AF5 AF 14 AF 18 AM8 AM 12 AM 13 AM 16 Total Use Day L M N I I 1 2 7 I 1 0 8 8 P 2 2 S 28 11 1 T V W 3 6 2 X Y 4 4 Z 1 18 20 2 2 1 2 4 4 R 1 1 14 55 49 2 1 1 2 2 10 4 1 24 13 1 3 3 1 1 1 1 1 38 21 # Dens Used 3 6 4 2 5 4 3 144 110 �221 Table 23. Number of foxes using dens and numbers of dens used, Montrose East, August-December, 1994. n Fox JF 176 AF23 AF22 AF21 AM 184 AM 48 AM 29 JM26 JM33 1 Total 1 C B A 0 E F G H 4 11 I I J K 2 2 L I 2 I 1 12 13 2 1 1 8 2 3 2 2 1 3 3 8 ?; .p =- 2 1 2 4 1 1 5 39 26 2 2 3 7 5 4 1 6 1 Total Dens Used 4 2 6 4 4 3 3 2 4 Avg. 3.6 103 T50N .z.o .E 5 6 1 3 1 1 e::: 0 N M 32 4 33 34 ;:!: v C\ c:: T15S . 7 8 7 •L 8 9 17 16 T51N 9 18 .M X y. • W •• 18 U· V 17 • Q G ••••• H F •S T· • D 16 • B .C 20 19 1~ • • N 21 A • F • 30 19 20 27 .• 1 ·29 28 ~ 0: 0\ T50N TSIN Figure 7. Den locations in Peach Valley. Southern (upper) Peach Valley on left, northern (lower) Peach Valley on right. . Den changes are thought to be associated with increased external parasite loads (e.g. fleas, ticks), shifts to richer prey areas (O'Neal et al. 1986), or as a mechanism to thwart coyote predation (Martin 1993). Egoscue (1956, 1962) reported whelping dens were often reused. He and O'Farrell (1987) note these are typically multi-entranced dens usually 1-2 miles (2-3 km) from each other. Egoscue (1975) believed carrying capacity of habitats for kit fox in Utah was primarily related to denning requirements, especially areas for whelping, more than prey availability. Our data on whelping dens is limited (6 located dens). Three had 3 entrances, 1 had four, and 2 had 2 entrances. �222 5 6 2 A • ~ I J •• 12 11 K • o • L • ... ~-_ -; . .. · 14 Figure 8: Location of dens used by foxes in Montrose East, 1994-95. Nwnbered dens were used by family groups May-August. Lettered dens were used by individuals located by Boyle August-December. The expanse of area and similarity of soils and habitat in the Gunnison and Colorado river drainages in west central Colorado suggests kit foxes are not lacking in suitable denning sites for whelping or escape burrows. Food resources may be more patchy but prairie dog colonies are widespread across the area and their burrows are known to provide refugia for many prey species including cottontails and mice (Clark et al. 1982). Movements and Home Range Estimates Boyle (1995) estimated home ranges of7 radio-collared kit foxes in the Montrose East drainage and 2 foxes in Peach Valley (Table 24). Animals at Montrose East averaged 1.4 mf (3.6 km') and in Peach Valley 2.9 mf (7.57.7 km'), We estimated home range averaged 2.3 mi? (6.0 knr') based on distances between denning sites for 8 kit fox in Peach Valley and Montrose East (Table 25). These are considerably larger than the estimates ofless than 1 square mile (2.6 knr') made by Morrell (1972). Home range sizes of 16 kit fox near �223 Table 24. Estimated home range size for 9 kit foxes, Peach Valley and Montrose East populations, OctoberJanuary 1994-95 (Boyle 1995). Animal Km2 # of locations Animal Mi2 Montrose East 56 JF 176 5.9 2.3 ADF22 4.7 52 1.8 ADF21 53 0.6 1.5 ADM 184 4.7 50 1.8 36 ADM 29 1.5 3.9 JM26 0.9 2.3 36 JM33 1.0 2.5 38 Peach Valley ADF7 3.0 7.7 22 ADF 190 2.8 7.5 18 Table 25. Estimated size of home range based on distances between denning sites for 8 kit foxes radiocollared for at least 7 months in the Peach Valley and Montrose East populations; Estimated Range Me Km2 Fox Months radioed # Dens Peach Valley 33 5 2.7 7 ADF5 25 8 ADF7 6 3.0 36 ADF14 7 3.0 8 ADM 8 28 7 3.0 8 24 6 7 ADM 12 2.7 Montrose East ADF21 35 7 1.2 3 JF 176 7 5 1.9 5 ADM 184 12 5 1.5 4 Pine Valley, Utah, varied seasonally and by sex (Daneke and Sunquist 1984). Males had.slightly larger summer home ranges 0.7 mf (1.9 km2) than females 0.5 me (1.4 km2). Males had larger winter ranges, 1.6 mi? (4.2 knr') than females (0.7 mi'). In the Desert Experimental Range, Utah, adult males had a home range size of 1.3 mi? (3.4 km") compared to 1.2 mf (3.0 knr') for adult females (O'Neal et al. 1986). In Arizona, home ranges of female kit foxes, 3.8 mi", (9.8 krrr') averaged 20% smaller than those of males, 4.7 mf (12.3 knr') (Zoellick end Smith 1992). Home ranges of kit fox in the San Joaquin Valley, California, averaged 4.5 mi? (11.6 knr') (White and Ralls 1993). Boyle (1995) estimated total minimum distance traveled and straight line distances traveled during single night movements of 5 individual foxes in the Montrose East group during October-December 1994. He estimated total minimum distance to average 3.8 miles (6.2 km) and maximum straight line distances to average 1.2 miles (2 km). However, movements of the observer might have caused foxes to increase their movements in response to his presence. A number of the field crew members (Link, Beck, Dent, Parmeter, Verbeck) all reported foxes moving away from them when they tried to track animals at night. Zoellick et al. (1989) reported much longer nightly movements, 7 miles (12 km) for females and almost 9 miles (14 km) for males in western Arizona. �~.~., .. (•...~ .•... ~ -, t , ;~1~,'~:-~ ~.... ... ... . ~""'" i,; -: • ; ~ ..' f ~ ' ". . • - .. . ,. . ' 1. " ',,.. ·.:i ~~. 1':"'- t.:',./. - J ..•.",f,,~'1'~:~•.~. ,.. ~ J..J' ,~.~"i...J9iCiI 'II'': ,". , *S~j.>.... E . ~:;:.J( .~~-'!. =~;.;..~~..•• • .•..~::..:~•• •.. .' ,. =,,"' ......0..: r. ~ . ~•••••. ;'" .•• . -- _•• - &'" • • 'T";'~~ " •• " ••.J;..•. _ •• .,_.; ,t,':'';'';'~~~ ~:::-;""~.~" ~ •••. 'c-..... .' ..' \ ii'S'-'..·. 1.\.i:..# .~~.. ".'ir.~" _~- .•.. ':If:!'"' ~ -' --- ' .. ";o'll'( 4. -.-~~ .•...••.•• ~,:':.. ;~'..':.':",..-. ~!:.!I{; • ~ . .,~..,.,. '._ • .r .:., ••• -.' • _._ ••••••• 0 •• . - _._ ;: ,:t.," .•••• .. ' .. '.',',. ,.,...,.. ~ - ---- ...~~ 1!i: ":$1~;.;"",:' <: .t ,>"";-: :,. .•~ •.•••••.~ ;ti.".... --:--- ~';~4 .~~.:. .~ .• , -.~\ - ~i.J.·· _,-: .; •• ~~..J"- ,:;. •.•..• " ,-:,">'~ '~ .... ...~'.7':~> - ...,..,...... ....,. '~~ -' ..- .:.;ff"::. . . _.:.-:.. ,;tIi!lf.'A. • ,.,Jr,!,-" ..•..<·~r.···' ..' •.• ~... :;~~ .",.,... - •••.••. .. :'.:'" ~~ ._.r .. .~,,'" .•...•.. ; ••~.'" '.. PO! ...u.t"f''':'''.:..J''' ~ ••.!:.-..:r,o;.~ <'" e :•r: .....•• -. . .••...••. :." .'. . ,.-:o! • . ..• • ">: ,: .•••.~ ,'. " .. _ ~~ --'ftm.. _ #.l9.~..::.-:.::..;~: .. ~t~~~:·:· Figure 9. Kit fox dens, clockwise from upper left: den in a deep wash, Peach Valley; den on rim ofa slope, Peach Valley; den in eroded clay-loam soil, Montrose East; den by road in Peach Valley. . �225 Disturbance Factors Research is needed on the extent to which human uses disturb kit foxes in western Colorado. Link (1995) observed that passing vehicles did not alter the foxes' behavior at 2 natal dens unless people stopped to watch them. The foxes would then casually retreat to their dens. Other crew members observed similar behaviors with the Montrose East foxes, a group that was exposed to vehicular traffic and to several private citizens who routinely filmed them at their natal dens. Link (1995) believed that foxes spent more time underground for longer periods on the weekends when peaks of human disturbance occurred in Peach Valley. Recreational use at all the kit fox areas in the Colorado and Gunnison drainages would probably be most intensive on weekends and during the summer. We believe that the relatively concentrated, constant monitoring done on foxes in the Montrose East group during Fall and Winter of 1994-95 was a contributing factor for animals leaving that area for much of 1995. Until proven otherwise we recommend ground surveillance of animals be done on an infrequent basis from as far a distance as possible to minimize disturbance to the animals. Spotlighting During the first 2 field seasons a total of 68 hours were spent spotlighting in the areas trapped by Link (1995) (Table 26). Since then field crews have dedicated most time to trapping with spotlighting conducted only in a few localized areas (Brown's Park, Rabbit Valley, Wells Gulch, Peach Valley, Montrose East) usually in association with radio-tracking of collared animals. Spotlighting effort was reduced in part because of restrictions imposed on the project by UNC personnel officials who found us in violation of fair labor standards in terms of crew work hours. Table 26. Spotlighting time and distance in road km surveyed for kit fox by study area, 1992-1993. Only 1 kit fox was observed, it was in Peach Valley (Link 1995). Distance (km) % Total Area Hours % Total 8 Browns Park 5.6 8 75 35 326 Grand Junction 25.3 38 129 14 Whitewater to Delta 9.0 13 127 14 Peach Valley 8.6 13 4 35 Gateway area 4.0 6 3 Sinbad Valley 2.3 3 24 8 74 Paradox Valley 4.0 6 7 67 Big Gypsum Valley 4.1 6 67 7 Disappointment Valley 4.7 7 924 Totals 67.6 Link (1995) located only 1 kit fox by spotlighting in Peach Valley in 8.6 hours of effort. Foxes at Corcoran Point, Peach Valley, and Montrose East when first located by radio-tracking could be found with spotlights but rapidly moved out of range of the lights (Verbeck, Dent, Eussen, Reddy, Watson, Fitzgerald field observations). One kit or swift fox was spotlighted in Browns Park by Link (1995). An additional 14 hours of spotlighting by Verbeck (1993); Dent (1994); Lechman and Prather (1995) failed to locate any animals. �226 OUTLINE OF MANAGEMENT ASSUMPTIONS AND NEEDS GENERAL ASSUMPTIONS (LITERATURE DERIVED) 1. Kit foxes need a large amount of relatively homogeneous habitat in which they can establish localized populations. Their occupancy of habitats is clustered. 2. Adequate sites for denning are probably more important than distribution of food resources 3. Local populations are prone to pronounced population cycles that seem to tie to localized weather patterns that influence prey species dynamics. 4. The species can tolerate at least moderate disturbance from humans in their habitat. 5. The species does not seem to disperse and colonize well. COLORADO POPULATION ASSUMPTIONS (FIELD AND LITERATURE DERIVED) 1. Foxes inhabiting the Gunnison and Colorado River Drainages probably recruit from populations in eastern Utah. . 2. Lack of sufficient males of breeding age may be a limiting factor to populations. 3. Lack of a significant number of females breeding at earlier than 22 months is contributing to limited population size. 3. Most litters are small compared to other studies. 4. Existing foxes are widely distributed in small family groups that hinder successful immigration or emigration and reproductive success. 5. General biology of Colorado kit foxes does not appear to be significantly different from other studied populations. 6. There appears to be habitat suitable for kit fox population increase in the Colorado-Gunnison River Drainage. 7. Urbanization over the next decade may shut off some corridors for movements of kit foxes from the Colorado to the Gunnison drainage. 8. With significant management and research inputs it may be possible to build a more viable population of kit foxes in westcentral Colorado. 9. Threats to kit foxes in western Colorado are a combination of the following factors: a. Urban growth that may shut off corridors for movements. b. Increasing risk of human disturbance to kit fox denning areas by recreational enthusiasts. c. Unknown effects oflivestock grazing and predator and/or rodent control activities on populations of kit foxes and prey species. d. Conflicting demands for uses on public domain lands to which kit foxes are largely confmed. Including growing conflicts between local control vs. federal control and consumptive uses vs. other uses of public lands. e. Any set of environmental conditions (non-human caused) that will place increased stress on the surviving kit foxes including: 1. Drought, changes in grazing pressure, diseases to prey species! or other factors that may decrease prey. 2. Increased pressure on populations by increased predation or disease - higher coyote mortality, rabies, distemper, etc. f. Political-social climate inside and outside the CDOW with respect to "value judgments" on importance and worth of management of predators like the kit fox - a species of low utilitarian value. 10. A minimal population target for the entire Colorado-Gunnison drainage should be 8 sub-populations of 15-25 individuals each with a reasonable chance for interchange among groups. 11. The only kit foxes in the state with any significant opportunity for management is the ColoradoGunnison River Drainage foxes. Foxes on the Ute Mountain Reservation are not subject to CDOW management, and foxes if present in McElmo Canyon or Browns Park are likely immigrants from Wyoming or Utah and may not exist over a large enough area to warrant management. �227 MANAGEMENT I: Secure Maximum Enforceable Statum RECOMMENDATIONS Protection Goal: Provide For maximum Enforceable Statutory Protection of the Species - In view of recent review and controversy over Wildlife Commission actions on Furbearers and the Governor's recent signing of SB167 - it is imperative that CDOW seek at a minimum "Threatened" or "At Risk" status for the kit fox. Objective: To maximize CDOW and Colorado Wildlife Commission authority over management and protection of the species. Strategy: Work within CDOW, Colorado Wildlife Commission and the Colorado Department of Agriculture framework to achieve a clear understanding of the status of the species and its need for protection and management. Including protection from any adverse methods for control of predators contemplated by the Colorado Department of Agriculture. II. Write a formal program for recoYety and maintenance of kit fox in western Colorado. Objective: To clearly evaluate and detail mechanisms for enhancing existing kit fox populations and to reach a reasonable timeframe for accomplishment of that task economically and biologically. Strategy: Work during the 1996-97 fiscal year to complete the work effort using as appropriate talents within and outside the CDOW. III. Detail the Recovety and Maintenance Program Objectives 1. Establish a minimum of 8 subpopulations, providing groups of animals with suffcient protection from human and environmental perturbation. 2. Take steps to evaluate, and manage the habitats those sub-groups live in to maximize quality for the animals and minimize disturbance factors. 3. Work within the structure of existing and potential sociall economic/ administrative groups to achieve management goals. 4. Establish a monitoring program to evaluate results of the enhancement/maintenance effort. 5. Conduct such research as needed to be able to understand demographics and necessary behavioral ecology of the species for its appropriate management. Strategies: Annual Population Level Target: a. Seek to secure 200 square miles (518 knr') in the Colorado River Drainage and 200 square miles (518 km") in the Gunnison Drainage as critical kit fox habitats with the species recognized and managed for in Land Use agency plans and in plans of wildlife damage control agencies. 1. Work out land use planning and wildlife management plans with the BLM. Including as needed local restrictions on recreational or grazing uses of important fox areas. 2. Work out predator and rodent control programs with the Colorado Department of Agriculture and USDA-APHIS. �228 RESEARCH RECOMMENDATIONS 1. More thoroughly investigate home range and population biology of the existing foxes in Peach ValleyMontrose East including a detailed assessment of how human disturbance disrupts the animals. 2. Determine some method for cooperation with the State of Utah to determine viability of populations in adjacent Grand County, Utah. ' 3. Investigate possibility of release of additional animals to populations in Peach Valley and Montrose East to see if reproductive success can be increased. 4. Develop (we recommend using the swift fox as a model) methods for better determining age of individuals and adequate techniques for census. The use of the camera system being used by Beck on black bears needs to be designed for use on smaller carnivores including kit arid swift foxes. We mayor may not accomplish that this coming year with the swift fox work on the Pawnee Grassland. LITERATURE QTED Armstrong, D. M. 1972. Distribution of mammals in Colorado. Monograph Museum of Natural History. University of Kansas 3: 1-415. __ . 1982. Mammals of the canyon country a handbook of mammals of Canyonlands National Park and vicinity, Utah. Canyonlands Natural History Association, Moab. 263pp. Beedy, E. C., V. K. Getz, and D. A. Airola. 1987. Status of the San Joaquin kit fox (Yulpes macrotis ~) in the Urban Kern River Parkway, Bakersfield, California. San Joaquin Valley Endangered Species Conference:47-53. Berry, W. H., 1. H. Scrivener, T. P. O'Farrell, C. E. Harris, T. T. Kato, and P. M. McCue. 1987. Sources and rates of mortality of the San Joaquin kit fox, Naval Petroleum Reserve #1, Kern County, California, 1980-1986. 33 pp. mimeo report to U. S. Dept. Energy. Boyle, S. 1995. Kit fox study, Montrose and Delta County report. Biologic Wildlife Research and Consulting to Kit Fox Project, Univ. Northern Colorado. 12 pp. plus figures. Brechtel, S., L Carbyn, G. Erickson, D. Hjertaas, C. Mamo, and P. McDougall. 1994. National recovery plan for the swift fox. National swift fox recovery team - Canadian Wildlife Service. Mimeo 49 pp. Briden, L. E., M. Archon, and D. L. Chesemore. 1987. Ecology of the San Joaquin kit fox (Yulpes velox macrotis) in western Merced County, California. California State University, Bakersfield; December 10. 11. San Joaquin Valley Endangered Species Conference: 81-87. Bunker, C. D. 1940. The kit fox. Science 92:35-36. Carbyn, L. N., and E. Klausz. 1995. Live trapping swift foxes in southeastern Wyoming and survival upon release in southern Alberta. Environment Canada report to Canadian Swift Fox Recovery Team. 39 pp. Clark, T. W., T. M. Campbell, III, D. G. Socha, and D. E. Casey. 1982. Prairie dog colony attributes and associated vertebrate species. Great Basin Nat. 42:572-582. Costello, D. F. 1954. Vegetation zones in Colorado. pp iii-x in Harrington, H. D. Manual of the plants of Colorado. Sage Books, Alan Swallow, Publisher; Denver. Cox, R L. 1994. Average pelt prices Department of Game and Fish, Santa Fe, New Mexico; 1987-88, 1989-90, 1992-93. Crumpacker, D. E., and B. L. Winternitz. 1985. Colorado wildlife workshop, species of special concern. University of Colorado, Denver. Published by Colorado Division of Wildlife. Cypher, B. L., and 1. H. Scrivener. 1992. Coyote control to protect endangered San Joaquin kit foxes at the Naval Petroleum Reserves, California. Proceedings 15th Vertebrate Pest Conference. University of California, Davis 15:42-47. Daneke, D., and M. Sunquist, M. 1984. Notes on kit fox biology in Utah. Southwestern Naturalist 29:361362. Disney, M., and L. K. Spiegel. 1992. Sources and rates of San Joaquin kit fox mortality in western Kern County, California. Transactions Western Section Wildlife Society 28:73-82. �229 Dragoo, J. W., J. R Choate, and T. P. O'Farrell. 1987. Intrapopulational variation in two samples of aridland foxes. Texas Journal of Science 39:223-232. ____J J. R Choate, T. L. Yates, and T. P. O'Farrell. 1990. Evolutionary and taxonomic relationships among North American arid-land foxes. Journal Mammalogy 71 :318-332. Egoscue, H. J. 1956. Preliminary studies of the kit fox in Utah. Journal of Mammalogy 37:351-357. __ 1962. Ecology and life history of the kit fox in Toole County, Utah. Ecology 43:481-497. __ . 1964. The kit fox in southwestern Colorado. Southwestern Naturalist 9:40. __ . 1975. Population dynamics of the kit fox in western Utah. Bulletin Southern California Academy of Science 74: 122-127. __ . 1979. Yulpes yelox. Mammalian Species. American Society Mammalogists 122: 1-5. Findley, J. S., A. H. Harris, D. E. Wilson, and C. Jones. 1975. Mammals of New Mexico. University of New Mexico Press, Albuquerque. Fitzgerald, J. P., C. Meaney, and D. M. Armstrong. 1994. Mammals of Colorado. Denver Museum of Natural History and University Press of Colorado. 467pp. __ . 1994. Furbearer management analysis: a report to the Colorado Division of Wildlife, Department of Natural Resources, Denver. 215pp. Galatowitsch, S. 1988. Colorado natural vegetation. Colorado Natural Areas Program, Denver, Colorado. Grieb, J. 1983. Black Footed Ferret Survey - annual report. Grand Junction. Ecological services: Colorado Division of Wildlife, Denver. Hadeen, K. D. 1990. Climatological data, annual summary Colorado. N.O.AA National Climatic Data Center 95. Hall, E. R 1981. The mammals of North America, 2nd. ed., John Wiley and Sons, New York, 2 vols. Halloran, A F., and F. E. Taber. 1965. Carnivore notes from the Navajo Indian Reservation. Southwestern Naturalist 10:139-140. Harrington, H. D. 1954. Manual of the plants of Colorado. Sage Books, Denver: Alan Swallow, Publisher. Harris, C. E., T. P. O'Farrell, P. M. McCue, and T. T. Kato. 1987. Capture recapture estimation of San Joaquin kit fox population size on Naval Petroleum Reserve #1, Kern County, California. U.S . . Department of Energy Topical Report, EG&GIEM Report No. EGG 10282-2149. 42pp. Hoffineister, D. F. 1986. Mammals of Arizona. The University of Arizona Press and Arizona Game and Fish Department, Tucson. 602 pp. Hunter, W. R 1981. Soil survey of Paonia area, Colorado, parts of Delta, Gunnison, and Montrose counties. United States Dept. Agriculture, Soil Conservation Service and Colorado Agricultural Experiment Station. 1, Linhart, S. B., and F. F. Knowlton. 1975. Determining the relative abundance of coyotes by scent station lines. Wildlife Society Bulletin 3: 119-125. Link, M. A 1995. Kit fox (Yulpes macrotis) distribution in western Colorado. Univ. Northern Colorado, Greeley. M.A Thesis. 115pp. Martin, G. 1993. A little fox's big troubles. Nature Conservancy. Marchi April: 10-15. McGrew, J. C. 1977. Distribution and habitat characteristics of the kit fox (Yulpes macrotis) in Utah. Utah State University, Logan. M. S. Thesis. 92pp. __ . 1979. Yulpes macrotis. Mammalian Species, 123. American Society of Mammalogists 123: 1-6. Mercure, A, K. Ralls, K. P. Koepfli, and R K. Wayne. 1993. Genetic subdivisions among small canids: mitochondrial DNA differentiation of swift, kit and arctic foxes. Evolution 47: 1313-1328. Miller, P. H. 1964. The ecological distribution of mammals in Colorado National Monument, Mesa County, Colorado. Oklahoma State University, Stillwater. M. S. Thesis. 133pp. ____J and C. J. McCoy, Jr. 1965. Kit fox in Colorado. Journal of Mammalogy 46:342-343. Morrel, S. H. 1972. Life history of the San Joaquin fox. California Fish and Game 58:162-174. __ . 1975. San Joaquin kit fox distribution and abundance in 1975. California Department ofFish and Game, Wildlife Management Branch, Administrative Report 75-327. O'Farrell, T. P. 1983. The San Joaquin kit fox recovery plan. United States Fish and Wildlife Service and United States Department of Energy, Portland, Oregon. 84pp. �230 __ . 1984. Conservation of the endangered San Joaquin kit fox, Yulpes macrotis mutica, on the Naval Petroleum Reserves, California. Acta Zoologica Fennica 172:207-208. and L. Gilbertson. 1986. Ecology of the desert kit fox, Vulpes macrotis arsipus, in the Mojave Desert of southern California. Bulletin. Southern California Academy Science 85: 1-15. __ . 1987. Kit fox. Pp 422-431 in M. Novak, J. A Baker, M. E. Obbard, and B. Mallock, eds. Wild furbearer management and conservation in North America. Ontario: Ministry of Natural Resources. O'Neal, G. T., J. T. Flinders, and W. P. Clary. 1986. Behavioral ecology of the Nevada kit fox (Yulpes macrotis neyadensis) on a managed desert rangeland. Current Mammalogy 1:443-481. Orloff, S. 1987. Survey techniques for the San Joaquin kit fox (Yulpes macrotis mutica) California State University, Bakersfield. San Joaquin Valley Endangered Species Conference 185-197. A W. Flannery, and K. C. Belt. 1993. Identification of San Joaquin kit fox (Yulpes macrotis mutica,) tracks on aluminum tracking plates. California Fish and Game 79:45-53. Robinson, W. B. 1953. Population trends of predators and fur animals in 1080 station areas. Journal of Mammalogy 34:220-227. Rohwer, S. A, and D. L. Kilgore, Jr. 1973. Interbreeding in the arid-land foxes, Yulpes yelox and macrotis. Systematic Zoology 22: 157-165. Samuel, D. E., and B. B. Nelson. 1982. The kit fox and the swift fox. Pp. 485 ..490 in J. A Chapman, and G. A Feldhamer, eds. Wild Mammals of North America: biology, management, and economics. Johns Hopkins University Press, Baltimore. Schitoskey, F., Jr. 1975. Primary and secondary hazards of three rodenticides to kit fox. Journal Wildlife Management 39:416-418. Scott-Brown, J. M., S. Herrero, and J. Reynolds. 1987. Swift Fox. Pp. 433-441 in M. Novak, J. A Baker, M. E. Obbard, and B. Mallock, eds. Wild furbearer management and conservation in North America. Ontario Ministry of Natural Resources. Sheldon, J. W. 1992. Wild dogs: the natural history of nondomestic Canidae. Academic Press Inc., San Diego. Smith, D. G. 1978. Notes on ecology and food of the kit fox in central Utah. Sociobiology 3:96-98. Thorton, W. A, and G. C. Creel. 1975. The taxonomic status of kit foxes. Texas Journal of Science 26:127136. Wayne, R K., W. G. Nash, and S. J. O'Brian. 1987. Chromosomal evolution of the Canidae. 2. Divergence from the primative carnivore karyotype. Cytogenetics Cell Genetics 44: 134-141. Weber, W. A, and R C. Wittman. 1992. Catalog of the Colorado flora: a biodiversity baseline. University Press of Colorado, Boulder. White, P. J., and K. Ralls. 1993. Reproduction and spacing patterns of kit foxes relative to changing prey availability. Journal Wildlife Management 57:861-867. . Wood, J. E. 1959. Relative estimates of fox population levels. Journal Wildlife Management 23:53-63. Woolley, T. P., F. G. Lindzey, and R Rothwell. 1995. Swift fox surveys in Wyoming - Annual Report. Pp. 61-79 in S. H. Allen, J. W. Hoagland, and E. D. Stukel, eds. Report of the swift fox conservation team. Mimeo Report, 170pp. Zoellick, B. W., T. P. O'Farrell, and T. T. Kato. 1987. Movements and home range of San Joaquin kit foxes on the naval petroleum reserves, Kern County, California. EG&GIEM report #EGG 10282-2184 for U.S. Dept. Energy. 38pp. N. S. Smith, and R S. Henry. 1989. Habitat use and movements of desert kit foxes in western Arizona. Journal Wildlife Management 53:955-961. ~ and N. S. Smith. 1992~ Size and spatial organization of home ranges of kit foxes in Arizona. Journal of Mammalogy 73:83- 88. _____J _____J v.. _____J �231 APPENDICES APPENDIX A. Table l. Total trap effort by counties and areas in which kit foxes were captured, western Colorado, 19921996. Does not include reca~tures. County Search Period Tra~ Nights Ca~tures Garfield (Western) Prairie Canyon Mesa Rabbit Valley Prairie Canyon to Mt. Garfied. MesalDelta Co. from Whitewater to Delta Delta County West of Gunnison Gorge Delta Airport MontroselDelta Peach Valley 8/17-22/94 5/27-31195 3/26-29/92 4/2-5/93 8/2-5/94 8/9-11/94 6/18/95 .3/29-4/15/92 4/25-28/92 9/15-23/92 4/11-13/93 10/20-28/93 7/11-8/14/94 5/31-6/18/95 6/21-27/95 6/27-30/95 8/11-14/95 10/30-11/9/95 4-28-5-16/92 5/2-9/92 6/21-27/94 7/7-17/94 1120-3-11195 7/30-8/4/95 8/7-10/95 10/16-12/12/95 12/14-16/93 11119/93-116/94 5-27-5-30/94 11116-19/95 5/13-31/92 6/10-25/92 9/25-30/92 1/14-17/93 2/23-28/93 4/19-23/93 8/29-9/2/93 9/12-30/93 12/5-8/93 4/5-24/94 5/2-6/2/94 6/6-9/94 7/12-29/94 9/1-7/94 96 60 84 24 90 45 60 156 48 150 30 132 463 153 67 53 47 204 192 93 119 138 160 46 48 240 60 245 39 80 198 141 76 48 144 24 81 150 6 139 215 34 258 54 1 0 0 0 2 1 0 0 0 0 0 0 4 0 0 0 1 1 0 0 0 2 0 0 0 1 0 1 0 2 1 3 0 2 1 0 2 0 0 0 0 4 0 --------------------------------------------------------------------------------------------- �232 Table l. (Continued) County Montrose East Search Period Trap Nights 1111-31/95 7/9-28/95 8/1-18/95 4/13-G/19/96 5/27-6/18/94 8/24-27/94 7/4-11/95 7/17-20/95 4/13-6/19/96 20 257 318 10 357 44 53 10 40 6099 Total Trap Nights Captures 0 5 0 0 9 4 0 0 0 47 Table A2. Summary of trapping effort in locations in which kit foxes were not captured, western Colorado 1992-1996. #Searches Year County & Area Trap Nights Moffat Browns Park Rio Blanco Blue Mtn.lMellen Hill Garfield DeBeque-Parachute Mesa National Monument Horsethief Canyon Reeder Mesa S.ofWhitewater Gateway Area Delta DryISawmill Mesas East of Austin West of Wildlife Area Lawhead Gulch Montrose S. Hwy 50, E Montrose Sinbad Valley Paradox Valley San Miguel Big Gypsum Valley Disappointment Valley McIntyre Canyon Montezuma County McElmo Canyon Total 4 93-95 636 1 93 72 1 94 172 1 1 1 1 1 92 94 94 92 92 78 30 76 69 114 1 1 1 1 92 92 92 95 21 87 96 96 2 1 1 94-95 92 92 90 168 141 1 1 1 92 92 93 174 141 77 92 60 2419 �233 APPENDIXB. Table 1. Locations of radio-collared and uncollared kit foxes trapped or observed in the lower Colorado River Drainagesz Mesa and Garfield Countiesz March 1992-June 1996. Status as Site & Date of Sex/Age Radio-collar Location of30 June Ear Tag # Freguency CaEtureffracking Rabbit Valley 8124/94 2/19/24 4/10/95 8/2/94 8/10/94 9/9/94 AM 194 151.244 AF196 AF198 151.221 151.034 TI0S,RI04W,S20 TI0S,RI04W,S20 TI0S,RI04W,SI7 TI0S,RI04W,SI2 TlOS,RI04W,S13 TIOS,RI04W,S13 Unknown Unknown T7S,RI05W,S12 Unknown Unknown Prairie Canyon 8/21/94 UAD153 151.082 JF126 150.834 Corcoran Point 7/15/94 12/19/94 3120/95 7/15/94 7/17/94 3120/95 4/9/95 7/19/94 12119/94 3/20/95 4/9/95 8/12/95 11/2/95 3/17/96 5116/96 1113/95 3/17/96 5/16/96 6/18/96 JM 128 AM 130 n. c. 150.468 AF 132 JM310 150.638 n. c. 150.189 AF311 151.186 Collar at Erivate residence T 10S,RI00W,S35 T 10S,RI00W,S35 T 1OS,Rl 00W,S35 TI0S,RI00W,S35 TI0S,RlOOW,S35 TI0S,RI00W,S35 TI0S,RI00W,S35 TI0S,RI00W,S35 TlOS,RI00W,S3 Tl OS,Rl 00W,S35 T 10S,RI00W,S35 T9S,RI00W,S35 T9S,RI00W,S34 TI0S,RlOOW,SIO Signal too weak to locate TIN,RlW,SIO TI0S,RlOOW,SIO TlN,RlE,S29 TINzRlWzS36 Unknown Unknown Unknown Unknown Unknown Dead Table 2. Locations of radio-collared and uncollared kit foxes trapped or observed in the Gunnison River Drainagez north of the river in MesallDelta Countiesz March 1992-June ·1996. Site & Date of Sex/Age Radio-collar Status as CaEtureffracking Ear Tag # Freguency Location . of30 June Cheney Reservoir 7111/94 7/11/94 1214/95 5116/96 6/18/96 MJ 101 150.851 AF Jaw Broken n.c. AM 312 151.083 moved to Wells Gulch T 13S,R98W,S30 T13S,R98W,S30 T3S,R2E,S24 T4S,R3E,S13 T4S,R3E,S13 Unknown Died in Rehab. Tl4S,R98W,S33 Tl4S,R96WzS36 Unknown Unknown Dead Delta Airport 11122/93 11119/95 * MJ 17 AM*33 Marked as ajuvenile in Peach Valley. 150.980 151.160 �234 Table 3. Locations of radio-collared and uncollared kit foxes trapped or observed in the Peach Valley area, March 1992-June 1996. Sex/Age Radio-collar Date of Status as Ear Tag # Freguency Location of30 June CaJ2turerrracking T51N,R9W,S29 Unknown 5/14/92 AM n. c. 5/31192 AF I 150.238 T5IN,R9W,S29 11123/92 TI5S,R94W,S27 Dead Unknown 6/25192 AM4 n. c. T50N,R9W.SI6 9/28/92 150.309 T50N,R9W,S9 Unknown AM2 T50N,R9W,S9 . AF5 150.338 9/28/92 4/6/94 150.956 T50N,R9W,S9 T51N,R9W,S32 4/14/95 150.873 T51N,R9W,S7 8/15195 T51N,R9W,S29-32 10/11195 TI5S,R97W,SI Unknown 12/4/95 Moved to Alkali Gulch T50N,R9W,S9 9128/92 AF6 150.379 T50N,R9W,S9 14114 150.850 3/2/93 T50N,R9W,S9 150.889 9/22/93 T50N,R9W,SI7 116/94 Dkad Moved PVto SE Montrose T49N,R8W,S5 2/10/95 T51N,R9W,S29 150.638 2/23/93 AF7 T50N,R9W,S5 10/26/93 T50N,R9W,S5 18/18 150.594 12/6/93 T50N,R9W,S5 151.009 915194 T50N,R9W,S5 2128/95 T51N,R9W,S29 Dead 4/14/95 T51N,R9W,S29 AM8 150.468 2/23/93 T50N,R9W,S5 10126/93 T50N,R9W,S5 150.189 12/6/93 9/9/94· T50N,R9W,SIO T51N,R9W,S29 151.095 147/148 1125/95 Dead T51N,R9W,S29 6128/95 150.708 T50N,R9W,S22 AM 12 4/21193 T50N,R9W,S8 150.037 15/15 9/24/93 T50N,R9W,S 17 10128/93 T50N,R9W,SI6 12/16/93 Dead T50N,R9W,S8 3126/94 T50N,R9W,SI6 150.813 AM 13 9120/93 T50N,R9W,SI6 12/14/93 T51N,R9W,S29 6/19/94 T51N,R9W,S29 Unknown 151.177 46147 1127/95 T50N,R9W,SI7 150.499 AM 16 9/30/93 Unknown T50N,R9W,SI7 11122/93 151.083 T51N,R9W,S31 AF 190 7/14/94 T51N,R9W,S31 917194 T51N,R9W,S29 12/19/94 T5IN,R9W,S29 1127/95 Dead? T51N,R9W,S29 Collar recovered no carcass 4/14/95 T50N,R9W,SIO 151.257 JM 192 7/18/94 T50N,R9W,SI0 9/9/94 Dead T50N,R9W,S21 11/18/94 --------------------------------------------------------------------------------------------- �235 Table 3. Continued. Date of Ca]2turefTracking 5/14/92 12/22/94 1/10/95 1/27/95 4/18/95 6/29/95 8/15/95 10/11/95 5/16/95 2/14/96 6/18/96 8/24/94 12/22/94 1/10/95 4/13/95 6/20/95 10/11/95 2114/96 4/9/96 12/4/95 2/14/96 7/26/95 2/14/96 4/9/96 7/26/95 12/25/95 7/27/95 2114/96 4/9/96 7/27/95 5/16/96 6/18/96 7/28/95 2/14/96 6/18/96 Sex/Age Ear Tag # 8/26/94 Radio-collar Fr~uency AM 30 moved ME to PV 30/151 151.209 collar in Selig Canal JM26/27 151.131 moved from ME to PV 151.056 (Moved to Alkali Gulch northwest of Delta) East of Olathe JF 306 150.728 JM305 150.920 Near Sweitzer Reservoir JF308 150.004 moved to ME moved back to PV JM307 n.c. 151.009 AF309 150.029 moved to ME moved between ME and PV Location 151.107 T49N,R9W,SI2 T49N,R9W,S12 T5IN,R9W,S29 T5IN,R9W,S32 T5IN,R9W,S32 T5IN,R9W,S30 T5IN,R9W,S29-32 T5IN,R9W,S29 T5IN,R9W,S30 T5IN,R9w,S29 T49N,R8W,S7 T49N,R8W,S7 T49N,R8W,S7 T50N,R9W,S9 T50N,R9W,S9 T5IN,R9W,S29 T50N,R9W,SI6 T50N,R9W,SI6 T 15S,R97W,S 1 Status as of30 June T49N,R9W,S7 Dead? Dead A? T50N,R9W,SI6 T50N,R9W,SI6 T50N,R9W,SI6 T50N,R9W,SI6 Tl5S,R95W,S33 T50N,R9W,SI6 T49N,R8W,S24 T51N,R9W,S30 T50N,R9W,SI6 T49N,R9W,S6 T49N,R9W,S6 T50N,R9W,S 1G T49N,R8W,S9 T49N,R8W,S 3 Dead Dead Dead Alive Alive �236 Table 4. Locations of radio-collared and uncollared kit foxes trapped or observed in the Montrose East area, March 1992-June 1996. Date of Sex/Age Radio-collar Status as Caetureffracking Ear Tag # Location of30 June Freguen£Y 5/29/94 AF21 150.947 T49N,R8W,S7 8126/94 T49N,R8W,S7 12/22/94 T49N,R8W,S6,7 1110/95 T49N,R8W,S6 4/17/95 52/52 T49N,R8W,S7 151.288 6127/95 T49N,R8W,S7 2/14/96 T49N,R8W,S7 6/18/96 T49N,R9W,S6 150.098 , Alive 5/29/94 AF22 150.403 T49N,R8W,S7 T49N,R8W,S7 8/25/94 1110/95 22/38 T49N,R8W,S7 Unknown 151.146 5/29/94 AF23 150.338 T49N,R8W,S7 T49N,R8W,S7 8/28/94 T49N,R8W,S7 9/15/94 7/7/95 151.042 T409N,R8W,S5 T49N,R8W,S7 2/14/96 T49N,R8W,S9 Dead 6/18/96 Unknown JM24 n. c. T49N,R9W,S7 5/29/94 JF 176 n.c. T49N,R9W,S7 5/30/94 150.037 T49N,R9W,S6 8/25/94 T49N,R9W,S7 1/10/95 Unknown questionable radio signal by Mack, Mesa County 2/14/96 T49N,R8W,S7 Unknown JF 178 n. c. 5/30/94 T49N,R8W,S7 5/31194 AF 180 150.940 T49N,R8W,S7 6/7/94 T49N,R8W,S9 Dead 7/28/94 T49N,R8W,S7 JF 182 n.c. 5/30/94 151.020 T49N,R8W,S7 8/28/94 T49N,R8W,S7 9/16/94 T49N,R8W,S8 Dead? 11111194 collar recovered no carcass T49N,R8W,S7 n.c. 6/3/94 AM 184 T49N,R8W,S7 8125/94 150.708 T49N,R8W,S7 1110/95 T49N,R8W,S7 151.237 4/18/95 T49N,R8W,S6 6/27/95 10/11195 T50N,R9W,SI8 T50N,R9W,S16 2/14/96 T50N,R9W,S26 5/16/96 151.160 T49N,R8W,S7 8127/94 JM33/34 T49N,R8W,S6-7 12/29/94 T49N,R8W,SI2 114/95 T 14S,R69W,S36 (Moved north of Delta) 10/11195 Not found radio probably dead. 2/14/96 151.107 T49N,R9W,S7 ADM29/30 8/26/94 T49N,R9W,S7 12122/94 T49NR9W,S 12 U 1110/95 ' 151.042 T49N,R9W,S7 ADM48/49 8/27/94 T49N,R8W,S8 1119/94 D(C) T49N,R8W,SI9 11/22/94 �237 APPENDIXC Summary of Characteristics of Trapped Areas: 1. Moffat County: Trapping took place in Browns Park, along the Vermillion Bluffs, in Powder Wash, and the margins of Irish Canyon. The area drains to the Green River. Trapping was conducted on Colorado Division of Wildlife, National Wildlife Refuge, or Bureau of Land Management property. Elevation ranged from 1650-1752 m in the trapped areas. Most of the area trapped was in sagegrassland, merging with pinyonjuniper at with higher elevations (Link 1995). 2. Rio Blanco County: Two areas: Blue Mountain (T2N, RI02W) and Mellen Hill (T3N, RI03W) were trapped. These are BLM lands at about 1650 m elevation dominated by sagebrush and draining to the White River. Washes usually had saltbush and greasewood cover. The Chevron Oil Company and Colorado Division of Wildlife have conducted numerous searches for black-footed ferret in the area but never observed kit fox (D. Sellars, Chevron Environmentalist, pers. commun.; CDOW 1986). The areas were trapped because of trapper harvest reports of kit fox from Rio Blanco County. 3. Mesa County and SW Garfield County: Grand Valley Area - This valley is an eastern extension of the Great Basin dominated by cold desert shrub lands that extend for hundreds of square km into eastern Utah. Most of the area is managed by the Bureau of Land Management for livestock grazing. Lands close to the Colorado river are urbanized or under intensive agriculture. Public lands, especially those close to Grand Junction and Fruita receive considerable use by recreationists involved in driving off-road vehicles, target shooting and similar pastimes. CDOW trapper returns have reported kit fox from Mesa County, Miller and McCoy (1965) reported on kit foxes killed in the valley, and CDOW personnel have reported observations of kit foxes in portions of the valley in the early 1990's. Those recordsjustified spending a considerable amount of project time trapping in the area. Colorado National Monument. Portions of the Monument, primarily along the Rim Rock Drive at average elevations of 1763 m were trapped. Most traps were placed on the margins of pinyon-juniper communities. The Monument was trapped on the basis of Miller's (1964) observations of kit foxes and a few more recent reports of unidentified small foxes occasionally seen in the area. NW of Gateway. A portion along the Dolores River to the Utah state line in parts ofTISS and RI04W was trapped. Cattle range over most of the area trapped. The elevation averaged 1388 m with cover consisting primarily of grasses, Russian thistle, blackbrush (Coleogyne sp. ), sagebrush, and some prickly pear cactus. The valley was trapped because it offered a narrow strip of potential kit fox habitat that was contiguous to potential kit fox habitat in Utah. 4. Garfield County - East of DeBeque Canyon: Portions of the upper Grand Valley from south and west of the town of DeBeque to Parachute were trapped. Most of the public lands are cold desert shrub communities interspersed with juniper woodlands. Private land intermingles with BLM range with private holdings used for irrigated agriculture or as pastures. The area was trapped on the basis of a 1980's report by a CDOW employee of possible kit fox in the area and a few trapper reports of foxes from unspecified locations in Garfield County. �238 5. Mesa and Delta Counties: Portions of the lower Gunnison River valley from Whitewater to Alkali Gulch west of Delta were trapped north of the river canyon. A few sites west and southwest of Delta and south of the Gunnison River and west of the Uncompaghre River were also trapped. Terrain varied from rolling hills, to mesas and large, open flats. Sparse pinyon-juniper occurs at higher elevations with most of area covered by grasses or low shrubs, primarily saltbush. The average elevation is slightly less than 1500 m. Most of the area is in BLM ownership while substantial areas bordering on U.S. 50 are in private ownership. We had little success in procuring permission to trap on private inholdings in the area. North and East of Delta, North of the Gunnison River. Two areas were trapped in this region bounded to the west by Alkali Gulch and to the east by private lands east of Lockhead Gulch. Most effort centered around BLM lands close to the Delta Airport in cold desert shrub communities. South and East of Delta, Delta County to Montrose, Montrose County. All areas in Peach Valley that could be accessed were trapped. Peach Valley drains to the Gunnison or Uncompaghre Rivers. Two general areas were trapped in this region: Peach Valley is centered east of the town of Olathe, south and west of the Gunnison River, and east of Highway 50 in Tl5S, T50N, R9W, and R94W. Peach Valley is .accessed by county roads as well as some rough four wheel drive roads. Dirt bike and horseback riders use much of the BLM land for recreation. Most of the northern portion and all of the western edge of Peach Valley is privately owned with much of it in irrigated agriculture. BLM land lies to the East (Fig. 14). The topography ranges from flats to hills and small mesas with slopes averaging between 25-45 degrees. Peach Valley had been selected for trapping based on vegetation and topographical features. In 1992 a local landowner directed Link (1995) to the first den of kit foxes located during the study. Soil surveys have not been completed for most of Peach Valley by the Soil Conservation Service. Its soils are similar to those described by Hunter (1981) for the Paonia area which included the northern and eastern edges of Peach Valley. Badlands, Billings, Chipeta, and Persayo soils are well-drained soils of Peach Valley, formed by weathered shale and characterized by light-brown colored soils made of sands. and silty, clay, loam soils (Hunter 1981). These clay-loam layers are about four inches thick and support vegetation including greasewood, shadscale, fourwing and mat saltbush, Indian ricegrass, wheatgrass and galleta (Hunter 1981). Silt contains mineral particles that range in diameter from the upper limit of clay (0.002 mm) to the lower limit of very fme sand (0.05 mm). Clay is made up of mineral soil particles less than 0.002 mm in diameter. Loam is soil material that is 7-27% clay particles, 28-50% silt particles, and less than 52% sand particles (Hunter 1981). The dens of kit fox found in Peach Valley were all found in soils of these types. 6. Montrose County: Montrose East - South and east of Flattop Mesa at the extreme end of Peach Valley lies a drainage we have termed Montrose East. It is a narrow, roaded drainage vegetated by sagebrush and saltbush. The area connects to Peach Valley via badlands east of or around the base of Flattop Mesa. We trapped the site because several local individuals reported a family of kit foxes using the area. Sinbad Valley in T49N and RI04W is a small mountain valley at approximately 1750 m elevation that drains eastward to the Dolores River. It is heavily vegetated with sagebrush, grasses, oak, prickly pear, and thistle (Link 1995). The valley is surrounded by dense stands of Pinyonluniper and is a major area . of deer winter range. Approximately 50% of the valley is private land most trapping was conducted on BLM lands. Habitat was the least similar of any trapped areas to habitats described in the literature for kit fox. There were no reports of kit fox from this area. �239 Paradox Valley was trapped in parts ofT46N, T47N, RI6W, and RI9W. It lies at an elevation of 15301630 m with the Dolores River the main drainage. Vegetation included large areas of grasslands, with PinyonJuniper and sagebrush stands at slightly higher elevation, or on well drained uplands, especially around the edges of the valley. Ground cover averaged over 50% (Link 1995). Private lands are interspersed with public lands in Paradox Valley. Private land owners were contacted and interviewed about kit fox presence, and permission was obtained to trap and spotlight private lands. A dead gray fox was collected by peregrine falcon researchers in the valley in the fall of 1992. We had no reports of kit fox from this valley. 9. San Miguel County: Big Gypsum Valley a drainage of the Dolores River Basin, was trapped in parts ofT44N, T45N, RI6W, and RI8W. It is a fairly flat valley at 1670-1780 m elevation. Vegetation includes sagebrush species (20%), various grasses, winterfat, horsebrush, some prickly pear and Russian thistle. Vegetation cover averaged 54% (Link 1995). The majority ofland is managed by the Bureau of Land Management with 10-20% of the valley owned privately. (Appendix C, Fig. 18). There were no records of kit fox from this area. Disappointment Valley was trapped in portions ofT44N, T43N, RI7N, and RI8N. The elevation of Disappointment Valley ranges from 1730-1800 m, with large flats vegetated by cold desert shrubs. The valley drains to the Dolores River. There were no records of kit fox from this area. McIntyre Canyon was trapped in portions ofT44N, RI9W, and R20W. This canyon lies between 18001900 m elevation on the west side of the Dolores River and can only be accessed by boat or by crossing the river when frozen (Appendix C, Fig. 20). Vegetation is similar to that in the Gateway area. There were no records of kit fox for this area. 10. Montezuma County McElmo Canyon was trapped in parts ofT36N, RI9W, and R20W. Elevations in the canyon range from 1500-1580 m. Fruit orchards, crop and pasture lands occupy the eastern portion or the canyon. The west bordering Utah, supports a cold desert shrub community (Link 1995) (Appendix C, Fig. 21). Red and gray fox have been seen in this valley (pers. commun. Tom Beck, Tozer families). Private land ownership is interspersed with public lands and permission was granted to trap on some privately owned lands. McElmo Canyon was selected for survey based on reports of two kit fox skulls from.the canyon by Egoscue (1964). ��241 Colorado Division Wildlife Research July 1997 of Wildlife Report JOB FINAL REPORT state of ~C~o~l~o~rLa~d~o~ Project Work No.W ~~-~1~3~5~-~R~--=1~0 Plan No. __~l~O~A~ Job No. __~2~ Period Covered: Author: James Personnel: July _ _ Mammals _ swift _ Swift Fox Investigations Colorado Research Fox Studies in 1, 1996 to June 30, 1997 P. Fitzgerald Colorado Division of Wildlife: R. Kahn, T.Beck, B.Gill. University of Northern Colorado: J. Fitzgerald, D. Finley, B. Roell, C. Gilin, L. Irby, J. Trefren, P. Stapp, T. Tombs, M. Henderson ABSTRACT The eastern plains live-trapping inventory was completed. Thirty plots in 11 counties were trapped in 1996-97 with 109 foxes (50 males, 57 females, and 2 sex unknown) captured from 25 of the plots. Captures averaged 4.4 foxes per plot (range 1-10). Since March 1995 we sampled 72 plots covering 1440 rni2 (3730 krn2) and have captured 243 foxes (118 male, 122 female, 3 sex unknown) from 51 (71%) of the plots. Most of our captures have been from large expanses of short grass prairie in the southern tier of counties. On the intensive study areas in Weld County we captured and marked 33 new individuals in 199697. Since October 1994 we have captured 122 foxes (40 females, 37 males on CPER; 24 females, 21 males on G1) on the 2 sites. capture success has been significantly lower on the CPER. CPER foxes are in significantly larger, more localized clusters than foxes on Gl. Fifty-four foxes have been found dead including 41 radio-collared animals. coyotes accounted for 66% of the deaths of radioed animals. Survivorship rates for 109 radio-collared foxes averaged .658 for 1995 and 1996 with no difference in survival between sexes or sites. Ten whelping dens, 6 on the CPER and 4 on G1 have produced 31 pups this year. Pup mortality is at least 54% based on radio-collared animals. Horne range estimates for 4 males and 3 females have averaged 4.2 rni2 (11 krn2) and 2.3 rni2 (6 krn2) respectively. Population estimation (program NOREMARK) using marked animals and infrared cameras yielded a mean of 25 foxes on the G1 site during January and February, 1 fox per 2 rni2 (5.2 krn2). ��243 SWIFT FOX rNVESTIGATIONS James IN COLORADO P. Fitzgerald P.N. OBJECTIVE Determine recommend habitats. the status and trend of swift fox populations in Colorado and conservation strategies to maintain existing populations and SEGMENT 1. Survey sites swift fox. in eastern Colorado OBJECTIVES which are believed 2 ..Conduct intensive research on populations of swift the Central Plains Experimental Range to: a. Measure factors to be inhabited by fox on and adjacent to recruitment in swift fox populations and identify potential which contribute to the regulation of recruitment; b. Develop estimates of swift fox home range size to environmental variables; size and relate home range c. Test the potential of automated photography systems to estimate fox numbers using mark/resight population estimation models; swift d. Collect, patterns and use analyze, and report on data describing characteristics of swift fox natal dens and den occupancy; e. Document incidents of swift fox mortality from coyote predation and evaluate the importance of coyote predation to the population biology swift fox. METHODS of AND MATERIALS The eastern plains survey of 72, 3x4 mi2 trapping plots (20 mi2 effective trapping area) started in March 1995 was completed. Plots were selected randomly and trapped an average of 4 days using 20 live traps placed at section corners on the grid. All captured foxes were sexed, weighed, and ear tagged prior to release. Studies on aspects of swift fox biology were continued on the Central Plains Experimental Range (CPER) and Pawnee National Grassland (G1) sites in Weld County using methods described in the 1995-96 an~ual report. We continued to live-trap and radio-collar foxes on both sites, monitor reproduction and survival, and obtain information on habitat use and den site characteristics. Additionally, 5 female and 4 male foxes on the CPER were live trapped and fitted with dummy collars in October and November of 1996. The dummy collars were made from industrial belting material with airplane cable antennas. Twenty guage shotgun shells filled with bb shot mimicked the radio-transmitters. The dummy collars and their antennae were painted in unique patterns to determine whether they could be used in conjunction with hair dye patterns to allow identification of individual foxes photographed with infra-red sensing cameras. Nyanza A black hair dye was used for painting distinctive patterns on individual foxes. The infrared recordercamera system was manufactured by Trailmaster (Lenexa, KS) and has been described by Beck (1955). We tested 20 camera units placed on a 1 mi grid system on the CPER in November. Cameras were run for 3, 4 night bouts with 4 �days between bouts. Attractant baits, varied with each bout, were placed between the cameras and the infrared transmitting unit. In late November and December we live trapped 10 females and 9 males on the G1 site and equipped them with distinctively colored functional radio-collars and dye marked them with unique patterns using the techniques described for the CPER trial. In January and early February 1997 we ran 4 camera bouts using 31 cameras placed at 2 mi intervals on an expanded G1 study area of approximately 62 mi2• Results of the camera, mark-resight test on G1 were analyzed by T. Beck using program NOREMARK (White 1996). Survivorship curves were generated for foxes captured on both sites since October 1994 using the Kaplan-Meier formula with staggered entry (Pollock et al. 1989). Nearest neighbor analysis (Manly 1991) was used to test for randomness in distribution of captured foxes. Efforts continued on mapping whelping dens and characterizing features at den sites including percent frequency of vegetation, habitat type, aspect and slope of den entrances. Food debris and scats were collected and scat are being analyzed using the point sampling method described by Cameron (1984) in his investigation of swift fox diets on the Pawnee National Grassland. RESULTS Eastern Plains Swift Fox Inventory: The field trapping phase of the eastern plains inventory was completed in January 1997. In 1996-1997, 30 plots in 11 counties were live trapped. One hundred and nine foxes, 50 males, 57 females, and 2 of unknown sex were captured. October and November were the months with highest capture success (Table 1). The second and third nights of trapping Table 1. Numbers of swift fox captured by year and month in eastern for the 1996-1997 field season. In 1997, January was the only month trapping was conducted. 1996-97 Sept Jan Jul Aug 4 5 3 3 100 156 79 4 3.2 2.5 (30 p1ots) Oct Colorado in which Nov Dec Totals 34 54 6 180 338 620 496 1969 1.7 10.1 8.7 1.2 4.5 ~vg 109 Captures Trap Nights Catch/ 100 Traps produced the highest yield of foxes (Table 2). Foxes were captured on 25 of the 30 plots (83%) with an average capture of 4.4 foxes per plot (range 1-10) about 1 fox per 4 mi2 (Table 3). Since March 1995 we have sampled 72 plots covering 1440 mi2• A total of 243 foxes (118 male, 122 female, 3 of undetermined sex) were captured from 51 (71%) of the plots. Most fox captures (62%) have been on sites dominated by short-grass prairie. Seven percent have been on sites with short grass prairie mixed with pinon-juniper and 7% have been on short grass prairie mixed with cropland. �245 Table 2. Fox captures by night of capture July 1996-June for the 1996-97 field season. 1997 Night 1 2 3 4 Totals Captures 21 34 40 14 109 Trap Nights 568 572 532 297 1969 6.1 7.7 5.1 4.5avg. Catch /100 Nights 3.7 Central Plains Experimental Range Investigations: In 1996-97 we captured and marked 33 new individuals, 10 female and 8 male foxes on the CPER and 10 females and 5 males on G1. Since October 1994 we have captured 122 foxes (40 females, 37 males on CPER; 24 females, 21 males on G1) and made 132 recaptures. We have had signific~nt differences (Z-test, p=<.OOl) in capture success between the G1 (9.8%) and CPER sites (3.9%). Nearest-neighbor analysis of foxes trapped on the two areas shows a significant difference from random distribution. The CPER foxes are in larger, more localized clusters in their dispersion pattern than are foxes on G1. Vegetation patterns probably account for most of this difference. One-hundred and nine foxes (52 males and 57 females) were fitted with radiocollars since October 1994. As of July 1 1997, 20% of our marked foxes are alive, 7 males and 5 females on the CPER and 5 males and 5 females on G1. We have lost radio contact with 44% (17 female, 11 male) of the CPER foxes and 40% (7 female, 9 male) of the G1 foxes. A few CPER collared foxes and one G1 female have moved off the study areas but the fate of most is unknown. We have had no marked foxes migrate between the two study areas located about 13 km apart. A total of 54 foxes have been found dead including 20 radio-collared males and 21 collared females (38% of collared animals). Twenty-five (39%) of the collared foxes on the CPER are dead (11 females, 14 males). Sixteen (37%) of the G1 foxes (10 females, 6 males) are dead. Coyotes were the primary cause of mortality (66%) for collared animals (27/41 deaths). Hunters illegally killed 4 (10%) marked animals, automobiles killed 3 others (7%). A cohort of 17 female and 13 male foxes were radio-collared on the CPER in fall and winter of 1994. Only 1 female and 3 males are still alive (13%) with an average survival of 949 days post capture. The fate of 15 animals, 10 females and 5 males is unknown after being radio-tracked an average of 226 days. Eleven animals are dead, 6 females and 5 males, after surviving an average of 193 days. Only 17 (8 male, 9 female) of our 109 marked animals were captured as pups while still at pupping dens. Thirteen of them (6 female, 7 male) were radio-collared. Three of the females are dead, one slipped its collar, the fate of 2 is unknown ..Four of the males are dead the other 3 are unaccounted for. Percent mortality is a minimum of 54% after an average tracking period of 168 days. Survivorship was estimated for the period October 1994 to July 1 1997 for the 109 radio-collared foxes (Table 4). The mean survival rate is .235 for the entire period with no significant difference between sexes or study areas. Annual survival for 1995 was .572 compared to .745 for 1996 (average = .658). Females marked in fall of 1994 and assumed capable of breeding in subsequent �246 Table 3. captures on swift fox live-trapping plots by latilong block and county, July 1,1996-January 9,1997. SGP= short-grass prairie, MGP= mid-grass prairie, SSP= sand-sage prairie, CL=cropland, CRP= conservation reserve, RIP=riparian, PJ=pinyon-juniper woodland, SLTB= saltbush. Latilong Block Plot # North-central Tier Limon 41 91 county Vegetation UNK CL SGP/CL M 0 0 F Li,ncoln Lincoln 0 0 1 South-central Tier Karval 38 Lincoln CL/MGP 1 0 1 77 Lincoln SGP/SSP 1 0 1 88 Lincoln SGP/MGP 1 1 2 17 Crowely SSP/SGP 4 2 29 Crowely SSP/SGP 2 6 8 43 44 57 115 Pueblo Pueblo Pueblo Pueblo SGP SGP sSP/SGP SGP/cholla 0 0 0 0 0 0 0 1 0 0 0 1 Southern Tier Kim 73 Las Animas SGP/cholla 1 2 3 La Junta 12 Bent SGP 4 1 5 27 37 116S otero otero otero SGP/SSP SGP/SSP SGP 5 2 0 2 3 5 7 5 5 Springfield 53 Baca SGP/CRP 0 1 1 Two Buttes 13 Bent SGP/MGP 4 6 10 66 92 119 Prowers Prowers Baca SGP/CRP SGP SGP 0 2 2 0 0 1 0 2 3 Tinidad 36 Las Animas SGP/SLTB 3 3 6 Walsenburg' 28 Pueblo SGP/PJ 1 1 2 29 56 64 68 80M 95S 104 Pueblo Huerfano Huerfano Heurf/LasAn Huerf/LasAn Huerfano Las Animas SGP/PJ SGP/PJ SGP/yucca SGP/PJ SGP SGP SGP 1 2 1 3 4 0 6 1 4 1 7 2 3 4 2 6 2 10 6 3 10 30 11 Counties 50 57 Las Animas Pueblo Totals Sex Totals 1 2 0 1 7 109 �247 years, had more than a .60 chance of surviving through the 1995 breeding season and slightly more than .50 chance of surviving into 1996. The likelyhood for a female surviving through 2 complete reproductive seasons drops to .331. During the 97 whelping period 10 dens were located, 6 on the CPER and 4 on G1. A total of 31 pups have been observed above ground (3.1 average). Search efforts continue for more litters. Pup production in 97 is similar to that reported for 1996 and higher than 1995. Home range estimates were made for 7 foxes (4 males, 3 females) tracked intensively in the summer of 1996 and from then to either their death or the end of this reporting period. Summer home range of the females averaged 1.3 mi2 (3.41 km") compared to 1.2 mi2 (3.2 km") for the males. Average home ranges increased over time to 2.3 mi2 (6.56 km2) for females and 4.2 mi2 (11km2) for males) . In January and February of 1997 we estimated population size on a 62 mi2 expanded G1 grid using 31 automated cameras. Over 4 camera bouts, averaging 4 days per bout, we obtained 469 photographs of foxes with 147 of them (31%) marked animals. Nineteen cameras (61%,) were visited by marked animals, 28 cameras (90%) were visited by unmarked individuals. Using program NOREMARK and 95% confidence intervals our most conservative estimate was 25 foxes (range 25-42) an average of 1 fox for every 2mi2 (5.2 km2). Camera results suggest the grid-trapping system is capturing over 44% of foxes available for capture. Minimum populations on the CPER and G1 sites are probably close to double our known numbers of marked individuals. Table 4. Survival rates for 109 radio-collared swift fox in northeastern Colorado, by season, sex, and site. October 14, 1994-July 1, 1997. Survival Period Oct 94-July 97 Oct 94-July 95 Oct 95-July 96 Oct 96-July 97 Winter 94-95* Winter 95-96* Summer 95** Summer 96** 1995 1996 Male 94-97 Female 94-97 G1 95-97*** CPER 95-97*** Rate 95% CI 0.235 0.691 0.696 0.613 0.765 0.696 0.816 0.905 0.572 0.745 0.247 0.215 0.293 0.300 0.147-0.323 0.544-0.838 0.553-0.839 0.441-0.784 0.623-0.909 0.553-0.839 0.692-0.940 0.779-1.000 0.427-0.717 0.590-0.900 0.095-0.401 0.071-0.359 0.140-0.446 0.152-0.448 * - December-March ** - June-September *** - Foxes were not radio-collared on G1 until 1995 DISCUSSION The trapping results from the eastern plains survey indicate are widely distributed across the eastern plains in suitable The largest blocks of short-grass prairie and correspondingly that swift fox prairie habitat. the largest �248 numbers of fox captured were plains. Insufficient trapping crop production to determine (Fox and Roy 1995, Roy 1996) may show considerable use of lands. from counties in the southern part of the eastern was conducted in areas with high agricultural the extent of use of such habitat. Kansas studies suggest that foxes on the Colorado-Kansas border smaller mosaics of native prairie in agricultural The intensive site studies provide for comparison with other reports. Our findings of foxes in distinct clusters is similar to reports by Cameron (1984), Loy (1981), and Covell (1992) in Colorado and Jackson (1997) in Kansas. We speculate the tendency for animals on the CPER to be in larger, tighter units than on G1 is probably the result of habitat differences. Portions of the CPER contain extensive saltbush and yucca communities and its margins are in closer contact with croplands or modified grasslands (crested wheat, alfalfa, wheatfields, etc.) taller and ranker than short-grass prairie. Vegetation on the G1 site is almost exclusively unbroken short-grass prairie. These habitat differences may in turn lead to differences in concentration and production of prey (stapp 1994). Home ranges for foxes in our study sites are considerably smaller than reports (average range 12~36 km2) from other investigators (Hines 1980, Hines and Case 1991, Roy 1996, Rongstad et al. 1989) , Coyotes have been reported by Covell (1992), Carbyn et al (1994), and Fox and Roy (1995) as the major cause of death in swift foxes. Our data support those observations. Covell (1992) and Fox and Roy (1995) suggested coyote mortality may impact fox populations and Covell suggested this in turn might justify coyote control programs. We do not believe that a case for coyote control can be made in northeastern Colorado where despite considerable coyote mortality our fox population appears stable with good survivorship and relatively high reproduction. Our average annual survival for adult animals is higher than the .53 reported by Covell (1992) and the .60 reported by Rongstad et al. (1989) at Pinon Canyon. Fox and Roy (1995) reported August-February survival for swift foxes in Kansas to be .611 on cropland and .365 on rangeland in 1994. Data on pup survival is very limited we can only demonstrate 54% mortality but cannot account for any pups still surviving on the study sites. This means we have out migration of individuals or higher mortality. Covell estimated pup survival at .22 while for the same area Rongstad et al (1989) estimated pup survival at 0.05. Considerable more research is needed on pup mortality and its impact on fox populations. At present we believe that the population of swift fox on the study sites in northern Colorado is stable. We believe the conservative estimate of 25 foxes per 62 mi2 (l'fox per 5.2 km2) obtained from the camera sessions is a realistic approximation of winter populations of foxes in northern Colorado. The figure is close to the 1 fox per 6.6 km2 estimate of Covell (1992) who did not discuss method for calculation. More studies are needed in other parts of the swift fox range to estimate populations for comparison with these data. Prepared by: James P. Fitzgerald University of Northern Colorado �249 LITERATURE CITED Beck, T. D. I. 1995. Development of black bear inventory techniques. Annual Report, Colorado Division of Wildlife. Mammals Research, 11 pp. Cameron, M. W. 1984. The swift fox (Vulpes velox) on the Pawnee National Grassland: Its food habits, population dynamics, and ecology. M.A. Thesis, University of Northern Colorado, Greeley. 110 pp. Carbyn, L. N., H. J. Armbruster, and c. Mammo. 1994. Swift fox reintroduction program in Canada - 1983-1992. pp 1-95 In: M. Bowles and C. J. Whelan, eds. Restoration of endangered plants and ani.ma.l,s , Uni versi ty of Cambridge Press. Covell, D. F. 1992. Ecology of the swift fox (Vulpes velox) in southeastern Colorado. M. S. Thesis, University of Wisconsin-Madison 111 pp. Fox, L. B. and C. C. Roy. 1995. Swift fox (Vulpes velox) management research in Kansas: 1995 annual Report. pp 39-47 In: Allen, S. H., J. W. Hoagland, and E. D. Stukel, eds. 1995 Report of the Swift Fox Conservation Team. 170 pp. and Hines, T. D. 1980. An ecological study of Vulpes velox in Nebraska. M. S. Thesis. University of Nebraska, Lincoln. 103 pp~ Hines, T. D. and R. M. Case. 1991. Diet, home zanq e , movements, and activity periods of swift fox in Nebraska. Prairie Nat. 3:131-138. Jackson, V. 1997. Denning Kansas. Final Report ecology of swift foxes (Vulpes velox) in to Kansas Department of Wildlife and Parks. western Loy, R. R. 1981. An ecological investigation of the swift fox (Vulpes velox) on the Pawnee National Grasslands. M. A. Thesis, University of Northern Colorado, Greeley. 64 pp. Manly, B. F. J. 1991. Randomization and Monte Chapman and Hall, London 281 pp. Carlo methods in biology. Pollock, K. H., S. R. Winterstein, S. m. Bunck, and P. D. Curtis. 1989. Survival analysis in telemetry studies. Journal Wildlife Manage. 53:7-15. Rongstad, o. J., T. R. Laurion, and D. E. Anderson. 1989. Ecology of swift fox on the Pinon Canyon Maneuver Site, Colorado. Final Report. Directorate of Engineering and Housing, Fort Carson, Colorado. 53 pp Roy, C. C. 1996. Swift fox (Vulpes velox) management and research in Kansas. 1996. pp 1-9 In: Luce, B. and F. Lindzey, eds. 1996 Annual Report of the Swift Fox Conservation Team. 110 pp Stapp, P. T. 1996. Determinants of habitat use and community structure of rodents in northern shortgrass steppe. Ph.D. Dissertation, Colorado State University, Fort Collins. 145 pp White, G. C. 1996. NOREMARK: Population estimation Wildlife Society Bulletin 24:50-52. from mark-resighting surveys. � https://cpw.cvlcollections.org/files/original/aeee57ae5230abbea81190c9ab258721.pdf 3f2ab7fe50c07ba315552e216aa7bc5b PDF Text Text 1 Colorado Division Wildlife Research April 1998 of Wildlife Report JOB PROGRESS State of Colorado Project: W-173-R-1 Job Title: Period Author: Peregrine Covered: Gerald Peregrine Falcon Inyestigations Job __ 1__ 2 Work Plan REPORT Falcon 1 January Restoration - 31 December 1997 R. Craig Personnel: Gerald Wildlife and James R. Craig and Catherine Wightman, Colorado H. Enderson, The Colorado college. Division of ABSTRACT In the 1997 peregrine falcon (Falco peregrinus ana~um) breeding season (1 April through 30 July 1997) 87 territories were occupied by 79 breeding pairs that fledged 93 young. A sample of 45 occupied sites was monitored and averaged productivity of 1.49 young per pair. Shell fragments representing 24 eggs were collected at 11 sites and were, on average, -8.4 % thinner than preDDT era eggs. contents of 13 nonviable eggs were collected and preserved for future analysis. This Job Progress Report represents a preliminary analysis and is subject to change. For this reason, information presented herein MAY NOT BE PUBLISHED OR QUOTED without permission of the author. ��3 PEREGRINE FALCON RESTORATION Gerald PROGRAM R. Craig P. N. OBJECTIVES 1. Annually Colorado survey all documented peregrine breeding sites throughout to establish the presence of nesting peregrines. 2. Annually document monitor a statistically significant their reproductive success. 3. Annually monitor 4. Investigate 5. Evaluate, 6. Document organochlorine population of breeding pairs to levels. dynamics. characterize, and protect pesticide sample and protect important breeding migration SEGMENT habitat. and wintering areas. OBJECTIVES 1. Whole, nonviable eggs which are encountered during eyrie visits will collected, preserved and submitted to the appropriate u.S. Fish and Wildlife Service approved laboratory for pesticide analysis. 2. Compile data from previous years and prepare final report of eggshell thinning and pesticide residues in Colorado peregrine falcons from 1974 through 1997. 3. Compile and analyze data on peregrine falcon recovery dynamics and prepare manuscripts and annual report. 4. Monitor selected when funded. peregrine falcon RESULTS eyries for occupancy be and population and productivity AND DISCUSSION Survey Effort Funding and personnel limitations necessitated a reduction from 4 teams in 1996 to 2 teams comprised of 2 observers each that conducted surveys in 1997. Although the teams were assigned a sample of sites to monitor for productivity throughout the season, they also attempted to visit all known sites as well as survey potential cliffs throughout the state as time permitted. One hundred and nine sites were on record at the beginning of the field season and the teams were able to check 99 of them. Time constraints and accessibility restricted them from visiting the 10 remaining sites. Fifteen potential nest cliffs were visited, and 5 previously undocumented pairs were located. By the end of the season 114 confirmed nesting territories had been recorded. Three of the new sites were reported to the Division by other agencies. �4 Territory Occupancy Breeding territory occupancy increased from 84 sites in 1996 to 87 in 1997 (Fig.1). Seventy-eight sites were occupied by adult pairs that attempted to breed (laid eggs). Time constraints did not permit documentation of breeding at 3 sites (sites 29, 67, and 111) and mixed pairs (an adult paired with an immature) did not breed at 5 other sites (sites 12, 37, 41, 61, and 108). The rate of occupancy remained at the 80% level which is generally considered appropriate for a stable population. I!!!!! !!i!! 100 - !i!!!!! !!!i!! ~----~--------------~~~ N:I'- _.. Elil 80 ------------------liiii-;iihiim- - 70 -----------------iii·I·lliIJ~~- ~ 60 ----------------uii-lll.itliiL b 50 --------------~!l-!ill- 00 40 ----------~-~ 30 - - - - ~ - £!!"j!!'~~ _ ._ lEI iil.. ••• Eli! I~I iiiB ii lID !i!I-~!I·I!-~!~i!U! iilil ::In iiiii:U1IIIil- -727374 757677 7879 8081 8283 84 8586 87 8889 9091 9293 94 95 96 97 YEARS • Occupied Sites I~I!IIVacant Sites Fig. 1. Occupancy of Colorado peregrine nests. Reproduction Resource limits necessitated implementation and testing of a sampling protocol in 1997. For the sample to be statisically powerful, the productivity of 40 breeding pairs was required to be .monitored. Since several pairs were likely not to breed, 45 sites were initially selected for periodic surveillance. Forty-one sites were occupied by adult pairs while mixed age pairs were present at 2 sites. Although site 102 produced young, fledging could not be confirmed and the outcome was known for 42. This sample yielded an average fledged brood size of 2.07 and productivity of 1.38. In addition to the sample sites, productivity was serendipitously obtained for 28 other sites. The combined 67 sites fledged an average brood size of 2.09 with a productivity of 1.34. These comparisons suggest that the sample poductivity (1.38) was an accurate representation of the population productivity (1.34). Since the larger number of sites (67), represents a greater proportion of the total population, the 1997 productivity of 1.34 is displayed in Figure 2. The survey teams in 1997 were able to visit 99 sites, confirm presence of pairs at 87 sites and document reproductive results at 67 sites. One hundred and twenty young were produced of which 90 fledged. The increasing number of occupied territories as well as funding and personnel constraints will require sampling in the future. Although site occupancy and eggshell condition are �5 parameters that also require monitoring, annual productivity may be the most sensitive measure of Colorado peregrine population viability. Should chemical contamination reoccur, reduced fledging success will be the first manifestation of population decline. The protocol developed and tested this year appears to be a viable method for monitoring Colorado's peregrine falcons. 2.6 2.4 2.2 - .~ 2 ~ 1.8 .~ 1.6 n <2 §0.8 ~ 0.6 7\. J\ ~ 1.4 1.2 ~ 1 J I ; : I / \ \ \/ .1 I!'( /" , 0.2 o - ./ -,V/~"\"Lr, /'\. - ~ GIJ''''- /~ ).. ~ =--------- t.~~ ~ ../7 I I - I 9l / '\ ...,,_/ 74 \/ ••• '\./ -~ 73 /'-... .1 '-./ 0.4 - "" /\ I \ 75 76 77 - 78 79 "\ 80 _j 81 82 I I 83 Total Productivity 84 85 86 87 Years -c>- 88 89 Unmanipulated 90 91 92 93 94 95 96 97 Productivity Fig. 2. Productivity of peregrine nest sites, Colorado 1973-97. Eggshell Condition Eggshell fragments were collected from 11 nesting sites in the course of visits to band young. Thickness measurements taken from eggshell fragments at 11 sites and whole eggs encountered at 13 other sites averaged -8.4 % (0.329 rom) with the thickest 0.6 percent (0.361 rom) and the thinnest -20.4 % (0.284 rom). Figure 3 shows eggshell thickness trends for all years through 1997. These values are highly variable due to small sample sizes, mixing of fragments from different eggs within the same clutch, and variation of thickness of fragments from the poles of the egg versus the waist. However, shell thicknesses have averaged less than -10% from the pre DDT-era since 1992. �6 0.390 ]' - -- - ---- ---- - -- -- - ---- 0.310 ";;' 0.350 ""u j = -£i u 88 0.330 0.310 0.290 J.Q 0.210 O.250 I--_r_-r-___.__,_---.-~_.__r____.__,____r-.__r__r__.___r___r__,r_~_r__.____.~__,~ 13 14 15 16 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 I Years .f\.iaximumand Mnimum Thicknesses - Average Thicknesses Fig. 3. Cumulative eggshell thickness of peregrine eggs, Colorado, 1973-97. Organochlorine Residue in Eggs The 11 nonviable eggs collected during the 1997 season have been preserved along with eggs encountered in 1991-1994. This collection of 53 eggs has awaited pesticide analysis by the Fish and Wildlife Service when funding was available. Analysis will occur at the Colorado college's facilities during the next segment and results will be reported at that time. Release Remedial management not been undertaken Prepared By: efforts such as recycling, since 1989. C ,f2 -~ Gerald R. Craig LSSRIV and Augmentation Efforts fostering, and hacking have �7 Colorado Division Wildlife Research April 1998 of Wildlife Report JOB FINAL REPORT state of Project: Work Plan Job Title: Period Colorado W-171-R-3 2 Job 1 July G.R. Craig, and Enhancement Program 2 Bald Eagle Nest Site Protection Covered: Personnel: : Bald Eagle Nest Site Protection 1987 - 31 December Colorado Division and Enhancement Program 1997 of Wildlife ABSTRACT Bald eagles occupied 29 Colorado nesting territories in 1997. Four new territories were discovered and one was not checked to confirm occupancy. Eighteen pairs hatched young and all pairs fledged 31 young. Productivity averaged 1.07 young per occupied territory Annual breeding pair inventories and production are summarized for the period 1974-97. ��9 BALD EAGLE NEST SITE PROTECTION Gerald SEGMENT AND ENHANCEMENT PROGRAM R. Craig OBJECTIVES 1. Annually visit all documented breeding sites to determi'ne the presence of bald eagles. Pairs at territories will be documented by DWMs and other field personnel. Previously unrecorded pairs will be revealed in the course of aerial eagle and waterfowl flights. DWMs will confirm actual incubation from ground visits. 2. Occupied'territories will be visited by DWMs periodically throughout breeding season to determine hatch of young, nesting failures, etc. 3. In May and June, a utility Worker will observe distance and endeavor to follow their movements foraging areas. Responses of eagles to various uses will be recorded. 4. In June, when the young are determined to be old enough to band, sites will be visited by Craig and Knight to place a federal band on one leg and a colored, alpha numeric marker on the other. The color markers will permit identification if the young return in subsequent years. During the same nest visit the following will be recorded: Physical parameters such as tree species, height, DBH, condition, and dominance. Nest condition, size, and location. Vegetative community and land use practices. In addition, prey remains, nonviable eggs, and eggshell fragments will be collected. 6. When necessary, remedial actions will be taken to stabilize nests that are threatened by wind throw. Should the tree be decadent and in danger of falling, an artificial nest base may be placed in a suitable, adjacent tree. Action will be taken only after it has been deemed desirable to encourage the eagles to nest at the same location. RESULTS the breeding eagles from a to locate important human activities and land AND DISCUSSION Territory Occupancy Bald eagle nesting activities in Colorado continue to expand (Fig. 1, Table 1). In 1997, 29 territories were occupied of which 4 (Larimer, Adams #2 and #3, and Montezuma #5) were first time nesting efforts. The Fremont site was not checked. �10 35 30 25 20 1S 10 S 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 9S 96 97 Years - Young Produced _ Breeding Pairs Fig. 1. Colorado bald eagle breeding pairs and productivity. Reproduction The lowest reproduction since 1980 was experienced in 1997. The overall productivity was 1.07 young per occupied territory. Although 25 pairs produced eggs, only 18 pairs fledged 31 young (1.72 young per successful pair) • The previous winter, the male at the Archuleta was discovered at the nest site with a fractured leg He was rehabilitated and released at the site several months later, but had been replaced. The new male was not successful and no eggs were produced. The Grand nest site that was destroyed in 1996 was relocated late in the season after the young had fledged. Only one was observed, so the count is minimal. Subadult females occupied nests at Larimer and Adams # 3. Although the Larimer female exhibited incubation behavior, eggs were not confirmed. The Adams #3 female abandoned the nest after protracted incubation and remains of an egg were located at the base of the tree. Eleven young were banded and color marked at 6 nests (LaPlata #3, Montezuma #3, Moffat #3 and #4, and Routt #3). Fish and Wildlife Service bands were affixed to the nestlings' right legs and red alpha-numeric bands with yellow vinyl flags were affixed to their left legs. Culmen length and foot pad length measurements were obtained from eaglets that were banded. Eaglets at 5 other sites were too old to band, landowner permission could not be obtained at Rio Blanco #4, and 2 nests were too unstable to climb. Land Status The 3 new territories (Larimer, Adams #2 and #3, and Montezuma #5) were on private property and livestock grazing is the primary land use. The Montezuma #5 site is on state property and is used for recreation. Disturbance by road construction during incubation probably caused this site to fail. �11 Banded Presence 1997. of bands or markers could Adults not be confirmed Nest Stabilization on any nesting adults in Efforts The Moffat #2 site was stabilized by cabling a weaker tree stem to a more substantial main stem. No other nest stabilization efforts were undertaken in 1997. The nest constructed by the new pair at Adams #3 rapidly deteriorated after abandonment. However, due to its proximity to a publIc road, the pair was left on their own to select a site and construct a nest in 1998. Dead or dying trees remained upright and occupied at Rio Blanco #1 and #3. The Archuleta site was successfully replaced in the fall of. 1996 and a pair continued to occupy the nest. An artificial nest base was not placed at the Jefferson site and the nest remains vulnerable to wind throw. /7 Prepared By: /) 0 ," /} ~ Gerald R. Craig LSSRIV �12 Table 1. Colorado Bald Eagle Nesting Efforts - 1997 Site Age of Birds Male Female Adult Adult Adams Co. #1 Adult Adult Jefferson Co. Adams Co. #2. Adult Adult Adult Imm Adams Co. #3 Adult Adult Jackson Co. Adult Adult Morgan Co. Adult Imm Larimer Co. Adult Adult Weld Co. #3 Archuleta Co. Adult Adult late. Gunnison Co. Adult Adult LaPlata Co. # 1 Adult Adult LaPlata Co. #3 Adult Adult Mineral Co. Adult Adult Montezuma Co.#2 IA Montezuma Co#3Adult Adult Montezuma Co#5Adult Adult Grand Co. Adult Adult Mesa Co.#2 Adult Adult Moffat Co # 1 Adult Adult Moffat Co. #2 Adult Adult Moffat Co. #3 Adult Adult Moffat Co. #4 Adult Adult MoffmCo.#5 IA Rio Blanco#l Adult Adult Rio Blanco#3 Adult Adult Rio Blanc0#4 Adult Adult Rio Blanco#5 Adult Adult Adult Adult Rio Blanc0#6 Rio Blanco#7 Adult Adult Routt Co. #1 Adult Adult IA Routt Co. #2 Adult Adult Routt Co. #3 29 27 Total Total pairs: 29 Egg laying pairs: 25 Successful pairs: 18 IA = Inactive Young Produced Young Fledged o o o o o o o 2 3 2 3 o o o o 1 1 1 2 1 1 2 o o 2 2 + o 1+ I o o 2 1 2 I 1 1 1 Comments 2 eggs failed to hatch, male disappeared after 30 days. Were feeding a chick, it died, found 2 infert. eggs. Pair incubating, failed Female 4 year old. Laid I egg, infertile? Pair attended nest, no eggs produced. Pair attended nest, no eggs produced. Pair failed prior to incubation. Male injured and returned 2 eggs, abandoned eggs, failed. Hatched 1 or 2 young, failed at 4 weeks. Nest found too late in season to determine production. 1 1 1 3 2+ 3 2 + o 3 1 2 3 1 2 2 32+ 2 31 Pair relocated to new nest, discovered at fledge Pair incubated, failed. �13 Colorado Division Wildlife Research April 1998 of Wildlife Report JOB FINAL REPORT state: Project Colorado No. Job Title: Work Plan: Period Author: W-164-R Forensic _2 Covered: William Personnel: Job: Inyestigations _i__ 01 July 1996 through 28 January 1998 J. Adrian William J. Adrian ABSTRACT A forensic slide series was developed to communicate contemporary information in the field of wildlife forensics. This was accomplished by obtaining existing slides and exhibits used for teaching wildlife forensics to field personnel, producing slides, title slides and exhibits that are not currently available to forensic laboratories and field personnel and disseminating this material to all contributing forensic scientists. The scope of this project covered all laboratory forensic techniques both at the professional laboratory personnel level and the wildlife officer level. The slide series describes how to submit all types of evidence to the laboratory and covers all field forensic techniques. ��15 FORENSIC INVESTIGATIONS William J. Adrian INTRODUCTION Wildlife forensics is a rapidly changing science. Publication of research findings is a necessary part of conveying information to the scientific community, but is often ineffectual for informing field and forensic technicians, and is subject to sUbstantial time delays. This project's goal is to communicate contemporary information in the field of wildlife forensics by obtaining existing slides and exhibits used for teaching wildli~e forensics to field and laboratory personnel, producing slides, title slides and exhibits that are not currently available to forensic laboratories and field personnel, then disseminating this material to all contributing forensic scientists. The scope of this project will cover all laboratory forensic techniques both at the professional laboratory personnel level and the wildlife officer level. The slides series will describe how to submit all types of evidence to the laboratory, and covers all field forensic techniques. P. N. OBJECTIVE To produce slides and exhibits to inform professional personnel of current forensic techniques, and provide instructional material for teaching field laboratory staff. and RESULTS The completed slide series will allow wildlife field and laboratory forensic personnel to learn about and exchange wildlife forensics techniques in a timely manner, and assist them in the use of these techniques. Additionally, it will help others in designing or developing similar techniques. OVerall, these benefits will result in more efficient use of time and resources. A complete slide series is available through the Colorado Division of Wildife Laboratory in Fort Collins, Colorado. Prepared by ��17 JOB PROGRESS State of: Colorado Migratory Project: W-166-R Work Plan: __l_: Job_22_ Job Title: Period Author: REPORT Monitor Covered: Michael Banding 01 January of Mallards through Bird Inyestigations in Colorado 31 December 1997 R. Szymczak Personnel: J. Broderick, R. Caskey, P. Creeden, R. Del Piccolo, J. Ellenberger, V. Graham, J. Gray, J. Gumber, T. Mathieson, J. Miller, M. Szymczak, and S. Yamashita, Colorado Division of Wildlife. ABSTRACT Ducks were wetland location September 1997. were banded; 362 trapped in modified Salt Plains bait traps and banded at 1 near Grand Junction in western Colorado in August and Three hundred and eighty-two mallards (Anas platyrhnchos) near Grand Junction and 20 near Yampa. ��19 PRESEASON MONITOR BANDING OF MALLARDS IN COLORADO INTRODUCTION In 1990, the Pacific Flyway Study Committee formulated a 5-year cooperative mallard and northern pintail (Anas acuta) preseason banding program that was endorsed by the Pacific Flyway Council. This program was designed to address banding needs throughout the western u. S., including Alaska, and in the provinces of British Columbia and Alberta. Through the first 5 years, about 9,000 ducks were banded in Colorado under this program. Following the 5th year of banding, an analysis of recoveries of all mallards banded during the 5-year period in the western u. S. and Canada showed that cohorts of mallards banded in southeastern Idaho, western Wyoming, northern utah, and western Colorado had similar recovery distribution properties. Since trapping and banding efforts in the 4-state region had been most successful in southeast Idaho and western Colorado, those 2 areas were selected for continued banding in relation to the Pacific Flyway Council efforts to establish a western mallard management unit. Banding activities in 1997 marked the second year of monitor banding activities. P. N. Objective Band mallards in Colorado's portion of Banding Reference Areas in the Pacific Flyway that will contribute information on harvest rates, survival rates, and distribution of harvest for use in Adaptive Harvest Management of western mallard populations. SEGMENT 1. OBJECTIVES Trap and band mallards in the Grand Junction/Delta/Olathe area in late August-early September using salt plains bait traps (Szymczak and Corey 1976). Banded ducks will be classified according to age and sex using accepted techniques (Carney 1964, Weller 1976: 35). Banding schedules and recapture reports will be submitted to the u. S. Fish and Wildlife Services' Bird Banding Laboratory. Band return reports will be summarized and remain on file with the Colorado Division of Wildlife. METHODS Trap Area Selection The selection of wetland locations for continued banding were based on number of birds banded per unit of effort during 1991 through 1995, and on the availability of personnel to operate the banding stations. The Walker State Wildlife Area (SWA) near Grand Junction and Markley's Pond near Olathe were selected sites. Trapping Ducks were trapped and banded near Grand Junction from 28 August through 6 September 1997. All birds were trapped in modified Salt Plains bait traps (Szymczak and Corey 1976) using whole shelled corn for bait. Traps were visited daily. Mallards were the target species. Banded birds were recorded by wetland site. Band numbers of all birds captured that were banded in previous years or outside the specific area of trapping were recorded. �20 RESULTS Trapping, Banding and Record Keeping Trapping occurred only on the Walker SWA near Grand Junction (Table 1). A trapping crew was not assembled to trap in the Uncomphragre River Valley. At Walker, 3 riverine sites were selected for trap sites. A t.ot.a L of 369 mallards was banded during trapping in western Colorado in 1997 (Table 1) with a reduced trapping effort compared to the previous 5 years. Immatures comprised 64 % of the sample. Females comprised 30% of the adult sample. Band Reporting and Record Keeping All band numbers of newly banded birds and recaptures were submitted to the U. S. Fish and Wildlife Service's Bird Banding Laboratory on standard forms. Computer files containing the number of birds banded by area, site, day, age and sex were constructed at the Colorado Division of Wildlife's Research Center. Table 1. Number of Mallards banded by age and sex during at the Walker SWA in western Colorado 1996-97. pre-season trapping Age/Sex Site Walker SWA Year AM AF 1M IF 1996 68 18 143 133 362 1997 93 40 120 116 369 LITERATURE Totals CITED Carney, S. M. 1964. preliminary keys to waterfowl age and sex identification by means of wing plumage. U. S. Dep. Inter., Fish and Wildl. Servo Spec. Sci. Rep.- Wildl. 82. 47pp. Szymczak, M.R., and J. F. Corey. 1976. Construction and use of the Salt Plains duck trap in Colorado. Colorado. Div. Wildl., Div. Rep. 6. 13pp. Weller, M. W. 1976. Molts and plumages of waterfowl. Pages 34-38 in F. C. Bellrose. Ducks, geese and swans of North America. Stackpole Books, Harrisburg, PA. Prepared by: �21 Colorado Division wildlife Research April 1998 of Wildlife Report JOB PROGRESS REPORT State of Colorado Project Work W-166-R __l__: Plan Job Title: Period Author: Migratory Covered: Inyestigations 24 Job Integrated Game Bird Waterbird 1 January Management through Studies 31 December 1997 James H. Gammonley Personnel: J. Gammonley, M. Szymczak, Colorado Laubhan, USGS Biological Resources Division Division of Wildlife; M. ABSTRACT Laboratory and data entry associated with field data collected during 1994-96 continued. Dissections and carcass nutrient analyses were performed on waterbirds collected at Russell Lakes State Wildlife Area (RLSWA) in the San Luis Valley (SLV), Colorado. Time-activity budget data and nesting were entered into computer files and checked for accuracy. Sites were chosen at RLSWA for habitat manipulations to determine if habitats and waterbirds respond as predicted. Summaries of results were prepared for incorporation into a revised management plan for RLSWA and management plans for other state wildlife areas in the SLV. A study plan was developed to monitor habitat conditions and waterbird use of wetland areas statewide in Colorado using methods for habitat measurements similar to those implemented in this study. In 1998, laboratory work will be completed and data analyses and manuscript preparation will continue. Results will be incorporated into management plans at RLSWA and other state wildlife areas in the SLV. ��23 INTEGRATED WATERBIRD MANAGEMENT STUDIES INTRODUCTION The San Luis Valley (SLV) is one of the most important breeding areas for waterbirds in Colorado (Ryder et al. 1979, CDOW 1989, Nelson and Carter 1990, Gilbert et al. 1996). A wetland ecosystem can be managed for habitats that maximize requirements for a narrow group of avian species, or for more diverse habitats that optimize resources for a variety of avian species. The latter approach, which embodies a philosophy of integrated waterbird management, better fits the philosophy of increased emphasis on managing landscapes for species diversity. One goal of the Colorado State Waterfowl Management Plan is to provide habitat of sufficient quality to maintain duck and goose populations at desired levels for maximum recreational opportunities (CDOW 1989). In addition, the SLV draft management plan for waterbirds recommends maintenance of diverse wetland habitats with 25% of the actively managed habitat on public lands managed for nongame waterbirds (Olterman 1995). In 1994, the CDOW initiated a study to examine resource use by both game and nongame breeding waterbird species on Russell Lakes state Wildlife Area (RLSWA) in the northwestern portion of the SLV. During 1994-96, foraging and nesting ecology of breeding mallards (Anas platyrhynchos), gadwalls (A. strepera), cinnamon teal (A. cyanoptera), redheads (Aythya americana), American avocets (Recurvirostra americana), killdeers (Charadrius vociferus), and Wilson's phalaropes (Phalaropus tricolor) were intensively studied at RLSWA. In 1997, efforts were initiated to manipulate habitats (hydrology, vegetation structure) and measure waterbird responses (foraging activity, nesting) on specific areas at RLSWA. P. N. OBJECTIVES 1. Map the location of wetlands State Wildlife Area. and wetland communities on Russell Lakes 2. Document the hydrologic regime and water, characteristics of each wetland type. 3. Identify the aquatic invertebrates associated with each wetland community, and document seasonal trends in invertebrate diversity, abundance and biomass. 4. Quantify the abundance, spatial and temporal use patterns, behaviors, and food habits of waterbirds in different wetland types. Relate the dynamics of endogenous lipid and protein reserves to food habits and migration and breeding ecology. 5. Determine the seasonal wetland habitat requirements for waterbirds, and consolidate these needs into a conceptual design for an optimum wetland community. 6. Determine the water management protocol and wetland development guidelines needed to produce the optimum wetland community. Prepare a wetland development and water management plan for Russell Lakes State Wildlife Area. soil and vegetation �24 SEGMENT OBJECTIVES 1. Conduct proximate analyses on carcass samples of mallards, gadwalls, cinnamon teal, redheads, American avocets, killdeer, and Wilson's phalaropes collected at Russell Lakes State Wildlife Area in 1995 and 1996, to determine the amount of nutrient reserves in each collected bird. 2. Determine food selection of waterbirds collected in 1, i.e., compare food use (gut contents) to availability (invetebrate and seed biomass at collection sites) for each species, with age, sex, reproductive status, molt intensity, size of nutrient reserves, and collection date as covariables. 3. Analyze time-activity budget and abundance data collected during 1994-96 for waterbird species in 1. Determine foraging habitat selection, and the influence of habitat features (cover type, water depth, vegetation structure) on food availability and use of foraging habitat. Model habitat management options that maximize foraging habitat for each species and optimize foraging habitat for all species. 4. Analyze nest habitat and nest success data collected during 1995-96 for each waterbird species listed in 1. Determine nest habitat selection, and the influence of habitat features (cover type, water depth, vegetation structure) and spatial attributes (distance to water; distance to roads, ditches, etc.; cover type patch size) on nest success. Model habitat management options that maximize nesting habitat for each species and optimize nesting habitat for all species. 5. Manipulate water and vegetation on selected sites at RLSWA in spring 1997. Following habitat manipulations, measure a) nest locations and nest success, b) locations used by foraging waterbirds, and c) invertebrate and seed biomass to see if patterns are as predicted in 2 and 4. 6. Select sites at other wetland complexes in Colorado that will be used starting in 1998 to measure water depth, conductivity, and vegetation structure (height and density) at random sites and at nest and feeding sites used by species listed in 1, to see if habitat selection patterns are similar to those observed at RLSWA. Develop a study plan for on-going monitoring and evaluation of wetland projects managed for waterbirds in Colorado. RESULTS Waterbird Carcass Analyses We continued dissections and proximate analyses of ,waterbird carcasses collected in 1994-96. Dissections have been completed on 216 of the 272 waterbirds collected. Nutrient analyses have been completed on 159 birds. Food Selection Preliminary results of food habits and food availability have been presented in previous reports (Gammonley 1995, 1996). Because age, sex, reproductive status, molt intensity, and size of nutrient reserves will be used as co-variables in the analyses of food selection by each species, final analyses will not be completed until all dissections and nutrient analyses are completed. �25 Foraging Habitat Selection We began transforming time-activity budget data, entering the data in a database suitable for statistical analysis, and checking the data. During 1994-96, over 1,550 individual, 10-min time-activity bouts were collected on the 7 waterbird species we studied at RLSWA. Data entry has been completed on 511 bouts. We recorded habitat variables (water depth, conductivity, vegetation structure, cover type) at 338 sites where we collected timeactivity data on birds that were foraging. This habitat data has been entered in a database and will be merged with the time-activity data when that database is completed. Habitat availability data from 330 random plots on RLSWA has been entered on computer files and checked, and a habitat map of the area has been completed (Gammonley 1995, 1996, 1997). Nest Habitat Selection and Nest Success Data on nest site habitat attributes and success of nests have been entered into computer files and checked for accuracy. Preliminary attempts have been made to use geographical information system software (ARC-INFO) to calculate spatial attributes of individual nest locations (e.g., distance to other cover types, distances to other nests). No additional work was done in 1997 on analyses of nesting data. Habitat Manipulations at RLSWA In 1997, we selected 3 sites for water level and vegetation management. One site was dominated by short emergent vegetation apparently preferred by nesting cinnamon teal, but this site had no use by nesting cinnamon teal during 1994-96. Based on our preliminary results, we speculated that maintaining shallow «5 cm) water at this site throughout the nesting period would attract nesting cinnamon teal. Water levels were maintained at desired levels, and 5 cinnamon teal nests, along with 1 mallard nest and 2 phalarope nests, were found in 1997. Vegetation structure also appeared to begin changing in 1997, and habitat measurements will be taken at this site in 1998. The second site was a short emergent site that in the past had been dominated by baltic rush (Juncus balticus), but had been dry for several years. Water control structures were placed along a ditch that will provide water to the site. The site will be seasonally flooded annually beginning in spring 1998, and vegetation response will be monitored. The third site is a large impoundment where relatively high duck nest densities were found during 1995-96. Preliminary data indicate that many ducks at RLSWA prefer to nest in dense cover over several cm of water. Our goal was to maintain vegetation structure in this impoundment, but de-water the impoundment, and compare subsequent nest densities to previous years. However, this large impoundment proved to have inadequate drawdown capabilities, and water levels were not reduced over a large proportion of the area until after the peak nesting period of most duck species at RLSWA. Three additional sites were selected for habitat management and monitoring beginning in 1998. Habitat Monitoring at other Wetland Sites in Colorado A pilot proposal was developed to monitor habitat conditions and bird use on wetland sites developed to benefit waterbirds throughout Colorado (Appendix A). Because the waterbird species studied at RLSWA have wide distributions in �26 Colorado, they characteirtics should allow for informative comparisons of habitat and habitat use patterns among wetland areas in the state. PLANS FOR 1998 Laboratory analyses of waterbird carcasses will be completed. Statistical analyses of all data sets will continue. Manuscripts will be prepared for publication in peer-reviewed technical journals. Results will be incorporated into a revised management plan for RLSWA, as well as management plans for other SWAs in the San Luis Valley. A new study to monitor waterbird habitats and use of habitats on wetland areas statewide will be initiated. LITERATURE Colorado Division of Wildlife. plan 1989 - 2003. Colorado 97pp. Gammonley, J. H. 1995. Division of Wildlife CITED 1989. Colorado statewide waterfowl management Division of Wildlife, Unpublished Report. Integrated waterbird Federal Aid Progress management Report. Gammonley, J. H. 1996. Integrated waterbird management Division of Wildlife Federal Aid Progress Report. Gammonley, J. H. 1997. Division of wildlife Integrated waterbird Federal Aid Progress management Report. studies. Colorado studies.Colorado studies.Colorado Gilbert, D. W., D. R. Anderson, J. K. Ringelman, and M. R. Szymczak. Response of nesting ducks to habitat and management on the Monte National Wildlife Refuge. Wildllife Monographs 131. 44pp. 1996. Vista Nelson, D. L., and M. F. Carter. 1990. Birds of selected wetland of the San Luis Valley. Colorado Division of Wildlife, Unpublished Report. 40pp. Olterman, J., ed. 1995. Division of Wildlife, The San Luis Valley Unpublished Report. waterbird 39pp. plan. Colorado Ryder, R. A., W. D. Graul, and G. C. Miller. 1979. Status, distribution, movements of ciconiforms in Colorado. proceedings Colonial Waterbird Group Conference 3:49-58. Preparedby~~~ James H. Gammonley LSSRm and �27 APPENDIXA. Study Title: A. Monitoring and Eyaluation of Wetland Deyelopment Projects in Colorado NEED: It is estimated that half of Colorado's wetlands have been destroyed in the last 100 years (Dahl and Johnson 1991). Many other wetlands in the state have been degraded or their functions have been greatly modified, particularly palustrine habitats with intermittently flooded, temporarily flooded, and seasonally flooded hydroperiods (Cowardin et al. 1979). These highly productive wetlands provide important habitats for waterfowl and other wetland-dependent wildlife (Fredrickson and Taylor 1982, Eldridge 1992, Mitsch and Gosselink 1986). In recent years, wetland conservation efforts have increased in Colorado, led by the Colorado Division of Wildlife (CDOW), Ducks Unlimited Inc. (DV), and the Partners for Wildlife (PFW) program of the U.S. Fish and Wildlfe Service (Chappell 1997). To date, more than 200 wetland conservation projects have been completed by these organizations in Colorado. The focus of most of these projects has been to protect, restore, enhance, or create habitat for waterfowl and other wetlanddependent birds (e.g., Colorado Division of Wildlife 1989, Colorado Division of Wildlife et al. 1996). Portions of Colorado are included in the Intermountain West Joint Venture (IWJV) and Playa Lakes Joint Venture (PLJV) of the North American Waterfowl Management Plan (NAWMP). Within the IWN, seven Focus Areas were established west of the continental divide where most IWN conservation efforts in Colorado would take place (Yampa/White River, Lower Colorado River, Gunnison River, North Park, Middle Park, South Park, and San Luis Valley). In 1995, CDOW created three additional Focus Areas in eastern Colorado (playa Lakes/Arkansas River, South Platte River, and Front Range) to facilitate wetland conservation efforts on a statewide basis. These ten Focus Areas collectively form the infrastructure for CDOW's Wetlands Initiative, a partnership between CDOW, DU, PFW, The Nature Conservancy, and the Colorado Department of Parks to spend approximately $10 million over the next three years on wetland conservation projects in Colorado (Chappell 1997). DU also has an ambitious Colorado Conservation Plan (Ducks Unlimited 1997) that focuses on conserving waterfowl habitat in the San Luis Valley, North Park, and South Platte River Focus Areas. A goal of all wetland conservation projects is to provide protection from further threats to wetland ecological functions. Naturally functioning wetland systems, requiring protection only, are relatively rare in Colorado, except at the highest elevations. Rather, natural or historic functions at most existing wetland sites have been lost or degraded. Consequently, many conservation projects have included structural developments, such as levees and water control structures, that provide greater hydrological control, allowing managers to more reliably restore and maintain historic wetland conditions (i.e., restoration projects). Usually, the broad goal of a wetland development project is to improve capabilities to emulate historic, natural wetland functions at the site, including hydrologic regimes (Fredrickson and Taylor 1982, Fredrickson and Reid 1990, Galatowitsch and Van der Valk 1996). Alternatively, developments can be used to produce wetland habitats at sites where they did not historically occur (i.e., creation projects), or to alter existing wetlands to produce greater benefits for specific wetland functions, such as providing habitat for wetland-dependent wildlife (i.e., enhancement projects) (Fredrickson and Reid 1986, Weller 1990). Many procedures have been developed to assess wetland functions (Harris 1988, Adamus and Brandt 1990, D' Avanzo 1990, Erwin 1990, Marble 1990, Kentula et al. 1992, Wodd Wildlife Fund 1992, Brinson 1993). Most of these approaches have focused on mitigation projects, but the assessment criteria developed for these programs can apply to functional assessment of wetland developments designed to improve waterbird habitat (LaGrange and Dinsmore 1989, Ringelman 1991, Weller et al. 1991). Here, wetland restoration, creation, and enhancement projects are collectively referred to as wetland developments. Despite the large number of wetland developments that have been constructed in Colorado, there has been no effort to track these projects and evaluate their success. Few records exist �28 of exactly what developments were constructed at each site. Specific management plans are lacking for most developed wetlands, and many sites are essentially unmanaged, even when management capabilities exist. Past wetland assessments in Colorado (U.S. Fish and Wildlife Service 1955, Hopper 1968) focused on key waterfowl breeding and hunting areas and therefore did not have statewide coverage, and did not examine outcomes of specific habitat management practices. A standardized monitoring program would provide important benefits, including to 1) provide a basis to evaluate the success of completed projects, 2) identify problem areas in wetland management in Colorado, 3) allow for learning about the effectiveness of specific wetland management techniques in providing resources for waterbirds, and 4) facilitate tracking of overall wetland resources in Colorado (Marble 1990, Kentula et al. 1992). The purpose of this project is to develop and implement a statewide monitoring program for wetland development projects in Colorado, assess the current status of wetland developments in various regions (Focus Areas) across the state, and develop guidelines for the selection, design, and management of future wetland development projects for waterbirds. B. OBJECfIVES: 1. To develop a standardized monitoring program for tracking the structural, biological, landscape, and management status of wetland development sites in Colorado over time. 2. To assess the current ecological functions of wetland development sites at individual project, regional, and statewide scales. 3. To develop regional (Focus Area) recommendations for the design and management of wetland development projects to provide habitats for waterbirds. C. EXPECfED RESULTS OR BENEFITS: This project will produce an initial assessment of existing wetland developments throughout Colorado, and provide the infrastructure for future periodic assessments of ecological functions of wetland developments on a regional or statewide. The project will result in guidelines for the design and management of wetland developments for waterbirds in different regions (Focus Areas), and recommendations for enhancing or rehabilitating existing wetland developments. Results will be compiled into reports, handbooks, and technical articles that will be available to wetland managers, landowners, and policy-makers. D. APPROACH: 1. Available information will be gathered for all past wetland development projects in Colorado that were funded by CDOW, DU, and/or PFW. All information will be entered into a computer database. Information fields will include Focus Areaaffiliation, specific location, date of project completion, wetland acres involved, upland acres involved, project costs, and party with primary management responsibility at the site. When specific biological objectives can be identified for a project (e.g., enhancing duck nesting cover), these objectives will be included in the database. Because most past projects were directed at providing habitat for waterfowl and other wetlanddependent birds, waterbirds will be a focus of the biological assessment of projects (see below). Projects will be stratified by Focus Area (10 strata) and age (completed in 1997 or later, 1994-96, or before 1994), and a random sample of projects from each stratum will be selected for monitoring as "reference wetlands" (Brinson and Rheinhardt 1996). Inclusion of randomly selected projects located on private land will be contingent on an oral or written agreement with the landowners (Fellows and BuhlI995). 2. Each selected project will be visited for an initial site assessment. Structural, biological, landscape, and management assessments will be made. Structural components will include identification and status of the water source; water availability; and condition and placement of levees, ditches, water control structures, spillways, fences, nesting islands and structures, and other constructed features at the development site. When possible, original engineering plans will be obtained for comparison �29 and interpretation of current conditions. Biological components will include measurements of the following habitat features: species composition, structure, and coverage of vegetation (Robel et al. 1970); water depth (mean, maximum,and range within a basin); hydroperiod (frequency and duration of flooding); water salinity and pH; and soil types (obtained from soil survey maps), salinity, and pH (Faulkner et al. 1989, Soil Conservation Service 1991, Natural Resources Conservation Service 1995). Habitat features will be compared to known habitat requirements and preferences of wetland-dependent bird species that occur in each Focus Area, and use of selected projects by waterbirds will also be quantified, consistent with the objectives of the wetland development project (e.g., nest success [Johnson 1979], estimates of abundance [Buckland et al. 1993], time-activity budgets [Altmann 1974 D. Landscape components will include status of surrounding land ownership and land use, public or private use of the site, wetland size and type, and distance to other basins of different wetland types (Kentula et al. 1992). The management component of the assessment will include an interview with the party responsible for management of the project site to determine the management history of the site (e.g., flooding cycle, grazing rotation, repairs to levees). If a management plan exists for a project, it will be included in the management assessment. All information from the initial site assessment will be consolidated into a site report for each selected project. Photo stations will be established at each site and a slide catalog of selected projects will be produced. 3. Recommended management practices will be developed for a sub-sample of projects in each Focus Area. Recommendations will be focused on water management (frequency, duration, and depth of flooding; management of salinity levels), and vegetation management (control of exotics; manipulation of vegetation structure and composition through water management, burning, grazing, or mechanical manipulation). Specific waterbird responses (e.g., nest success, foraging activity, species diversity) will be measured as appropriate on each of these projects. Additional management practices may include upland vegetation management and control of public use. Management practices will be implemented and projects will be re-assessed at least once each year to determine the success of management practices. 4. A wetland development manual will be prepared, describing wetland characteristics, project design considerations, and recommended management and monitoring practices in each Focus Area. The report will include information on plant species responses to different management scenarios (e.g., drawdown dates, flooding depths), conservation and management concerns specific to each Focus Area (e.g., biologically critical and threatened wetland types, water quality issues, noxious weeds of concern), and summaries of waterbird use patterns on different wetland types in each Focus Area. Assessments of project components will be used to identify functional strengths and weaknesses of individual developments (Marble 1990, Kentula et al. 1992, Brinson 1993) for groups of birds that depend upon palustrine wetlands in Colorado, including dabbling ducks, diving ducks, shorebirds, rails, large waders (herons, egrets, bitterns, and ibis), and marsh-nesting passerines (Andrews and Righter 1992). �30 E. lIME SCHEDULE: fim: Al2l2rQach 1-3 1-3 1-4 3-4 1998-99 1999-2000 2000-01 2001-02 F. PERSONNEL ASSIGNMENTS: 1998-99 James H. Garnmonley Michael R Szymczak 1999-2000 James H. Gammonley Michael R Szymczak 2000-01 James H. Garnmonley Michael R Szymczak 2001-02 James H. Garnmonley Michael R Szymczak G. MQnths 4 4 4 4 4 4 4 4 ESTIMATED COSTS: 1998-99 1999-2000 2000-01 2001-02 H. SUPERVISION AND COOPERATION: Project Leader: Principal Investigator: Cooperation: I. $77,012 $77,012 $77,012 $77,012 Clait E. Braun James H. Garnmonley Ducks Unlimited Inc., U.S. Fish and Wildlife Service, USGS Biological Resources Division, private landowners. LOCATION OF WORK Statewide in Colorado, with individual projects selected on public and private lands within each of the 10 wetland Focus Areas. Data management, analyses, and writing will be conducted at the CDOW Wildlife Research Center, Fort Collins. J. RELATED FEDERAL AID STUDIES: Colorado W-166-R, Work Plan 31, Job 1: Integrated waterbird management studies; Work Plan 10, Job 1: Cooperative management programs. �31 LITERATURE OTED Adamus, P. R, and K. Brandt. 1990. Impacts on quality of inland wetlands of the United States: a survey of indicators, techniques, and applications of community level biomonitoring data. Environmental Protection Agency, EPA 600/3-90/073, Cincinnati, Ohio, USA. Altmann.J. 1974. Observational study of behaviour: sampling methods. Behaviour 49:227-267. Andrews, R, and R Righter. 1992. Colorado birds: a reference to their distribution and habitat. Denver Museum of Natural History. 442pp. Brinson, M. M. 1993. Changes in the functioning of wetlands along environmental gradients. Wetlands 13:65-74. ~ and R Rheinhardt. 1996. The role of reference wetlands in functional assessment and mitigation. Ecological Applications 6:69-76. Buckland, S. T., D. R Anderson, K. P. Burnham, and 1. L. Laake. 1993. Distance sampling: estimating abundance of biological populations. Chapman and Hall, London, UK. Chappell, A. 1997. The Colorado Division of Wildlife wetlands program. Unpublished report, Colorado Division of Wildlife, Denver. Colorado Division of Wildlife. 1989. Colorado statewide waterfowl management plan 1989-2003. Colorado Division of Wildlife, unpublished report. 98pp. __ -" U.S. Fish and Wildlife Service, and U.S. Bureau of Land Management. 1996. San Luis Valley waterbird plan. Unpublished report. 47pp. Cowardin, L. M., V. Carter, F. C. Golet, and E. T. LaRoe. 1979. Classification of wetlands and deepwater habitats of the United States. FWS/OBS-79/31, U.S. Fish and Wildlife Service, Washington, D.C. 131pp. Dahl, T. E., and C. E. Johnson. 1991. Status and trends of wetlands in the conterminous United States, mid1970's to mid-1980's. U.S. Department of Interior, Fish and Wildlife Service, Washington, D.C. 28pp. D' Avanzo, C. 1990. Long-term evaluation of wetland creation projects. Pages 487-496 in J. A. Kusler and M. E. Kentula, editors. Wetland creation and restoration: the status of the science. Island Press, Washington, D.C. Ducks Unlimited. 1997. Colorado conservation plan. Ducks Unlimited, Inc. Unpublished report. 18pp. Eldridge,1. 1992. Management of habitat for breeding and migrating shorebirds in the midwest. U.S. Fish and Wildlife Service, Fish and Wildlife Leaflet 13.2.14. 6pp. Erwin, K. L. 1990. Wetland evaluation for restoration and creation Pages 429-449 in J. A. Kusler and M. E. Kentula, editors. Wetland creation and restoration: the status of the science. Island Press, Washington, D.C. Faulkner, S. P., W. H. Patrick, Jr., and R P. Gambrell. 1989. Field techniques for measuring wetland soil parameters. Soil Science Society of America Journal 53:883-890. Fellows, D. P., and T. K. Buhl. 1995. Research access to privately owned wetland basins in the prairie pothole region of the United States. Wetlands 15:330-335. Fredrickson, L. H., and F. A. Reid. 1986. Wetland and riparian habitats: a nongame management overview. Pages 59-96 in 1. B. Hale, L. B. Best, and R L. Clawson, editors. Management of nongame wildlife in the midwest: a developing art. North Central Section of The Wildlife Society. __ and __ . 1990. Impacts of hydrologic alteration on management of freshwater wetlands. Pages 7190 in 1. M. Sweeney, ed. Management of dynamic ecosystems. North Central Section of The Wildlife Society, West Lafayette, Indiana, USA. __ '_ and T. S. Taylor. 1982. Management of seasonally flooded impoundments for wildlife. U.S. Fish and Wildlife Service Resource Publication 148. 29pp. Galatowitsch, S. M., and A. G. An der Valko 1996. Characteristics of recently restored wetlands in the prairie pothole region. Wetlands 16:75-83. Harris, L. D. 1988. The nature of cumulative impacts on biotic diversity of wetland vertebrates. Environmental Management 12:675-693. Hopper, R M. 1968. Wetlands of Colorado. Colorado Division of Wildlife Technical Publication No. 22. 89pp. u.s. �32 Johnson, D. H. 1979. Estimating nest success: the Mayfield method and an alternative. Auk 96:651-661. Kentula, M. E., R P. Brooks, S. E. Gwin, C. C. Holland, A. D. Sherman, and J C. Sifueos. 1992. An approach to improved decision making in wetland restoration and creation. Edited by A. J. Hairston. U.S. Environmental Protection Agency, Environmental Research Laboratory, Corvalis, 'Oregon, USA. 151pp. LaGrange, T. G., and J J Dinsmore. 1989. Plant and animal community responses to restored Iowa wetlands. Prairie Naturalist 21 :39-48. Marble, A. D. 1990. A guide to wetland functional design. Lewis Publishers, Boca Raton, Florida, USA. 222pp. Mitsch, W. L, and 1G. Gosselink. 1986. Wetlands. Van Nostrand Reinhold, New York, USA. 539pp. Natural Resources Conservation Service. 1995. Field indicators of hydric soils in the United States. Version 2.0. U.S. Government Printing Office, Washington D.C. Ringelman, J. K. 1991. Evaluating and managing waterfowl habitat. Colorado Division of Wildlife Report No. 16. 46pp. Robel, R J, J N. Briggs, A. D. Dayton, and L. C. Hulbert. 1970. Relationships between visual obstruction measurements and weight of grassland vegetation. Journal of Range Management 23:295-297. Soil Conservation Service. 1991. Hydric soils of the United States 1991. U.S. Department of Agriculture Soil Conservation Service, U.S. Government Printing Office, Washington, D.C. U.S. Fish and Wildlife Service. 1955. Wetlands inventory: Colorado. U.S. Departrment of Interior, Office of River Basin Studies, Albuquerque, New Mexico, USA. 33pp. Weller, M. W. 1990. Wetland management techniques for wetland enhancement, restoration and creation useful in mitigation procedures. Pages 517-528 in J A. Kusler and M. E. Kentula, editors. Wetland creation and restoration: the status of the science. Island Press, Washington, D.C. ___, G. W. Kaufmann, and P. A. Vohs. 1991. Evaluation of wetland development and waterbird response at Elk Creek Wildlife Management Area, Lake Mills, Iowa, 1961-1990. Wetlands 11:245-262. World Wildlife Fund. 1992. Statewide wetlands strategies. Island Press, Washington, D.C. 268pp. �33 Colorado Division Wildlife Research April 1998 of Wildlife Report JOB PROGRESS State of: Colorado Project: Work W-166-R Plan: Cooperative Covered: Authors: Migratory Bird Inyestigations __lQ_: Job_l_ Job Title: Period REPORT James Management 01 January H •.Gammonley through Programs 31 December and Michael 1997 R. Szymczak Personnel: Matthew Reddy and Robert Sanders, Ducks Unlimited, Inc.; William Noonan, U. S. Fish and Wildlife Service; and James H. Gammonley, Alex Chappell, and Michael R. Szymczak, Colorado Division of Wildlife. ABSTRACT Recommendations for wetland habitat improvements and/or management were provided for public and private land managers throughout Colorado. Proposals for funding projects with Duck Stamp funds were evaluated and rated. Presentations on wetland ecology in relation to waterfowl were given at workshops. Work with the Wetland Focus Area committees continued in relation to wetland project development proposals for the Great Outdoors Colorado Wetland Legacy grant, the Intermountain West Joint Venture, and the Playa Lakes Joint Venture. Responsibilities as Colorado's representative on Pacific Flyway study Committee and Council and Central Flyway Technical Committee, including Pacific Flyway Consultant to the U. S. Fish and Wildlife Service's Regulation Committee were fulfilled. No post~breeding Canaqa goose (Branta canadensis) banding was conducted during this segment. . ��35 COOPERATIVE MIGRATORY BIRD MANAGEMENT PROGRAMS In 1988, the Colorado Division of Wildlife (CDOW) created the Migratory Game Bird Program unit (~BPU) within the Terrestrial Wildlife Section. This administrative change combined all individuals having statewide responsibilities for research and management of migratory game birds. Members of the MBPU worked in concert to improve migratory bird management in Colorado. This job was created to allow team members to participate in these management programs. In November 1993, project personnel assumed additional responsibility for leading and administering the Duck Stamp wetland development program. Since 1993, personnel of the MBPU have taken on additional responsibilities with wetland programs in Colorado. In July 1996, the MBPU was dissolved with most of the responsibilities being transferred to the Avian Program. This report covers activities of the migratory bird segment of the Avian Program. P. N. OBJECTIVES 1. Continue to aid the formation of Wetland Focus Area committees in the state, monitor the functioning of these committees, and aid in obtaining funds for proposed projects by serving as chairman of the state-wide oversight committee and state coordinator for the Intermountain West Joint venture. 2. Advise public land management agency personnel, including area biologists and district wildlife managers, on the potential benefits to migratory birds of acquisition and/or development of wetland areas. Activities may include on-site inspection, formulating plans for the collection of biological data, recommending wetland enhancement developments, or review of development plans. Provide information on wetland habitat development to private land managers. 3. Prepare and present information programs on the principles of migratory bird management. Preparation may include literature review, construction of charts, graphs, and tables as photographic slides or posters, and rehearsal of presentations. 4. Attend flyway Technical Committee and Council meetings and flyway special workshops as assigned. Compile information on current status of Colorado's migratory bird population and hunting season results through consultation with CDOW biologists and managers. Prepare reports for presentation at meetings and workshops. Serve on committees as assigned by flyway Technical Committee and/or Council Chairman. Attend seiected meetings in Colorado that address migratory bird management programs, at which in-depth biological expertise would be of value. 5. Provide methodology to migratory birds and wetland managers for sampling the biological parameters of interest. Literature review may be required to develop appropriate methodology. Parameters of interest may include breeding pairs, nesting densities, nesting success, fledging success, vegetation composition and density, invertebrate composition and density, or population survival. 6. Assist in collecting information that will enable waterfowl and wetland managers to make decisions on population and wetland management. �36 SEGMENT 1. OBJECTIVES Use the knowledge and skills of the federal aid supported members of the Migratory Bird Unit to facilitate wetland and waterfowl management and informational programs within Colorado. a. Continue to aid the formation of Wetland Focus Area committees in the state, monitor the functioning of these committees, and aid in the design and procurement of funding for proposed wetland conservation projects by serving as Colorado Division of Wildlife representative on the Wetland Initiative Partnership Committee, chairman of the Waterfowl Habitat Project Review Committee, state coordinator for the Intermountain West Joint Venture, and as a member of the Playa Lakes Joint Venture Management Board. b. Advise public land management agency personnel, including area biologists and district wildlife managers, on the potential benefits to migratory birds of acquisition and/or development of wetland areas. Activities may include on-site inspection, formulating plans for the collection of biological data, recommending wetland enhancement developments, or review of development plans. Provide information on wetland habitat development to private land managers. c. Prepare and present information programs on the principles of migratory bird management. Preparation may include literature review, construction of charts, graphs, and tables as photographic slides or posters, and rehearsal of presentations. d. Attend flyway Technical Committee and Council meetings and flyway special workshops as assigned. Compile information on current status of Colorado's migratory bird population and hunting season results through consultation with CDOW biologists and managers. Prepare reports for presentation at meetings and workshops. Serve on committees as assigned by flyway Technical Committee and/or Council Chairman. Attend selected meetings in Colorado that address migratory bird management programs, at which in-depth biological expertise would be of value. e. Participate in cooperative migratory the Central and Pacific Flyway councils. bird studies sponsored through f. Provide methodology to migratory bird and wetland managers for sampling the biological parameters of interest. Literature review may be required to develop appropriate methodology. Parameters of interest may include breeding pairs, nesting densities, nesting success, fledging success, vegetation composition and density, invertebrate composition and density, or population survival. g. Assist Basin. with Canada goose trapping and banding in the upper Gunnison �37 RESULTS The Wetland Initiative (WI), Wetland Focus Area Committees (WFC), Joint venture and Waterfowl Habitat Project Reyiew Committee (wePRC) Actiyities As members of the WI project planning and CDOW Wetland Program team, migratory bird researchers assisted in preparing the final grant application and developing project selection criteria; consulted on further development and refinement of wetland proposals; served on the committee to select project proposals for funding; and helped develop project development packages for some of the selected projects. Szymczak continued to communicate with all WFC in the state and met with committees for the South Platte River (2), southeast Colorado(1), the San Luis Valley (1), North Park (2), Lower Colorado/Gunnison (1), and Front Range (1). As Colorado's representative on the Playa Lakes Joint Venture Management Board, Szymczak attended 1 meeting and approved funding for projects in Colorado, Oklahoma, and Texas. As chairman of the WHPRC, Szymczak chaired 1 committee meeting for ranking and funding proposals submitted for the 1997-98 funding year; informed proposal submittees of the outcome of their funding request; periodically monitored progress of project planning, construction, and money flow for new and previous years funded projects; coordinated Site Specific Agreements and fund reimbursement with the Ducks Unlimited Inc. MARSH program; served as the Project Officer for wetland development contracts formulated with Ducks Unlimited, the Bureau of Land Management, and the U. S. Fish and Wildlife Service. Specific projects were funded along with non-project specific funding support for the USFWS Partners For Wildlife program. Informational Programs Gammonley: (1) participated in a workshop on wetland ecology and management at Alamosa-Monte Vista National Wildlife Refuge for U.S. Fish and Wildlife Service and CDOW biologists; (2) visited wetland sites and made recommendations for developments, management, and/or monitoring programs at Roots Reservoir, Higel SWA, Rio Grande SWA, Jackson Lake SWA, Elliott SWA, and XY Ranch SWA; (3) assisted with writing the wetland (moist-soil impoundment) portion of the XY Ranch SWA development plan; (4) visited wetland projects constructed through the U.S. Fish and Wildlife Service Partners for Wildlife program along the South Platte River, Lower Colorado River, and in southeastern Colorado, and discussed management of these projects; (5) assisted in duck nest searches at Hebron Wildlife Area in North Park, and made recommendations on improving the nest search methodology and water management on wetlands on the area; (6) gave a presentation on waterfowl identification and ecology to the Fort Collins chapter of the Audubon Society; (7) gave a presentation on waterfowl identification and wetland management to CDOW trainees; (8) presented a I-day field lecture and workshop on wetlands and waterbirds in the San Luis Valley for an undergraduate course in "Rocky Mountain Fauna from Western State College; (9) was asked to write (and began work on) a chapter on wildlife use of palustrine wetlands for a book, "Ecology and management of wetlands in the Intermountain West being edited by staff from the University of Wyoming; and (10) was asked to be an associate editor for Wetlands, the journal of the Society of Wetland Scientists. N N , �38 Szymczak visited existing and potential wetland sites and made recommendations for development and/or management for: Partners For Wildlife (U. S. Fish and Wildlife) projects on private land in southeast Colorado and in the Grand Junction/Montrose area; Elliot state Wildlife Area (SWA), XY Ranch SWA, and Russell Lakes SWA. Szymczak also gave waterfowl identification presentations to CDOW Trainees and Longmont Explorer Scout Post. Waterfowl Technical Committee and Council Meetings In January, Szymczak attended the winter meeting of the Pacific Flyway study Committee (PFSC) at which the main topic of discussion was Adaptive Harvest Management (AHM) and the proposed changes to the regulation packages. In March, Gammonley attended the Central Flyway Waterfowl Technical .Committee (CFWTC) and Central Management Unit Technical Committee (CMUTC) meetings while Szymczak attended the PFSC and Western Management Unit Technical Committee (WMUTC) meetings. Major items of direct relevance to Colorado included a review of analyses of neck collar observations of shortgrass prairie population Canada geese (CFWTC), completion of a revision of the management plan for the Hi-line population of Canada geese (CFWTC), review of the Rocky Mountain Population of greater sandhill crane management plan revisions and allocation of harvest for 1997 (PFSC), development of a Central Flyway early season teal harvest strategy (CFWTC), review of the current status of the pintail interim harvest strategy (PFSC), compilation of the Four-Corners band-tailed pigeon 1996 harvest report and recommendation for 1997 seasons (WMUTC) discussion of mourning dove survey methods and population trends (CFWTC, PFSC) , revision of the North American Waterfowl Management Plan (CFWTC, PFSC), and review of Central and Pacific Flyway recommendations for duck harvest regulation packages under the AHM approach. Szymczak represented Colorado on the Pacific Flyway Council (PFC) at the March Council meeting at which recommendations from the PFSC were reviewed and early hunting season recommendations were forwarded to the U. S. Fish and Wildlife Service. In July, Gammonley attended the CFWTC and Central Flyway Council sessions while Szymczak attended the PFSC meeting and again served on the PFC. At the CFWTC and PFSC the status of waterfowl populations was reviewed along with characteristics of the 1996-97 waterfowl hunting season harvest, and proposed 1997-98 hunting season recommendations were formulated and forwarded through the Council to the USFWS Regulation Committee. A major topic of discussion at the PFSC was the development of continental pintail and western mallard models including the establishment of the Continental Divide as the demarcation between the western and mid-continent mallard populations. other important topics pertinent to Colorado were kill permits for Rocky Mountain greater sandhill cranes and the planned revision of the Rocky Mountain Canada goose population management plan. Subsequently, Gammonley and Szymczak made recommendations to CDOW regulations personnel on the structure of waterfowl hunting seasons in Colorado, and reviewed final selections for hunting season structure and regulations for migratory game birds. In addition to flyway meetings, Gammonley participated in the Central Flyway waterfowl parts collection wing-bee in February. He also reviewed and provided comments on the CMU mourning dove management plan, and served on a CMUTC subcommittee to review and rank proposals submitted for Webless Migratory Game Bird Research (WMGBR) grants. Gammonley also attended a meeting of the ad hoc subcommittee to revise the management plan for midcontinent population sandhill cranes, and submitted a WMGBR proposal for a �39 study on spring food availability and energetics of Rocky Mountain population greater sandhill cranes in the San Luis Valley of Colorado. Szymczak attended a meeting in Oregon for further discussions and decisions on the western mallard management unit. At the winter meeting of the CFWTC in December attended by Gammonley, major items of direct relevance to Colorado included a review of analyses of neck collar observations of short-grass prairie population Canada geese, work on a revision of the management plan for the high-line population of Canada geese, and development of Central Flyway recQmmendations for duck harvest regulation packages under the Adaptive Harvest Management (ARM) approach. Cooperative Canada No Canada segment. Goose geese Banding were banded under the cooperative program during this DISCUSSION Project personnel provide useful information in planning and evaluating waterfowl management and habitat enhancement programs in Colorado and educating land management agency personnel about the habitat requirements of waterfowl. With increasing emphasis on wetland habitat in Colorado, and the initiation of new programs with expanded responsibilities for project personnel, wetland-related objectives of this job will receive more emphasis in the near future. Colorado now has 10 Wetland Focus Area Committees functioning in the state that will require coordination and expertise in wetland project planning. The resources provided by project personnel will insure that money raised through the Colorado Duck Stamp program or any other funding initiative will be spent in accordance with the objectives of the program. Conducting and/or formulating surveys and banding efforts and informing management agency personnel about aspects of waterfowl and wetland ecology provides a valuable service to management agencies, the waterfowl resource and, in some cases, the hunting public. Continued participation on Pacific and Central Flyway committees ensures that Colorado will remain informed on migratory bird matters, have input in migratory bird hunting regulations, and influence habitat programs affecting migratory game birds. Prepared by, .~.J 1!!~ Michael R. Szymczak LSSR IV James H. Gammonley LSSR III ��41 Colorado Division Wildlife Research April 1998 of Wildlife Report JOB PROGRESS state of ColOrado Project Work Migratory W-166-R 22 Plan Job Title: Period Personnel: Michael Game Birds Inyestigations 2 Job Migratory Covered: Author: REPORT Game 01 January Bird Publications through 31 December 1996 R. Szymczak James H. Gammonley Wildlife and Michael R. Szymczak, ABSTRACT No articles Prepared by were published ~~7?~ Michael R. Szymczak Researcher/Scientist IV during this segment. Colorado Division of ��43 JOB PROGRESS State of: Colorado Project: W-167-R Work Plan: _--",1~_: Job Job Title: Period Authors: Ayian Thomas 01 January Research 24 Evaluation of Habitat Eastern Colorado Covered: REPORT Development through E. Remington for Ring-necked 31 December and Warren Pheasants in 1997 D. Snyder Personnel: C. E. Braun, T. J. Davis, M. J. Emerson, M. A. Etl, K. M. Giesen, E. T. Gorman, L. K. Haynes, R. W. Hoffman, K. D. Johnson, J. L. Mekelburg, K. L. Martin, M. L. Molarsky, T. E. Remington, W. D. Snyder, M. L. Trujillo, A. E. Vitt, J. D. Wieland, B. T. Weinmister, J. A. Yost, D. J. Younkin; Colorado Division of Wildlife. ABSTRACT Expenditures under the Pheasant Habitat Improvement Program (PHIP) increased from about $299,000 in 1995 to $318,000 in 1996, but declined to about $297,000 in 1997. Most habitat developments were either plum thickets (225 of 509), or sorghum food plots (175), although, as in past years, most (84%) PHIP expenditures went toward establishment of plum thickets. PHIP has now resulted in the establishment of 954 plum thickets in 6 years. Average counts of crowing male ring-necked pheasants (Phasianus colchicus) did not differ (~ > O.O~) between treatment and control blocks. Counts averaged about 14 calls per station in 1993, 17 in 1994, 13 in 1995, 14 in 1996, and 20 in 1997. Hunter pressure (opening weekend only) in 1996 was similar to 1995, but increased about 79% in 1997. Harvest rates on opening weekend increased about 78% to 0.14 birds per hour in 1996 and 0.16 birds per hour in 1997. Hunters were most successful when hunting sorghum food plots created under the PHIP program where harvest rates were over 3 times higher than the average for other cover types. Eighty-two hens were captured and radiomarked for survival analysis, while 33 hens radiomarked in 1994, 1995, and 1996 were also relocated for survival analysis. The sample size of birds available for survival estimates was lIS. Annual survival of radio-marked hens (1 Nov - 31 Oct) was 29.2%, an increase from the 15% in 1995 and 23.8% in 1996 but still substantially below the 41% survival in. 1993-94. The intermediate survival rate was attributed to generally poor height and quality of wheat stubble and declining quality in Conservation Reserve Program (CRP) fields. Most mortality was due to predation. The percentage of wheat in Phillips county harvested with a stripper header increased from 12 to 19.5%. Most (89%) stripped-wheat stubble was treated with herbicides or mechanically cultivated which decreased its cover value to pheasants. OVer half (54%) of conventionally-harvested wheat stubble was sprayed with herbicides, undercut, or disced in the fall which, because of short height, essentially eliminated these fields as pheasant habitat. ��45 EVALUATION OF HABITAT DEVELOPMENT FOR RING-NECKED PHEASANTS IN EASTERN COLORADO Thomas E. Remington and Warren D. Snyder INTRODUCTION Pheasants are pursued by more hunters than any other small game species in Colorado (83-88% of small game license buyers). In a recent survey, 74% of pheasant hunters rated their hunting trips in Colorado as poor (45%) or fair (29%), while only 10% rated their trips as very good or excellent. Lack of birds and places to hunt were identified as the most significant reasons why some hunters did not hunt pheasants in Colorado. Small game license sales in Colorado have declined by about 90,000 (45%) in the last 10 years. It is apparent that if the Division of Wildlife is gOing to increase license sales and hunting participation, pheasants will be a key species. presumably, recruitment and retention of hunters will increase if the quality of pheasant hunting is improved, i.e., increases in pheasant numbers and places to hunt. Previous research has indicated that over-winter survival of pheasants is the most critical factor limiting pheasant populations. The Pheasant Habitat Improvement Program (PHIP) was created to establish overwinter survival cover within historically good pheasant range in eastern Colorado. The program was conceptually designed to overcome significant obstacles to developing habitat, mainly a lack of manpower and a burdensome contractual system (costs of administering contracts exceeded costs of developments). Under PHIP, the Division of Wildlife contracts with individual Pheasants Forever chapters in eastern Colorado to contact landowners and develop habitat on private lands following specific guidelines. Each chapter develops contracts with individual landowners and pays them when the habitat work is completed and verified. Division of Wildlife personnel inspect habitat developments and verify completion and compliance with guidelines. A new method of harvesting wheat has potential to increase pheasant survival. Stripper headers strip the grain from the stalks rather than cutting and thrashing the stalks to separate the grain, which leaves much taller stubble. Pheasant survival from July through April is largely dependent upon the height and cover value of wheat stubble (Snyder 1985). We trapped, radiomarked, and released a sample of hens in paired blocks of stripped and conventionally-cut wheat to assess their survival in each cover type. P. N. OBJECTIVES To determine if habitat developments offered through the Pheasant Improvement Program increase pheasant survival, breeding density, harvest within selected northeast Colorado study areas. Habitat and pheasant SEGMENT OBJECTIVES 1. Work with Pheasants Forever chapters, management personnel, landowners to develop annual sorghum plantings, disturbance switchgrass, or plum thickets and to develop stripped-wheat in other locations. and tillage, test plots �46 2. Evaluate Pheasants Forever chapter and landowner acceptance and consider modifications as suggested or needed. 3. Evaluate 4. Quantify hunting pressure control sites. 5. Quantify extent of pheasant conditions. 6. Monitor spring breeding densities pheasant crow-count surveys. 7. capture and radiomark up to 100 hen pheasants within stripped wheat areas, and up to 100 hens in conventionally-cut wheat areas. 8. Relocate hen pheasants periodically to quantify survival rates within stripped wheat and control areas and ascertain habitat use. 9. Analyze quality of annual plantings as survival cover. and pheasant harvest within treatment use of treatments data and prepare progress of program in treatment and under winter snow and control areas using report. METHODS Procedures used during this work segment to evaluate quality of annual plantings, quantify hunting pressure and pheasant harvest, conduct crowing counts, and capture and radio-mark hen pheasants were described in previous progress reports (Remington and Snyder 1994, 1995, 1996, 1997). The PHIP Habitat Project Guidelines for participating Pheasants Forever chapters were modified slightly from 1996 and are attached (Appendix A). The most significant changes were the addition of linear plum thickets along drainages (at a reduced payment rate), addition of a payment rate for mowing thicket plantings, and an increase in the payment rate for replanting seedlings from $0.25 to $0.40. We continued to compare both survival and nest success of hens using wheat harvested with stripper headers and conventional headers. Complexes of stripped-wheat fields were identified for use as treatment areas. Following wheat harvest in July 1997, we mapped all the wheat stubble in Phillips and Sedgwick counties where stripper headers were prevalent. Stubble was classified along road-side transects as to header type used and how the stubble was treated post-harvest; i.e., untreated, sprayed with herbicides, or undercut. We trapped hens from CRP and conventionally-harvested stubble .in areas we had trapped in previous years, and moved most of them into strippedwheat fields in stripped-wheat study areas. Research in Missouri (Wilson et al. 1992) indicated most translocated hen pheasants stayed within 1-2 miles of their release point. Study areas and release points were established so that hens moving 1-2 miles would still have access to stripped-wheat fields. Translocated hens that were relocated in or near stripped wheat fields were considered stripped wheat "treatment" hens. "Control" hens were hens that were trapped and radiomarked but not translocated, or translocated hens which moved from stripped wheat fields and subsequently were relocated primarily in conventionally-cut wheat or CRP fields. We designed an experiment to evaluate the impact of mowing on growth and survival of first-year plum and juniper seedlings. Weeds between rows in �47 thickets and windbreaks were mowed with a rotary mower pulled behind a quad runner in late-June through July. Weeds growing around seedlings in slits through the weed barrier fabric were killed with Glyphosate (RoundupR). A 1:7 mixture of Roundup was mixed in a hand sprayer, the nozzle of which was covered with a sponge which was used to swab weeds with the herbicide. In a few cases when rain was imminent weeds were pulled from around shrubs. Plums and junipers were evaluated at the time of mowing; dead plants were marked with white paint while those in poor condition were marked with green paint. This was done so that shrubs already dead or perhaps dying at the time of mowing could be eliminated from the analysis (since they couldn't or were unlikely to benefit from mowing) thereby eliminating some of the statistical variance. Growth and survival of plums and junipers will be evaluated in spring 1998 after buds have opened and live trees can be distinguished readily from dead. RESULTS Rainfall and Weather Patterns The survival and quality of habitat developments and the quality of wheat stubble, CRP, and other cover types is ultimately dependant on the amount and timing of precipitation. Winter precipitation was light, with little persistant snow cover. Wheat survival was good however as heavy rains the previous July through September had recharged soil water profiles. March through May precipitation was about half of normal levels throughout the area. Above average rainfall in June salvaged much of the wheat in northeastern Colorado and prevented substantial mortality on plums and junipers in PHIP plantings. Rainfall was adequate and fairly well distributed through the remainder of the growing season and germination and growth of sorghum food plots was good. An unusually heavy snowstorm accompanied by heavy winds occurred on 25 and 26 October. This broke over some sorghum and filled in much of the wheat stubble. Snow persisted in wheat stubble for a week or more before melting. Annual precipitation averaged between 15 and 17 inches which is about average (Table 1). Table 1. Monthly precipitation (inches) and departure from 30-year (in parentheses) at 4 U.S. Weather Service stations in the Pheasant Improvement Program area, 1997. Month Jan Feb Mar Apr May Jun JuI Aug Sep Oct Nov Dec Totals Holyoke 0.19 (-0.27) 0.63 (+0.24) 0.27 (-0.93) 1.15 (-0.54) 1.85 (-1.50) 4.89 (+1.70) 1.35 (-1.38) 1.32 (-0.59) 0.24 (-0.97) 3.l9 (+2.47) 0.06 (-0.54) 0.73 (+0.34) 15.87 (-2.31) 8Insufficient or partial data. Yuma 0.50 (+0.12) 0.81 (M)8 0.00 (M) 0.60 (M) 1.96 (-1.05) 3.06 (+0.19) 1.97 (-0.76) 4.38 (+2.71) 1.41 (+0.09) 2.23 (+1.45) 0.00 (M) 0.45 (+0.14) 16.92 (M) Akron4E 0.52 (+0.19) 0.53 (+0.22) 0.07 (-0.92) 0.93 (-0.42) 2.24 (-0.95) 3.11 (+0.51) 1.13 (-1.69) 3.47 (+1.44) 1.06 (+0.06) 2.40 (+1.71) 0.26 (-0.31) 0.44 (+0.08) 15.72 (-0.16) average Habitat Sterling 0.37 (+0.04) (M) 0.30 (-0.74) 0.71 (-0.66) 3.58 (+0.42) 4.40 (+1.49) 2.53 (M) 2.44 (+0.57) 0.53 (-0.50) 1.78 (+0.99) 0.00 (-0.49) 0.40 (M) 16.64 (M) �48 PUlP Expenditures Six Pheasants Forever chapters in eastern Colorado participated in the Pheasant Habitat Improvement Program (PHIP) in 1997 and received funds from the Division of Wildlife for habitat development. Contracts were signed for a total of $290,000 with chapters in Logan County (Sterling, $15,000 and Fleming, $25,000), Phillips County ($80,000), Washington County ($55,000), Yuma County ($80,000), and Morgan County ($35,000). These 6 chapters expended $280,695 in PHIP funds for habitat development (Table 2). Shrub thickets with accompanying juniper windbreaks continued to be the most popular habitat option, accounting for 204 of 378 habitat developments (54%) (Table 3). About 85% of PHIP funds were expended on plum thickets and juniper windbreaks, 8% on sorghum or other food plots, 2% on switchgrass plantings, and 4.6% on custom work such as site preparation, replants, mowing etc. (Table 3). Expenditures declined in 1997 by about 12% from 1996 spending levels (Table 4); the first decline in spending since the program began. This was caused by the loss of the Burlington Chapter (which lost their Pheasants Forever charter), reductions in the scale of shrub thicket plantings in Yuma and Logan counties, and loss of sorghum food plots in CRP as CRP acreage expired and was not renewed. Spending should increase above 1996 levels in spring 1998 with the addition of chapters new to the PHIP program in Flagler, Cheyenne Wells, Lamar, and Springfield. Table 2. Expenditures under PHIP during 1997 by Chapter and habitat type. Cha ter Phillips Yuma Washington Morgan Sterling Fleming Totals $62,991.77 $45,579.00 $65,990.10 $30,858.22 $16,358.51 $18,023.11 $239,800.71 3,125.00 360.00 2,751.00 917.00 2,291.16 10,390.00 3,280.00 2,303.21 2,061.00 2,653.60 300.00 1,120.00 1,100.00 212.00 400.00 800.00 2,926.00 1,965.16 72.00 16,115.00 4,972.00 6,447.21 6,450.00 6,909.92 $72,435.93 $66,266.81 $72,301.26 $32,242.22 Habitat Shrub thickets Food Plots Sorghum Weeds Switchgrass Custom labor Replants Totals Shrub Thickets 1,393.00 474.00 $16,758.51 $20,690.11 $280,694.84 and Windbreaks The ·PHIP program has now resulted in the establishment of 958 plum thickets in 6 years (Table 5). Chapters continued to use diverse means to plant shrub thickets. The Frenchman Creek chapter (Fleming, Logan County) once again planted all of their thickets with over 50 chapter and community volunteers participating. Phillips County chapter volunteers planted most of their thickets, but subcontracted about 20 to Windbreaks Diversified. All Washington County plantings were sub-contracted to local Future Farmers of America (FFA) and Young Farmer chapters. Morgan County used FFA chapters for the first time this spring along with the Brush Prairie Baseball team. Yuma County plantings were completed by an FFA Chapter and Yuma County Soil Conservation District personnel. An Explorer Scout Troop from Sterling assisted the Northeastern Colorado (Sterling) Chapter. �49 Survival and growth of seedlings on most sites has been good to excellent. Plums in many sites that were planted during 1992-94 are root sprouting between the rows to begin forming -closed-canopy thickets. Survival of Rocky Mountain junipers has been even higher than that of plums at most sites. Over 200 shrub sites were inspected during fall/winter of 1995-96 to assess where fair to poor survival mandated replanting. Replanting of junipers was completed on most sites where it was needed, but time and manpower constraints prevented replanting plums on all sites. Replanting continued in 1997, but more remains to be done. Shrub survival was good on most 1997 sites, although extended drought iriearly spring increased mortality on sites where soil preparation was inadequate or other problems occurred. Some planting crews were cutting the roots back on bare root plums to facilitate planting. This practice greatly increased mortality, as did leaving the tar paper covering on Rocky Mountain juniper seedlings. Two juniper windbreaks around thickets in Yuma county had to be almost completely replanted because of extensive mortality in late summer. The felt "pot" had not been removed and junipers were unable to send out roots and died when moisture within the soil ball was depleted. Chapters were informed in person, in field trips, and via advisories in the PHIP guidelines not to cut root masses back any further (roots and crowns are cut back on large plums at the nursery) and to remove tar paper from juniper seedlings prior to planting. Sorghum Food/Cover Plots Sorghum food/cover plots were planted on 91 sites totaling 545 acres in 1997 (Table 3). This was a reduction of 84 sites totaling 115 acres. Sites planted by CDOW temporary personel were planted using surface planters with 30-inch row spacings. Sites planted by landowners were typically drilled on a 12- to 15-inch row spacing. Planted sites were subsequently cultivated to reduce grass and weed competition. Germination and early growth was good to excellent in all plots. Pheasant broods were flushed from several plots in August and September. Heavy, wet snow accompanied by strong winds broke over most sorghum stalks in late October. Stems generally broke over 12-15 inches above ground level but plots continued to provide fair to good cover and food for pheasants (Table 6). Sorghum plots planted by landowners using drills with a 12-inch row spacing had good growth but thin stalks. Cover value was good until the October snowstorm when sorghum in these plots blew over at or near ground level. Cover for the remainder of the fall and winter was poor, although pheasants used and were harvested from these food plots. Landowners in the future will be encouraged to plug every other row in their drills to alter spacing to 24-inch rows, thereby reducing competition among sorghum plants and increasing stalk strength and ultimately cover value. Thirty sites were disced (by landowners) and allowed to develop into annual weeds. No measurements were made of cover quality in these plots, however visual inspections of many of these showed most developed into excellent stands of tall annual weeds which held up to the October snowstorm quite well. switchgrass switchgrass was planted on numerous plots that had been planted to sorghum in previous years within Conservation Reserve Program (CRP) fields. Satisfactory stands were attained on nearly all, but herbicides were not used to reduce weed competition so growth was marginal. Switchgrass was planted in several other locations with generally excellent establishment and first-year growth. �so Switchgrass was replanted been attained in previous on several years. tracts where satisfactory Table 3. Pheasant habitat planted and/or contracted chapters during 1997 in northeastern and east-central Pheasant Habitat Improvement Program. Habitat/Chapter Shrub Thickets and Windbreaks Washington County Phillips County Frenchman Creek (Fleming) Yuma County N. E. Colorado (Sterling) Morgan County Subtotal Sorghum Food/Cover Plots Washington County Phillips County Frenchman Creek (Fleming) Yuma County N.E. Colorado (Sterling) Morgan County Subtotal Disturbance Tillage (Annual Forbs) Washington County Phillips County Frenchman Creek (Fleming) Yuma County N. E. Colorado (Sterling) Morgan County Subtotal Switchgrass Plantings Washington County Phillips County Frenchman Creek (Fleming) Yuma County N. E. Colorado (Sterling) Morgan County Subtotal Totals .as 204 2 15 7 58 2 .i 91 11 3 0 14 0 ...1 30 0 13 5 7 0 _Q 25 378 had not by Pheasants Forever Colorado through the Number Plantings Acres 50 58 21 38 12 stands Payment ($) 33.5 29.0 9.4 20.9 7.9 13.4 114.1 65,990.10 62,991.77 18,023.11 45,579.00 16,358.51 30,858.22 239,800.71 7.5 81.5 20.0 270.0 10.0 27.5 545.5 300.00 3,125.00 800.00 13,670.00 400.00 1,100.00 17,575.00 28.0 12.0 0.0 82.0 0.0 5.3 127.3 1,120.00 360.00 0.00 3,280.00 0.00 212.00 4,972.00 0 55.35 23.60 45.21 0.00 0.00 124.16 0 2,751.00 1,393.00 2,303.21 0.00 0.00 6,447.21 911.1 �51 Table 4. Pheasant type and Pheasants Habitat Forever Habitat/Chapter 1992 Pll.!m Thi~k~t~LWinQQr~gk~ Phillips 23,454 Yuma 8,730 Sterling 6,123 Washington Burlington Fleming Morgan Subtotals 38,307 Swit~hgrg~~ ~lgnting~ Phillips Yuma Sterling Washington Fleming Subtotals Sorohl.!m ~lgnting~ Phillips Yuma Sterling Washington Fleming Morgan Subtotals Di~tl.!rQgn~~ Tillgg~ Phillips Yuma Sterling Washington Morgan Subtotals ($ ) by habitat Improvement Program expenditures Chapter, northeastern Colorado, 1992-97. Totals 1993 1994 1995 1996 1997 59,661 27,525 11,992 41,760 5,556· 52,982 36,992 23,263 58,030 12,319 77,866 47,083 16,642 65,513 17,738 9,888 65,990 45,579 16,359 65,990 183,586 234,730 62,088 50,457 19,059 67,513 10,320 36,075 ~Q.f!27 266,176 146,494 2,800 1,676 754 20,030 76 1,692 5,160 150 888 700 5,333 4,800 240 1,275 1.1~Q 12,798 342,041 216,366 93,438 298,603 45,933 18,023 63,986 ~1. 72~ 3Q.f!~f! 239,801 1,109,094 6,447 36,074 9,002 3,574 1,975 ~.~~3 53,168 25,163 3,125 13,670 400 300 800 1.1QQ 17,575 48,985 113,221 35,266 39600 2,552 l.2~~ 203,823 377 1,720 360 3,280 6,472 22,020 3,797 2,070 212 34,571 5,230 21,795 6,898 360 3,530 7,388 17,385 22,102 9,472 700 14,640 30,322 7,075 1,100 8,218 29,377 8,8S6 560 5,257 14,220 2,075 1,300 1,752 11,278 49,659 53,137 47,011 1,170 1,610 520 3,615 5,250 1,200 300 590 6,640 1,005 360 3,520 1,072 180 ~~~ 470 2,751 2,303 ° 0 l......lll 1,120 _2.li 3,300 Iall Kb~atLHQ-lill :rib~gt Phillips Yuma 1,069 Washington Sterling 1.QQQ Subtotals 2,069 10,365 300 400 60 760 8,235 5,132 150 90 1QQ 640 360 2,567 4,972 300 1,979 150 ;!..1QQ 3,829 360 CUl:ltom Horka Phillips Yuma Sterling Washington Morgan Subtotals Totals 54,954 1,591 3,290 2,382 1,257 6,264 1,253 8,520 10,537 .4,549 221,028 277,930 298,680 3,020 90 1,313 163 2,983 6,680 517 3,791 12~ 11,150 317,854 3,208 4,715 4,891 _____:n_ 12,886 280,690 aReplanting shrubs, repairing fabric, applying fertilizer, discing, etc. 17,833 10,571 3,062 15,942 23~ 47,642 1,170,446 �S2 Table under 97. 5. Number and type of plantings the Pheasant Habitat Improvement Habitat! Chapter Elum ThicketslHindb~eaksa Phillips Yuma Sterling washington Burlington Fleming Morgan Subtotals 1992 1993 1994 1995 1996 1997 Totals 26 7 5 51 30 20 31 6 41 36 25 46 14 63 38 15 50 17 8 54 40 19 52 11 32 58 38 12 50 21 293 189 96 229 48 61 _l1 ....£S. _fl 138 162 19l 225 204 958 11 7 6 63 1 9 22 1 4 2 44 40 2 7 13 7 153 56 21 9 _.2. _2 _ll 24 73 29 102 25 253 2 15 38 113 131 52 4 110 169 46 8 55 171 64 5 42 85 13 12 12 337 629 215 31 19 _ll 15 58 2 2 7 __ 7 55 300 333 295 175 91 1,250 5 9 2 19 14 7 2 6 24 8 7 5 22 3 1 1 6 3 14 39 89 20 10 38 Switchg~ass Elantiogsb Phillips Yuma Sterling Washington Fleming Subtotals SQrghym Elanting~ Phillips Yuma Sterling Washington Fleming Morgan Subtotals DiQtyrQgn~e Tillage Phillips Yuma Sterling Washington Morgan Subtotals completed by Pheasants Forever chapters Program, northeastern Colorado, 1992- 16 Tall wheatllfQ-Till Wheat Phillips Yuma 8 Washington Sterling _2 Subtotals 13 42 45 31 5 2 1 1 3 1 ---.l..l 11 7 ____2_ ____2_ 30 141 5 12 4 ____!t _i 8 8 1 aWoody plantings consist of 0.1 to 0.3 acre plum thickets by three-row juniper or juniper/plum windbreaks. bMany of the 1996 switchgrass plots had previously seeded back to grass within CRP fields. 30 and most are accompanied been sorghum plots that were �53 Table 6. Vegetation characteristics of sorghum plots planted within blocks in 1997 and sampled in ,ebruary 1998, northeastern Colorado. VOR Treatment block N Pauli Clarkville 3 3 Means Coyer Yalue of Wheat Height (ft) S.D. (dm) S.D. 1.4 1.6 0.3 0.2 1.2 1.1 1.5 0.25 1.15 0.15 0.1 0.0 treatment CanoQYcover (%) Sorghum S.D. Forbs S.D. 33.4 38.5 25.6 4.2 10.0 3.8 14.4 3.3 36.0 14.9 12.2 3.6 Stubble stripper headers were first used in Phillips County in 1992. By 1996, 12.4% of wheat harvested in Phillips County was combined with a stripper header. Wheat harvested with stripper headers increased to about 19.5% in 1997. Stripped wheat has the potential to provide tremendous cover for pheasants because it is so tall (Remington and Snyder 1996, 1997). However, only 11.1% of stripped-wheat acreage was left untreated after harvest. Most (89%) stripped-wheat fields were treated to control weeds; either with herbicides in spring to control mustard (10%), post-harvest with herbicides (75%), or by discing or undercutting (4%). Landowners in this area purchased stripper headers primarily to facilitate a crop rotation where 2 crops are grown in 3 years. In this cropping system dryland corn or sunflowers are planted into the wheat stubble. A fall application of herbicides is used to conserve moisture for the spring crop. .Herbicide application and undercutting both significantly reduce the cover value of stripped wheat for pheasants (Remington and Snyder 1997), although both treatments still leave stubble with higher VORs and height than does conventionally-combined stubble. The cover ·value of conventionally-cut wheat stubble has declined as semi-dwarf varieties of wheat are planted almost exclusively, and as fall herbicide use or mechanical cultivation become more prevalent throughout northeastern Colorado. In Phillips County during 1997, only 46% of wheat combined with conventional headers was left untreated, while 41% was sprayed with herbicides and 12% was undercut or disced in fall. Semi-dwarf varieties of wheat seldom yield adequate stubble height for pheasant survival when harvested by modern combines; this problem is exacerbated when herbicides control weed growth that can mitigate short stubble height. Pheasant Crowing Census Average counts of crowing males did not differ between treatment blocks (Table 7; ~ = 0.95 and 0.50 using high count of replicate average of replicate counts, respectively). Average high counts 1995, 14 in 1996, but increased to 20 in 1997. and control counts and were 13 in �54 Table 7. Pheasant crowing census data among treatment northeastern Colorado, spring 1997. and control blocks, Count 1 Block Holyoke SE Mailander Kurtzer Clarkville Pauli Y-W Co. Line Fleming Kuntz otis Curve Means 14.7 11.2 42.0 27.0 22.1 16.6 22.5 17.1 8.5 Paoli NE Haxtun NE Paoli South st. Pete Kelly Yuma Co. Lonestar Platner Washington W. Means 5.7 21.5 16.6 25.9 31.4 10.0 22.1 23.4 10.5. Overall ~ Obtained Hunter Average of count.s" 2 32.7 23.5 Highest count Treatments 14.7 11.2 42.0 27.0 22.1 16.6 27.6 17.1 8.5 20.7 ± 10.3 Controls 5.7 21.5 16.6 25.9 31.4 10.0 22.8 23.4 10.5 18.6 ± 8.5 mean using Pressure 14.7 11.2 42.0 27.0 22.1 16.6 32.7 17.1 8.5 21.3 ± 10.8 14.7 11.2 42.0 27.0 22.1 16.6 34.7 17.1 8.5 21.5 ± 11.1 5.7 21.5 16.6 25.9 31.4 10.0 23.5 23.4 10.5 18.7 ± 8.5 5.7 21.5 16.6 25.9 31.4 10.0 24.8 23.4 10.5 18.9 ± 8.6 20.0 the highest count per station among High count/ at.at.Len" ± 9.5 counts before averaging. and Success During opening weekend 387 hunters were contacted and interviewed to ascertain their success. This was a substantial increase from 1996 when 299 hunters were contacted. Expressed as contacts per checker, hunter pressure was 59, 36, 43, and 77 over the last 4 years. By this measure hunter participation increased 79% over 1996. Harvest rates were about 18% higher than last year, 0.160 (1997) vs. 0.136 birds harvested per hour of hunting effort (Table 8). Harvest rates almost tripled last year over 1994. Success remained high this past season despite considerably more hunters pursuing pheasants opening weekend. Hunters expended 7 hours of effort to harvest a rooster, or on average 1.2 roosters were bagged per 8-hour hunting day. Harvest has now exceeded, on average, a bird per hunter for two years in a row when it had not previously been that high since we began collecting harvest information in 1992. Hunter effort and harvest were recorded and summarized by habitat type (Table 8). Hunters were most successful in sorghum food plots and stripped wheat and least successful in wheat stubble and CRP. Harvest rates in sorghum food plots were 2-4 times higher than harvest rates in other, more commonly hunted cover types such. as creek bottoms, CRP, or wheat stubble. This confirmed earlier hypotheses that sorghum food plots could concentrate pheasants and make them more vulnerable to hunters (Remington and Snyder �55 1996). Hunters were also quite successful hunting stripped wheat, although this cover type sustained only 2% of total hunter effort. Success by Cover type was generally similar to results from the 1996 survey, although corn stubble more than doubled in productivity. The northern edge of many corn circles in Yuma county, and to a lesser extent Phillips and Logan counties could not be harvested because 5-6 feet of snow drifted in during the October snowstorm and either persisted to harvest or left fields too wet to harvest. These unpicked areas adjacent to picked corn stubble held birds, were very attractive to hunters, and undoubtably increased success. Corn harvest was delayed enough by wet conditions that stalks in portions of fields that were picked were not cultivated by opening weekend which also enhanced conditions for hunters. Table 8. Colorado, Pheasant hunter effort, 8-9 November 1997. Cover Hours Total Sorghum Stripped wheat Creekbeds, weeds CRP Wheat stubble Corn stubble Weighted average 127 31 499 327 308 206 1,498 flush and harvest rates in northeastern Flush/hr Bag/hr 1.01 0.52 0.66 0.55 0.49 0.75 0.63 0.38 0.26 0.16 0.12 0.09 0.19 0.16 % crippled Daily bag 11.0 20.0 12.0 16.7 15.6 11.1 13.6 3.0 2.1 1.3 1.0 0.7 1.6 1.2 % 8.4 2.0 33.0 21.6 20.4 13.6 lAssumes a 8-hour hunting day Pheasant Trapping and Survival Annual mortality (1 Nov 1996- 31 oct 1997) of hen pheasants was relatively low for the second straight year. Mortality was less than 10% of the hens alive at the beginning of the month for 8 of 12 months, compared to 7 of 12 in 1996 and only 3 of 12 during 1994-95 (Fig. 1). The pattern of mortality observed was typical of patterns identified over the last 4 years (Fig. 1); increasing mortality from November through January, low mortality in February through April followed by peaks in May and June or July. Mortality was exceptionally high in October 1997 (21% vs. a 4 year average of about 7%), but declined to a more average 12% in November and 8% in December. Much of the mortality in late October and early November was due to enhanced predation following the heavy snow when roosting and escape cover was filled with snow. Predation accounted for almost all mortality as in past years, although causes of predation have not been tabulated or summarized. Night-trapping began on 9 October and concluded on 23 December. Trapping success was hampered by the late-October snowstorm which prevented us from trapping for almost 3 weeks, and by loss of two temporaries to permanent jobs in December. We captured and radiomarked 82 new hens. We also continued to follow 33 hens trapped in previous years that were still alive with functioning radios as of 1 October (21 from 1996, 11 from 1995, and 1 from 1994). One hundred and fifteen hens were available for survival estimation. Of these, 70 were moved a distance of from 5 to 75 km and released into stripped-wheat fields. Most translocated hens stayed within 1-2 km of their �56 release point, although some birds left the stripped wheat fields and will be used as controls. Impact of method of wheat harvest on pheasant survival will be evaluated after radios are followed through April 1998. LITERATURE CITED Remington, T. E., and W. D. Snyder. 1994. Evaluation of habitat development for ring-necked pheasants in eastern Colorado. Colorado Div. Wildl., Prog. Rep., Fed. Aid Proj. W-167-R. Apr. pp. 1-1.9. ___________, and 1995. Evaluation of habitat development for ring-necked pheasants in eastern Colorado. Colorado Div. Wildl., Prog. Rep., Fed. Aid Proj. W-167-R. Apr. pp. 1-13. ___________, and 1996. Evaluation of habitat development for ring-necked pheasants in eastern Colorado. Colorado Div. Wildl., Prog. Rep., Fed. Aid proj. W-167-R. Apr. pp. 45-66. ___________, and 1997. Evaluation of habitat development for ring-necked pheasants in eastern Colorado. Colorado Div. Wildl., Prog. Rep., Fed. Aid Proj. W-167-R. Apr. Snyder, W. D. 1985. Survival of radio-marked hen ring-necked pheasants in Colorado. J. Wildl. Manage. 49:1044-1050. Wilson, R. J., R. D. Drobney, and D. L. Hallett. 1992. Survival, dispersal, and site fidelity of wild female ring-necked pheasants following translocation. J. Wildl. Manage. 56:79-85. Prepared b~ {. <e,,~ - /--- Thomas E. Remingt LSSRIV Prepared by _ Warren D. Snyder LSSRIV �25 20 15 ~-. 10 5 o NOV JAN MAR MAY JUL SEP . 1994 NOV JAN MAR MAY JUL SEP 1995 NOV JAN MAR MAY JUL SEP 1996 NOV JAN MAR MAY JUL SEP NOV 1997 Fig. 1. Monthly mortality (%) of radio-marked hen pheasants, October 1993 through November 1997. U1 ._J ��59 SHRVBTHlCKEISANDSUPPLEMrnNTAL¥aNDBREAKS Thickets (> 0.1 ac) may be planted with or without a windbreak, but windbreaks will not be funded if planted alone. Plantings Will be eligible for funding only in farmed areas within less than 0.1 mile of cultivated cropland and greater than 0.1 mile away from occupied dwellings. If placed in pastured grass they must be at the pasture edge adjacent to farmed cropland, and the windbreak and thicket must be fenced as one unit (using permanent fencing smaterials) to exclude livestock. PUlP does not pay for fencing. If placed in CRP, thickets must be within 0.1 mile of cultivated cropland. Plantings must not be disturbed for 10 years. Planting sites must be inspected and approved by a Colorado Division of Wildlife person before being prepared and planted. Maximum Funded: No more than 1 thicket (with or without windbreak) can be planted per 80 acres. Each thicket/windbreak must be at least 1/4 mile from another thicket/windbreak (including old plantings). Size: Shrub thickets must be at least 1I1Oth acre (4,300 if) in size, and contain at least 8 rows (excluding windbreak rows). Twelve hundred (1,200) feet is the maximum linear feet of fabric funded per thicket. Smaller plum thickets (4-row minimum and 125 ft. maximum length) will be approved but cannot be accompanied by a funded windbreak. Two-row plantin~ of wild plum along drainages adjacent to cropland must be pre-approved by the PHIP Coordinator. Maximum length is 1,320 ft. x 2 row = 2,640 ft.! site. See reduced payment rate below. Windbreaks, when used, must be to the north and west side of the thicket and will be funded up to 900 linear feet total if straight and 1,200 linear feet total if L-shaped. They must include at least 3 rows, one of which must be juniper/cedar. Spacing between the thicket and windbreak should be 100 feet (range 50 ft. min.; 120 ft. max.). Mulching: Woven polypropylene fabric such as Lumite 994 or Eartbmat is required on all plantings. Minimum fabric width is 6 ft. (3 ft. on each side of the row). Drip systems will not be cost shared. Payment Rate: Payment will be $O.55l1inear ft. of fabric to the maximums listed above (900 & 1200 lin. ft.) completed by PF Chapters, subcontracted to approved groups, or to approved private contractors. Wire staples (2 x 12-inch, 10 or 12-gauge) are required in the center of fabric at every R.M. juniper and every other plum seedling (placement at every seedling is preferred) and must be fully inserted. Dirt (only as a supplement to staples) should not be closer than at 10-yard intervals. Rocks can replace staples in rocky sites. Payment will be $O.30lIinear ft. for 2-row linear plantings along drainages. Band application of approved herbicides along the exterior edge of the fabric or June - July mowing will be funded at $O.Oll1inear ft. of fabric. Wicking of weeds at slits adjacent to seedlings will be funded at $0.005 (1/2 cent)lIinear ft. of fabric. Fertilizer will not be funded. The payment rate may be reduced up to $O.12l1inear ft.•for poor site preparation, improper planting, excessive slit size, and not properly securing fabric edges. These plantings are expensive and long-term, therefore, quality work is essential. A $0.01/ linear ft. incentive payment may be made for sites where quality in site selection, site preparation, and planting has been emphasized. Planting Dates: Between March 20 and May 15. Pre-PIant Treatment: Sites must be tilled, preferably in fall and then rototilled in spring. Tillage must be to bare soil with little residue remaining and must be deep enough to kill existing vegetation. Approyed Species: American Plum (bare root), Rocky Mt. Juniper, E. Red Cedar (potted). Golden willow may be used in sites previously inundated by water. One or two rows (maximum) may be planted to choke cherry or sumac (quail bush) within thickets. �60 Between-row Spacing: A maximum of 10 ft. spacing will be permitted (8 ft. recommended) for shrub thickets. A maximum of 15 feet will be permitted for wind breaks. In-row Spacing: A maximum of 8 ft. (6 ft. recommended) will be permitted for shrub thickets, and 12 feet for evergreens within wind barriers. Replacement Planting: Payment will be at the actual cost of seedlings (State Forest Service price less quantity discount) plus $0.40 per seedling for labor (this includes travel time between sites), and $0.11 per wire staple (1 or 2 per seedling). Labor will not be funded for landowners replanting their own sites. SUPPLEMENTAL Purpose: PAYMENTS FOR CUSTOM SITE PREPARATION To prepare planting sites when the landowner does not have the proper equipment. Treatment: Breaking out small tracts within CRP or sodded waste areas with a mold-board plow, sweep plow, heavy disc, rototiller, or combinations of these, to completely destroy existing vegetation for reseeding to switchgrass, planting sorghums, or site preparation for thickets and windbreaks. This does not include previously farmed (planted) areas. Tillage must be to a depth of at least 6 inches . . Payment Rate: Payment rate will be $18.00/acre for preparation of sites greater than 1 acre in size, which may involve two to three treatments. The payment rate will be $24.00/acre for preparing small sites (less than I acre) for shrub thickets/windbreaks where a rototiller is used because of the extra time needed per acre. Equipment transportation to and between will be paid at! $15.00lhour. ·PERENNIAL GRASS AND GRASS-LEGUME PLANTINGS Switchgrass provides tall cover that stands well over winter. Small, unfarmed tracts containing short, sodded grasses, are recommended for revegetation to switchgrass. Other shorter, 0001- season grasslegume mixtures may be used in roadsides where snowdrift is a problem. This practice is funded only in farmland (not rangeland) settings. Payment Rate: $6O.00/acre as a one-time payment for sites up to 5 acres, $5O.00/acre for additional area to 10 acres, and $35.00/acre above 10 acres (25 acres maximum). Payment will be $12.00/acre for heavy double discing to destroy sod, and $6.00/acre for each subsequent tillage (if needed) or harrowing. Payment will be $12.00/acre for applying either atrazine at 1 qt.lacre or Roundup at 1.5 pints/acre onto tilled soil. Use of herbicides instead of tillage to destroy perennial vegetation will not be funded. Preplant Soil Preparation: Adequate tillage to completely destroy existing perennial vegetation and to establish a firm, weed-free seed bed is required. A partial kill will not work. We recommend planting a tall-sorghum mix (See payment rates in subsequent section) the first year following tillage. Switchgrass can be seeded into the residual sorghum without tillage during the subsequent spring if the perennial vegetation was previously destroyed. If not, shallow tillage prior to drilling switchgrass is essential. Atrazine herbicide, at 1 qt./acre, applied either preplant or pre-emergent to suppress annual weeds, is strongly recommended to establish switchgrass. Roundup herbicide (1.5 pints/acre) can be applied to kill annuals just before switchgrass seedlings emerge but is not as effective as atrazine. Planting Procedures: Procedures outlined in the Division's Game Information Leaflet #113 should be reviewed. About 20 pure live seeds/if (2 - 3 lbs/acre) should be planted using a drill with double-disk furrow openers, l-inch depth bands, and packer wheels. If atrazine herbicide is not used, up to lIb/acre of an adapted dryland alfalfa (Spredor III) and up to 112 lb of sweet clover should be added. �61 Approved Species: In plots, switchgrass should comprise at least 75% of the live seed (alfalfa and sweet clover are approved additions). Within roadsides, switchgrass is the priority species where snowdrift is not a problem. Where these can not be used the tallest wheatgrasses (tall or intermediate) the roadsite site will allow, should be used in combination with alfalfa (1 to 2Ibs/acre). Planting Dates: Warm-season grasses including switchgrass: legume Mixtures: March 15 - July 15 March 15 to May 25: Cool-Season Grass- Plot Duration: Grass and grass-legume plantings must remain ungrazed and undisturbed for at least 7 years. Roadsides should remain unmowed unless essential to reduce snowdrift. If essential, mowing should be delayed until 1 August and restricted to the road shoulder. Prescribed burning, thinning tillage, or other rejuvenation treatments may be applied after 7 years. DISTURBANCE TILLAGE AND TALL WIW ANNUALS Wild sunflowers, kochia, pigweed, and other tall annuals which attain 4 to 6 ft. height stand better through winter than other herbaceous vegetation, and provide excellent cover for broods, protection from blizzards and predators, and supplemental food. This is the most effective and least expensive approach for increasing pheasants and other upland game birds. Fallow land that is left idle usually converts to annual grasses or dog-hair stands of weeds by the 2nd year following tillage. Therefore, at least one tillage in mid-spring is usually needed to promote growth of tall annuals and a second thinning tillage is sometimes needed. Maximum Funded: 14 acres/0.25 section, 28 acresllandowner and section. Plots larger than 3 ac. should be at least 0.25 mi. apart. Funding Rate: $3O.00/year for patches 0.1 to 0.5 acres in size, patches larger than 0.5 acre are considered 1 acre. $4O.00/acre/year for sites up to 5 acres (7 ac in pivot corners). $3O.00/acre/year for additional acres up to 10 (6th-l0 acres). Seeding wild sunflower or other approved wild annuals at 2 to 4 lbs. per acre will be funded at direct seed costs (see seed sources below). Plot Dimensions: Short, relatively wide patches, which will not be easily inundated by drifting snow, are preferred. Placement: Adjacent to woody cover when possible. Draw bottoms that already contain weeds and above average moisture are ideal. Sites containing noxious perennials should be avoided. Specifications: Initial tillage with a disk plow or mold-board plow is needed in sites containing perennial grass to destroy all perennial cover, preferably, immediately after the ground has thawed in early March. Large clod size is preferred to retain thin stands of annual forbs. Initial tillage in subsequent years should be conducted prior to May 1. A second thinning tillage may be used prior to the 1st of June. Spring tillage is needed each year to retain tall annuals. Annual grasses usually dominate if tillage is not used each spring. Wild sunflowers can be drilled or broadcast and harrowed at low rates to help establish tall annuals, if they are not already present. Known sources in Colorado include the Arkansas Valley Seed Company - Denver & Longmont, and Sharp Bros. Seed Company - Greeley. Retention: Tall annuals must remain undisturbed through March of the following year. Sites should be prepared for the next year's growth during mid- to late-April if weedy cover exists. �62 ANNUAL SURVIVAL PLANTINGS - SORGHUMS & DRYLAND CORN APPLICATION: On CRP, other cropland, or tilled wasteland. Annual sorghums are recommended the first year that wasteland is broken out before it is converted to switchgrass during the 2nd growing season. When applied within CRP fields, NRCS specifications for CP-12 must be used (see supplement). Dryland com is primarily applicable in center-pivot corners. MAXIMUM FUNDED: least 114 mile apart. 1 plot/80-acre field, 2 plots/16O acres, 4 plots (28 ac.)/section. Plots must be at PAYMENT RATE: $4O.00/acre/year for 1 - 5 acres (7 acres within center pivot corners) and $25.00/acre/year for additional acreages in tracts larger than 5 acres (12-acre maximum). $3O.00/acre/year if planted after June 20th. $7.50/acre for application of 30 lbs of nitrogen/acre. $18.00/acre will be paid for breaking out sod in CRP or heavily sodded sites and supplemental discing prior to planting. Landowners can not be paid for labor/tillage on their own land other than for breaking out sod. PLACEMENT: Plots should be within or near cropland and placed crosswise to prevailing winds. SPECIFICATIONS: Preplant Soil Preparation: Initial treatment: vegetation in early spring prior to annual growth. Adequate tillage to destroy existing perennial Subsequent years: Preferably minimum tillage shredding of old materials as needed prior to April 25. Annual application of nitrogen at 30 - 40 lbs.lac. is recommended. Plot Dimensions: Minimum total plot width shall be 150 feet. Wider strips are preferred to reduce impacts of drifting snow. (See restrictions on dimensions in CRP). Row Spacing: Sorghums - 15 to 30 inches; Dryland com - 30 to 36. Seed Specifications: Sorghum Patches - At least 60% (75% preferred) of an adapted tall forage sorghum that will stand well with minimal lodging and will mature before frost. Up to 40% can be adapted varieties of grain sorghum. These can be mixed or planted in separate rows (i.e., 2 rows of grain sorghum to 6 rows of forage sorghum. These sorghums should equal a minimum of 75 % of the total weight. Maximum amounts for other grains include: Dryland com (25%), safflower (25%), sunflowers (10%) and proso millet (10%). Addition of 1 to 2lbs.lac. of wild sunflower seed is recommended (Source: Arkansas Valley Seed Company - Denver). Dryland Com Plots - Early maturing dryland varieties adapted to NE Colorado. Seed, from these varieties, that is one year removed from purchased hybrid can be used to reduce seed cost. Planting Dates & Rates: Sorghums - Between April 25 and June 15; Mid to late May is recommended. Plantings conducted after June 15 may be assessed a $1O/acre payment reduction. Sorghums should be planted at 48 lbs.lacre (30-inch rows) and at higher rates if drilled . . Dryland com - Between April 25 and May 15. Plantings after June 1 will not be accepted for payment. Seeding for dryland varieties should be from 10,000 to 13,000 seeds/acre. At least one cultivation is needed for com and fertilizer should be used. �63 Plot Duration: following year. 1 year. Sorghum and corn plantings must remain undisturbed through March of the SUPPLEMENT FOR SORGHUM PLANTINGS WITHIN eRP NRes Notification: The CRP contract must be amended at the local NRCS office prior to implementing CP-12 and breaking out food plots within CRP. This requires filling out a one-page form at your NRCS office. The FSA must be advised of the change for their records. Dryland corn is not approved for plots within CRP. Winter cover-food plots within CRP must be replicated until the end of the CRP contract. If the farmer wishes to discontinue this practice he must reestablish grass (required by the NRCS). Payments will be made annually. Reimbursement will be at $4O.00/acre to cover reseeding grass unless Division of Wildlife personnel do the work. Maximum Funded: 1 plot/80-acre field, 2 plots/l60 acres. Plots must be at least 114 mile apart. Maximum Size: The maximum size is 3 acres per site. CRP fields must contain at least 40 acres to be eligible for a CP-12 food plot. Plot Dimensions: Plantings may be up to 200 feet wide (100 ft in sandy soils). Typical3-acre plots measure 198 x 660 feet. Where a 100 ft maximum is required a 3O-ft wide buffer of untilled grass is left between two 99 x 660 ft parallel strips to obtain a 3- acre plot. Smaller plots should have reduced length to retain at least the ISO ft. minimum width. For example, a plot 99 ft wide x 440 ft. long equals 1 acre and two adjacent plots will exceed the minimum width requirement. Placement: Preferably within 50-100 yds of edge and near cropland, but location can vary depending on soil, wind, and moisture, and location of other winter covers if they occur. Sorghum plantings are not permitted in soils containing free lime (shows effervescence), or soils that are deep sands or choppy sands. Plot Duration: One year. Sorghum and corn plantings in CRP must remain undisturbed through March of the following year. ��65 Colorado Division of wildlife Wildlife Research Report April 1998 JOB PROGRESS REPORT State of: Colorado Project: W-167-R Ayian Research Work Plan: Job Title: ~J~0~b~.1~9~ Implications of Habitat Loss and Fragmentation strategies for Gunnison Sage Grouse Period COvered: 01 January through 31 December _ on Conservation 1997 Author: Sara J. Oyler-McCance Personnel: Clait E. Braun, Colorado Division of Wildlife; Oyler-McCance, Colorado State University Sara J. ABSTRACT DNA was successfully extracted from all samples. A subset of the samples was used to screen microsatellite primers. Primers which looked promising were used in polymerase chain reaction (PCR) reactions and parameters in those reactions were optimized to produce clear products. screening revealed 4 microsatellites for use in this study. A PCR product from each individual sage grouse (Centrocercus minimus) from southwestern Colorado and 20 individuals each from 5 sage grouse (C. urophasianus) populations in northern Colorado was amplified for each of the 4 microsate1lites used in the analysis. In every microsatellite there were fewer alleles among the small-bodied b~rds than among the large-bodied birds, and 3 of 4 microsatellites showed evidence of a lack of gene flow between large and small-bodied sage grouse. A portion of the control region of the mitochondrial DNA was also seqUenced for all birds. We found 17 hap10types among the large-bodied birds, 4 of which were represented in almost every 1arge~bodied population. Small-bodied sage grouse had only 3 haplotypes, 1 of which was common among the large-bodied birds and 2 which were unique to the small-bodied birds. Geographic information system (GIS) coverages of the present distribution of sage grouse in southwestern Colorado, present distribution of sagebrush-steppe habitat in southwestern Colorado, and paved roads in southwestern Colorado were created. A user-friendly model continues to be developed which will allow a user to investigate different conservation strategies for Gunnison sage grouse. Additional coverages will be added and development of conservation strategies will continue in 1998-99. ��67 IMPLICATIONS OF HABITAT LOSS AND FRAGMENTATION ON CONSERVATION STRATEGIES FOR GUNNISON SAGE GROUSE Sara J. Oyler-McCance INTRODUCTION There are serious knowledge gaps which impede development of a conservation plan for Gunnison sage grouse. First, it is not known how much sage grouse habitat has already been lost and how much might be lost in the future given human population growth and land development. This is essential information if a balance between human growth and sage grouse conservation is to be achieved. Second, little is known about landscape level habitat requirements of sage grouse living in fragmented habitats. Questions such as: how large must a habitat patch be to support sage grouse?, can a well-connected network of small patches support a sage grouse population?, and are some interpatch matrices more detrimental to sage grouse than others? need to be answered before a landscape level conservation plan can be developed. Third, little is known of the movement of young sage grouse, as only one study has addressed this issue. Dunn and Braun (1985) measured natal dispersal of sage grouse in contiguous but altered habitats of northwestern Colorado and found average dispersal distances of 8.8 km for juvenile females and 7.4 km for juvenile males. It is not known, however, whether Gunnison sage grouse move among fragmented habitats (across distances up to 300 km) or whether some populations in southwestern Colorado are truly isolated. The amount of inbreeding in small, isolated populations is also unknown. Young (1994) measured inbreeding within the Gunnison population, yet that population is large compared to other populations of this species. Knowledge of movement among patches and the amount of inbreeding would provide essential information about potential inbreeding effects and aid in any conservation plan which addresses reintroductions. P. N. OBJECTIVES The objectives of this study are to (1) develop a habitat-based model which can be used to predict sage grouse occupancy of patches in southwestern Colorado, (2) investigate gene flow among isolated populations using rnicrosatellites as a molecular marker as well as sequencing the control region of the mitochondrial genome, (3) determine the level of inbreeding within populations using the aforementioned techniques, (4) document the loss of sagebrush-steppe habitat using aerial photography, and (5) develop a spatially explicit model which integrates information gained in the previous portions of the study to be used to assess potential conservation strategies of Gunnison sage grouse. SEGMENT OBJECTIVES 1. Review literature pertinent to the objectives of this study. 2. Finish extracting DNA from southwestern sage grouse samples. �68 3. Finish screening microsatellite primers for loci polymorphic in sage grouse. 4. Using the primers found to be effective, amplify micro satellites for all individuals in southwestern Colorado and 20 individuals each from 5 populations in northern Colorado. 5. Sequence the control region of the mitochondria for all southwestern sage grouse samples. 6. Continue developing GIS coverages for use in GIS-based model. 7. Order aerial photos for documentation of the loss of sagebrush steppe habitat. 8. Prepare annual report. METHODS Genetic Analysis DNA Extraction Fifty J..lIof blood or the bottom 2 em of the feather shaft were placed into 500 J..lIof 10mM Tris-HCI pH 8.0. Forty J..lIof 0.40 Proteinase K was then added and the mixture was vortexed. Twenty-six J..lISDS was then added and samples were then rotated for 3 hours at 50 C. The digested material in each tube was then extracted 3 times with an equal volume of phenol.chloroform.isoamyl alcohol (25:24:1), and once with chloroform:isoamyl alcohol (24:1). One hundred J..lIof IOmg/ml RNase was added, the mixture was vortexed, and incubated for 3 hours at 37 C. The material was then extracted twice with an equal volume of phenol:chloroform:isoamyl alcohol (25:24:1), and once with chloroform:isoamyl alcohol (24:1). The resulting aqueous solution of genomic DNA was ethanol-precipitated, and resuspended in TE (Tris-HCII0mM, EDT A ImM pH 8.0), to a concentration of approximately 300 ug/ml, Microsatellite Analysis Microsatellite primers were obtained from two sources. The first group of primers were those designed for use with poultry chickens and were obtained from Hans Cheng at the Poultry Research Group. The second group of primers were designed for use with red grouse (Lagopus lagopus scoticus) and were obtained from Stuart Piertney at the University of Aberdeen. Microsatellite primers were radioactively labeled for later visualization on autoradiography film. The T4 Polynucleotide Kinase (PNK) labeling procedure was used. One primer (either the forward or reverse primer) was chosen and radioactively labeled using the following established procedures. In a 150 J..lIEppendorftube, 1 J..lII0 M primer, 1 J..lIIOXBuffer, 0.25 J..lIT4 PNK' 0.25 J..lIp33 A-ATP, and 7.5 J..lIH20 were mixed and incubated at 37 C for 15 minutes. The reaction was stopped by heating to 70 C for 10 minutes. Screening micro satellite primers consisted of choosing 4 - 12 DNA samples (representing at least one individual from each �69 population, plus at least one individual from Cold Springs Mountain to use as an outgroup) and amplifying the micro satellite region using PCR. PCR reactions were performed in a PerkinElmer DNA thermal cycler. Approximately 30 ng of genomic DNA (in a 1 III volume) was used in each reaction. Each PCR reaction was performed in a total reaction volume of 25 Ill, using 1.25 III of the forward and 1.25 III of the backward primer at 1.0 11M,dNTPs at 25 11Meach nucleotide, and 0.5 U Taq polymerase in a Ix Taq buffer (67 mM Tris-HCl pH 8.0, 6.7 mM MgS04, and 16.6 mM AmS04, 10mM beta-mercaptoethanol). The reaction was overlaid with 2 drops of light mineral oil to prevent evaporation, and amplified for 35 cycles with a touchdown thermal profile which starts with a 2 minute denaturation at 94 C, then cycled 35 times through the touchdown: 30 seconds of a denaturation (94 C), and 30 seconds of annealing (stepping from 60 C to 50 C). When 35 cycles had been completed, a 20 minute extension at 74 C occured. PCR products were then electrophoresed at 55 watts for 2 hours through 6% poly-acrylamide gels as described in Sambrook et al. (1989). Audioradiograms were made of each dried acrylamide gel by exposure to X-ray film (Fuji RX). Screening the primers revealed 4 micro satellites that were optimal for this study. These 4 primers, (LLSTl, LLSD3, LLSD4, and LLSD8) were designed by Piertney and Dallas (1997). Each micro satellite was then amplified for all individuals from southwestern Colorado as well as approximately 20 individuals from each of 5 northern Colorado populations (Blue Mountain, Cold Springs Mountain, Eagle, North Park, and Middle Park). DNA from these northern Colorado individuals was obtained from Nate Kahn and Tom Quinn at the University of Denver. PCR products were then electrophoresed at 55 watts for 2 hours through 6% poly-acrylamide gels as described in Sambrook et al. (1989). Audioradiograms were made of each dried acrylamide gel by exposure to X-ray film (Fuji RX). Individuals were assigned genotypes (corresponding to micro satellite fragment length) based on banding patterns on the audioradiograms. The distribution of allele frequencies for each population was then recorded. Mitochondrial DNA Sequencing Analysis Two primers, 16775L and H521, were chosen to amplify a portion of region I of the control region of mitochondrial DNA which was approximately 500 base pairs long. These primers are described by Quinn and Wilson (1993). We used a third primer, 418H (Quinn and MindellI996), to conduct a semi-nested-Pf'R.' PCR amplifications were performed in a Perkin-Elmer DNA thermal cycler. For the initial double-stranded PCR, 1 III of DNA template (approximately 30 ng of genomic DNA) was combined with 24 III of cocktail mixture consisting of: primers 16775L and 521H at 1.0 11M, dNTPS at 250 11Mfor each nucleotide, and 0.5 U Taq polymerase in IX Taq Buffer (67mM TrisHCL pH 8.0, 6.7 mM MgS04, 16.6 mM AmS04, 10 mM beta-mercaptoethanol). Two drops of ultraclean light mineral oil were added to each reaction tube to prevent evaporation. The samples were then amplified for 30 cycles using the following thermal profile: denaturation, 92 C for 40 seconds, annealing, 55 C for 1 minute; extension, 72 C for 2 minutes. A final extension period of 5 minutes was performed at the completion of the 30 cycles. The samples were then stored at 4 C. Five III of each double stranded product was then electrophoresed through a Nusieve 2% agarose gel in IX TA (0.04 M Tris-acetate buffer) and visualized with ethidium bromide under UV light. �70 A plug of the double stranded PCR product was then removed from each band and melted in 100 H20 at 65 C for 2 minutes. Two III of this diluted double stranded PCR product were then used as template for the single stranded PCR reaction. This template was then used in a 50 III single stranded nested PCR, using primers 16775L and 418H in the aforementioned cocktail. The single stranded products were amplified for 35 cycles using the following thermal profile: denaturation, 92 C for 40 seconds, annealing, 55 C for 1 minute; extension, 72 C for 2 minutes. A final extension period of 5 minutes was performed at the completion of the 35 cycles. The single stranded PCR products were then stored at 4 C. III of ultraclean Each single stranded PCR product was cleaned using three distilled H20 washes in Millipore Ultrfree-MC-30 microcentrofuge tubes. The single stranded DNA was then recovered from the Millipore filters using 35 III of distilled H20. The clean single stranded PCR products were then sequenced using a Sequenase 2.0 kit (USBiochemicals). Products from the sequencing reaction were then electrophoresed through a 6% straight poly-acrylamide gel at 55 watts for 2.5 hours as described in Sambrook et al. (1989). The gels were then dried and autoradiograms were made of each dried gel by exposure to X-ray film (Fuji RX). For each individual, the DNA sequence (141 base pairs) was then read and a haploptye was assigned. Spatially Explicit Model A GIS coverage of the distribution of sage grouse was developed by digitizing known locations of sage grouse populations in southwestern Colorado. A GIS coverage of the current distribution of sagebrush-steppe habitat was obtained from Suzanne Noble who had classified sagebrush-steppe habitat from a series of satellite images. A coverage of paved roads was created by downloading digital line graph files containing transportation information from the EROS data center. These files were then linked together in ARCIINFO and the appropriate information was accessed and saved as a separate coverage. Additional information (e.g., genetic data, census data) continues to be added so that an interactive program can be developed allowing the user to investigate different conservation strategies for Gunnison sage grouse. A program is being written in C language to interface with the GIS coverages. RESULTS Genetic Analysis Microsatellite Analysis Because we amplified 4 different micro satellites we obtained results for 4 different micro satellite loci. The distribution of alleles for each microsatellite are shown in Tables 1 - 4. For micro satellite LLST1, there were 5 different alleles in the large-bodied birds and only 2 alleles in the small-bodied birds (Fig. 1). Although the allele frequencies for Gunnison Basin, Crawford, Eagle, Cold Springs, and Blue Mountain are similar, suggesting there could be gene flow among the small and large-bodied birds, the results from the next 3 micro satellites suggest otherwise. The results for micro satellite LLSD8 showed 4 alleles in the large-bodied birds and only 2 in the small-bodied birds (Fig. 2). Little variation exists at this locus among the small- �71 bodied birds, with only 1 allele being dominant and represented in all populations. The 3 dominant alleles (137, 143, and 157) among the large-bodied populations were found in similar frequencies, suggesting some amount of gene flow among these populations, yet not between the large and small-bodied populations. Like LLSD8, micro satellite LLSD3 was more variable in large-bodied birds (Fig. 3), with 3 dominant alleles (133, 137, 141) well represented in all populations. Only 2 alleles were common to all small-bodied populations (133, 135). Microsatellite LLSD4 was the most variable micro satellite with 33 different alleles among all populations (Fig. 4). While there is variability among the small-bodied birds (10 different alleles), they remain less variable than the large-bodied populations (30 different alleles). Four alleles were dominant and found in all large-bodied populations (187,189, 193, 195). Only 1 of those alleles (193) was found at all among small-bodied birds and there were 3 alleles (199, 201, 217) which were unique to the small-bodied birds. We conducted F tests for each loci to determine whether the distributions of alleles were significantly different between the large and small-bodied birds. Three microsatellite loci showed a significant difference between the two groups of birds (LLSD3 F= 5.95, P<O.OOI; LLSD4 F=2.51, P<O.OOI; LLSD8 F=6.80, P<O.OI) and 1 did not (LLSTI F=0.983, P>0.05). Genetic distances and inbreeding coefficients will be calculated for these data to determine gene flow and inbreeding among the small-bodied sage grouse. Mitochondrial DNA Sequencing Analysis The distribution ofhaplotypes among a119 populations in Colorado are shown in Table 5. In total, there were 19 different haplotypes across all individuals. We found the 5 large-bodied populations all had a number of different haplotypes in each population while the small-bodied grouse populations had only 2 or 3 haplotypes per population (Fig. 5). Among the large-bodied populations, we found 4 dominant haplotypes (A, B, C, and D). Haplotypes B, C, and D were common in all large-bodied populations and haplotype A was found in all but 1 large- bodied population. In the small-bodied populations, only 1 of these haplotypes, A, was found. Further, the G and AI haplotypes were unique to the small-bodied birds. To test whether the distribution ofhaplotypes from the large-bodied populations differed from the distribution ofhaplotypes from the small-bodied populations, we used an F test. We found there was a statistically significant difference between the distribution ofhaplotypes in the large and small-bodied populations (F = 4.478, P<O.OOI). These data will be combined with the micro satellite data and used to document gene flow and inbreeding coefficients. Spatially Explicit Model GIS layers will continue to be developed as the data becomes available (e.g., genetics data and data from analysis of habitat loss). DISCUSSION An examination of the distribution and frequency ofhaplotypes found in this study allows �72 some inferences to be made about recent patterns of gene flow, and the recent evolutionary history of sage grouse. Among the small-bodied birds, relatively little genetic variation was found (3 mitochondrial haplotypes and 15 micro satellite alleles), while large-bodied birds were found to have relatively high genetic variability (17 mitochondrial haplotypes and 44 microsatellite alleles). If gene flow between the large and small-bodied birds was occurring we would expect from the mitochondrial data to find an array ofhaplotypes among the small-bodied birds similar to those found in every large-bodied population. Similarly, we would expect from the micro satellite data to find the alleles dominant in the large-bodied birds to be present among the small-bodied birds. The extreme differences in distributions of alleles and haplotypes between the large and small-bodied sage grouse support the idea that there is no gene flow between two and is consistent with Braun and Young's (in review) proposal that the small-bodied sage grouse of southwestern Colorado are a separate species. Further, the low genetic variability among smallbodied sage grouse may have important implications for the conservation of that species. Genetic distances between all pairs of populations will be calculated so that amount of gene flow among populations (particularly among the small-bodied birds) can be elucidated. LITERATURE CITED Dunn, P.O., and C. E. Braun. 1985. Natal dispersal and lek fidelity of sage grouse. Auk 102:621-627. Piertney, S. B., and J. F. Dallas. 1997. Isolation and characterization ofhypervariable micro satellites in the red grouse Lagopus /agopus scoticus. Journal of Molecular Ecology 6:93-95. Quinn, T.W., and A.C. Wilson. 1993. Sequence evolution in and around the control region in birds. Journal of Molecular Evolution 37:417-425. . Quinn, T.W., and D.P. Mindell. 1996. Mitochondrial gene order adjacent to the control region in crocodile, turtle, and tuatara. Molecular Phylogenetic Evolution 5:344-351. Sambrook, J., E. F. Fritsch, and T. Maniatis. 1989. Molecular cloning: a Laboratory Manual. 2nd edition. Cold Spring Harbor Laboratory Press, New York. Young, J. R 1994. Sexual selection of sage grouse. Ph.D. Dissertation, Purdue Univ., West Lafayette, IN. 123 pp. Prepared by: _-,~.q-._-t}.,__Oc~~A---= Sara J. Oyler-McCance I_14_<!_~_~_~__;.::__ _ �Table I. Allele distributions for microsatellite LLSTl among 9 populations of sage grouse in Coloroado. Crawford Gunnison Cold Springs Dove Creek Dry Creek CRl 154 157 DYCl 154 154 DVCl 154 154 CSl 154 154 CR2 154 154 DYC2 154 154 DVC2 154 154 CS2 GB3 154 154 CR3 154 154 DYC3 154 154 DVC3 154 154 GB4 154 157 CR4 154 154 DYC4 154 154 DVC4 154 154 GBl 154 GB2 154 Blue Mountain BM2 154 154 North Park Middle Park Eagle NPl 154 154 MPl 154 154 EGl 154 154 NP2 154 163 MP2 154 157 EG2 154 154 154 154 BM3 CS3 154 157 BM4 154 154 NP3 154 163 MP3 154 154 EG3 154 157 CS4 154 154 BM5 154 154 NP4 154 163 MP4 154 157 EG4 154 157 157 GB5 154 157 CR5 154 154 DYC5 154 154 DVC5 154 154 CS5 154 154 BM6 154 157 NP5 MP5 154 154 EG5 157 GB6 154 154 CR6 154 157 DYC6 154 154 DVC6 154 154 CS6 154 154 BM7 154 154 NP6 154 154 MP6 154 154 EG6 154 154 GB7 154 157 CR7 154 154 DYC7 154 154 DVC7 154 154 CS7 154 154 BMB 154 154 NP7 154 163 MP7 154 154 EG7 154 154 GBB 154 157 CR8 154 157 DYC8 154 154 GB9 154 154 CR9 154 154 DYC9 GGl 154 154 CR10 154 154 DYC10 154 GG2 154 154 CR11 154 157 DYCll 154 GG3 154 157 CR12 154 157 DYC12 154 GG4 157 157 GG5 DVC8 154 154 CS8 154 157 BM9 154 154 NP8 154 154 MPB 154 154 EGa 154 154 DVC9 154 154 CS9 154 154 BM10 154 154 NP9 154 154 MP9 154 163 EG9 154 157 154 DVC10 154 154 CS10 154 154 BM11 154 157 NP10 154 154 MP10 154 DVCll 154 154 CSll 154 154 BM12 154 154 NPll 154 157 MPll 154 154 EGll 154 154 154 DVC12 154 154 CS12 154 157 BM13 154 157 NP12 154 154 MP12 151 157 EG12 154 157 EG10 CR13 154 154 DYCFl 154 154 DVCFl 154 154 eS13 154 154 BM14 154 157 NP13 154 157 MP13 154 157 EG13 154 154 CR14 154 154 DYCF2 154 154 DVCF2 154 154 CS14 154 157 BM15 154 157 NP14 154 157 MP14 154 154 EG14 154 157 154 157 DYCF3 DVCF3 154 154 CS15 154 157 BM16 154 157 NP15 154 154 MP15 154 154 EG16 154 154 154 154 eS16 154 154 BM17 154 154 NP16 154 154 MP16 151 157 EG18 154 154 BM18 154 154 NP17 154 157 MP17 154 157 EG19 154 157 DYCF6 154 154 eS18 154 157 BM19 154 154 NP18 154 154 MP18 154 154 EG20 157 157 DYCF7 154 154 eS19 154 154 BM20 154 154 NP19 154 154 MP19 151 154 EG21 154 157 GG6 154 154 CR15 GG7 154 157 CR16 DYCF4 154 154 eS17 DYCF5 GG8 154 157 CEFl GG9 154 154 FM2 GG10 154 157 FM3 GGll 154 154 FM4 DYCF8 154 154 eS20 154 157 BM21 154 154 NP20 154 154 MP20 154 154 EG22 154 154 GG12 154 154 FM5 DYCF9 154 154 CS21 154 154 BM22 154 154 NP21 154 154 MP21 154 157 EG23 154 154 GG13 154 154 DYCF10 154 154 eS22 BM23 154 154 NP22 154 157 EG24 154 154 eS23 BM24 154 154 NP23 BM25 154 154 NP24 154 157 NP25 154 157 GG14 154 157 GG15 154 154 eS24 154 157 GG16 154 157 eS25 157 157 GG17 154 157 eS26 154 157 GG18 154 154 GG19 154 157 GG20 154 154 -.J W �...,J "'" Table 2. Allele distributions for microsatellite Dove Creek Dry Creek Crawford Gunnison LLSD8 among 9 populations of sage grouse in Coloroado. Cold Springs Blue Mountain GBl 143 143 CRl 143 143 DYCl 143 143 DVC1 143 143 CSl 137 137 BM2 GB2 137 143 CR2 143 143 DYC2 143 143 DVC2 143 143 CS2 137 143 GB3 143 143 CR3 143 143 DYC3 143 143 DVC3 143 CS3 143 GB4 143 143 CR4 143 143 DYC4 143 143 DVC4 143 CS4 GB5 143 143 CR5 143 143 DYC5 143 143 DVC5 143 GBS 143 143 CR6 143 143 DYC6 143 143 DVC6 143 GB7 143 143 CR7 143 143 DYC7 143 143 DVC7 143 GB8 143 143 CR8 143 143 DYC8 143 143 DVC8 143 GB9 143 143 CR9 143 143 DYC9 143 143 DVC9 143 GGl 143 143 CR10 143 143 DYC10 143 143 DVC10 143 GG2 143 143 CRll 143 143 DYC11 143 143 DVC11 143 GG3 143 143 CR12 143 143 DYC12 143 143 DVC12 143 1G 1G. 1G 1G 1G lG lG 1G lG lG GG4 143 143 CR13 143 143 DYCFl 143 143 DVCF1 143 GG5 143 143 CR14 143 143 DYCF2 143 143 DVCF2 GG6 143 143 CR15 143 143 DYCF3 DVCF3 GG7 143 143 CR16 143 143 DYCF4 143 143 GG8 143 143 FMl DYCF5 CS17 GG9 143 143 FM2 DYCF6 CS18 137 GG10 143 143 FM3 DYCF7 143 143 CS19 GGll 143 143 FM4 DYCF8 143 143 CS20 GG12 143 143 FM5 DYCF9 143 143 CS21 GG13 143 143 CRFl GG14 143 143 CS23 GG15 143 143 CS24 GG16 143 143 CS25 GG17 143 143 CS26 143 GG18 143 GG19 143 143 GG20 143 143 143 143 DYCF10 137 143 North Park Middle Park Eagle NPl 137 143 MPl 137 143 EGl 137 143 BM3 NP2 137 157 MP2 137 157 EG2 137 143 157 BM4 NP3 137 137 MP3 137 143 EG3 137 143 137 143 BM5 137 157 NP4 137 157 MP4 137 157 EG4 137 157 CS5 137 157 BM6 137 137 NP5 137 157 MP5 143 157 EG5 137 157 CS6 143 157 BM7 137 137 NP6 137 157 MP6 137 157 EG6 137 157 CS7 137 157 BM8 137 163 NP7 137 143 MP7 137 143 EG7 137 137 CS8 143 137 BM9 137 157 NP8 137 137 MP8 137 157 EG8 137 137 CS9 157 157 BM10 143 143 NP9 143 1557 MP9 137 137 EG9 137 157 CS10 137 143 BMll 137 137 NP10 137 137 MP10 137 143 EG10 csn 137 137 BM12 137 157 NPll 137 157 MP11 137 143 EG11 137 137 CS12 137 157 BM13 137 157 NP12 137 157 MP12 137 157 EG12 137 137 143 CS13 137 143 BM14 137 137 NP13 137 137 MP13 137 157 EG13 137 157 143 lG CS14 137 157 BM15 137 137 NP14 137 143 MP14 137 143 EG14 137 157 143 143 CS15 137 157 BM16 137 157 NP15 137 143 MP15 137 137 EG16 137 157 CS16 157 157 BM17 143 157 NP16 137 137 MP16 137 143 EG18 137 143 BM18 137 157 NP17 143 143 MP17 137 157 EG19 137 143 143 BM19 137 143 NP18 143 157 MP18 137 137 EG20 137 157 137 143 BM20 137 157 NP19 137 157 MP19 137 143 EG21 137 137 157 157 BM21 137 157 NP20 137 ·143 MP20 137 143 EG22 143 157 137 137 BM22 157 157 NP21 137 137 MP21 EG23 143 157 BM23 137 157 NP22 137 157 EG24 137 143 BM24 137 143 NP23 137 143 BM25 137 137 NP24 137 143 NP25 143 143 CS22 143 143 �Table 3. Allele distributions for microsatellite LLSD3 among 9 populations of sage grouse in Coloroado, Dove Creek Dry Creek Crawford Gunnison Cold Springs Blue Mountain GBl 135 137 CRl 133 133 DYC1 135 135 DVCl 133 135 CS1 133 137 BM2 GB2 133 135 CR2 133 133 DYC2 133 135 DVC2 135 135 CS2 133 141 BM3 GB3 133 133 CR3 133 133 DYC3 135 135 DVC3 135 135 CS3 133 133 BM4 141 GB4 133 135 CR4 133 133 DYC4 133 135 DVC4 135 135 CS4 133 133 BM5 GB5 133 133 CR5 133 135 DYC5 135 135 DVC5 133 133 BM6 133 133 CR6 133 135 DYC6 133 133 DVC6 135 135 CS5 GB6 133 . 135 CS6 133 133 GB7 133 135 CR7 133 133 DYC7 135 135 DVC7 135 135 CS7 133 GBB 135 135 CR8 133 133 DYCB 135 135 DVC8 135 135 CS8 GB9 133 135 CR9 133 133 DYC9 135 135 DVC9 135 135 CS9 GG1 133 135 CR10 .133 133 DYC10 133 135 DVC10 133 133 GG2 133 135 CRll 133 133 DYC11 135 135 DVC11 133 GG3 133 133 CR12 133 133 DYC12 135 135 DVC12 GG4 133 135 CR13 133 135 DYCF1 135 135 GG5 133 135 CR14 133 133 DYCF2 135 GG6 135 135 CR15 133 133 DYCF3 GG7 135 135 CR16 133 133 GG8 133 133 FMl GG9 133 135 FM2 133 GG10 133 135 FM3 GG11 133 135 FM4 GG12 133 133 GG13 133 133 GG14 133 133 133 141 North Park NP1 133 Middle Park 133 NP2 Eagle MPl 133 153 EG1 133 MP2 141 153 EG2 133 141 MP3 133 141 EG3 133 137 141 153 NP3 133 143 133 133 NP4 133 133 MP4 133 153 EG4 133 137 133 137 NP5 133 133 MP5 133 133 EG5 141 153 BM7 141 141 NP6 133 153 MP6 133 137 EG6 133 137 137 BM8 141 141 NP7 133 141 MP7 141 153 EG7 133 141 133 141 BM9 133 133 NP8 133 133 MP8 133 141 EG8 133 141 133 135 BM10 133 141 NPS 133 133 MPS 135 141 EG9 133 137 CS10 133 137 BMll 133 137 NP10 1M 141 MP10 133 141 EG10 133 141 135 CS11 133 133 BM12 133 141 NP11 133 133 MPll 133 133 EG11 133 137 135 135 CS12 133 133 BM13 133 141 NP12 1M 153 MP12 133 141 EG12 133 133 DVCF1 135 135 CS13 133 133 BM14 133 141 NP13 1M 153 MP13 133 137 EG13 133 137 135 DVCF2 135 135 CS14 133 133 BM15 133 133 NP14 133 141 MP14 133 153 EG14 133 137 133 133 DVCF3 135 135 CS15 133 133 BM16 133 133 NP15 133 141 MP15 133 133 EG16 133 137 DYCF4 135 135 CS16 135 141 BM17 133 141 NP16 133 153 MP16 133 141 EG18 133 153 DYCF5 135 135 CS17 BM18 133 153 NP17 1~ 153 MP17 133 153 EG19 133 137 133 DYCF6 135 135 CS18 135 141 BM19 133 141 NP18 133 141 MP18 133 135 EG20 133 137 133 133 DYCF7 133 133 CS19 133 141 BM20 133 141 NP19 133 133 MP19 133 137 EG21 133 141 133 133 DYCF8 133 133 CS20 133 133 BM21 133 137 NP20 133 141 MP20 133 133 EG22 133 137 FM5 DYCF9 135 135 CS21 133 141 BM22 153 153 NP21 133 133 MP21 CRFl DYCF10 CS22 BM23 133 153 NP22 133 153 133 141 CS23 BM24 133 141 NP23 133 141 BM25 133 133 NP24 133 141 GG15 135 137 CS24 GG16 133 135 CS25 GG17 133 135 CS26 GG18 133 137 GG19 133 135 GG20 135 135 EG23 EG24 NP25 -.J U1 �-..J 0\ Table 4. Allele distributions for microsatellite LLSD4 among 9 populations of sage grouse in Coloroado. Dove Creek Dry Creek Crawford Gunnison 203 DYCI 191 203 Cold Springs Blue Mountain DVCl 199 201 CSI 195 197 BM2 DVC2 191 201 CS2 195 243 BM3 GBI 191 191 CRI 191 GB2 191 201 CR2 203 203 DYC2 GB3 191 191 CR3 191 191 DYC3 191 205 DVC3 191 201 CS3 189 189 BM4 GB4 191 225 CR4 193 193 DYC4 203 207 DVC4 201 201 CS4 193 225 GB5 191 191 CR5 191 225 DYC5 215 215 DVC5 191 199 CS5 195 243 GB6 191 191 CR6 193 203 DYC6 215 215 DVC6 201 201 CS6 189 GB7 203 225 CR7 191 203 DYC7 191 191 DVC7 201 201 CS7 GB8 191 203 CR8 191 203 DYC8 191 191 DVC8 201 201 CS8 GB9 191 215 CR9 191 203 DYC9 191 215 DVC9 191 201 CS9 GGI 191 191 CR10 191 191 DYC10 191 217 DVC10 191 201 CS10 GG2 191 191 CRll 191 225 DYC11 DVCll 191 201 CSll GG3 191 203 CR12 193 203 DYC12 DVC12 191 191 CS12 193 193 North Park 207 MPI 207 207 EGI 185 195 215 MP2 193 193 EG2 185 329 189 215 MP3 189 189 EG3 189 215 197 NP3 BM5 191 289 NP4 191 205 MP4 193 215 EG4 189 245 BM6 185 329 NP5 203 215 MP5 183 209 EG5 215 267 195 BM7 281 329 NP6 205 245 MP6 EG6 187 267 193 203 BM8 189 357 NP7 203 245 MP7 193 205 EG7 219 297 195 215 BM9 267 321 NP8 193 205 MP8 183 189 EG8 189 197 BM10 189 195 NP9 203 207 MP9 187 187 205 391 BMll 189 191 NP10 207 207 MP10 187 309 EG10 195 243 BM12 195 195 NPll 205 215 MPll 189 189 EGll 215 267 197 391 BM13 189 195 NP12 205 205 MP12 183 207 EG12 187 225 203 DVCFl 191 191 CS13 197 215 BM14 309 329 NP13 205 245 MP13 189 189 EG13 191 203 DVCF2 199 201 CS14 197 215 BM15 189 193 NP14 205 205 MP14 189 193 EG14 CS15 197 215 BM16 223 267 NP15 191 245 MP15 189 193 EG16 195 267 191 217 CS16 183 197 BM17 185 323 NP16 195 205 MP16 183 195 EG18 195 215 BM18 187 289 NP17 205 239 MP17 189 205 EG19 187 267 NP18 191 205 MP18 189 193 EG20 187 193 187 NP19 187 189 MP19 187 189 EG21 189 267 329 NP20 209 245 MP20 EG22 195 267 MP21 EG23 187 205 EG24 187 215 191 CR13 191 191 DYCFl 191 215 CR14 193 203 DYCF2 GG6 215 225 CR15 193 203 DYCF3 GG7 191 191 CR16 203 203 DYCF4 GG8 191 205 FMl DYCF5 CS17 GG9 191 191 FM2 DYCF6 CS18 187. 193 BM19 GG10 191 205 FM3 DYCF7 217 217 CS19 197 255 BM20 187 GGll 193 219 FM4 DYCF8 191 191 CS20 189 255 BM21 189 GG12 191 191 FM5 DYCF9 CS21 BM22 NP21 GG13 191 203 CRFI DYCF10 CS22 BM23 NP22 GG14 191 203 CS23 BM24 NP23 GG15 191 191 CS24 BM25 NP24 GG16 191 191 CS25 GG17 193 .215 CS26 191 191 205 GG20 191 191 EG9 .191 191 191 329 187 GG4 GG19 Eagle 187 GG5 GG18 Middle Park NPI . NP2 DVCF3 NP25 �Table 5. Distribution of mitochondrial DNA haplotypes among 9 populations Haplot)l2e Population A Gunnison Basin Crawford Dry Creek Dove Creek Cold Springs Blue Mountain Middle Park North Park Eagle of sage grouse in Colorado. 38 2 4 11 3 1 4 2 B C D E G H L S X Z AA AC AD AE 2 15 6 7 8 7 5 2 10 1 9 6 15 2 1 1 3 1 2 AI AL AM 8 2 1 2 3 4 AF 1 . 1 1 1 1 2 1 3 -..I -..I �78 North Park Eagle Cold Springs 157 157 Blue Mountain 157 Middle Park 103 151 154 157 ~"" ~l;;%5;;J%ii%t#;/~~ lS. , Y.;»'_'» ....• ;-' >, >~-,' 157 ~~ilt$f±j~t;j~~ild~~J0 lS. Dry Creek Dove CreeR54 Crawford 154 Gunnison Basin Figure 1. Microsatellite allele frequencies for microsatellite LLST 1. Eagle Cold Springs 143 157 ~J&';'"' ...... North Park 143 157 '''w..... .. ~~.us~~;.;w;:N' ' .... .. .... 137 137 143 ~"." 137 Blue Mountain 163 Middle Park 143 157 143 137 137 137 143 Dry Creek 143 143 Dove Creek 143 Crawford Figure 2. Microsatellite allele frequencies for microsatellite LLSD8. Gunnison Basin �79 Eagle Cold Springs North Park H3 153 153 141 135 137 133 141 133 133 Blue Mountain Middle Park 153 153 141 135 133 137 141 133 137 135 133 135 Dry Creek Gunnison Basin 135 Dove Creek 133 Crawford Figure 3. Microsatellite allele frequencies for microsatellite LLSD3. North Park Eagle 239 Cold Springs 187 245 189 19\93 195 203 225 215 Middle Park Blue Mountain 207 209215 309 205 357 323329 321 309 289 281 183 187 185 195 267 223197 195 193 191 217 205 215 219 225 215 ~oo 191 ~«/.~~~~ -:.~, ' ,~~~-:. , " Dry Creek 191 ~:i 2m ",;<;,~ •.y/.'•.•. n" 201 199 Dove Creek Gunnsion Basin 193 Crawford Figure 4. Microsatellite allele frequencies for microsatellite LLSD4. �80 Eagle Cold Springs H L " B Z AC North Park A A '!W¢"'" ,,14 ? ~ x ~ o Blue Mountain Middle Park L At II " c , 'N~/'''''' ~ E ) 0 G A AI A Dry Creek A Dove Creek Gunnison Basin G Crawford Figure 5. Mitochondrial DNA haplotypes among sage grouse populations in Colorado. �81 Colorado Division Wildlife Research April 1998 of Wildlife Report INTERIM state of: W-167-R Work 12 Plan: Period REPORT Colorado Project: Job Title: FINAL Ayian Job Genetic Diversity Colorado Covered: 01 January Research 18 Among populations of Merriam's through 31 December 1997 Author: Richard Personnel: Richard W. Hoffman, Colorado Division Dujay, Colorado state University Wild Turkeys in C. Dujay of Wildlife; Richard C. ABSTRACT Blood samples were collected from 333 Merriam's wild turkeys (Meleagris gallopayo merriami) trapped (n = 312) and/or harvested (n = 21) between September 1995 and February 1998. Samples (n = 301) were obtained from 5 East Slope and 5 West Slope flocks in Colorado and represented 9 different counties. In addition, 32 samples were obtained from birds captured on the Mogollon Rim in north-central Arizona. DNA was successfully extracted from 331 of the 333 samples. All samples were assayed and DNA concentrations quantified using spectrophotometry. A protocol for non-isotopic Restriction Fragment Length Polymorphism (RFLP) analysis was developed for turkey blood .using the M-13mp18 phage probe and Pst 1 endonuclease. Seventy-seven bands between 33.05 and 1.61 kilo-bases were resolved for the entire sample. The mean number of bands resolved per flock and per individual were 39.5 and 10.2, respectively. Not all bands appeared in every flock. Certain bands were exclusive to one flock and occurred in every individual tested from that flock. There were differences among flocks, but when each flock was compared to the combined sample for the entire state, no differences were detected. Likewise, the Colorado birds did not differ from the Arizona birds. Birds from the Spanish Peaks flock had the greatest genetic diVersity, whereas birds from Parachute Creek were genetically distinct and may be experiencing inbreeding depression or cross breeding with domestic/game farm turkeys. No detrimental genetic conditions Were identified within the other flocks tested. A dissertation incorporating the data collected in this study has been submitted to the Department of Biology at Colorado State University in partial fulfilment for the Doctor of Philosophy degree. The dissertation will be submitted as the final r~ Prepared by, in 1999. ~ Research C. Dujay Tech I ��83 Colorado Division Wildlife Research April 1998 of Wildlife Report JOB FINAL REPORT state of: Colorado Project: W-167-R-5 Work Plan: Job Title: 13 Personnel: Job: 10 Movements, Reproductive Success, and Habitat Use by Introduced Plains Sharp-tailed Grouse Period Covered: Author: Ayian Research 01 January 1994 through 31 July 1997 Kenneth M. Giesen Jim Aragon, Clait E. Braun, Kenneth M. Giesen, Colorado Division of Wildlife Chuck Loeffler, ABSTRACT A total of 76 plains sharp-tailed grouse (Tympanuchus phasianellus jamesi) was trapped in southeastern Wyoming and transplanted onto Raton Mesa in Las Animas County, Colorado in April 1995 and 1996. Twenty males and 22 females were fitted with radio transmitters prior to release. Weather conditions at the time of release in 1995 were characterized by blizzard conditions; in 1996 weather was more moderate with cool temperatures and less than 30% snow cover on the Mesa. Documented mortality of radio-marked birds was 43% (18 of 42) within 60 days post-release and signals from another 16 birds (5 males, 11 females) were lost due to long distance dispersal or radio failure. Maximum individual dispersal from the release site ranged from 1.2 to 41.6 km with 13 of 30 birds moving ~ 5.0 km. Home ranges of 5 birds surviving at least 60 days ranged from 0.32 to 117.12 km2• Height density indices within grasslands used by radio-marked birds ranged from 1.24 ± 0.66 dm to 3.45 ± 1.35 dm. Sightings of unmarked birds and'survival of radiomarked birds indicates that spring through fall habitat on Raton Mesa meets the minimum requirements for this species, although dispersal and overwinter survival of birds may prevent sharp-tailed grouse from establishing and maintaining a population on Rat,on Mesa. No additional transplants of sharptailed grouse to Raton Mesa are recommended at this time. �84 RECOMMENDATIONS This is the second unsuccessful effort in recent years to establish a viable population of sharp-tailed grouse in Las Animas County near or on Raton Mesa. Although there was little follow-up or evaluation of the initial transplant in the late 1980's, all information suggests that transplant was a failure since no lekking sites were established and no successful reproduction was documented. Because the recent (1995-96) transplant was also unsuccessful, despite documentation of some lekking behavior and successful nesting, it is likely that either Raton Mesa is too small to sustain a viable population of sharp-tailed grouse, the habitat isn't suitable, or the transplant protocols were inadequate. Most mortality or disappearance of transplanted sharp-tailed grouse occurred shortly following their release, and in 1995 may have been exacerbated by weather conditions at the time of their release. Suitable habitat appears to be limiting on Raton Mesa, especially wintering habitat comprised of deciduous shrubs or agricultural crops preferred by plains sharptailed grouse within the core of their current range. Further, historically high levels of livestock grazing on Raton Mesa in the last 100 years may have changed both the composition and structure of the grassland habitats in this area making it unsuitable for sustaining a viable population of sharp-tailed grouse. Because of the failure of two recent transplants of sharp-tailed grouse to Raton Mesa, no further transplants are recommended to this site at this time. Further, evaluation of additional release sites for sharp-tailed grouse along the Front Range of Colorado should evaLuate potential wintering habitat as well as nesting habitat, and all habitat components should be available within a few kilometers to reduce dispersal or seasonal migration. Because few transplants of sharp-tailed grouse have been successful, future transplants need to be treated as management experiments and evaluated, and additional transplant methodologies, including releasing hens with chicks in summer should also be considered. �85 MOVEMENTS, REPRODUCTIVE SUCCESS, AND HABITAT INTRODUCED PLAINS SHARP-TAILED GROUSE USE BY Kenneth M. Giesen IHTRODtlCTIOH Plains sharp-tailed grouse historically occurred in suitable foothill and riparian habitats along the Front Range of Colorado (Bailey and Neidrach 1965, Hoag and Braun 1990). Sharp-tailed grouse populations declined with human settlement and were extirpated from most of their range in eastern Colorado by the late 1800's. Although the historical breeding population of sharp-tailed grouse in Douglas County continues to decline, small breeding populations and winter migrants or transients have been reported in recent years from Yuma, Logan, and Weld counties (Hoag and Braun 1990, C. E. Braun unpubl. data). However, the total breeding population of plains sharp-tailed grouse in Colorado remains small «300 birds) and most populations are associated with privately-owned lands and, therefore, subject to land management activities which may have detrimental consequences. Plans to increase distribution and populations of plains sharp-tailed grouse in Colorado will rely primarily on transplants (Braun et al. 1992). While numerous transplants of prairie grouse have been attempted in North America, few have been successful (Toepfer et ale 1990, Rodgers 1992, Hoffman et al. 1992). A previous transplant of sharp-tailed grouse into Las Animas County northeast of Raton Mesa was attempted with a total of 85 males and 83 hens being released over a 3-year period (1987-89). This transplant was not successful in establishing a breeding population, and little follow-up of released birds was conducted to collect important data on survival and movements of the released birds. Habitat conditions were thought to be better on the top of Raton Mesa that at lower elevations where they had been released previously. Thus, this study was initiated to establish a population of plains sharp-tailed grouse on Raton Mesa and to document movements, mortality, and reproduction following an experimental transplant. Recommendations for transplant methodology as well as selection of release sites were evaluated during this study, as these two factors potentially have the greatest effect on the success of transplants. P. N. OBJECTIVES The objectives of this project were to trap and transplant plains sharptailed grouse into selected sites along the Front Range of Colorado and evaluate transplant success. Population characteristics of the transplanted population including movements and home range size, timing and causes of mortality, and habitat use will be compared to those described in the literature for native and transplanted prairie grouse. Results of this study will assist in developing transplant protocols for future transplants of prairie grouse. Specific objectives were to: 1. Transplant up to 80 plains sharp-tailed grouse from southeastern Wyoming into suitable habitats along the Front Range of Colorado. 2. Radiomark up to 40 sharp-tailed grouse in the transplanted population and monitor movements, habitat use, reproduction, and timing and causes of mortality. �86 3. 4. Conduct a pre-release evaluation of the habitat at the selected release site. Analyze data and prepare a final report. METHODS contact was made in 1994-96 with personnel of the Wyoming Game and Fish Department to obtain the necessary permits for trapping plains sharp-tailed grouse in southeastern Wyoming for transplant into Colorado. Active dancing grounds in Platte and Goshen counties, Wyoming were located as potential trapping sites and permission from affected landowners was obtained. Male and female sharp-tailed grouse were trapped on dancing grounds using walk-in funnel traps (Toepfer et al. 1988, Schroeder and Braun 1991). Captured birds were classified to age and sex (Henderson 1967) and fitted with seriallynumbered aluminum bands on the right leg and colored plastic bandettes on both legs (yellow in 1995, red in 1996). Battery-powered transmitters (weight 1213 gms) were attached with a necklace (Amstrup 1980) to selected birds to facilitate monitoring of movements and survival after release. Birds were held in captivity 1-4 days before they were transported by vehicle or helicopter to the top of Raton Mesa. Birds were released 5-10 minutes after arrival by opening the boxes and allowing the grouse to walk or fly from the site. In 1995 wheat grain was scattered at the release site to provide supplemental food but there was no indication it was used by the grouse. In both years, an automatic recording of sharp-tailed grouse display vocalizations and dancing sounds was programmed to broadcast for 1-2 hours at sunrise and sunset near the release site in an attempt to attract birds to the release site and begin displaying. A hand-held telemetry receiver and 3-element Yagi antenna were used to locate radio-marked grouse. Grouse were approached until flushed and information on location and flock size was recorded. A portable Global Positioning Satellite (GPS) receiver was used to record UTM locations of birds and minimum convex polygon home ranges were calculated using the McPaal software package (M. Stuwe and E. E. Blohowiak, Conserv. Res. cent., Natl. Zool. Park, Smithsonian Inst., Front Royal, Virginia, 1985). Vegetative cover (height-density) in the general release area was measured using a Robel pole (Robel et al. 1970). Attempts were made in April and May 1995-97 to locate dancing grounds by systematically searching ridges, knolls, and other suitable habitats on top of Raton Mesa. Efforts were made to search locations near radio-marked birds and look for tracks and other sign of display activity. RESULTS 1995 Release A total of 43 (20 males, 23 females) sharp~tailed grouse was captured in Platte and Goshen counties, Wyoming and released onto Raton Me~a in Las Animas County, Colorado in April 1995. These birds were trapped on 4 dancing grounds (Baker Swale = 6 males, 4 females; Gladys'= 4 males, 1 female; Thomas Jeffersons = 3 males, 12 females; Grange = 7 males, 6 females). Twenty-one birds (20 males, 1 female) were released on 11 April, 5 hens were released on 15 April, and 17 hens were released on 21 April. Ten male and 12 female sharp-tailed grouse were fitted with radio transmitters prior to release. Weather conditions at the time of release were less than ideal. Unusually cold and wet weather in March and April resulted in 100% snow cover (depth> 1.0 m) at the release site and adjacent areas on Raton Mesa when birds were released. These weather conditions persisted into late May. Snow �87 conditions and restricted access on Raton Mesa hampered monitoring birds for several weeks following release. There was little evidence of lekking behavior by released birds in 1995. No dancing grounds were located although signs of male display were observed on snowfields in late April and early May. Tracks indicated only 1-2 males were displaying and all signs of display were within 500 m of the release site. There was no evidence that the automatic tape player with sharp-tailed grouse display sounds attracted any birds to the site. Mortality Mortalities of 12 released birds (7 males, 5 females) were documented from 11 to 60 days post-release (Table 1). Mortalities were documented from 1.24 to 11.16 km from the release site with most occurring within 30 days of release. Four birds (1 male, 3 females) were not located (no radio signals received) following their release and it is suspected that most dispersed from the study area and likely died soon after. Several attempts to locate missing radio signals from aircraft were unsuccessful. The high initial mortality (minimum 60%) was likely caused by the severe weather conditions at time of release and the total snow cover at the release site. Most of the known causes of mortality were attributed to raptor depredation and may have been influenced by the condition of the birds following capture, captivity (1-4 days), and lack of food availability following release. . Table 1. Fates of radio-marked sharp-tailed Las Animas County, Colorado, 1995. Bird # Age Sex Max Dise (m) 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 2+ 0925 1045 1184 1254 1275 1334 1534 1655 1684 1745 1765 1775 1805 1835 1845 1875 1885 1904 1945 1955 1965 1994 a b F M M M F F F M F M M F M M F F M 1- F 2+ 2+ 2+ 2+ M F F F 2,530 2,250 11,160 7, 000 (est. ) 6, 000 (est.) n.d. b 7,530 8,410 8, 000 (est.) 1,400 1,240 8,200 1,230 n.d. 3,580 6,024 2,220 2,120 3,560 n.d. n.d. 1,750 grouse released on Raton Fate Home Range (km2) 1.607 0.476 0.137 Maximum distance from release site. No data, bird not located after release Mesa, Raptor kill, 3 May Raptor kill, 2 May Raptor kill, ? May Mortality signal, May Mortality signal, Jun No signal postrelease Last signal 29 Jun Radio fell off, 13 Sep Mortality signal, May Raptor kill, 2 May Raptor kill, 2 May Alive in Sep Radio fell off, 14 Jun No signal postrelease Last signal 2 May Mortality, 18 May Mortality signal, 3 May Mortality, 2 May Raptor kill, 3 May No signal postrelease No signal postrelease Last signal 2 May onto Raton Mesa. �88 Movements and Home Range The distribution of movements from the release site was bimodal with one group of birds staying relatively close to the release site while another group established ranges ~ 6.0 km from the release site (Table 1). High mortality of some birds soon after release may have skewed innate movement patterns, although some birds dispersed> 6.0 km and died away from Raton Mesa within 1-3 weeks after release. Some of these birds were not located or the radios recovered although mortaliuy signals were received for >30 days. Although sample sizes were small, it did not appear that post-release movements were related to sex of the grouse, with both males and hens showing long dispersal movements. Minimum convex polygon home ranges were calculated for 3 grouse (2 males, 1 female) which were located periodically for> 60 days after release. Home ranges were relatively small (0.137 - 1.697 km2, Table 1) and may have been affected by the few locations of each bird. However, consecutive locations at 1-2 week intervals showed that birds were usually within 200-400 m of their previous location. Habitat Height-density of vegetation was measured during June-August at 150 points within 3 pastures (50 measurements/pasture) where radio-marked sharptailed grouse were regularly located. The greatest height-density was measured at the release site pasture (5.52 ± 1.62 dm). Two grazed pastures in New Mexico where sharp-tailed grouse occurred during May-September had mean height-density measurements of 1.56 ± 0.77 dm and 1.95 ± 1.17 dm. However, height-density was quite variable in these pastures with 33% having heightdensity measurements ~ 2.25 dm, and 15% ~ 3.0 dm. The height-density measured in 1995 likely reflected the high precipitation the areas received during March-June. 1996 Release A total of 33 (20 males, 13 females) sharp-tailed grouse was captured in Platte and Goshen counties, Wyoming and released onto Raton Mesa in Las Animas County, Colorado in April 1996. One additional male died during the trapping process. Birds were trapped on 4 dancing grounds (Baker Swale = 1 male, 8 females; Kennedys'=10 males,S females; Thomas Jeffersons = 4 males; Grange = 5 males). Twenty birds (16 males, 4 females) were released on 5 April, 3 hens were released on 12 April, and 4 males and 6 hens were released on 19 April. Ten male and 10 female sharp-tailed grouse were fitted with radio transmitters prior to release. In contrast to 1995, weather conditions at time of release and during the month following were characterized by mild temperatures and average precipitation. Snow cover on Raton Mesa was <30 %, compared to 100 % in 1995, which allowed better access for monitoring birds. The automatic tape player with sharp-tailed grouse display vocalizations appeared to attract some sharp-tailed grouse to the site in 1996. The location was changed from 1995 to a site along a pipeline road where signs of display were observed the previous year. In May, the recorder was moved to a knoll about 400 m south where tracks and fecal droppings indicated regular display by several birds. While no display was observed, both radio-marked and unmarked birds were regularly flushed from this site in late April and early May. �89 Mortality Depredation of 6 released birds (3 males, 3 females) was documented from 5 to 35 days post-release (Table 1). One hen was found freshly killed by vehicle traffic 61 days post-release, 5 birds (2 males, 3 females) dispersed off Raton Mesa and could not be located, and signals from 2 birds (1 male, 1 female) were not heard following their release. Mortalities were documented from 1.31 to 41.62 km from the release site with most occurring within 30 days of release. Several attempts to locate radio signals from missing birds using aircraft were unsuccessful. Known mortality and dispersal from Raton Mesa resulted in high loss of radio-marked birds following release (at least 14 of 20 birds). While depredation by avian and mammalian predators was high, dispersal from the release site was likely a greater cause of mortality following release of translocated birds. No sharp-tailed grouse known to have dispersed from Raton Mesa was known to return and the habitat surrounding Raton Mesa likely lacked food resources or adequate cover. Mortality attributed to depredation may have been influenced by the condition of the birds following capture, captivity (1-4 days), and lack of familiarity with food resources and escape cover following release. Table 2. Fates of radio-marked sharp-tailed Las Animas County, Colorado, 1996. Bird I Age 2+ 2+ 2+ 2+ 2+ 2+ 12+ 2+ 2+ 2+ 2+ 2+ 12+ 2+ 2+ 12+ 2+ 1383 1497 1566 1649 1779 1819 1889 0455 1799 1195 1958 1698 1751 1589 1518 1539 1395 1859 1343 1678 a b Sex M M M M M F F M F M F F F F F F F M M M Max Dist" (m) 9,550 n.d. b n.d. 4,670 2,350 n.d. 6,619 3,984 2,350 n.d. 7,680 n.d. n.d. n.d. 1,310 n.d. 41,620 7,590 2,390 2,204 grouse released on Raton Fate Home Range (km") 2.39 0.31 117.12 0.32 Maximum distance from release site. No data, bird not located after release Mesa, Raptor kill, 10 May Signal from off Mesa Signal from off Mesa Radio fell off, 23 Apr Depredation, 1 May Signal from off Mesa Radio fell off, 28 Aug Raptor kill, 2 May Mortality, 1 May No signal postrelease Last signal 3 Jul Signal from off Mesa No signal post release No signal postrelease Mammalian kill, 9 May Signal from off Mesa Roadkill, 19 Jun Radio fell off, 28 Aug Raptor kill, 24 Apr Last signal 6 Jun onto Raton Mesa. �90 Movements and Home Range The distribution of movements from the release site in 1996 was bimodal with some birds staying relatively close to the release site while others dispersed off Raton Mesa where none was relocated alive (Table 2). High mortality of some birds soon after release may have skewed innate movement patterns, although some birds dispersed> 6.0 km but eventually returned to the vicinity of the release site. Some of these birds were not visually located or the radios recovered although mortality signals were received for >30 days. Although sample sizes were small, it did not appear that mortality or post-release movements were related to sex of the grouse, with both males and hens showing long dispersal movements. Minimum convex polygon home ranges were calculated for 4 grouse (1 male, 3 females) which were located periodically for >60 days after release. Home ranges of 3 birds were relatively small (0.31 - 2.39 km2, Table 1) and may have been affected by the few locations of each bird. The largest home range resulted from a long dispersal movement from the release site into New Mexico (where the bird was struck by a vehicle and killed). Consecutive locations at 1-2 week intervals showed that birds were usually within 200-400 m of their previous locations. One hen initially dispersed >6.6 km south from the release site but returned after 5 weeks to within 1.1 km of the release site where she nested. She incubated a clutch of 11 eggs of which 5 eventually hatched in late June. Her movements for the remainder of the summer were within 400 m of the nest site, and at least 3 chicks survived to 60 days-of-age. Habitat Height-density of vegetation was measured during June-August at 200 points within 4 pastures (50 measurements/pasture) where radio-marked sharptailed grouse were regularly located. The greatest height-density was measured at the release site pasture (3.45 ± 1.35 dm). Overall the two pastures in Colorado had 58 of 100 VOR measurements ~3.0 dm and 23 of 100 VOR measurements ~4.0 dm. Pastures in New Mexico where sharp-tailed grouse occurred during May-September had mean height-density measurements of 1.59 ± 0.54 dm and 1.24 ± 0.66 dm, reflecting greater livestock grazing. However, height-density was quite variable in these pastures with 21\ having heightdensity measurements ~ 2.25 dm. The average height-density measured in both Colorado pastures in 1996 was slightly less than in 1995 and likely reflected the high precipitation the areas received during March-June 1995 and the drier conditions in 1996. DIScuSSIQH Most transplants of prairie grouse in North America have been unsuccessful (Toepfer et al. 1990), thus, each transplant should be well documented and thoroughly evaluated. Consecutive year transplants of prairie grouse in spring, similar to the methods used in this study, have been successful in establishing populations of greater prairie-chickens (Tympanuchus cupido) in Colorado (Hoffman et al. 1992,' Beauprez 1994) when habitats were suitable. Although limited survival and reproduction by the transplanted sharp-tailed grouse were documented, and grassland habitat conditions available in summer (as reflected by VOR measurements) were apparently suitable, other habitat factors may have been limiting. There were no agricultural fields on or adjacent to Raton Mesa, thus, winter foods with which the sharp-tailed grouse were familiar in Wyoming may have been lacking. �91 Further, few deciduous shrubs used as food by plains sharp-tailed grouse in Colorado, including sumac BbYa spp. and snowberry Symphicarpos spp. (Hoag 1989) were available on top of Raton Mesa, where the dominant shrub was cinquefoil (Potentilla spp.). Results from both years indicate high mortality of released birds following the transplant and long dispersal movements from the release site. The long movements may result from birds seeking suitable habitats or trying to return to their native area. Hoffman et al. (1992) recorded post-release movements up to 29 km for greater prairie-chickens (Tympanuchus cupido) transplanted in spring, and a transplanted lesser prairie-chicken (Tympanuchus pallidicinctus) was documented returning nearly 300 km to the original trap site within 6 months (unpublished data). It appears that relatively few sharp-tailed grouse survived and successfully reproduced, thus, there is little chance that a population will become established on Raton Mesa as a result of this transplant effort. High winds and the location of the release site relative to the edge of the mesa likely resulted in some birds flushing from the mesa top and landing in forested habitats where they perished or were depredated. Because the elevation on top of Raton Mesa is 300-600 m above the surrounding terrain, any sharp-tailed grouse leaving the mesa top may have perished before finding their way back. No birds were known to leave the top of the mesa and return. The west, north, and east sides of Raton Mesa are surrounded by forests and steep cliffs; conditions which may have hindered return movements to the mesa top. Lack of familiarity with food resources and escape cover combined with several days of captivity, may have weakened the birds and increased their susceptibility to predation. It is not known whether the radio-marked birds had higher mortality than those not marked as reported in other studies (Marks and Marks 1987). Sharptailed grouse without radios were observed regularly following the release indicating actual survival may have been higher than indicated by radio-marked birds. While few nesting attempts were documented, observations of small flocks of birds in late summer and fall suggest that reproduction may have occurred but was not detected. Also, the number of birds observed in summer and fall which were not radio-marked suggests that some reproduction and recruitment occurred each year. One factor not studied was the availability of winter food for this population. ,Waste grain from wheat fields was available in Wyoming where these grouse were captured but was lacking on Raton Mesa and adjacent areas. If the grouse released onto Raton Mesa dispersed into New Mexico onto agricultural areas in winter, they may not have returned to breed near their release site. However, other than the one documented road-killed bird, no reports of sharp-tailed grouse were received from New Mexico. Access was a major problem which hindered monitoring of the released birds. Further, attempts at aerial monitoring were unsuccessful due to logistics in scheduling flights, unfavorable weather, and lack of radiotracking experience by the pilot and observer. It is likely that many birds which disappeared following release dispersed from the site and were not located, despite perhaps surviving for several months (as did the bird found in New Mexico nearly 42 km from the release site). MAHAGEMEHT IMPLICATIONS Because of the lack of success of the two recent transplant efforts on or near Raton Mesa, I recommend no further transplants occur to this area unless additional suitable winter habitat and food resources become available. Although the Division of Wildlife owns and manages two state properties at �92 this site (Lake Dorothy, James John), neither provides suitable year-round habitat for plains sharp-tailed grouse, and the potential for management as sharp-tailed grouse habitat is limited. Some privately owned lands nearby, especially in New Mexico, are lower in elevation and with proper management could potentially support populations of sharp-tailed grouse. Any additional transplants of sharp-tailed grouse into Colorado should be considered experimental and detailed evaluation of methods and causes of success or failure documented. Seasonal habitat needs of sharp-tailed grouse should be evaluated at each potential release site prior to release, and careful documentation of movements, habitats used, and causes of mortality recorded following release. LITERATURE CITED Amstrup, S. C. 1980. A radio-collar for game birds. J. Wildl. Manage. 44:214217. Bailey, A. M., and R. J. Niedrach. 1965. Birds of Colorado. Denver Mus.Nat. Hist., Denver, CO. Vol. 1, 454 pp. Beauprez, G. M. 1994. Movements, reproductive success, and habitat use by introduced greater prairie-chickens in northeastern Colorado. M. S. Thesis, Univ. Northern Colorado, Greeley. 100 pp. Braun, C. E., R. B. Davies, J. R •.Dennis, K. A. Green, and J. L. Sheppard. 1992. Plains sharp-tailed grouse recovery plan. Colorado Div. Wildl., Denver. 33 pp. Henderson, F. R., F. W. Brooks, R. W. Wood, and R. B. Dahlgren. 1967. Sexing of prairie grouse from crown feather patterns. J. Wildl. Manage. 31:764769. Hoag, A. W. 1989. Plains sharp-tailed grouse status and habitat use in Douglas County, Colorado. M. S. Thesis, Colorado State Univ., Fort Collins. 45 pp. _______ , T. W., and C. E. Braun. 1990. Status and distribution of plains sharp-tailed grouse in Colorado. Prairie Nat. 22:97-102. Hoffman, R. W., W. D. Snyder, G. C. Miller, and C. E. Braun. 1992. Reintroduction of greater prairie-chickens in northeastern Colorado. Prairie Nat. 24:197-204. Marks, J. S., and V. S. Marks. 1987. Influence of radio-collars on survival of sharp-tailed grouse. J. Wildl. Manage. 51:468-471. Robel, R. J., J. N. Briggs, A. D. Dayton, and L. C. Hulbert. 1970. Relationships between visual obstruction measurements and weight of grassland vegetation. J. Range Manage. 23:295-297. Rodgers, R. D. 1992. A technique for establishing sharp-tailed grouse in unoccupied range. Wildl. Soc. Bull. 20:101-106. Schroeder, M. A., and C. E. Braun. 1991. Walk-in traps for capturing prairie-chickens on leks. J. Field Ornithol. 62:378-385. Toepfer, J. E., R. L. Eng, and R. K. Anderson. 1990. Transplanting prairie grouse: what have we learned? Trans. North Am. Wildl. and Nat. Resour. Conf. 55:569-579. __________ , J. A. Newel, and J. Monarch. 1988. A method for trapping prairie grouse hens on display grounds. Pages 21-23 in A. D. Bjugstad, Tech. Coord. Prairie chickens on the Sheyenne National Grasslands. u.S. Dep. Agric., For. Servo Gen. Tech. Rep. RM-159. Prepared by Kenneth M. Giesen Wildlife Researcher �93 Colorado Division Wildlife Research April 1998 of Wildlife Report JOB PROGRESS State of: Colorado Project: Work W-167-R Plan: __ Job Title: Period REPORT ..• 1•...• 3..____ Ayian : Job Research 11 Evaluation of Columbian Sharp-tailed Opportunities in Western Colorado Covered: 01 January through Author: Richard W. Hoffman Personnel: Richard W. Hoffman, 31 December Colorado Grouse Reintroduction 1997 Division of Wildlife ABSTRACT Efforts for this reporting period focused on reviewing literature, conducting lek surveys, conducting meetings and gathering information to assist in the preparation of a conservation plan. ��95 EVALUATION REINTRODUCTION OF COLUMBIAN OPPORTUNITIES SHARP-TAILED IN WESTERN COLORADO Richard W. Hoffman INTRODUCTION Use of common names and misidentification of blue grouse (Pendragapus obscurus) and sage grouse (Centrocercus urophasianus) by early explorers have made it difficult to ascertain the precise distribution of Columbian sharptailed grouse (Tympanuchus phasianellus columbianus) in Colorado (Rogers 1969, Giesen and Braun 1993). However, historical records suggest this subspecies may have occurred in at least 22 counties in western Colorado (Bailey and Niedrach 1965, Rogers 1969). Recent surveys indicate viable populations are restricted to Moffat, Routt, and Rio Blanco counties, with possible remnant populations in Mesa and Montrose counties (Giesen and Braun 1993). Similar reductions in the distribution of Columbian sharp-tailed grouse have occurred throughout western North America (Miller and Graul 1980). This decrease in distribution resulted in designation as a Category 2 species (U. S. Pep. Inter. 1989). Factors responsible for the reduction in distribution include conversion of native rangeland to cropland, excessive grazing by livestock, vegetative succession due to fire suppression, herbicide treatments, mineral exploitation, and urban development (Meints et al. 1992, Giesen and Connelly 1993). These factors have had the most pronounced impact on nesting, brood rearing, and winter cover through loss of native grasses and deciduous shrubs (Giesen and Braun 1993). Cover types used by Columbian sharp-tailed grouse tend to be structurally and vegetatively diverse with an extensive deciduous shrub component (Meints et al. 1992, Giesen and Connelly 1993). In Colorado, Columbian sharp-tailed grouse occur in mountain shrub communities interspersed with grasslands, small aspen (Populus tremuloides) stands, and riparian zones (Giesen 1987). serviceberry (Amelanchier spp.) is an essential element of these communities and usually grows in association with one or more of the following deciduous shrubs: Gambeloak (Ouercus gambelii), common chokecherry (Prunus yirginiana), snowberry (Symporicarpos spp.), and sagebrush (Artemisia spp.) (Giesen 1987). Wheat is the primary agricultural crop within the range of sharptails in western Colorado. Wheatfields may be used during late summer and fall after the wheat has been harvested. These fields are usually snowcovered and unavailable during winter. Much of what is known about Columbian sharp-tailed grouse in western Colorado has resulted from studies in the northwest portion of the state (Dargan et al. 1942, Rogers 1969, Giesen 1987). Little is known about sharptailed grouse in southwestern Colorado other than they once occurred there and may still exist in low densities on the north end of the Uncompahgre Plateau (Rogers 1969, Giesen 1985). It has been 10 years since the last intensive effort to conduct lek surveys for Columbian sharp-tailed grouse in western Colorado. Another intensive effort is needed because changes in land use practices have occurred since then including implementation of the Conservation Reserve Program, additional mining and development activities, and alteration of grazing practices. Perhaps the most important action in the last 10 years affecting the need for current population and distribution data has been the petition to list Columbian sharp-tailed grouse as "threatened" or "endangered" in the lower 48 conterminous United States pursuant to the Endangered Species Act (Carlton 1995). This action is of special significance in Colorado because Idaho and Colorado are the only states that allow hunting �96 of Columbian sharp-tailed grouse and that still have adequate populations to provide transplant stock for future restoration programs. Opportunities for management of sharptails in western Colorado may be limited because much of the occupied habitat occurs on private lands. The most extensive areas of public lands within the historic distribution of Columbian sharp-tailed grouse are in southwest Colorado. The last confirmed sighting of sharptails on these lands was in 1985 (Giesen 1985). Before a reintroduction program can be implemented, current habitat conditions and status of sharptails on these lands must be evaluated and management strategies formulated based on the outcome of the evaluation. It is likely that any effort to restore sharptails in western Colorado will require a commensurate effort to restore and protect habitat. P. N. OBJECTIVES Objectives of this project are to (1) form a sharp-tailed grouse working group with broad citizen, community, and agency representation, and in cooperation with this group, prepare a conservation plan for Columbian sharptailed grouse in Colorado, (2) conduct intensive lek surveys of Columbian sharp-tailed grouse in northwest Colorado, (3) ascertain presence or absence of sharptails on historic range in southwest Colorado, (4) identify potential reintroduction sites within the historic range of Columbian sharp-tailed grouse, (5) evaluate existing habitat conditions on these sites based on the habitat suitability index model described by Meints et ale (1992), and (6) cooperate with other western states in preparing conservation strategies for Columbian sharp-tailed grouse. SEGMENT OBJECTIVES 1. 2. 3. 4. 5. 6. 7. 8. Review literature pertinent to the objectives of this study. Identify participants interested in preparing conservation plan. Organize and conduct public meetings. Form working group and conduct regularly scheduled meetings to develop management strategies and prepare conservation plan. Prepare conservation plan in collaboration with working group. Conduct leks surveys in Moffat, Routt, and Rio Blanco counties. Conduct surveys to ascertain presence or absence of sharptails in Mesa, Montezuma, and Montrose counties. Compile data, analyze results, and prepare progress report. RESULTS AND DISCUSSION Segment Objectiye 1 - Literature on all aspects of the biology and ecology of sharp-tailed grouse was reviewed. Literature searches were conducted through Current Contents and the Fish and Wildlife Reference Service. Efforts were made to review draft and final conservation plans and strategies prepared by other states and to talk with the people involved in preparing these documents. Efforts also were made to review all documents pertaining to the petition to list the Columbian Sharp-tailed grouse as threatened or endangered. Segment Objectiyes 2-4 - Discussions were held with the Human Dimensions section of the Colorado Division of Wildlife and with other CDOW personnel with experience in organizing and conducting public meetings. In addition, stakeholder documents (i.e., manuals, plans, and strategies for effective stakeholder involvement) obtained from the Human Dimensions section were reviewed. �97 The CDOW maintains a database of external publics and key contacts in other agencies that are routinely invited to participate in stakeholder meetings. This list was reviewed and names were added or deleted as deemed necessary. Meetings were held with CDOW field personnel to determine their interest in participating in the working groups and to obtain names of local landowners, businesses, and organizations that should be invited to participate. A letter was drafted to organize informational meetings at several locations in western Colorado. Meetings were held with local landowners, and officials of the Natural Resource Conservation Service, u.S. Forest Service, and the following coal mining companies: Colowyo, Trapper, Peabody, Seneca, Twentymile, and Pittsburg/Midway. To date, no official working group has been formed. Attempts were made to join existing sage grouse working groups in northwest and southwest Colorado, but members of these groups were opposed to the idea. This has created problems in terms of generating interest from other agencies, private individuals, and businesses because they are already involved with other working groups and there is only so much time they can devote to this activity. Segment Objective 5 - The recommendation was that personnel of the Colorado Division of Wildlife prepare the conservation plan rather than a working group. Once a draft plan is prepared, then a working group should be formed to review, revise, and finalize the plan. Information (literature, unpublished data ••etc.) has been gathered to use in the preparation of the plan. Writing of the plan will begin in June 1998. Segment Objective 6 - Traditionally, lek counts were designed to provide information on average number of males per lek and average number of birds per lek. These estimates, when collected consistently over long periods, were presumed to provide trends in population size. However, Kobriger (1975), concluded that lek counts have little value in measuring population size or documenting population trends because of inconsistent lek attendance patterns within and among years. Cannon and Knopf (1981) recommended replacing lek counts with lek surveys (i.e., number of active leks) based on the observation that when populations increase, males respond by forming more leks instead of increasing the average number of males on each lek. These findings suggest that lek surveys should include two components: (1) surveys of known lek sites to ascertain status (active or inactive), and (2) searches for new leks. Between 1 April and 6 June 1997, 45 days were spent searching for Columbian sharp-tailed grouse leks in extreme east-central Moffat County and most of Routt County, except areas south of Oak Creek. Efforts focused on searching for new leks. Area 10 personnel concentrated their efforts on verifying the status of known leks. A~tempts were made to count the total number of birds per lek, and if possible, the number of males per lek. The count data were collected to compare with similar data collected between 1964 and 1985 and reported by Giesen (1987). Accurate counts were difficult to obtain on many leks because the birds were obscured by vegetation or there was no vantage point from which to observe the entire lek. Also, due to the vast area that needed to be searched and the large number of known leks that needed to be checked, there was insufficient time to conduct more than one count per lek. Lek Surveys Seventy-seven sharptail grouse leks are listed in the WRIS database for northwestern Colorado; 57 in Routt County and 20 in Moffat County. Examination of the database revealed .the following duplicate entries: Fish �98 Creek and Missy, Gray's Divide and Soash, Long's Gulch and Wiseman's # 2, RCR27-9SE and Annan's #1, and RCR27-9SENW and Annan's #1. In addition, there are 2 Cottonwood Creek leks listed in the database. Only one is a valid lek site and it is actually on Dry Fork Elkhead Creek. This is the same lek Rogers (1969) called Cottonwood Creek. The other Cottonwood Creek lek does not exist. Also, there are 4 previously documented leks not listed in the database: 3 were found within the past 2 years (1995-96) and 1 was an historic lek that somehow never was entered into the database. Excluding the 5 duplicates and 1 invalid listing, and adding the 4 previously documented leks, the database now contains 75 lek sites of which 53 are in Ro~tt County and 22 in Moffat County (Table 1). Of the 53 known lek sites in Routt County, 45 were checked in 1997. Fifteen were classified as inactive and 30 were classified as active, including 2 leks (Salt Creek 1 and 2) that combined into 1 lek (Table 1). Nine of 22 lek sites in Moffat County were checked in 1997 and 6 were active (Table 1). Rogers (1969) reported finding 21 lek sites in Routt County. These sites are identified in the database as historic leks. Of the 20 historic leks checked in Routt County during 1997, 13 were still active (Table 1). Rogers (1969) listed 10 leks he located in Moffat County. Three were checked in 1997, but only one was active and it (Elkhead Road #3) is actually in Routt County (Table 1). TABLE 1. 1997 COLUMBIAN SHARP-TAILED GROUSE LEK SURVEYS - STATUS OF PREVIOUSLY DOCUMENTED LEKS. Active - Routt Annan's 1* Annan's 2* Barnes Dry Elkhead Ridge* Dry Fork Elkhead * Eckman Park 1 (Energy Fuels) Elkhead Road 3* Elk Mountain 2 (Sam's)* Elk Mountain 4 (Clark's Pasture)** Foidel Creek* George's Gulch ( Salt Creek 1&2)*** Hayden Divide* Heidel Hinkle Hocket (not in database) Horton Knoll 1 Maneotis McKinney Ranch* Morgan Creek Mud Springs* Rock Creek 2 Sage Creek* Smiths* Soash (Gray's Divide) Twentymile Twentymile Cliffs 4 (Scott) Woods Yellowjacket Road (Sidney Peak)* Yellowjacket 2 (Willie Ranch) * historic lek sites ** possible replacement for Elk Mt 3 ***two leks combined Into one Inactive - Routt 80 Road California Park Road 1* California Park Road 2 Dry Gulch 2* Dry Gulch 3 Elk Mountain 1* Elk Mountain 3* Elk River Cemetery* Gillilands* Green Acres Hicks Milner Robinson Sherrod-Sandelin Yellow jacket 1* Not Checked - Routt Califomia Park County Airport Dinwiddie Fish Creek (Missy) Five Pines Mesa Horton Knoll 2 Rock Creek 1* Slater Park 1 (not in database) Wymans Active - Moffat Buck Mountain 1 (not in database) little Buck 1 little Buck 2 long Gulch (Wiseman's 2) Villards Wilderness Ranch 1 (not in database) Inactive Elkhead .Elkhead Elkhead - Moffat Road 1* Road 2* Road 4 Not Checked - Moffat Baker'S Peak 1 Cedar Hill Gulch Fly Creek Fortification Rocks lies Dome (Stinking Gulch)* Mcinturf Mesa Morapas Gas Field* (not in database) Nolands Pelleys* Schneiders* Taylors* Wingate* Wiseman's 1* �99 At least 35, and possibly 42, new lek sites were found in 1997 (Table 2); all were in Routt County. Sixteen new leks were found in CRP, 15 on mine reclamation lands, 9 in sage/grassland communities, and 2 in grass pasture land. TABLE 2. 1997 COLUMBIAN SHARP-TAILED GROUSE LEK SURVEYS - NEW LEK LOCATIONS. Lek Name County Bloomquist Calf Creek Deep Creek Dry Creek Eckman Park 2 Eckman Park 3 Eckman Park 4 Eckman Park 5 Eckman Park 6 Elk Creek 1 Elkhead Finger Rock Gnat Hill Hightail Hillberry Hoffman Homestead Homestead Ditch Little Hunter Middle Creek Moming Crow North Giant Penny Pleasant Valley RCR33b Ricks Rogers Saddle Mountain 1 Saddle Mountain 2 Shivers Smuin Gulch Gravel Pit Smuin Gulch Oil Well 1 Stokes Gulch Twenty-mile Cliffs 1 Twenty-mile Cliffs 2 Twenty-mile Cliffs 3 Twenty-mile Cliffs 5 TumerCreek Warrick Pasture Wolf Mountain Ranch Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt USGS Quad Wolf Mountain Quaker Mountain Clark Blacktail Mountain Rattlesnake Butte Rattlesnake Butte Rattlesnake Butte Rattlesnake Butte Rattlesnake Butte Milner Quaker Mountain Quaker Mountain Breeze Mountain Rattlesnake Butte Mount Harris Cow Creek Rattlesnake Butte Cow Creek Clark Cow Creek Mad Creek Mad Creek Rock Springs Blacktail Mountain Cow Creek Rattlesnake Butte Rattlesnake Butte Cow Creek Cow Creek Rattlesnake Butte Hayden Hayden Breeze Mountain Rattlesnake Butte Rattlesnake Butte Rattlesnake Butte Rattlesnake Butte Wolf Mountain Hooker Mountain Hooker Mountain Legal 7N86W4SW 8N88W14NE 8N85W30SW 5N84W32NW 4N86W18NW 4N86W18NE 4N86W18SE 4N86W8SE 4N86W33SW 6N86W28NE 8N88W13NE 8N88W11SW 6N89W20SE 4N87W12NE 5N87W6SW 5N86W12SW 4N87W12SW 5N86W24NE 8N85W19NW 5N86W13SE 7N85W8NW 7N85W6NW 8N88W29SE 4N84W9NW 6N85W30SW 5N86W26NW 5N87W36SW 6N85W19NW 6N86W23SE 4N87W11SE 6N89W21NE 6N89W23SE 5N89W3NW 5N86W30SE 5N87W25SE 5N86W19SW 5N86W20SE 7N86W5SW 7N88W11SW 6N87W4SE UTMX 327400 311850 333550 344050 322500 323250 323200 325300 326150 327500 313600 311050 297200 320900 313700 330900 320350 332150 333650 331900 334650 333250 307200 345350 333250 329500 321250 332750 332600 319750 298750 301800 298850 323600 322350 322950 325850 325250 311250 318350 UTM Y 4495050 4502800 4498950 4467900 4465700 4465600 4464600 4466200 4467450 4479850 4502100 4503150 4481150 4467100 4476400 4474250 4466450 4471750 4500500 4472650 4494550 4495550 4498650 4466650 4478950 4470050 4467800 4481650 4480900 4466350 4482050 4481500 4477200 4468600 4469500 4470700 4470600 4495650 4493900 4485350 Comments found by M. Middleton not sure if lek site not sure if lek site not sure if lek site not sure if lek site not sure if lek site not sure if lek site satellite lek of 20mi 2 satellite lek of 20mi #4? found by M. Middleton found by Haskins not sure if lek site Lek Counts Counts were obtained for 48 leks (Table 3). Total birds per lek averaged 12.4, whereas the average number of males per lek was 10.9. The number of leks counted between 1964 and 1985 varied from none to 26; average birds per lek ranged from 5.4 to 14.0 (Giesen 1987). �100 TABLE 3. 1997 COLUMBIAN SHARP-TAILED GROUSE LEK COUNTS. Lek Name Annan's 2 Bloomquist Calf Creek Deep Creek Dry Creek Dry Elkhead Ridge Eckman Park 1 Eckman Park 2 Eckman Park 3 Eckman Park 4 Eckman Park 5 Eckman Park 6 Elk Creek 1 Elkhead Elkhead Road 3 Elk Mountain 2 Elk Mountain 4 Foidel Creek George's Gulch Hightail Hillberry Hinkle Hoffman Homestead Homestead Ditch Maneotis McKinney Ranch North Giant RCR33b Ricks Rogers Saddle Mountain 1 Shivers Smuin Gulch Gravel Pit Smuin Gulch Oil Well 1 Twentymile Cliffs 1 Twentymile Cliffs 2 Twentymile Cliffs 3 Twentymile Cliffs 4 Twentymile Cliffs 5 TumerCreek Woods Yellowjacket Road Total Leks Counted Total Birds Counted Average Birds/Lek SO Range Date Males Total Birds Lek Status 05/13/97 05/08/97 05/07/97 05/30/97 05/09/97 05/05/97 05/14/97 05/14/97 05/14/97 05/14/97 05/14/97 05/14/97 05/20/97 05/07/97 04/25/97 06/02197 06/02197 04/30/97 05/29/97 04130197 OS/28/97 04/15/97 05/16/97 04/30/97 05/15/97 05/15/97 04/22197 05/22197 04/23/97 05/18/97 04/29/97 04/23/97 04/30/97 05/08/97 05/29/97 04/15/97 05/17/97 05/17/97 05/17/97 05/03/97 06/02197 05/08/97 04/07/97 16 9 16 9 12 6 10 4 20 21 20 10 10 13 15 10 8 6 35 11 24 15 11 5 11 6 7 15 10 19 53 10 16 5 6 14 10 3 11 4 10 25 16 17 historic new new new new historic known new new new new new new new historic historic historic? historic historic new new known new new new known historic new new new new new new new new new new new known new new known historic 6 9 4 17 18 17 10 10 11 12 6 11 13 10 5 11 7 15 7 19 44 10 15 5 6 14 10 3 10 4 8 20 14 3 3 43 562 13.1 9.1 3-53 36 409 11.4 7.2 3-44 Recommendations 1. Efforts should be made in 1998 to visit Table 1 that were not checked in 1997. 2. all known Comments Energy Fuels minimum count minimum count minimum count minimum count Clark's Pasture Salt Creek 1 and 2 may be satellite lek may be satellite lek Scott minimum count Sidney Peak lek sites listed in Revisit all leks in 1998 that were identified as inactive in 1997 (Table 1); 1997 lek sites that remain inactive in 1998 should be deleted from the database of active leks. Another database should be created for leks that become inactive. �101 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. Efforts to locate new leks in 1998 should be focused in eastern Moffat and southern Routt counties. Count 40 leks per year. These should be referred to as the count leks. The objective should be to count as many of the same leks each year as possible. The initial sample of count leks should be randomly-selected from the list of all known active leks (Tables 1 and 2). If a count lek becomes inactive, it should be replaced with another lek randomlyselected from the list of known active leks (excluding the count leks). Results of lek surveys and lek counts should be recorded on standardized forms. An example form and instructions are included with this report. Devise a standard system for naming leks possibly based on the USGS quadrangle and closest landmark to where the lek is located. Verify that the following new leks located in 1997 are active lek sites: Morning Crow, Penny, Pleasant Valley, Saddle Mountain 2, Smuin Gulch Oil Well (confirm location and verify whether it is the same as the Heidel lek) , Stokes Gulch 1, Stokes Gulch 2, Twenty-mile Cliffs 3, and Wolf Mountain Ranch. Record the major habitat type for all known lek sites. Habitat classifications should include but are not limited to the following types: Conservation Reserve fields, mine reclamation, mountain shrub, sagebrush steppe, hay/pasture, agricultural field (alfalfa or wheat stubble), and gra~sland. Confirm location of lek in Slater Park. This lek appears on U.S. Forest Service records, but is not listed in the database nor has it been surveyed in recent years. Confirm location of lek found by B. Postovit on Senneca mine property, and two leks found by J. Monarch on Colowyo mine property in Axial Basin. Add these leks to the database. Delete the following lek sites from the database: Cottonwood Creek - located in T8NR87W7NW (Hooker Mt Quad); according to Jim Haskins there has never been a lek found at this location. Gray's Divide - same as Soash lek; birds displayed at a slightly different location one year and it was identified as a new lek rather than an old lek that has shifted locations. Wiseman's #2 - same as Long's Gulch lek; similar situation as Gray's Divide and Soash leks. Wiseman's #2 is the original name, however, Long's Gulch better describes the location and should be the name retained in the database. RCR 27- 9SE and RCR 27- 9SENW - both are the same as Annan's #1, which tends to shift locations in some years. Leks located within 1 km of known leks that are found to be inactive during the same year should not be classified as new leks. If the known lek is active and another lek is located within 1 km, it can be classified as a new lek provided it is at least 0.5 km from the known lek and has 4 or more males on the lek. otherwise, it should be classified as a satellite lek. Expand lek searches on mine reclamation lands. Work cooperatively and share information with the mining companies and the consultants they hire to conduct surveys. The lek listed as "not named" should be changed to Five Pines Mesa. In addition to the new leks listed in Table 2, add the following 4 leks that were documented prior to 1997 but were never entered into the database: Morapas Gas Field - historic lek found by Rogers (1969); not sure of exact location; believe it is on the Thornburg Quad in T3NR91W17NE or 16NW, Moffat County; could not find Morapas Gas Field on the map; could be same as Thornburg Gas Field. �102 Rocket - located by Jim Haskins in 1996; active in 1997; in Routt on Mount Harris Quad in T6N87W31NE; UTM = 314950, 4478650. Buck Mountain Moffat County 4406700. County located by Mike Bauman in 1996 (?); active in 1997; on Slide Mountain Quad in T9N89W36SE; UTM = 303350, Wilderness Ranch in Moffat County 4531050. in - located by Mike Bauman in 1996 (?); active in 1997; on Baker's Peak Quad in T11N89W16SW; UTM = 298850, LITERATURE CITED Bailey, A. M., and R. J. Niedrach. 1965. Birds of Colorado, Vol. 1. Denver Mus. Nat. Hist., Denver, CO. 454pp. Cannon, R. W., and F. L. Knopf. 1981. Lek numbers as a trend index to prairie grouse populations. J. Wildl. Manage. 45:776-778. Carlton. J. C •• 1995. Petition for a rule to list the Columbian sharp-tailed grouse, Tympanuchus phasianellus columbianus, as "threatened" or "endangered" in the conterminous United States under the Endangered Species Act, 16 U.S.C. Sec. 1531 et seq. (1973) as amended. Biodiversity Legal Foundation, Boulder, CO. 52pp. Dargan, L. M., H. R. Shepherd, and R. N. Randall. 1942. Data on sharp-tailed grouse in Moffat and Routt counties. Colorado Game, Fish, and Parks Dep, Sage Grouse Survey, Vol 4, Denver. 28pp. Giesen, K. M. 1985. Inventory of Columbian sharp-tailed grouse in western Colorado. Colorado Div. Wildl., Unpubl. Rep, Fort Collins. 6pp. Giesen, K. M. 1987. Population characteristics and habitat use by Columbian sharp-tailed grouse in northwest Colorado. Pages 251-279 in Wildlife Res. Rep., Part 2. Colorado Div. Wildl., Fed Aid Proj. W-152-R, Apr. 1987. Giesen, K •.M., and C. E. Braun. 1993. Status and distribution of Columbian sharp-tailed grouse in Colorado. Prairie Nat. 25:237-242. Giesen, K. M., and J. W. Connelly. .1993. Guidelines for management of Columbian sharp-tailed grouse habitats. Wildl. Soc. Bull. 21:325-333. Kobriger, G. D. 1975. Correlation of sharp-tailed grouse population parameters. North Dakota Outdoors 25(5):10-13. Meints, D. R., J. W. Connelly, K. P. Reese, A. R. Sands, and T. P. Hemker. 1992. Habitat suitability index procedure for Columbian sharp-tailed grouse. Univ. Idaho For., Wildl., and Range Exp. Stn. Bull. 55. 27pp. Miller, G. C., and W. D. Graul. 1980. Status of sharp-tailed grouse in North America. Pages 18-28 in P. A. Vohs and F. L. Knopf, eds. Proceedings of the prairie grouse symposium. Oklahoma State Univ., Stillwater. Rogers, G. E. 1969. The sharp-tailed grouse in Colorado. Colorado Game, Fish, and Parks Tech. Publ. 23. 94pp. Robel, R. J., J. N. Briggs, A. D. Dayton, and L. C. Hulbert. 1970. Relationships between visual obstruction and weight of grassland vegetation. J. Range Manage. 23:295-297. U. S. Department of the Interior. 1989. Endangered and threatened wildlife and plants; annual notice of review; proposed rules. Fed. Register 54:560. Prepared by Richard W. Hoffman LSSR IV �103 Instructions for conducting Grouse Lek Surveys and Counts These instructions and the accompanying form are an attempt to standardize information we collect. Please follow the steps outlined below. 1. Conduct surveys from 30 minutes before to two hours after the sunrise. 2. Timing of breeding activities can vary 2-3 weeks from one year to the next depending on spring conditions. In most years, the best time to conduct surveys is from 10 April to 15 May. Only one visit per lek is necessary if birds are observed on the lek. If birds are not found, then at least one and preferably two more visits should be made to the lek to confirm that it is inactive. Complete a form each time you visit the lek. 3. These instructions are specific for Columbian sharp-tailed grouse, but the form can be used for other species of grouse. Therefore, be sure to record the species being surveyed in the appropriate space. 4. Provide the complete date (mm/dd/yy). 5. Lek names can be confusing. Sometimes the lek name has no relevance to where the lek is located. Over the years, lek names may change or old leks that shift locations are given new names. Therefore, it is important when recording the lek name to be as complete and specific as possible. If the lek has been referred to by other names, also list those names. Be sure to include numbers that are part of the lek name; i.e., Eckman Park #2, Eckman Park #2 •••etc. 6. Give the official Yampa •••etc. name of the district; i.e., Steamboat South, Hayden, 7. For lek status, check either established (previously documented) or new. If the lek is established, check whether it is active or inactive (this is the most important priority for established leks), and list the USGS quad and county in which the lek is located. If the lek is new, provide the UTM coordinates, indicate whether the coordinates were read from a map or determined with a GPS unit, list the USGS quad and county in which the lek is located, and identify if the lek is on public or private land. 8. If the lek is active, attempt to make three counts at five minute intervals. First attempt to count the total birds present on the lek, then classify them as males, females, and unknown birds. It may be difficult to distinguish males from females especially if the males are not displaying. Topographic and vegetative features also may make it difficult to observe and count all birds on the lek. If it is apparent that you will not be able to obtain an accurate count, then flush the birds, record the total count, and check the space that indicates the birds were flushed 9. Provide whatever comments you feel are necessary to interpret the data that was gathered, such as weather conditions, presence of predators, and additional information about the location of the lek. �104 I 998 GROUSE LEK SURVEY FORM SPECIES: _ DATE: ,LEK NAME: ______________________________________________________________ _______________________________________________ _____________ OBSERVER(S): DWM DISTRICT: _ LEKSTATUS: ESTABLISHED ACTIVE NEW INACTIVE UTM Y (NORTHING) __________ UTM X (EAsTING) USGS QUAD UTM COORDINATESDETERMINEDBY GPS MAP COUNTY _ LAND STAlUS: PUBLIC TlME TOTAL MALES PRIVATE TOTAL FEMALES TOTAL UNKNOWN TOTAL BIRDS HIGH COUNTS: MALES UNKNOWN FEMALES TOTAL BIRDS __ FLUSH COUNT * ~_ • THE FLUSH COUNT IS ONLY NECESSARYWHEN BIRDS ARE FLUSHED INADVERTENTLYOR THE OBSERVER IS UNABLE TO OBSERVE BIRDS ON THE LEK AND OPTS TO FLUSH THE BIRDS TO OBTAIN A COUNT. COMMENTS: _ _ �105 Colorado Division Wildlife Research April 1998 of Wildlife Report JOB PROGRESS State Colorado of: Project: Work W-167-R Plan: Job Title: Period Covered: Author: Personnel: REPORT Clait Upland 1 22 : Job Upland Bird Research 01 January Bird Research through Publications 31 December 1997 E. Braun Clait E. Braun, K. M. Giesen, Colorado Division of Wildlife R. W. Hoffman, and T. E. Remington, ABSTRACT The following articles were published in 1997: Braun, C.E. 1997. Identification of Utah's The Wildlife Society. Abstract. sage grouse taxa. Utah Chapter, Braun, C.E. 1997. Long-term monitoring of montane species: white-tailed ptarmigan, 1966-96. Western Section, The Wildlife Society. Abstracts: 4. Braun, C.E. 1997. The status of sage grouse in southeast Utah and southwestern Colorado. Utah Chapter, The Wildlife Society. Abstract. Commons, M. L. 1997. Movement and habitat use by Gunnison sage grouse (Centrocercus minimus) in southwestern Colorado. M.N.R.M. Thesis, Manitoba, Winnipeg. 108pp. Commons, M.L., R.K. Baydack, and C.E. Braun. Centrocercus minimus use of fragmented Colorado. Wildlife Biology 3:283. Univ. 1997. Gunnison sage grouse habitats in southwestern Connelly, J.W., and C.E. Braun. 1997. Long-term changes in sage grouse Centrocercus urophasianus populations in western North America. Wildlife Biology 3:229-234. Giesen, K.M. 1997. Demography and population changes in white~tailed ptarmigan in Rocky Mountain National Park, 1975-1996. Proc. Rocky Natl. Park All Scientists Conf. Abstract. Mtn. �106 Giesen, K.M. 1997. Seasonal movements, home ranges, and habitat use by Columbian sharp-tailed grouse in Coloradq. Colorado Div. Wildl. Spec. Rep. 72. 16pp. Giesen, K.M., and G.D. Kobriger. 1997. Status and management of sharp-tailed grouse Tympanuchus phasianellus in North America. Wildlife Biology 3:286. Hoffman, R.W. and G.M. Beauprez. chickens Tympanuchus cupido 3:283. 1997. Reintroduction of greater praLrLein northeastern Colorado. Wildlife Biology Hoffman, R.W., M.P. Luttrell, W.R. Davidson, and D. H. Ley. 1997. Mycoplasmas in wild turkey living in association with domestic Wildl. Dis. 33:526-535. Martin, K., P.B. Stacey, and C.E. Braun. maintenance of population stability Wildlife Biology 3:295-296. 1997. Demographic in grouse - beyond fowl. J. rescue and metapopulations. Oyler, S.J., C.E. Braun, and K.P. Burnham. 1997. Use of a habitat-based model to predict sage grouse Centrocercus urQphasianus occupancy of patches in southwestern Colorado. Wildlife Biology 3:282. Quinn, T.W., N.W. Kahn, J.R. Young, N.G. Benedict, S. Wood, D. Mata, and C.E. Braun. 1997. Probing the evolutionary history of sage grouse centrocercus urophasianus populations using mitochondrial DNA sequence. Wildlife Biology 3:291. Remington, T.E., and R.W. Hoffman. 1997. Costs of detoxification of xenobiotics in conifer needles to blue grouse Dendragapus obscurus. Wildlife Biology 3:289. Prepared by Clait E. Braun Avian Prowam Manager �107 Colorado Division Wildlife Research April 1998 of Wildlife Report JOB PROGRESS State of: Colorado Project: W-167-R Upland Work Plan: 26 Job Title: Analysis Period Covered: Author: REPORT Clait Personnel: 1 Job of Upland 01 January Bird Research Bird PQPulation through 31 December Trends 1997 E. Braun Clait E. Braun, Kenneth M. Giesen, Richard W. Hoffman, E. Remington, Colorado Division of Wildlife and Thomas ABSTRACT The following Braun, C.E. reports 1997. were published Sage grouse counts, Blue Mountain, 1997. Sage grouse counts, Cold Spring 1997. Sage grouse counts, Gunnison 1997. Sage grouse counts, Lower Moffat 1997. sage grouse counts, North 1997. Sage grouse harvest report, Eagle, 1997. Sage grouse harvest report, Gunnison 1997. Sage grouse County, 1997. harvest report, Lower Basin, Park, 1997. 1997. 1997. Basin, Moffat harvest report, Middle 1997. Sage grouse harvest report, Yampa grouse 1997. 1997. County, Sage grouse sharp-tailed 1997. Mountain, 1997. Giesen, K.M. 1997. Columbian Colorado, 1976-97. Hagen, in 1997: Park, Area, harvest 1997. and western Routt 1997. 1997. data, northwest C.A., and C.E~ Braun. 1997. Habitat use and seasonal movements sage grouse in the Piceance Basin, Rio Blanco County, Colorado. Colorado Div. Wildl. Unpubl. Rep., Fort Collins. 21pp. of �108 Hoffman, R.W. 1997. 1997. Analyses of statewide Hoffman, R.W. 1997. Columbian sharp-tailed for northwest Colorado. Unpubl. Rep., blue grouse wing collections grouse lek surveys and lek counts Fort Collins. 9pp. Larison, J.R. 1997. Multiple-metal stress in an avian herbivore: the reproductive and physiological effects of chronic exposure to toxic metals. Cornell Univ., Unpubl. Rep. 18pp. Prepared by Clait E. Braun Avian Program Manager for �109 Colorado Division Wildlife Research April 1998 of Wildlife Report JOB PROGRESS State of: Colorado Project: W-167-R REPORT Ayian Research Work Plan: Job: Job Title: Evaluate Population Trends of Selected Migratory Birds in Colorado Period Covered: 01 July through 31 December Species of Neotropical 1997 Author: Kenneth H. Giesen Personnel: Gerald D. Craig and Kenneth M. Giesen, Colorado Division Wildlife, Michael F. carter, Colorado Bird Observatory of ABSTRACT Evaluation of current programs for monitoring nongame or passerine birds in Colorado was initiated following reports of apparent rangewide declines of many Nearctic-Neotropical migratory species. Recent efforts in Colorado to monitor the distribution and population status of nongame birds in Colorado resulted in several efforts documenting species distribution, but current methods of monitoring population trends (BBS routes, MAPS stations) were found to lack statistic vigor or adequate sample sizes. Discussions with the Colorado Bird Observatory (CBO) and statisticians resulted in developing a population monitoring protocol based on distance sampling methods. Initial efforts using distance sampling methodologies will be initiated through a contract with the Colorado Bird Observatory and evaluated in 1998. ��III EVALUATE POPULATION TRENDS OF SELECTED SPECIES OF NEOTROPICAL MIGRATORY BIRDS IN COLORADO INTRODUCTION There is widespread concern that populations of many species of NearcticNeotropical migratory passerine birds have declined in the last 30 years (Robbins et al. 1986, Robbins et al. 1992). The primary source of avian population trend data in North America is the Breeding Bird Survey (BBS), a volunteer-based population monitoring program administered cooperatively by the U. S. Geological Survey and the Canadian Wildlife Service. Although BBS data monitors population trends for many avian species, some species are not sampled adequately because of low densities or geographic distribution. Further, when trends are detected, especially declines in populations, causes for observed changes may not be readily apparent. Also, there is some criticism of BBS data because it is based on population indices, and assumptions about those indices (i.e., density-dependent singing rates, annual constancy of singing rates and detection, etc.), and competence of volunteer observers (proper identification of birds by sight and calls) as well as observer hearing acuity, have not been adequately addressed. Although many Colorado passerine birds are monitored annually with BBS routes, some species are not represented or sampled in numbers too small to ascertain population status and trend, even if we assume BBS indices reflect actual population trends (Colorado Bird Observatory 1991). Distribution of Colorado Birds has been documented in recent years using a latilong approach (Kingery and Graul 1918, Bissel et al. 1982, Kirigery 1988). A breeding bird atlas project has recently been completed which will provide documentation of-avian distribution, nesting, and relative abundance for most of Colorado's breeding birds (Kingery, in press). However, there is no statewide program for monitoring the population status of most Colorado avian species other than game species and some threatened and endangered species, although monitoring of all species is integral to the Division's Long Range Plan (CDOW 1994). Numerous techniques for both intensive and extensive monitoring programs to document status and population trend of avian species are available and their usefulness and shortcomings have been evaluated (Ralph and Scott 1981, Sauer and Droege 1990, Ralph et al. 1993, Ralph et al 1995). Recent monitoring efforts in Colorado and elsewhere have used banding programs similar to those of the Monitoring Avian Productivity and Survival (MAPS) (Desante 1992) or the Breeding Biology Research and Monitoring Database (BBIRD) (Martin et ale 1991). Both programs are time and labor intensive, and many assumptions about the techniques have not been adequately addressed, thereby limiting inferences to a specific site rather than to larger areas. Recently, there has been an emphasis to measure actual densities of animals using distance sampling methods rather than rely on untested population indices to monitor population trends (Buckland et al. 1993). P. N. OBJECTIVES The primary objectives of this study are to evaluate current monitoring programs for Nearctic-Neotopical passerine birds in Colorado, to test and implement a statistically valid population monitoring program for Colorado's breeding passerine birds, and coordinate monitoring of passerine birds in Colorado with other agencies. A second objective of this study was to develop a study plan to test survey methodology on selected species of passerine birds in Colorado. �112 SEGMENT OBJECTIVES 1. Review literature appropriate to monitoring Neotropical migratory and literature on ecology and biology of selected avian species. 2. Develop and implement a program to monitor trends in breeding abundance for Colorado's breeding bird population. 3. Select study areas and initiate research on abundance and productivity of selected species of Neotropical migratory birds in Colorado. 4. Monitor abundance of breeding birds, nest success, and fledging success of selected species of Neotropical migratory birds in Colorado. 5. Compile data and prepare annual progress birds bird report. METHODS Literature concerning monitoring of avian populations and detecting population trends was reviewed and discussed with personnel of the Colorado Bird Observatory (CBO) and other agencies involved in bird monitoring programs in Colorado (U. S. Forest Service, Bureau of Land Management, U. S. Geological Survey - Biological Resources Division). Mist-netting data collected in 1996 and 1997 by CBO personnel using MAPS protocols (DeSante 1992) was evaluated by Drs. David R. Anderson and Kenneth P. Burnham of Colorado State University. Banding data were analyzed to estimate sample sizes needed to reasonably detect population trends. Following analysis of existing data a meeting was held to discuss the results, assumptions of the existing protocol, and design of a more appropriate bird monitoring program for Colorado. RESULTS AND DISCUSSION BBS routes and MAPS stations are the two primary statewide bird monitoring programs in Colorado. Although BBS routes have been conducted in Colorado since 1965, there are too few routes, and poor distribution of routes in Colorado to adequately detect statistical trends in population for most breeding birds (CBO 1997). Further, BBS population trends are based on indices of abundance (number of individuals seen or heard) and there have been no statistical evaluation to date showing the relationship between these indices and actual populations for most passerine bird species. MAPS protocols (Ralph et al. 1993) were used in Colorado by the CBO and other agencies in 11 areas throughout Colorado in 1997 for monitoring avian population trends. OVerall, 1,270 point counts were conducted and 8,704 individuals of 169 species were detected, mist netting resulted in 3,238 captures of 73 species, and 334 nests of 32 species were located (Table 1). CBO provided summary information and the larger data sets were analyzed by Dave Anderson and Ken Burnham to calculate annual adult survival and the potential to use the information to measure population trends. On 20 November the Division of Wildlife hosted a meeting with D. Anderson and K. Burhnam along with the CBO to discuss analysis of the current MAPS data collected in Colorado. Several problems with existing MAPS protocol and resulting data were identified. One major problem was that MAPS sites were not randomly located, thus inferences to any population parameters are restricted to tnat specific �113 Table 1. Results Observatory) Station of Colorado MAPS Points Ii A. M. Bailey Apishapa SWA Bodo SWA Comanche NG Grand Mesa Ken Caryl Ranch Lone Dome SWA Middle Park Monte Vista NWR Pawnee NG Rocky Mountain NP Totals 0 100 100 250 103 0 98 105 230 250 34 1,270 stations, 1997. Detected Ii 0 625 908 1,472 558 0 734 799 2,182 1,234 192 8,704 (Data from Colorado Captures Ii 440 0 1000 0 601 680 0 140 0 0 377 3,238 Bird Nests Ii 46 0 73 58 16 10 0 102 0 0 29 334 banding or survey site. In Colorado, these sites are not representative as most were in areas of public lands that were easily accessible and were protected from grazing, fires, logging, and other typical land management practices that likely affect avian demography and populations. Thus, use of MAPS data may not reflect avian population trends on a statewide basis. The second problem with MAPS data, especially the constant-effort mist netting portion, is that samples of most species captured were not sufficient to calculate reasonable estimates of annual survival and productivity. For the best data, the 95% confidence intervals around survival estimates were quite large, and, for most species, the estimate covered the entire universe (survival from 0.0 to 1.0) and, thus, provided no useful data. Only 1 species on 1 site was sampled adequately, and that occurred by combining males and females (in reality, one might expect each gender to have different demographic parameters). Given the cost of establishing and operating a single MAPS station, and the small samples obtained for most sepcies, the data obtained did not appear cost effective. The decision resulting from the November meeting and analysis of existing data was to terminate MAPS as a method for monitoring avian population trends in Colorado and replace it with a system based on distance-measured transects and point counts (Buckland et al. 1993). The CBO developed a new protocol (Monitoring Birds 2001) which will establish 30 permanent transects in each of 12 habitat types in Colorado and have 15 points along each transect. Distance to each bird observed will be recorded and will result in density estimates for each species. Additionally, populations of some species (e.g., owls, some uncommon species) will need to be tracked using special techniques. After 3-5 years, sufficient data should be available to estimate baseline population abundances and trends may be come apparent within 15-20 years. Management efforts can then be focused on declining species. Annual analysis of data collected using the new protocols will be completed to ascertain which species lack adequate sample sizes to calculate densities and trends. Additional survey efforts or different techniques will be used to monitor populations of these species. �114 LITERATURE CITED Bissell, S. J., C. Chase III, H. E. Kingery, and W. D. Graul. 1982. Colorado bird distribution latilong study. Colorado Div. Wildlife, Denver. 78 pp. Buckland, S. T., K. R. Anderson, K. P. Burnham, and J. L. Laake. 1993. Distance sampling: estimating abundance of biological populations. Chapman & Hall, New York. 446 pp. Colorado Bird Observatory. 1997. 1996 reference guide to the monitoring and conservation status of Colorado's breeding birds. Colorado Bird Observatory, Colorado Div. Wildl., Great outdoors Colorado Trust Fund, and Partners, Denver. Colorado Division of Wildlife. 1994. State of Colorado, Dep. Nat. Resour., Div. Wildl. Long Range Plan. Denver. 34 pp. DeSante, D. F. 1992. MAPS (Monitoring Avian Productivity and Survival) General Instruction. Institute for Bird Populations, Point Reyes, CA. Kingery, H. E. 1988. Colorado bird distribution latilong study. Colorado Div. Wildlife, Denver. 81 pp. Kingery, H. E, and W. D. Graul. 1978. Colorado bird distribution latilong study. Colorado Div. Wildlife, Denver. 58 pp. Martin, T. E., C. Paine, C. J. Conway, W. M. Houchachka, P. Allen and W. Jenkins. 1997. BBIRD Field Protocol. U. S. Geological Survey, Biol. Resour. Div., Montana Coop. Wildl. Res. Unit, Univ. Montana, Missoula. 64 pp. Ralph, C. J., and J. M. Scott, editors. 1981. Estimating numbers of terrestrial birds. Studies in Avian Biology No.6. Cooper Ornithol. Soc., Allen Press, Inc., Lawrence, KS. 630 pp. Ralph, C. J., J. R. Sauer, and S. Droege. (Eds.). 1995. Monitoring bird populations by point counts. U. S. Dep. Agric. Forest Serv., Gen. Tech, Rep. PSW-GTR-149. Albany, CA. 187 pp. Ralph, C. J., G. R. Geupel, P. Pyle, T. E. Martin, and D. F. DeSante. 1993. Handbook of field methods for monitoring landbirds. U. S. Dep. Agric., Forest Serv., Gen. Tech. Rep. PSW-GTR-144. Albany, CA. 41 pp. Robbins, C. S., D. Bystrak, and P. H. Geissler. 1986. The breeding bird survey: its first fifteen years, 1965-1979. U. S. Pep. Inter., Fish and Wildlife Servo Resour. Pub. 157. 196 pp. Robbins, C. S., J. R. Sauer, and B. G. Peterjohn. 1992. population trends and management opportunities for neotropical migrants. Pages 17-23 in P. M. Finch and P. W. Stangel (eds). Status and management of Neotropical migratory birds. U. S~ Pep. Agric. Forest Serv., Rocky Mountain Forest and Range Exper. Sta., Gen. Tech. Rep. RM-229. Fort Collins, CO. 422 pp. Sauer, J. R., and S. Proege, editors. 1990. Survey designs and statistical methods for the estimation of avian population trends. U. S. Pep. Inter., Fish and Wildl. Serv., Biological Rep. 90(1). 166 pp. Prepared by: ~~~~~~~~~._~~~~,~ Kenneth M. Giesen Wildlife Researcher _ �115 Colorado Division Wildlife Research April 1998 of Wildlife Report JOB PROGRESS State of Colorado Project: W-167-R Work Plan __~2~9~__ Job Title: Period Covered: Author: Gerald Personnel: Wildlife Ayian Research 1 Job Identify REPORT Distribution and Reproductiye 1 July - 31 December Status of Mountain Plovers 1997 R. craig Gerald R. Craig and Kenneth M. Giesen, Colorado Division of ABSTRACT Literature searches were conducted. The current status of mountain plover (Charadrius montanus)was reviewed and evaluated at a Division sponsored workshop attended by agencies within the species' breeding range. Suggestions for additional investigations were developed from this workshop and incorporated into a preliminary study plan. This Job Progress Report change. For this reason, QUOTED without permission represents a preliminary analysis and is subject information presented herein MAY NOT BE PUBLISHED of the author. to OR ��117 IDENTIFY DISTRIBUTION AND REPRODUCTIVE STATUS OF MOUNTAIN PLOVERS INTRODUCTION The mountain plover (Charadrius montanus) occurs as a summer resident on the eastern Colorado plains in Weld, Bent, Cheyenne, Crowley, and Kiowa counties and a small population also exists in Fremont and Park counties. Historically, plovers were numerous in eastern Colorado (Bailey and Niedrach 1965), but populations have declined over the past 2 decades. In 1968, Graul and Webster (1976) estimated 21,000 birds occurred in Weld County alone. In a little more than 2 decades later, Knopf (1991) assessed the national population at less than 10,000 individuals of which approximately half may breed in Colorado. It is estimated that the species is undergoing an annual decline of 3.7%, the primary cause of which appears to be conversion of native prairie grasslands to cultivation. In Colorado, the mountain plover is currently classified as a Species of Special Concern. The species is a Federal Candidate Species and a petition was recently submitted for listing under the Endangered Species Act. To date, pioneering efforts by Graul (1975) and Knopf and Rupert (1996) have described the plover's natural history and breeding requirements on the short grass prairie of eastern Colorado. Although the species has declined throughout its range, the recently completed Colorado Breeding Bird Atlas suggests that mountain plovers may be more widespread than previously known. Additional investigations are warranted in Fremont and Park counties as well as the San Luis Valley to gain a better understanding of the extent and status of intermountain populations. Finally, standardized methods need to be developed and tested to accurately inventory and monitor productivity . P. N. OBJECTIVES The objectives of the initial planning are to (1) review literature pertinent to monitoring status and trends of mountain plovers, (2) evaluate existing efforts which monitor mountain plovers in Colorado, (3) expand reproductive monitoring efforts to other regions of the state, particularly South Park and portions of southeastern Colorado, and (4) develop a detailed study plan for monitoring mountain plover reproductive status and population trends. SEGMENT OBJECTIVES 1. Review literature pertinent to this study. 2. Interview and cooperate with other authorities currently investigating mountain plovers in Colorado. 3. Formulate a study plan to monitor and evaluate population trends. 4. Develop strategies to assure secure populations. �118 RESUL TS AND DISCUSSION A literature search was undertaken on mountain plovers with particular emphasis on plovers nesting in Colorado and the surrounding states. Records were stored in the "Notebook"database. On 3-4 December, 1997 the Colorado Division hosted a workshop on mountain plovers at Denver. Fifty-eight participants representing federal and state biologists and land managers from California, Colorado, Kansas, Montana, Nebraska, New Mexico, Oklahoma, Texas, Utah, and Wyoming were in attendance. The group reviewed distribution and population status throughout the plover's range and then focused on future needs for habitat descriptions, census methods and regional coordination. In addition to updating the current state of knowledge about mountain plovers, the workshop identified several focus areas. Statewide population inventories and trends are obvious areas of responsibility, however, as well as recent findings that suggest plovers are nesting on cultivated fields and recently burned prairie. A study plan was developed and submitted for peer review. The plan proposed the following objectives: . 1. Develop, implement, and evaluate a statewide monitoring program using Division field personnel. 2. Investigate contributions of burned grasslands to plover productivity. 3. Investigate contributions of fallow cultivated fields to plover productivity. At present, plover conservation measures have not been initiated. Strategies to protect and enhance plovers will be developed and tested as more information is accumulated in the course of this investigation. LITERATURE CITED Bailey, A.M., and RI Niedrach. 1965. Birds of Colorado, Denver Mus. Nat. Hist., . Denver, CO. Vol I. Graul, W.D. 1975. Breeding biology of the mountain plover. Wilson Bull. 87:6-31. Graul, W.O., and L.E. Webster. 1976. Breeding status of the mountain plover. Condor 78:265-267. Knopf, F.L. 1991. Status and conservation of mountain plovers: the evolving regional effort. Report of research activities, U.S.Dep. Inter., Fish and Wildl. Serv., National Ecology Research Center, Fort Collins, 9pp. Knopf, F.L., and IR Rupert. Productivity and movements of mountain plovers breeding in Colorado. Wilson Bull. 108:28-35. Prepared By: C R G-...tJ Gerald R Craig LSSRIV . � https://cpw.cvlcollections.org/files/original/9603a52c210cd112aa59fd55a1761d43.pdf 88ee5ed929728fa1ac8d45e73a8dd4b4 PDF Text Text 1 Colorado Division Wildlife Research July 1998 of Wildlife Report JOB state of Project Colorado No. Work Package No. __~0~6~6u2~ Author: Covered: Tanya July 1, 1997 - REPORT Cost Center W-153-R-11 Task No. Period PROGRESS Mammals _ 3430 Program Preble's Meadow Conservation Jumping Mouse Preble's Meadow Conservation Jumping Mouse June 30, 1998 Shenk ABSTRACT A draft conservation plan, entitled 'Conservation Assessment and Preliminary Conservation Strategy for Preble's Meadow Jumping Mouse (Zapus hudsonius preblei), was completed and submitted to the .USFWS on January 9, 1998 during the Open Comment Period for the Proposed Listing of Preble's meadow jumping mouse as endangered. The conservation plan was divided into two parts. The first part, a conservation assessment summarized and evaluated available information on the taxonomy, distribution, and ecology of Z. h. preblei and identified potential threats to the conservation of the mouse. The second part, a preliminary conservation strategy outlines the goals and objectives of a conservation strategy for Z. h. preblei and summarized prioritized research needs to provide necessary information to develop sound conservation strategies. From this list of prioritized research needs two study plans were completed and research begun on both starting in May 1998. These research plans are designed to investigate (1) temporal and spatial variation in the demography of Preble's meadow jumping mouse and (2) habitat use and distribution of Preble's meadow jumping mouse in Larimer and Weld Counties, Colorado. ��3 Preble's Meadow Jumping Mouse Tanya Shenk P.M. Develop and implement Colorado. Conservation Plan OBJECTIVE a conservation plan SEGMENT for Preble's meadow jumping mouse OBJECTIVES 1. Complete a literature search for all known, relevant information develop a conservation plan for Preble's meadow jumping mouse. 2. Complete an inventory meadow jumping mouse. 3. Evaluate known information on Preble's meadow jumping context of providing information necessary to develop for conservation of Preble's meadow jumping mouse. 4. Define and prioritize needed research components to develop sound strategies for conservation of Preble's meadow jumping mouse. S. Begin study design for research to address a high priority need to develop sound strategies for conservation of Preble's meadow jumping mouse. 6. Complete a draft of current conservation RESULTS in research plan being for Preble's conducted meadow to on Preble's mouse sound in the strategies jumping mouse. AND DISCUSSIQN 1. Complete a literature search for all known, relevant information to develop a conservation plan for Preble's meadow jumping mouse. A literature search was completed, the information was summarized and presented in the draft conservation plan (see Appendix A). 2. Complete an inventory of current research being conducted on Preble's meadow jumping mouse. There are four research projects being conducted on Z. h. preblei, with a fifth planned to begin spring of 1998 (Table 1). Each of these projects is designed to provide further information on the ecology or demography of Preble's meadow jumping mouse. However, if research methodologies were standardized and coordinated across all the projects, comparability and quality of the data would be greatly enhanced. Such a coordinated effort would maximize data quantity, quality, and comparability over varying conditions throughout the range of Z. h. preblei in the most effective and efficient way possible. Comparable information gained across the ecological range of Z. h. preblei would then provide more useful information to develop sound management strategies for conservation of the mouse. To achieve such a coordinated research effort all project leaders must agree to follow data collection protocols, establish 'ownership' of .data and future publications, and work cooperatively to share equipment and technical assistance during peak data collection periods. To facilitate such a mutual effort will require the participation of all �4 project leaders conducting field studies of Z. h. preblei as well as cooperative investigators, and specialized data analysts with expertise in the study design, analysis, and interpretation of mark-recapture and telemetry data. Therefore, I organized a Preble's Meadow Jumping Mouse Research Working Group with the following objective: To explore the idea of coordination of research efforts across multiple principal investigators to enhance comparability and quality of data across the range of Z. h. preblei for use in developing sound management strategies for conservation of the mouse. Such an effort would maximize data quantity, quality, and comparability over varying conditions throughout the range of Z. h. preblei in the most effective and efficient way possible. To make this work, all principle investigators must agree to follow data collection protocols, establish 'ownership' of data and future publications, and work cooperatively on the possible sharing of equipment and technical assistance during peak data collection periods. This working group needs to involve all interested principle investigators involved in field studies of Z. h. preblei as well as cooperative investigators, and specialized data analysts with expertise in the study design, analysis, and interpretation of mark-recapture and telemetry data. The first meeting of the working group (November 12, 1997) was a success with all principle investigators supportive of the idea of a coordinated, cooperative research program. The second meeting of the Research Working Group is scheduled for February 4, 1998 where we will outline what research questions will be addressed at which sites and draft standardized protocols for data collection. A proposed agenda for the Research Working Group includes the following tasks, task leaders, and a timetable to initiate a cooperative research effort: Task Task Leader, Affiliation Date Identify all potential participants: project leaders, cooperating investigators, data analysts. T. Shenk, Colorado Division of Wildlife January 1998 Prioritize questions. All project leaders* January 1998 Identify sites to best address prioritized research needs. All project leaders January 1998 Design studies specific to research question to be addressed at each site. All project leaders, cooperating investigators, data analysts JanuaryFebruary 1998 Evaluate needs at each site to conduct specified research. All project leaders March 1998 Identify discrepancies between needs and available funds, equipment, and personnel currently allocated for each site. All project leaders March 1998 research Attempt to balance discrepancies. All project leaders, cooperating investigators * To date: M. Bakeman, C. Meaney, March T. Ryon, 1998 R. Schorr, T. Shenk �5 3. Evaluate known information on Preble's meadow jumping mouse in the context of providing information necessary to develop sound strategies for conservation of Preble's meadow jumping mouse. Known information on the ecology of Preble's meadow jumping mouse was summarized and presented in the Conservation Plan (see Appendix A). 4. Define and prioritize needed research components to develop sound strategies for conservation of Preble's meadow jumping mouse. The following outline is a complete list of research needs to provide 'information to develop sound management strategies for conservation of Z. h. preblei. Research needs are not listed in order of priority, but listed in a logical manner to provide completion of needs. Demographic studies: Information on the population dynamics of Preble's meadow jumping mouse is necessary to determine which areas support populations where the rate of population growth is ~ 1. Key parameters to estimate include: Survival: • estimates of survival, including ~ annual ~ over-summer ~ over-hibernation • investigate possible factors affecting survival, including ~ weight sex age abundance (i.e., density dependent response) habitat features: stream reach, vegetation composition weather ~ predation ~ disease Approach: mark-recapture techniques Recruitment: • estimate recruitment (as an alternative to reproductive parameters listed below) • investigate possible factors affecting recruitment, including ~ weather ~ habitat features Approach: mark-recapture techniques Population structure: • estimate sex ratios • estimate age ratios Approach: mark-recapture techniques Abundance: • estimate abundance • investigate factors affecting abundance, including ~ habitat features (see under habitat use) Approach: mark-recapture techniques for closed populations Immigration, emigration: • estimate rates of immigration and emigration �6 • investigate possible factors affecting immigration, emigration • habitat features (see under habitat use) • abundance (i.e., density-dependent response) Approach: estimate rates from mark-recapture data, identify existence with radio telemetry Reproduction: • estimate number of litters per year • estimate number of young per litter • estimate age at first reproduction • estimate juvenile survival • investigate possible factors affecting reproduction, • habitat features: food availability, nest site availability, cover availability • abundance (i.e., density-dependent response) • weather Approach: from telemetry work including Dispersal Studies: Dispersal is a key process in metapopulation and to maintain genetic diversity between isolated populations. parameters to evaluate include: Population parameters • who disperses • time of dispersal • estimate rate Approach: estimate rate with mark-recapture, from both telemetry and mark-recapture data document theory Key who and when Habitat parameters through what habitat • end point descriptions (disperses from what to what) • landscape features (connectivity with other riparian strips, corridor use, overland use) Approach: document where from both telemetry and mark-recapture data Distribution Studies: Conservation of Z. h. preblei should maintain populations throughout the range of its natural variation and to try to identify ecological limits for the subspecies. Key concerns include: Range-wide • conduct distribution: trapping surveys to better define the eastern boundary of the range of Z. h. preblei • conduct trapping surveys in areas of potential h. preblei with Z. princeps Approach: determine from trapping surveys Habitat preblei sympatry Studies: To identify and define habitat requirements studies should be conducted to address the following: Habitat use: • estimate distances • describe landscape sites, geology) • determine seasonal traveled: daily, seasonally features (connectivity with other use of Z. of Z. h. potential �7 describe hibernacula describe nest sites • estimate distance to nearest open water from other habitats used • describe hydrology (water quality, flow) in areas of use • evaluate effects of abundance on habitat used (i.e., density dependent responses) Approach: estimate from telemetry work Physiological Studies: Physiological studies on the mechanisms driving habitat selection. will provide information Physiological requirements: • estimate dependency of z. h. preblei on open water • determine energetic requirements to survive hibernation Approach: estimate from laboratory studies Systematic Studies: To better define the relationship of Z. h. preblei to other subspecies of Z. hudsonius and other species of Zapus the following studies should continue. Molecular systematic relationships: • further explore genetic relationships among different preblei populations • further explore genetic relationships among different subspecies of Z. hudsonius • further explore genetic relationships among different of Zapus. Approach: explore through laboratory studies Systematic relationships: • link genetic relationships to systematic hudsonius. Approach: explore through museum studies studies Z. h. species of Z. Community Studies: Composition of the community where Z. h. preblei occur could help explain ecological tolerances of the subspecies, providing insight to the mechanisms determining its distribution. Small mammal assemblages: • comparison of small mammal assemblages in areas where populations of Z. h. preblei occur and areas where they do not • species composition • relative abundance Approach: estimate from trapping surveys There are currently five research projects on Z. h. preblei planned to begin spring of 1998. Each of these projects is designed to provide further information on the ecology or demography of Preble's meadow jumping mouse. However, if research methodologies were standardized and coordinated across all the projects, comparability and quality of the data would be greatly enhanced. Such a coordinated effort would maximize data quantity, quality, and comparability over varying conditions throughout the range of Z. h. preblei in the most effective and efficient way possible. Comparable information gained across the ecological range of Z. h. preblei would then provide more useful information for use in developing sound management strategies for �8 conservation of the mouse. Therefore, a research group was formed including all principal investigators of the five studies as well as other interested personnel. This research group has developed standardized protocols to be followed in all studies. These include protocols on techniques, data collection, and data analyses for (1) mark-recapture, (2) radio-telemetry, (3) habitat use, and (4) genetic sampling. 5. Begin study design for research to address a high priority need to develop sound strategies for conservation of Preble's meadow jumping mouse. There are four components of Z. h. preblei ecology that are currently unknown and yet key to any sound conservation strategy for the subspecies. These are (1) detailed demographic studies estimating survival and reproduction and determining the factors influencing each of the parameters, (2) detailed studies evaluating movements and dispersal habitat of individuals within and among populations, (3) detailed studies to define hibernation needs, primarily descriptions of suitable hibernacula criteria and food requirements for sufficient fat storage prior to immergence, and (4) better defined distributions of the mouse. The following describes the research I will be conducting beginning Spring 1998 to address these needs. Demography Study: Information on the population dynamics of Preble's meadow jumping mouse is necessary to determine which areas support populations where the rate of population growth is stable or suggests an increasing population. Key parameters to estimate include survival (annual, over-summer, and over-hibernation), reproduction, dispersal rates, and density. A combination of mark-recapture techniques and radio-telemetry studies will be used to estimate these parameters and evaluate possible factors affecting such parameters. Intensive trapping efforts will be conducted in June and August of 1998 and in May of 1999 within a given population. All animals captured will be permanently marked with PIT (passive integrated transponders) tags. Select mice will also be fitted with radio transmitters. Survival estimates and evaluation of possible factors affecting survival, including weight, sex, age, abundance (i.e., density dependent response), habitat features (stream reach, vegetational composition), and weather will be obtained from data collected from mark-recapture efforts. Dispersal is a key process in meta-population theory and to maintain genetic diversity between isolated populations. The key objectives are to estimate rate of dispersal and to determine who disperses, time of dispersal, and describe suitable dispersal habitat. These estimates will be obtained from analyses of both the markrecapture data and from following radio-collared individuals. Estimates of immigration and emigration rates will be obtained from data collected during the mark-recapture efforts as well as an evaluation of possible factors affecting these rates including habitat features and abundance (i.e., density-dependent response). Data on who disperses, when dispersal occurs, and dispersal habitat will be obtained by following radio-collared animals. From the mark-recapture data collected during the trapping sessions, abundance and density estimates can also be made. Other ancillary information which can be obtained from data collected during �9 the tapping sessions include estimates of sex and age ratios within the populations sampled. Reproduction parameters such as number of litters per year, number of young per litter, and age at first reproduction will be estimated from radio-collared females. An analysis of variance will be conducted to evaluate possible factors affecting reproduction, including habitat features such as food availability, nest site availability, and cover availability as well as the possible effects of abundance (i.e., density-dependent response) and weather. A complete study plan entitled "Temporal and spatial variation in the demography of Preble's meadow jumping mouse (Zapus hudsonius preblei) is included in Appendix B. Distribution, Habitat Use, and Monitoring study: Conservation of Z. h. preblei should maintain populations throughout the range of its natural variation and to try to identify ecological limits for the subspecies. Key concerns include range-wide distribution, habitat use, and population persistence at a given site. To address these concerns a study will be designed where a sampling frame of suitable habitat will be constructed. The starting point for developing the sampling frame will be to determine the feasible distributional range of Z. h. preblei. A conservative approach will be used for elevational and directional limitations. Once the potential distribution map is developed, all but potentially suitable habitat within the feasible range of the subspecies will be eliminated. From this sampling frame, random sites will be selected to conduct trapping surveys. Therefore, all successful trapping sites will provide further information on the distribution of the mouse. Information on habitat will be collected at all sites surveyed, regardless of trapping success for Preble's meadow jumping mouse. Comparisons of habitat variables from both successful and unsuccessful sites will be used to better define suitable habitat for the subspecies. Habitat variables recorded will include both site level characteristics such as vegetational composition, cover, and composition of 'other small mammal species as well as landscape level characteristics including connectivity with other potential sites, geology, hydrology, and distance to development. Data collected on movements of radiotelemetered mice will also provide information on seasonal use of different habitats, dispersal corridor and end point habitat characteristics, and descriptions of nest sites and hibernacula. Repeated annual visits to the randomly selected sites will provide information on population persistence at a given site. Such a monitoring scheme will be necessary for evaluating the continued status of the mouse as well potentially evaluating the efficacy of any conservation planning efforts. Genetic tissue samples will also be collected from mice captured at these new locations. Future genetic analyses should help to better define the relationship among (1) different populations of Z. h. preblei, (2) other subspecies of Z. hudsonius and (3) other species of Zapus. A complete study plan entitled "Habitat use and distribution of Preble's meadow jumping mouse (Zapus hudsonius preblei) in Larimer and Weld Counties, Colorado" is included in Appendix C. 6. Complete a draft conservation plan for Preble's meadow jumping mouse. The draft conservation plan, entitled 'Conservation Assessment �10 and Preliminary Conservation strategy for Preble's Meadow Jumping Mouse (Zapus hudsonius preblei), was completed and is attached as Appendix A. The document was submitted to the USFWS during the Open Comment Period for the Proposed Listing of Preble's meadow jumping mouse as endangered. prepared by ~';: s-!:!"J__ �Table 1. Current and proposed research projects for Preble's Marking method Study location Principle Investigato r Pit tags Telemetr y meadow jumping mouse. Parameters Surviva 1 Reproducti on Recruitme nt to estimate Abundanc e Dispers Habita Populatio al t Uset n structure Stomach content analyses Rocky Flats Tom Ryon x x x x x AOA* Air Force Academy Rob Schorr x x x x x AOA Boulder Open Space Carron Meaney x x x AOA Lyons Mark Bakeman x x AOA Cherokee Park ? Tanya Shenk x x x x x x x x AOA Tanya Shenk x x x x x x x x AOA Douglas (Plum Creek)? cty ... * AOA As opportunity ----- - arises I-' I-' ��13 APPENDIX A CONSERVATION ASSESSMENT AND PRELIMINARY CONSERVATION STRATEGY FOR PREBLE'S MEADOW JUMPING MOUSE (Zapus hudsonius preblei) prepared by Tanya Shenk Colorado Division of Wildlife 317 West Prospect Fort Collins, Colorado 80526 January 1998 �14 EXECUTIVE SUMMARY In 1994 a petition was submitted to the U. S. Fish and Wildlife Service from the Biodiversity Legal Foundation (USFWS 1997a) to list Preble's meadow jumping mouse (Zapus hudsonius prebleit under the Endangered Species Act (ESA). A Proposed Rule to list Preble's meadow jumping mouse as endangered under the ESA was submitted by the U. S. Fish and Wildlife Service on March 25,1997 (USFWS 1997a). Following the Proposed Rule to list the species as endangered, the Colorado Division of Wildlife agreed to develop a species conservation assessment and preliminary conservation strategy which would provide a framework for conservation efforts. This document is the result of that agreement between the Colorado Division of Wildlife and The U. S. Fish and Wildlife Service. Therefore, the purpose of this document is to summarize the current known ecology and status of Z. h. preblei, to outline goals for conservation of the subspecies, to prioritize what information is most needed to develop sound conservation strategies to meet those goals, and to suggest future research required to provide such information. This document will provide the Colorado Division of Wildlife with a prioritized list of research needs to direct ecologists and managers toward obtaining the necessary information for developing, implementing, and evaluating strategies for conserving the Preble's meadow jumping mouse. Should the Final Decision list the species as either threatened or endangered, this document could also provide scientific information for any Habitat Conservation Plan(s) to be developed under the ESA. Any conservation plan for Z. h. preble; should address what is needed for recovery of the subspecies, including how much habitat is needed and where, what information and conservation actions are needed, whether restorations are needed, and what biological information is needed to manage or conserve the mouse. Some of the basic information required to defme these needs includes the distribution of the mouse and its habitat, population dynamics (survival, reproduction, and dispersal), minimal habitat requirements, genetic variation between and among populations, physiology, hibernation requirements, and resilience of populations to human alteration of their habitat. This document attempts to summarize what is currently known on each of these topics, prioritizes what information is still most needed to develop conservation strategies based on the conservation goals, and suggests future research required to provide missing information. A review of the studies conducted on Preble's meadow jumping mouse shows that there is insufficient information to fully address defming range-wide ecological requirements, limiting factors, limits of species tolerance, or population status. Most work to date has focused on geographic distribution (presence or absence of Z. h. prebleii, taxonomy, and habitat descriptions of sites where mice have and have not been captured. Although available information is limited, it is clear that measures must be taken to begin developing a conservation strategy for Z. h. preblei. The primary objectives necessary to develop a successful conservation strategy for Preble's meadow jumping mouse are: 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. Document the present distribution of Z. h. preblei. Identify populations of Z. h. preble; where the rate of population growth ~ 1. Protect populations of Z. h. preble; where the rate of population growth ~ 1. Maintain current range of natural variability of Z. h. preblei. Identify ecological requirements for sustaining viable populations of Z. h. preblei throughout its range of natural variability. Protect habitats to sustain existing populations of Z. h. preble; where the rate of population growth z 1. Promote protection and management of habitat for conservation of Z. h. preble; in all currently or recently occupied habitat, and in habitat suitable for restoration of mouse populations. Monitor the status of populations of Z. h. preblei throughout its known range to detect changes in local distribution. Identify threats to the conservation of Z. h. preblei. Eliminate or minimize threats to conservation of Z. h. preblei. �15 11. 12. 13. Integrate Preble's meadow jwnping mouse conservation strategy objectives with management and habitat objectives of other Front Range riparian species. Promote scientific management of Preble's meadow jwnping mouse. Promote public support for conservation efforts and scientific management of Preble's meadow jwnping mouse through public education. In order to achieve the stated objectives outlined above, the following actions must occur: 1. 2. 3. 4. 5. . 6. 7. 8. 9. Conduct population studies to estimate and determine what factors influence demographic parameters and rate of population change. Conduct studies to better defme the ecological requirements of Z. h. preble; (food, cover, water, hibernation requirements, etc.). Conduct research to evaluate the effects of potential threats on survival, reproduction, dispersal, and abundance of Preble's meadow jwnping mouse. Implement habitat and population management practices that emphasize the conservation of Preble's meadow jwnping mouse throughout the natural variability of its range. Develop survey protocols to monitor trends in the distribution of Z. h. preble; and protocols to store and analyze survey data. Conduct trapping surveys to better defme the boundaries of the range of Z. h. preblei. Implement research on Preble's meadow jwnping mouse biology to better understand the mechanisms driving the ecological requirements. Further explore genetic relationships among different Z. h. preble; populations, among different subspecies of Z. hudsonius, and among different species of Zapus. Develop educational programs to promote positive public support for Preble's meadow jwnping mouse conservation. The criterion to be used to evaluate the success of this program will be the maintenance of local selfsustaining populations of Preble's meadow jwnping mouse which are geographically distributed throughout the range of the subspecies and throughout the natural variation of its ecological requirements. The docwnent is divided into two parts. The first part, a conservation assessment will (1) summarize available information on the taxonomy, distribution, and ecology of Z. h. preblei; and (2) identify potential threats to the conservation of the mouse. The second part, a preliminary conservation strategy, will (1) outline the goals and objectives of a conservation strategy for Z. h. preblei; and (2) summarize prioritized research needs to provide necessary information to develop sound conservation strategies. ACKNOWLEDGMENTS This docwnent could not have been prepared without generous help from all members of the 'Z. h. preble; Conservation Docwnent Working Group' (Mark E. Bakeman, Carron A. Meaney, Chris Pague, and Peter Plage). Members gave freely of their time and expertise and readily shared their data, anecdotal observations, impressions, and scientific interpretations. This working group truly exemplified how scientists should interact when working toward a common goal. Additional scientific contributions came from David M. Armstrong, Alan B. Franklin, Lawrence A. Riggs, Thomas R Ryon, Robert Schorr, Rick Schroeder, and Bruce Wunder. Technical support during this effort was provided by Alan B. Franklin, David Lovell, Francie Pusatari, and Michael Wunder. Logistical support was provided by Steven Nesta, Douglas Robotham, Judy Sheppard, and John Stover. �16 CONSERVATION ASSESSMENT INTRODUCTION The following is a summary of information collected primarily from the five studies conducted on Z. h. preblei. The longest and most varied studies have been conducted on the Rocky Flats Environmental Technology Site (1990 through the present) including distribution studies, habitat surveys, location ofhibernacula, fat requirements for hibernation, and genetic studies conducted by M. Bakeman, A. Deans, F. Harrington, T. Ryon, R Stoecker, and B. Wunder. Range-wide distributional and vegetation surveys have been conducted on sites other than Rocky Flats Environmental Technology Site by (1) Meaney et al. (1995, 1996, 1997), funded by the Colorado Division of Wildlife, (2) City of Boulder Open Space and Boulder County Open Space, (3) Ensight Technical Services of Boulder, Colorado funded by the Colorado Department of Transportation, (4) the Colorado Natural Heritage Program, (5) U. S. Forest Service, Medicine Bow National Forest, Wyoming, and (6) numerous other environmental consulting firms. An evaluation of historical Z. h. preblei sites was conducted by Ryon (1996). Genetics studies were conducted in 1995 by B. Wunder and in 1996 and 1997 by Biosphere Genetics, Inc. Where data were not available for Z. h. preblei, information from studies of other subspecies of Z. hudsonius was provided. In particular, studies conducted by Quimby (1951) and Whitaker (1963) provided substantial information on the species. Caution must be used in extrapolating information reported for other subspecies of Z. hudsonius to Z. h. preblei, although, such information is useful as a general framework from which to work until such information becomes available for Z. h. preblei itself Possible threats to the conservation of Preble's meadow jumping mouse were derived from both general principles of conservation biology and information specific to Zapus hudsonius in general, or the subspecies itself. TAXONOMY Description Quimby (1951) described Z. hudsonius as follows: 'A mouse-like rodent with greatly enlarged hind feet and an exceptionally long tail. The forelegs are relatively short. The ears are somewhat conspicuous. The body is clothed in moderately long, somewhat dense hair of a rather coarse texture and several colors. The dorsal portions are marked by a broad stripe of brownish hairs many of which are tipped with black giving the region a grayish-black appearance. The sides are bright yellowishorange, whereas the underparts and feet are white. The tail is bicolor, dark above and light below, and sparsely covered with hair which is longer on the terminal part. The mammae are eight, and quite prominent in lactating females. The male genitalia are inconspicuous except during the breeding season when the scrotal sac becomes enlarged. The testes enlarge and may be either abdominal, inguinal, or scrotal during this period.' The skull of Z. hudsonius is small and light with a narrower braincase and smaller molars than in Z. princeps (from Fitzgerald et al. 1994 p. 18). However, Fitzgerald et al. (1994) urge caution in distinguishing the two species 'of Zapus in Colorado. Such caution is further warranted by the recent conflicting identifications of mice based on genetic characteristics (see Genetics section below). E. A. Preble (1899) first collected the specimen that is, now, the holotype for Z. h. preblei at Loveland, Colorado in 1895 and assigned it taxonomically to Z. h. campestris. Krutzsch (1954) revised the taxonomy of Z. hudsonius and assigned the meadow jumping mice from Colorado and southeastern Wyoming to their own distinct subspecies. Krutzch (1954) described this subspecies as having fewer black-tipped dorsal hairs and a less distinct dorsal band; smaller cranial measurements; a narrower interorbital constriction; smaller, less inflated auditory bullae; narrower incisive foramina; and a more inflated frontal region (Fig. 1) than Z. h. campestris. Krutzch named this subspecies Z. h. preblei in honor of the collector of the holotype (and previous reviewer of genus). �17 Figure 1. Preble's meadow jumping mouse (Z. h. prebleii. Note the long tail curves around handlers index fmger and the dark dorsal stripe. Also note the large hind foot and small first digits on front and rear feet (insert). Photographs by Norm Clippinger. Measurements for meadow jumping mice in Colorado are: total length 187-255 mm; length of tail 108-155 mm; length of hind foot 28-35 mm (Fitzgerald et al. 1994). Body weights of meadow jumping mice are variable, not only for different animals, but for the same individual depending on activity or season (Quimby 1951). Such variability arises from sex, age, reproductive status, preparation for entering hibernation, and condition on emergence from hibernation. Mean body weights reported for adult Preble's meadow jumping mice are 199 (n = 37; Meaney et al. 1996),23.5g (n = 5, se = 1.2g; Colorado Natural Heritage Program, unpublished data);22.7g (n = 30, se = 0.7g; M. Bakeman unpublished data 1997), 20.3 g (n = 26, se = 0.5g; Meaney et al. 1997). Two species of ectoparasites have been identified on Preble's meadow jumping mice (R Schorr, personal communication). These includes fleas (Megabothris abantis, (identified by K. Gage of the Center for Disease Control) and a larval form of a tick, either Ixodes sculptus or l. kingi. Genetics The family Zapodidae (jumping mice) consists of small to medium-sized mice with enlarged hind feet and exceptionally long tails. Four living genera are recognized in this family, two of which, Zapus and Napaeozapus, are found in North America (Hall 1981 ). There are three living species of the genus Zapus: Z. trinotatus (Pacific jumping mouse), Z. princeps (western jumping mouse), and Z. hudsonius (meadow jumping mouse). Z. h. preblei is one of twelve living subspecies of the species Z. hudsonius (Fig. 2a, Krutzsch 1954, Hafner et al. 1981). Z. h. preble; was first described by Krutzsch (1954) from a specimen collected by E. A. Preble in 1895 near Loveland, Colorado. Genetics work using DNA sequencing of the mitochondrial DNA non-coding or re D-Ioop.IE region was conducted by Biosphere Genetics, Inc., to help determine whether populations of Z. h. preblei constitute one or more distinct evolutionary units. Genetic samples were collected from livetrapped mice, following a protocol specified by Biosphere Genetics, Inc., adapted and updated from the U. S. Fish and Wildlife Service standardized protocol. Sampling in 1996 and 1997 by all field crews (project leaders M. Bakeman, C. Meaney, T. Ryon, B. Stoecker, Colorado Natural Heritage Program) �18 was performed on meadow jumping mice presumed to be Z. h. preblei, yielding samples from 72 individual mice. Twenty genetic samples were also prepared from specimens provided either directly or indirectly by five different museums and university cooperators. Samples analyzed from livetrapped mice came from 23 locations in Colorado and two locations in Wyoming (Table 1). Samples analyzed from specimens provided by museums and cooperators represented reference material for species and subspecies from Colorado, Indiana, Minnesota, Nebraska, New Mexico, and Wyoming (Table 1; Fig. 2a and b). Preliminary DNA sequencing work in 1996 using a 550 basepair fragment of the mitochondrial DNA non-coding region examined 16 individuals from seven locations in Colorado and found two populations from Las Animas County (Lake Dorothey area) to be distinct from five other populations which ranged from Boulder County south to EI Paso County. As more samples from additional trapping locations and museum specimens were brought in to the analysis in 1997, a smaller segment of the non-coding region included within the original 550 base-pair fragment was used to avoid problems with non-specificity of one of the primers. DNA sequences were obtained for 433 basepair-Iong fragments in all samples analyzed in both years. Phylogenetic relationships among the samples were inferred from maximum parsimony analysis. Strength of these phylogentic relationships were evaluated using bootstrap analysis (see Riggs et al. 1997, Appendix A for detailed methodology). Analysis of the mitochondrial DNA sequence data indicates that mice sampled from southeastern Albany County, Wyoming, south along the Front Range of the Rocky Mountains to western Las Animas County, Colorado (Purgatoire Campground, San Isabel National Forest), form a coherent genetic group (Fig. 3, L. Riggs et al. 1997). This group of samples are distinct from samples obtained from mice from three other populations. Genetic samples from mice captured in the Dorothey Lakes area of southern Colorado (Las Animas County) group together and are most closely allied with Z. h. luteus, a subspecies described from New Mexico (Riggs et al. 1997). The single genetic sample collected from Weld County, Colorado (Lone Tree Creek) and six samples obtained from Warren Air Force Base in Laramie County, Wyoming are most similar to reference samples of Z. princeps from Colorado (Larimer County) and New Mexico (Taos County) (Riggs et al. 1997). The sequence data in:dicate that the samples from specimens identified as Z. h. luteus and samples from specimens of Z. princeps are more closely allied with each other than either is with the samples defining the Z. h. preblei group. This closer alliance of the samples of Z. h. luteus with Z. princeps rather than Z. h. preblei conflicts with results of Hafner et al. (1981), based on a combination of pelage, morphologic, and genetic data, which support a closer alliance with Z. hudsonius. Phylogenetic analysis indicates that the group of samples referred to as Z. h. preblei cannot at this point be distinguished clearly from four reference samples of Z. h. campestris from Weston County, Wyoming (two samples), Custer County, South Dakota (two samples), one sample of Z. h. pallidus from Garden County, Nebraska, or from two samples of Z. h. intermedius from an unspecified county in Minnesota (Fig. 3, L. Riggs et al. 1997). A suggestion from the analyses is that Z. hudsonius from Indiana (presumed to be Z. h. american us) may have shared a common ancestor with the progenitors of samples from Z. h. luteus and Z. princeps more recently than with the Z. h. preblei group and perhaps other subspecies described for the north central states (Riggs et al. 1997). A more complete biosystematic evaluation of jumping mice is needed to clarify and further refine relationships among populations of the group referred to as Z. h. preblei as well as to other subspecies and species of the genus Zapus. Such' an evaluation requires detailed analyses of pelage, morphometric, and genetic data from sufficient numbers of individuals to adequately represent the populations of interest. However, the mitochondrial DNA non-coding (D-Ioop) sequence data available at this time are consistent with the view that a geographically contiguous set of populations previously recognized as Preble's meadow jumping mouse form a homogenous group recognizably distinct from other nearby populations and from another geographically-adjacent species of the genus (Riggs et al. 1997). Therefore, given the genetic data available, Riggs et al. (1997) conclude Preble's meadow jumping mouse is a distinct population of Z. hudsonius. �19· a. '. :z. 1. Z h. tICIId;cvs Z h. aiMccnsls 3. Z h. tl11Ierlcanus 4. Z h. campestrts Z h. canadensis s. 6. 7. 8. 9. 10. 11. 12. Z It hudson;us Z h. Inlermedius Z h. /adas Z h. paJlldus Z h. preblel Z h. tenellus Z h. lul= b. -, Figure 2. Range of Z. hudsonius and its 12 subspecies (a) and Z. princeps (b) (modified from Hall 1981). Stars .indicate locations where reference samples were collected from specimens at either museums or from university cooperators for genetic analysis. Reference samples for Z. h. luteus include two from Otero County, New Mexico, and two samples from Sandoval County, New Mexico. Reference samples for Z. h. campestris include two from Weston County, Wyoming, and two samples from Custer County, South Dakota. Two reference samples of Z. h. intermedius came from Minnesota. Two samples of either Z. h. americanus or Z. h. intermedius came from Indiana. One sample of Z. h. pal/idus came from Garden County. Nebraska. Reference samples for Z. princeps princeps include one sample from Taos County, New Mexico, and four samples from Larimer County, Colorado. �20 Table 1. List of specimens from which samples were taken for genetic analyses. Each specimen is identified by a genetic code name, general location, and origin of the specimen. Code Location BAD BOS BVC CCF CCR CGA CZP DC DMNH Origin of Specimen Badwater Creek, Natrona County, WY hudsonius South Boulder Creek, Boulder County, CO Beaver Creek, EI Paso County, CO Warren Air Force Base, Laramie County, WY Chicorica Creek, Lake Dorothey State Park, Las Animas Co., CO Medicine Bow NF, Albany County, WY Neota Creek, Larimer County, CO U. S. Air Force Academy, EI Paso County, CO San Isabel National Forest, Las Animas County, CO LPC LTC MRD RBC RF ROX SCR STV WFS WHR WPC WRN WRP ZAHU East Plum Creek, Douglas County, CO Hay Gulch, Elbert County, CO Coal Creek, Jefferson County, CO Lake De Smet near Buffalo, Johnson County, WY hudsonius Lone Pine Creek, Larimer County, CO Lone Tree Creek, Weld County, CO Marshall Road, Dry Creek Ditch #2, Boulder County, CO Rabbit Creek, Larimer County, CO Rocky Flats Environmental Technology Site, Jefferson County, CO Roxborough State Park, Douglas County, CO Smith Creek, U. S. Air Force Academy, EI Paso County, CO St. Vrain, Boulder County Open Space, Boulder County, CO West Fork Schwacheim Creek, Lake Dorothey, Las Animas Co., CO Woodhouse Ranch, Douglas County, CO West Plum Creek, Douglas County, CO Monument Creek, EI Paso County, CO Ralston Creek, Jefferson County, CO Indiana ZHCSDC Custer County, SD ZHCWYW Weston County, WY ZHLNMO ZHLNMS ZHPNEC Otero County, NM SandovalCounty,NM Cherry County, NE ZHPNEG Garden County, NE ZPPCOG Grand County, CO ZPPNMT Taos County, NM EPC HAY JCC LDS Museum specimen identified as Z. Z. h. preblei survey Z. h. preblei survey Z. h. preblei survey Museum specimen identified as Zapus Z. h. preblei survey From B. Wunder identified as Z. princeps Z. h. preblei survey Museum specimen identified as Z. princeps princeps Z. h. preblei survey Z. h. preblei survey Z. h. preblei survey Museum specimen identified as Z. Z. h. preblei survey Z. h. preblei survey Z. h. preblei survey Z. h. preblei survey Z. h. preblei survey Z. h. preblei survey Z. h. preblei survey Z. h. preblei survey Museum specimen identified as Zapus Z. h. preblei survey Z. h. preblei survey Z. h. preblei survey Z. h. preblei survey Museum specimen, may be either Z. h. intermedius or Z. h. americanus Museum specimen identified as Z. h. campestris Museum specimen identified as Z. h. campestris Museum specimen identified as Z. h. luteus Museum specimen identified as Z. h. luteus Museum specimen identified as Z. h. pallidus Museum specimen identified as Z. h. pallid us Museum specimen identified as Z. p. princeps Museum specimen identified as Z. p. princeps �21 MRD97&lMNH9716 ROX9701 B059706 EPC9701 ZHCS0C2 _ 70 L051 --- RSS1076. Figure 3. Graphical representation of the genetic relationships among the single most-representative individuals from eachpopulation or reference group of jumping mice sampled as determined by Riggs et al. (1997). The more branching that occurs between the center circle and the edge of the circle, the more dissimilar that sample, or group of samples is from the rest of the samples. The numbers on the lines emanating from the center circle are bootstrap values. Bootstrap values greater than 90 indicate those relationships which are strongly supported. Each specimen location can be derived by matching the sample code with those in Table 1. Numbers following the alphabetical code in this figure denote the sample number used in the analyses from that location. �22 DISTRIBUTION The meadow jumping mouse (z. hudsonius) is broadly distributed across North America from the Atlantic to Pacific coasts, extending south into the United States to Alabama and Georgia and west across the Great Plains to the base of the Rocky Mountains (Fig. 2a). In general, it is a common inhabitant of moist, grassy and herbaceous fields. Eleven living subspecies have been described (Whitaker 1972). Hafner et al. (l981) describe a twelfth subspecies, Z. h. lute us. Z. h. preblei occurs only in eastern Colorado and southeastern Wyoming (Krutzsch 1954, Long 1965, Armstrong 1972). From its limited ecological and geographic distribution, Fitzgerald et al. (1994) suggest it is an Ice Age relict, once widespread in tallgrass prairie across the eastern plains of Colorado but now restricted to scattered localities on the Colorado Piedmont. Similar relict populations of meadow jumping mice in the White Mountains of Arizona and the Sacramento Mountains and Rio Grande Valley of New Mexico are described as the subspecies Z. h. luteus (Hafner et al. 1981). Numerous surveys have been conducted since 1990 to establish the current distribution of Preble's meadow jumping mouse (Fig. 4). Surveys have been funded by the U. S. Fish and Wildlife Service, Rocky Flats Environmental Technology Site, Colorado Division of Wildlife, U. S. Air Force Academy, Warren Air Force Base, Colorado Department of Transportation, City of Boulder Open Space and Real Estate Department, and Boulder County Parks and Open Space Department (Armstrong et al. 1997, P. Plage, personal communication). From 1990 to 1997 such surveys have yielded captures of Preble's meadow jumping mouse, based on field identification and supported by the genetic analyses, at the following sites: 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. Lone Pine Creek, Larimer County, Colorado Rabbit Creek, Larimer County, Colorado St. Vrain Creek and associated tributaries in Boulder County, Colorado City of Boulder Open Space, along South Boulder Creek and its tributaries, Boulder County, Colorado Coal Creek, Jefferson County, Colorado Rocky Flats Environmental Technology Site, Jefferson County, Colorado White Ranch Park, Ralston Creek, Jefferson County, Colorado Plum Creek drainages including Indian Creek, West Plum Creek, and East Plum Creek, Douglas County, Colorado Roxborough State Park, Douglas County, Colorado Hay Creek, Elbert County, Colorado Monument Creek on the U. S. Air Force Academy (AFAC) and tributaries of Monument Creek off the AFAC including Smith Creek, Pine Creek, and Jackson Creek, EI Paso County, Colorado Medicine Bow National Forest, Albany County, Wyoming Two more sites yielded mice that were identified as Z. h. preblei in the field but were genetically found to be more closely allied with Z. princeps (see Genetics section). These sites include: 1. 2. Warren Air Force Base, Laramie County, Wyoming Lone Tree Creek, Weld County, Colorado Similarity of the habitat at these sites compared to those where Z. h. preblei (as determined genetically) were found suggests the possibility of areas of sympatry or parapatry between the two species of Zapus. According to Fitzgerald et al. (l994) the distributions of Z. princeps (Fig. 2b) and Z. hudsonius (Fig. 2a) do overlap. The area of overlap occurs in eastern Wyoming. The distribution of Z. hudsonius in Colorado is now known to be larger than shown in Fitzgerald et al. (l994). The boundaries are currently as far south as Las Animas County (based on genetically identified specimens of Z. h. �23 Goshen Platte 0 Albany 0 0 0 ~":l ~ -c 0 • Wyoming Colorado ~ Laramie I!l Larimer Weld Boulder o Adams Denver Arapahoe • Elbert Pueblo Las Animas a New Mexico Figure 4. Distribution of historical occurrence (0) of Z. h. preblei, sites trapped since 1990 where Z. h. preblei were found (.), sites trapped since 1990 where Z. h. preblei were not found (0), sites where mice were found and identified as Z. h. preblei in the field but were genetically identified as Z. princeps (l!I), and sites where mice were found and identified as Z. princeps but were genetically found to be Z. h. preblei (D). �24 preblei). Captures of Z. princeps and Z. hudsonius (as identified in the museum) have occurred as close as 8 miles within the same drainage (Armstrong 1972). Z. princeps were reported captured in 1981 (Olson and Knopf 1988) at the Lone Pine site in Larimer County, Colorado, where Z. h. preblei were captured this year. Because neither specimens or genetic samples were taken in the 1981 study, identification of those mice will remain in question. The discrepancy may be explained by field misidentification or Meaney et al. (1997) also suggest this discrepancy might be explained by displacement of Z. princeps with Z. h. preblei sometime in the 16 intervening years between trapping efforts. Although assumed to have different ecological requirements, genetic evidence presented here suggests further investigation of possible distributional overlap between Z. h. preblei and Z. princeps. Several notable results from the genetic analysis of specimens acquired from the Denver Museum of Natural History may affect the location of the southern distribution of Z. h. preblei. One site yielded mice identified as Z. p. princeps in the field but were found to be genetically more closely allied with Z. h. preblei. This site is the most southern location to date of Z. h. preblei and is located at: 1. Purgatoire Campground, San Isabel National Forest, Las Animas County, Colorado Also from Las Animas County were mice collected and identified as Z. h. luteus (Jones 1996), an identification also supported by the genetic analysis. These mice were from: 1. 2. Lake Dorothey State Wildlife Area, Chicorica Creek, Las Animas County, Colorado Lake Dorothey State Wildlife Area, West Fork Schwacheim Creek, Las Animas County, Colorado These sites suggest a more northerly distribution of Z. h. luteus, currently known only from limited areas in New Mexico and Arizona (Hafner et al. 1981). Identifying the southern boundary of Z. h. preblei clearly needs further study. Trapping surveys have also provided evidence that a number of historically occupied sites are no longer inhabited by the mouse. Ryon (1996) trapped eight sites of historically occupied sites with no captures of Z. h. preblei. The eight historic sites had suffered either direct disturbances or increasing isolation as a result of past land use, affecting vegetation structure and composition (Ryon 1997). Thus, although the perimeter of the range of Z. h. preblei mayor may not have remained the same, specific locations within the range have definitely changed and may have decreased. The pattern of successful and unsuccessful capture sites (Fig. 4) also suggests that development of the Denver metropolitan area may have created a north-south severing of the range of Z. h. preblei. ECOLOGY The ecology of Preble's meadow jumping mouse has not been studied in detail. However, from the limited information available there appear to be a number of similarities in its natural history and ecological requirements to populations of Z. hudsonius studied in more detail in the eastern and midwestern United States. Therefore, select information available on eastern subspecies of Z. hudsonius is presented below to provide some framework for and possible limitations in ecological tolerances for Z. h. preblei. However, it is critical to acknowledge the limitation and possible misleading effects of extrapolating biological information, natural history, and ecological requirements from one subspecies to another, across the geographic range of a species. Demography Reproduction: Meadow jumping mice have been observed to produce up to three litters per season (Whitaker 1963). Breeding peaks appear to occur in early to mid-June and August with a possible third litter in September (Whitaker 1963). Juvenile Z. h. preblei have been observed in June, August, and September (Meaney et al. 1996, 1997, PTI 1996a, M. Bakeman unpublished data, T. Ryon unpublished data), suggesting two litters per year. Z. hudsonius typically have litters of 5-6 young per �25 litter (Quimby 1951). Age of first reproduction is unknown for Z. h. preblei, however, females of Z. hudsonius have been observed to give birth at 3 months (i.e., females born in June have been observed to give birth in August of the same year). Gestation period is approximately 18 days (Quimby 1951). Young remain dependent on the female for approximately 18 days (Quimby 1951). No evidence of male parental care exists for Z. hudsonius (Whitaker 1963). Survival: No information exists on survival rates for populations of Z. h. preblei. Whitaker (1963) observed an over-winter loss of 67% in a New York population of meadow jumping mice. Most of this loss was.assumed to be from insufficient fat stores to survive hibernation. Besides insufficient fat storage prior to hibernation, other observed mortality factors in Z. hudsonius include predation (Whitaker 1963, Poly and Boucher 1997) and cannibalism (Sheldon 1934). Other assumed mortality factors for Z. h. preblei include starvation, exposure, and disease. Population Structure: Armstrong et al. (1997) reported an overall sex ratio for all captured Preble's meadow jumping mice of 51.6 males: 48.4 females; approximately 86.0% of captures were identified as adults. However, Armstrong et al. (1997) suggested that these data be interpreted with caution because of possible differences in field techniques. Longevity: Very few individuals of Z. h. preblei have been permanently marked. Therefore, recapture information, necessary to determine longevity, is minimal. Several recaptures have yielded adults surviving through two years, indicating a longevity of at least three years (T. Ryon, unpublished data). One recapture history from Rocky Flats Environmental Technology Site recorded an adult male in 1996, two years after it was first captured as an adult in 1994, iildicating survival of at least four years (PTI 1996b). Quimby (1951) found that only a low percentage of Z. hudsonius lived two years or more, but gave two records of mice that lived for at least two years under natural conditions. Whitaker (1963) reported a female living at least two years. Dispersal: Dispersal information will be key to any conservation strategy designed for Preble's meadow jumping mouse. Key factors include (1) which segment of the population disperses, (2) when do they disperse, (3) through what habitat do they disperse, (4) how far will individuals disperse (i.e., what is the maximum distance that separates adjacent populations) and (5) how critical is dispersal (both into and out of a population) to the persistence of a given population. The only data available on dispersal and/or movement for Z. h. preblei are from marked mice at Rocky Flats Environmental Technology Site (T. Ryon unpublished data). Two mice, an adult female and an adult male, were observed approximately 1.6 kilometers from previous locations (incidences occurred separately). Each of the locations were in the same drainage (Woman Creek). Population Persistence: For purposes here, population persistence is defined as the presence of Z. h. preblei at the same site for multiple years. The majority of recent locations of meadow jumping mice have been the result of survey efforts that focused only on determining presence of Z. h. preblei. Once survey protocols (USFWS 1997b) are met [i.e., a minimum of 400 trapnights (one trap set for one night = 1 trapnight) conducted] sites are typically not revisited eliminating the possibility of determining population persistence at these sites. Thus, these and other sites not yet surveyed mayor may not have persistent populations. Areas where trapping was conducted over multiple years as part of further research, yielded three sites in Colorado that support persistent populations: Rocky Flats Environmental Technology Site, the U. S. Air Force Academy near Colorado Springs, and Boulder Open Space on South Boulder Creek. Besides establishing the presence of Z. h. preblei over multiple years, other considerations of persistence include (1) presence in consecutive years versus a 'blinking' in and out of a population as would be expected in a metapopulation [a set of local populations which interact via dispersal of individuals moving among populations and where local extinctions and recolonizations occur (Levins �26 1970)], (2) how populations are sustained in a given area (e.g., source or sink populations), (3) maximum abundance supportable by a given area, and (4) fluctuations in abundance of a population over years. If Z. h. preblei exist in areas as part of a metapopulation or as a series of source-sink populations it would be critical to conserve and protect all key sub-population areas as well as critical habitat for dispersal. Detailed population studies, including estimating and determining factors affecting survival, reproduction, and dispersal rates, must be conducted to determine if Z. h. preblei occurs as either metapopulations or in source-sink populations. There have been no such studies conducted on Z. h. preblei or Z. hudsonius elsewhere to provide any supporting or refuting evidence of this hypothesis. Consistently low abundance in a given area due to limiting ecological requirements (e.g., size of area of suitable habitat) or fluctuations in population abundance, including years of low abundance, must be considered in any conservation strategies for Z. h. preblei because of the threat of complete loss of the population due to catastrophic or extreme environmental conditions. Some evidence exists for such fluctuations in population abundance at Rocky Flats Environmental Technology Site. Trapping surveys on the Woman Creek drainage yielded the following captures of Z. h. preblei: seven in 1993 (EG&G 1993), zero in 1994 (T. Ryon, unpublished data), one in 1995 (T. Ryon, unpublished data), two in 1996 (PTI 1996a), and 33 in 1997 (T. Ryon, unpublished data). Trapping effort was consistent from 1995-1997. Repeated trapping surveys conducted over different years intermittently yielded successful captures of Z. h. preblei along the St. Vrain Creek and lower Coal Creek (three in 1989, zero in 1992, zero in 1994, zero in 1996, one in 1997, M. Bakeman, unpublished data). If protocols in each year were the same, these data also suggest populations may undergo fluctuations in abundance. Hibernation Jumping mice of the genus Zapus are true hibernators, spending much of their lives in hibernation. Meadow jumping mice spend approximately 7 months (~210 days) per year in hibernation (Quimby 1951) whereas estimates for Z. princeps indicate that some populations (e.g., in the western mountains of Utah) spend up to 300 days per year in hibernation (Cranford 1983). Males emerge prior to females (Bailey 1923, Bailey 1929, Hamilton 1935, Quimby 1951, Whitaker 1963) with the earliest annual recorded dates for Z. hudsonius males being April 25-May 16 and females May 4-26. Trapping surveys for Z. h. preblei were not designed to provide estimates for dates of immergence or emergence. However, such dates might be approximated by earliest spring and latest fall capture dates. The earliest spring capture date recorded for an adult male was May 5 in 1993 at Rocky Flats Environmental Technology Site (M. Bakeman unpublished data) and for an adult female, May 21 in 1996 also at Rocky Flats Environmental Technology Site (PTI 1996b). Latest fall capture date for an adult male was October lOin 1994 at South Boulder Creek (ERO Resources 1995) and September 28 in 1995 at the VanFleet Parcel on South Boulder Creek (Armstrong et al. 1997). A juvenile male was captured as late as October 26 and a female juvenile on October 27 both in 1995 at Rocky Flats Environmental Technology Site (M. Bakeman, unpublished data). Whitaker (1963) reported a 67% loss of individuals over hibernation and that average body mass of individuals emerging from hibernation was greater than the average for mice entering hibernation. Because no mice are known to store food in their hibernacula, this indicates that the lighter individuals died during hibernation and only those entering with higher masses survived. All the energy they use during hibernation and the periodic arousals (the energetically most expensive part of hibernation) must be the fat they carry into hibernation (B. Wunder, personal communication). Thus, the ability to put on sufficient fat for overwinter survival during hibernation is a critical factor in the life history of these mice. Jumping mice hibernate in underground burrows (Quimby 1951, Whitaker 1963). They are excellent burrowers and create their own hibernacula, Meadow jumping mice are generally solitary hibernators, however, there have been occurrences of more than one mouse found in a single hibernaculum. One hibernaculum, located on Rocky Flats Environmental Technology Site, used by Z. �27 h. preblei has been located (Armstrong et al. 1997). This site was 9m above a creek bed 0NaInut Creek); it had a thick cover of chokecherry (Prunus virginianay and snowberry (Symphoricarpos spp.), the mouse was found in a leaf litter nest 30cm beneath the ground in coarse textured soil (Armstrong et al. 1997). Four possible hibernacula were located by tracking radio-telemetered mice at the U. S. Air Force Academy in fall 1997. These sites are located 7, 12,29, and 31m from a creek bed (R Schorr, personal communication). There was no consistency among sites in aspect (N/NW~ S/SE, E, and none [level ground]). Three sites were in vegetation dominated by coyote willow (Salix exigua), one site was in vegetation dominated by snowberry and mullein (Verbascum thapsus). However, all four hibernacula appear to be below coyote willows. These four U. S. Air Force Academy sites have not been disturbed to protect any hibernating mice and therefore are only possible hibernacula because there is no confirmation a mouse is actually hibernating there. Confirmation of a true hibernaculum cannot be made until a chamber, or nest is located. These sites may also possibly be locations of radios discarded by the mice or dead mice. Behavior Jumping mice, including Preble's meadow jumping mouse, are primarily nocturnal or crepuscular but can be observed during the day. Unlike most other nocturnal small mammals Preble's meadow jumping mice do not spend the day underground. The mice can be observed during the day, often under shrubs attempting to remain still (M. Bakeman, T. Ryon, R Schorr personal communication). The shrubs may provide either cover from predators or thermo-regulation, As their common name suggests, the saltatorial ability of jumping mice is well developed. Although longer jumps have been observed the typical behavior is an initial jump of 60 - 90cm followed by rapid jumps of approximately 30cm (Quimby 1951). The more common mode of locomotion, however, is to crawl through or under grass and other vegetation flattening their bodies close to the ground and proceeding quadrupedally (Quimby 1951). Similar jumping and crawling behavior has been observed in Preble's meadow jumping mice as well as running, jumping, and climbing through shrub canopies (M. Bakeman, C. Meaney, T. Ryon, R Schorr personal communication). Given their preponderance for riparian habitats, it is not surprising to find that the mice are also strong swimmers (M. Bakeman, C. Meaney, R Schorr, personal communication). Both swimming and jumping abilities apparently serve more commonly as mechanisms for retreat from either predation or perceived threats rather than as typical mechanisms for locomotion. The mouse is also adapted for digging and creates nests of grass, leaves, and woody material (Harrington et al. 1996). All nests found at the Rocky Flats Technology Site, with the exception of the hibernaculum, were above-ground, generally at the base of willow (Salix spp.) clumps (M. Bakeman, personal communication). There does not appear to be any social structure among individuals within a population of jumping mice. No evidence exists for parental care by the male or any interaction between males and females after breeding occurs. Adults of the species Z. hudsonius are essentially solitary individuals. Except when young, jumping mice are usually silent. On occasion adults have been heard to emit chirps and other sounds (Quimby 1951, Whitaker 1963). No observations have been made of vocalizations by Preble's meadow jumping mice. Drumming noises can be produced by vibrating the tail rapidly against a surface (Svihla and Svihla 1933, Sheldon 1938, Whitaker 1963). Tail rattling, appearing to be a nervous reaction when placed in a capture bag, has been observed in Preble's meadow jumping mouse (M. Bakeman, personal communication). Quimby (1951) and Whitaker (1963) reported that mice in the genus Zapus often washed their feet, faces, and especially their tails. The tail was grasped in the forepaws, and passed completely through the mouth, whereas the hands and feet were washed by means of the forepaws. Bailey (1926) found that jumping mice would reach as high as it could on grass stems, bite them off and pull them to the ground, repeating the procedure until the head was reached and eaten. This process would result in a pile of pieces of grass stem, often with the rachis and glumes on top. Whitaker (1963) found similar behavior in his study of meadow jumping mice in New York. M. �28 Bakeman (personal communication) found piles of pieces of grass stem in areas where Preble's meadow jumping mice were known to occur, however these piles may also have been created by individuals of the species Microtus. Food Preferences Specific food habits are unknown for Preble's meadow jumping mouse. Armstrong et al. (1997) summarized what is currently known about food habits of meadow jumping mice in general as follows: Studies of food habits in central and eastern United States indicate they are governed by . availability more than preference (Whitaker 1963). Grass seeds of several species are probably the most important component of the diet, and mice will shift to those species that have available seed. Invertebrates and fungi are also readily eaten. Mice feed on both adult and larval invertebrates, especially Coleoptera (beetles). Invertebrate feeding is very important in the spring as mice emerge from hibernation, and may consist of half of the diet at that time. Mice also feed on various species of fungi, which are often encountered during burrowing activity. As the growing season progresses, graminoid seeds dominate the diet. There is no reason to believe food habitats of one subspecies should differ greatly from those of other subspecies. This belief is further supported by the indirect evidence available that food habits of Z. h. preblei are similar to that described above (i.e., the observation of the piles of grass stems observed in areas where the subspecies is known to occur [see Behavior section)) .. As true hibernators, meadow jumping mice do not cache food in their hibemacula, Therefore, it is assumed they require high quality food for high fat accumulation prior to immergence. Preble's meadow jumping mice have been observed to increase body weight by 10% in the 2-3 weeks prior to immergence. Laboratory studies show the mouse can gain mass at rates up to 1.0 grams per day (B. Wunder, personal communication). This ability to increase fat reserves at such a rate is used by the mice for preparation to enter hibernation. However, for the mice to successfully gain enough fat prior to hibernation to ensure a high probability of survival throughout hibernation, food of sufficient quality must be available. No specific information is available on what foods Preble's meadow jumping mice eat to meet these ecological requirements. However, from late August through early September, when adults begin to gain weight, there are many species of grarninoids that have available seed. In most years, grasshoppers are also readily available and probably dominate invertebrate biomass in most habitats. Preble's meadow jumping mice are most often found near open water. Dependency on open water has not been established, however work is currently in progress to establish if such a need exists (B. Wunder, personal communication). Based on trapping records at Rocky Flats Environmental Technology Site, trapping success at some sites declines dramatically after creekflow ceases, and it is suspected there may be a shift in site use because of the absence of open water (M. Bakeman personal communication). Habitat Use Landscape scale: The habitat matrix within the range of Z. h. preblei is mixed grasslands adjacent to the Colorado Front Range along the Piedmont and along the base of the Laramie Mountains in Wyoming and extends to the Colorado plains. Within this matrix, Preble's meadow jumping mice occur along stream drainages that contain patches of suitable vegetation. Suitable habitat appears to have at least two major components. The first component is a supply of open water, at least in part of the active season (M. Bakeman, C. Meaney, personal communication). Secondly, areas where Preble's meadow jumping mouse has been found have dense cover (M. Bakeman, C. Meaney, personal communication). �29 If Preble's meadow jumping mouse behaves as a metapopulation in the classical sense of a set of local populations linked by infrequent dispersal then habitat includes not just one area of suitable habitation but also areas suitable for nearby mouse populations. These suitable areas must also be linked by dispersal habitat. If the mice are dependent on dense riparian habitat for dispersal as well as for areas to reproduce, persistence of discrete populations would require a mosaic of suitable discrete riparian patches interconnected with dispersal corridors of similarly dense riparian vegetation. If mouse populations function in a source (populations where growth rate <!: 1) and sink (those populations where growth rate < 1, maintained through immigration) system, it will be critical to identify and protect those populations serving as sources. Thus, for a source-sink population critical habitat will include those areas that support source population, dispersal habitat to sink areas will be less critical. If local mouse populations are functionally discrete, such a mosaic of interconnected areas of suitable habitat would provide a buffer for local, and source, populations against deleterious stochastic events by providing the opportunity for local population failures to be 'rescued' by immigration from other populations. Areas of suitable habitat must also provide requirements to survive throughout the life cycle. These requirements must provide necessities for both the active period and hibernation periods. During the active period suitable habitat must provide requirements for daily survival, reproductive activities (breeding, nesting, and rearing of young to independence), and dispersal. The hibernation period requires sufficient food supplies to assure fat storage prior to hibernation and suitable hibernacula (see Hibernation above). Habitat providing all seasonal and life cycle requirements may or may not occur in a single contiguous area If not in a contiguous area, habitat patches must occur in a mosaic of usable areas where suitable corridors exist for seasonal movement among sites. Site scale: Based on studies of Z. h. preblei and Z. hudsonius elsewhere, Z. h. preblei apparently occurs mostly in undergrowth consisting of grasses, forbs, or both in open wet meadows and riparian corridors, or where tall shrubs and low trees form an overstory and provide adequate cover (Armstrong et al. 1997). Meadow jumping mice are widespread in abandoned grassy fields, but is often more abundant in thick vegetation along ponds, streams, and marshes or in rank herbaceous vegetation of wooded areas (Whitaker 1963). Preble's meadow jumping mice have been trapped in natural riparian areas as well as areas altered by anthropogenic influence including ditches and wetlands adjacent to interstate highways, cement-lined ditches with tall cover, ditches along driveways and moderate road use, and moderate cattle grazing (M. Bakeman, personal communication). The majority of sites where Z. h. preblei have been found consist of multistoried cover but the species composing the cover vary greatly (Armstrong et al. 1997, Meaney et al. 1997). Vegetation composition of the dense cover varies considerably and includes both native and non-native species (Meaney et al. 1996, 1997, Armstrong et al. 1997, M. Bakeman, personal communication). The herbaceous understory can be primarily grasses or forbs or a mixture of the two. Few of the sites, however, are dominated by fewer than two understory species. The tall shrub canopy at most sites is willow (several species), although scrub oak, birch, and alder occur in sites south of the Palmer Divide (Armstrong et al. 1997). Ponderosa pine is the most common tree at higher elevations. The mouse appears to tolerate weedy or exotic species in areas that are structurally diverse and species rich; nearly every successful site contained Canada thistle (Armstrong et al. 1997). Thus, the mouse does not appear to have an affinity toward any single plant species but instead favors sites that are structurally diverse and provide adequate cover and food throughout its life cycle. Preble's meadow jumping mouse mice are typically not found in upland areas away from riparian habitats but are most often captured where either ground water daylights to seep springs or on main water channels (M. Bakeman, T. Ryon, personal communication) suggesting a dependence on open water, at least during their active periods. Movement of Z. h. preblei on Woman Creek at Rocky Flats Technology Site suggests mice move along corridors of shrub cover, generally Salix exigua (PTI 1996a, T. Ryon, unpublished data), suggesting dispersal habitat is similar to habitats used for other activities. �30 SMALL MAMMAL ASSEMBLAGE Preble's meadow jumping mice are only one component of the small mammal community inhabiting areas where they were captured. These mice were more often found at sites with high species richness and abundance of small mammals (Armstrong et al. 1997, Meaney et al. 1997). The variety and relative abundance of other small mammalian species trapped during surveys for Preble's meadow jumping mouse provide some framework for comparison of small mammal assemblages at sites where Z. h. preblei occurred and sites where no Preble's meadow jumping mice were captured (Table 2). All trapped sites were either historical sites of known occurrence or apparently suitable habitat for Z. h. preblei, providing a common basis for comparison. Three small mammal species (Spermophilus variegatus, Peromyscus nasutus, and Rattus norvegicus) were not captured where Z. h. preblei occurred but were captured in unsuccessful sites. However, total capture occurrences of these three species were only one or two sites and the species comprised an extremely small percentage of the species caught in those areas (Table 2) providing too little information for speculating on possible interactions between these species and Z. h. preblei. Table 2. Small mammal community analysis from sites where Preble's meadow jwnping mice (Zapus hudsonius preblei) were captured (n = 28) and not-captured (n = 35). Reported here are nwnber of occurrences a given species was found in areas with and without Preble's meadow jwnping mice captures (no) and mean percentage and standard error (se) of capture that given species contributed to total small mammal captures at a given site. Data are swnmarized from Armstrong et al. (1997) and Meaney et al. (1997). All sites were either areas of historical occurrence of Preble's meadow jwnEing mice ~n = 82 or occurred in areas of aEEarently suitable habitat. Z. h. preblei Captured Z. h. preblei Not Captured Species no % Capture no mean (se) % Capture mean (se) Zapus hudsonlus preblei 28 6.65 (1.38) Spermophilus variegatus 0 0 2.0 Peromyscus nasutus 0 0 3.0 Rattus norvegicus 0 0 2 9.5 (8.2) 1.0(-) 0 0 Mustelasp. 35 Chaeotodipus hispidus 2 1.4 (0.2) 5 3.0 (0.8) Mus musculus 3 5.8 (4.6) 17 18.4 (6.0) Sorex cinereus 4 1.1 (0.7) 5 2.0 (0.5) Microtus longicaudus 5 15.8 (6.3) 3 9.6 (4.3) Neotoma mexicana 5 6.1 (1.7) 8 12.5 (3.6) Reithrodontomys 5 3.6 (1.4) 11 3.7 (0.6) Microtus spp. 8 18.3 (6.4) 4 5.7 (4.3) Microtus ochrogaster 18 16.0 (3.7) 25 13.0 (3.5) Microtus pennsylvanicus 21 26.2 (4.4) 32 25.8 (4.4) Peromyscus maniculatus 27 50.4 (3.4) 32 54.8 (5.2) megalotis �31 The higher occurrence and percent composition of capture of Mus musculus, the house mouse, in areas where Z. h. preblei was not caught might suggest degradation of habitat for Z. h. preblei in those areas or possibly competition between the two species (Ryon 1996). The house mouse, thrives in areas of human habitation and croplands, but also occurs in abandoned fields and ditch banks where they may displace native rodents (Fitzgerald et al. 1994). Ryon (1996) also reported the presence of domestic cats (Felis catus) at sites where Z. h. preblei historically occurred but were not found during his study. Contrarily, C. Miller (personal communication) reported house cats along South Boulder Creek where Preble's meadow jumping mice are known to occur. Caution must be used when percentage of capture is used as an index.to the composition and relative abundance of small mammals within a given area Captures may be influenced by trap placement, bait, season, or trapping protocol. Percentage of capture for a given species may also be biased because of either trap-shy or trap-happy individuals within a species or because some species, in general, tend to be easier or harder to trap. Preble's meadow jumping mice appear to favor 'clean' traps. Once traps have been used and soiled by individuals of other species, the probability of capturing a meadow jumping mouse in that trap appears to decrease (M. Bakeman, A. Deans, F. Harrington, T. Ryon, personal communication). For example, of the 60 captures of Z. h. preblei by M. Bakeman (unpublished data) in 1997 all were captured in traps that had only previously been occupied by Preble's meadow jumping mice. Evidence to the contrary, however, also exists. On at least one occasion, R Schorr (personal communication) caught a Preble's meadow jumping mouse in a trap previously occupied by Sorex sp. If one does assume no trapping biases occurred, where captured, Z. hudsonius was one of the less common species in the areas trapped (Table 2). If such rarity in areas of occurrence is genuine then this combined with both its limited distribution and absence from historically occupied areas supports the need for immediate protection of the subspecies and its habitat. POSSIBLE THREATS TO CONSERVATION .The range of Z. h. preblei corresponds largely to the rapidly developing I\ront Range Urban Corridor that runs from Colorado Springs, Colorado to Laramie, Wyoming. Therefore, possible threats to Z. h. preblei survival may include the widespread destruction and modification of riparian corridors and wet meadow habitats by human land uses (USFWS 1997 a). Agricultural, residential, commercial, industrial, and recreational development are likely the cause of such impacts. Specific activities such as water diversion, stream channelization, and sand, gravel and aggregate mining have impacted habitats required by the mouse Other activities such as overgrazing and development of recreational trails may also impact Preble's meadow jumping mouse habitat. Armstrong et al. (1997) summarized results of habitat studies conducted on Z. h. preblei, including the following summary of disturbance elements. Over the past 100 years the [riparian habitat used by Z. h. preblei] has become increasingly urbanized as influenced by grazing, water diversions, wetland conversion, and real estate development. Compton and Hugie (1993) identified agricultural, residential, and commercial development as habitat impacts and suspected grazing as having a negative influence on meadow jmnping mouse habitat. Ryon (1996) identified alterations to habitat at eight historic meadow jumping mouse sites. These alterations include water diversion, highway construction, gravel mining, grazing, and real estate development. Although suitable habitat exists within this landscape unit, the anthropogenic influence is changing the unit as a whole and fragmenting and degrading riparian areas in general. The effect of these land use practices on mouse distribution is poorly understood. Several investigators have been puzzled at the lack of jmnping mice at sites with seemingly well structured habitat, and the presence of mice at a few sites with less than optimal conditions. Land use history of an area may provide clues as to whether mice are at a site or if they can be �32 restored. The reviewers of the [Report on Habitat Findings of the Preble's Meadow Jumping Mouse, edited by M. Bakeman] list six potential impacts that were thought to playa role in distribution of Z h. preblei including trails, grazing, mining, development, haying, and riparian hydrology. In order to threaten conservation of Z. h. preblei a feature or features of their ecological requirements must be altered in such a way as to make the area unusable to the mouse or limit its usefulness to such an extent as to deplete the areas' potential for supporting a viable population of Z. h. preblei. Therefore, a threat is defined as any activity thathas the potential to negatively alter any ecological requirements of Z. h. preblei. Effects of the potential threats, listed below, may be permanent or temporary. Management strategies may be developed to offset or eliminate effects from various activities. However, such management strategies are only as good as our knowledge of the full complement of ecological requirements necessary to maintain a viable population of Z. h. preblei. The following defines such ecological requirements, provides specific examples of activities that could negatively alter these requirements, and suggests how these negative impacts would affect the mouse. Vegetation composition and structure Vegetation composition and structure affects many of the ecological requirements of meadow jumping mice including food, cover, nest sites, and suitable dispersal corridors. Vegetation composition in areas inhabited by Preble's meadow jumping mouse must include species that would provide not only daily nutritional requirements but also high quality foods necessary for the quick, high fat storage that occurs prior to hibernation. Cover may provide concealment from predators, areas for thermoregulation, and raw material for above-ground nests. Above ground nests appear to be associated with woody (tree and shrub) structures (M. Bakeman, personal communication). Dispersal habitat must provide sufficient cover and food to assure successful dispersal from one area to another. Activities which could threaten vegetation structure include: grazing, development (including residential, commercial, industrial, and recreational), fire, sand, gravel and aggregate mining. Grazing may alter vegetation structure by either decreasing or eliminating sufficient tree, shrub, and/or tall grass cover. Heavy grazing is especially hard on woody plants. Species composition may also be altered by grazing, eliminating critical food resources. Development of an area may impact species composition, abundance, and density. Developments such as cutbanks would eliminate all vegetation. Short-term effects of fire may be deleterious if above-ground nests (especially with young) are destroyed. Longterm effects may be minimal, if the population can withstand minor losses. Mining operations for aggregate material generally eliminates most vegetation, affecting both structure and composition. Possible deleterious effects on populations of Z. h. preblei include decreased survival, reproduction, or ability to disperse. Riparian Hydrology Hydrology includes physical presence of open water, water quality, seasonality of available open water and amount of water flow. Activities which could threaten suitable hydrologic regimes for populations of Z. h. preblei include water diversion projects, water management projects including irrigation regimes and ditch maintenance, and pollution. These activities could result in loss or increase of surface water supplies, alter flooding events which may be necessary to maintain vegetation composition and structure, and produce siltation in catchment areas. Such activities do not actually have to occur on areas of suitable Z. h. preblei habitat but could occur upstream or downstream of the site. Mining operations can indirectly affect habitat by altering downstream hydrological flows of both surface water and groundwater. Urbanization may affect water quality and stream flows through siltation of catchment areas, presence of pesticides in water, increased nutrient load, and vegetation composition. At Rocky Flats Technology Site, Preble's meadow jumping mice are most often captured where either ground �33 water daylights to seep springs or on main channels fed by seep springs (T. Ryon, personal communication) suggesting a dependence on open water, at least during their active periods. Possible deleterious effects on populations of Z. h. preble; if riparian hydrology is altered include decreased survival or reproduction. Habitat structure Habitat structure includes size of suitable area, connectivity of suitable sites to other suitable sites through corridors, and inclusion of all ecological requirements within a given area Thus, altering habitat structure could result in fragmentation of a suitable site, isolation.of a site to other suitable sites, degradation of a suitable site, or total loss of useable habitat Activities which could threaten habitat structure include: aggregate mining, grazing, real estate development, pollution, and fire. Movement of Preble's meadow jumping mice on Woman Creek at Rocky Flats Technology Site suggests mice move along corridors to patches of the most suitable habitat. Most captures occur in areas of shrub cover, generally Salix exigua (PTI 1996a, T. Ryon, personal communication). Possible deleterious effects on populations of Z. h. preble; include decreased abundance due to a decrease in suitable habitat, loss of inter-connectivity of sites that may be necessary to maintain viable populations either in a source-sink or metapopulation situation, and may also result in decreased gene flow through isolation. Distribution Distribution can be defined on several scales: range perimeter, distribution within the range, and distribution within connected suitable habitat. As discussed above (see Distribution section), based on a comparison of currently known sites and historical sites of occurrence of Z. h. preble; it is clear distribution (within the historical range) has already been altered. Activities which could alter distribution of Z. h. preble; include real estate development and sand, gravel and aggregate mining by severely degrading or completely eliminating required habitat. Possible deleterious effects on Z. h. preble; include loss of populations throughout the complete ecological range of suitable habitat and further fragmentation within the range of Z. h. preblei. These effects could (1) eliminate critical populations of Z. h. preble; to maintain possible metapopulations or source populations or (2) restrict gene flow among populations. Geomorphology Geomorphology is defined as the description of features of the earth's surface as explained by the underlying dynamic and structural geology. Important geomorphological features that may help delineate habitat for Z. h. preble; include stream and floodplain geometry. Steepness of stream banks may affect vegetation composition ans structure. Underground structure along stream banks may provide critical hibemacula sites. Activities that would affect geomorphology include development and mining of sand, gravel, and aggregate material. Preble's meadow jumping mice have rarely been captured in steep-sided drainages (M. Bakeman, personal communication). Possible deleterious affects of altering the geomorphology in areas used by Preble's meadow jumping mouse include decreased survival due, possibly, to insufficient availability of suitable hibernacula, Animal Community Composition Animal community composition includes the ocuurance and abundance of all other species in Preble's meadow jumping mouse habitat. The primary activity that may affect populations of Z. h. preble; is development. In particular, the introduction of the house mouse and the house cat may alter the suitability of a site for Preble's meadow jumping mouse; the house cat as a new and effective predator, the house mouse as a possible �34 competitor for resources. Other possible effects of altering the animal community is the introduction of new diseases and parasites to Preble's meadow jumping mouse. The possible deleterious effects of altering the animal community within areas of habitat for Preble's meadow jumping mouse include decreased survival through increased predation, disease, parasites, or competition for resources. CONSERVATION STRATEGY INTRODUCTION This preliminary conservation strategy describes the current status and the goal, objectives, strategies, and research that should be implemented to maintain and restore viable populations of Z. hudsonius throughout its range and to conserve and manage Preble's meadow jumping mouse habitat. This is intended to be a preliminary planning effort based on the best scientific data available to date. This strategy may also serve as a starting point for more comprehensive conservation planning efforts regarding Z. h. preblei. . This strategy is designed to be implemented through the cooperation of state, federal, and municipal government agencies, and involve partnerships with private conservation organizations, individuals, and landowners. The outlined conservation strategy objectives are not necessarily listed in order of priority, but in a logical manner that provides for completion of information needs prior to addressing specific actions. CURRENT STATUS On March 25, 1997 the U. S. Fish and Wildlife Service (USFWS) published a proposed rule in the Federal Register (62 FR 14093) to list Preble's meadow jumping mouse (z. h. preblei) as 'endangered' under the Federal Endangered Species Act (ESA). Final ruling on the proposed listing is scheduled by March 25, 1998. There are three possible outcomes depending on progress of conservation efforts and knowledge gained prior to the decision date: list as endangered, list as threatened, or withdraw the proposal. The primary reason for the proposed listing of Preble's meadow jumping mouse under Section 4 of the ESA is 'present or threatened destruction, or modification of its habitat or range' as well as lack of meaningful protection. The listing decision will be made by the U. S. Fish and Wildlife Service solely on the basis of the best scientific data available. As stated in USWFS 1997b: "All Federal agencies have responsibility under section 7(a)(4) of the ESA to protect proposed endangered and threatened species and habitats on which they depend. For projects where a Federal nexus exists (Federal permit, Federal funding, projects on Federal land) and there is potential affect on Z. h. preblei or its habitat, the Federal action agency should contact the USFWS. In addition, the USFWS encourages all Federal agencies to review their properties and projects and make funds available to conduct Z. h. preblei surveys in all potential habitat. It is also imperative to make project proponents aware of the presence of Z. h. preblei should it be listed prior to the time all actions related to any project are completed. Section 9(a)(l) of the ESA prohibits the take (i.e., harass, harm, pursue, hunt, shoot, kill, wound, trap, capture, or collect, or to attempt to engage in any such conduct) of federally endangered or threatened species, except as provided in sections 6(g)(2) and 10 of the ESA. If Z. h. preblei is listed while a project is impacting it or its habitats, the ESA would enter into effect to protect Z. h. preblei. A project occurring in poor or marginal habitat but having potential to disrupt a travel corridor (such as a road crossing a creek) is also of concern as well as the question of secondary impacts from proposed projects. Projects removed from potential Z. h. preblei habitat that have the potential to adversely impact the habitat may also require a survey. For example, a residential or commercial development upslope from a creek supporting potential Z. h. preblei habitat may significantly increase runoff or otherwise impact the hydrology, and thereby the habitat present on the creek." Therefore, even prior to a decision on the proposed listing, there is a �35 great need to provide information on the ecology and ecological requirements necessary for conservation of Z. h. preblei. The species Z. hudsonius was designated as Colorado nongame wildlife (Colorado Division of Wildlife Regulations, Chapter 10, Article N, #1004 A 6), which provides a legal protection against taking. It is also a Colorado Division of Wildlife Species of Special Concern, which is an administrative classification rather than a legal one. However, there are current efforts within the Colorado Division of Wildlife to list the subspecies as State Endangered (J. Sheppard, personal communication), defined as any species or subspecies of native wildlife whose prospects for survival or recruitment within the state are in immediate jeopardy. The Colorado Natural Heritage Program (1997) lists the mouse as an S2 species: imperiled in the state because of rarity (6 to 20 occurrences) or because of other factors demonstrably making it very vulnerable to extirpation from the state. Conservation of Preble's meadow jumping mouse was also listed as a specific task to be addressed by both the State of Colorado and the Department of the Interior in the Memorandum of Agreement (94SMU-058), signed by Governor Roy Romer and Interior Secretary Bruce Babbitt on November 29, 1995, between the State of Colorado and the Department of the Interior concerning 'Programs to Manage Colorado's Declining Native Species.' The Wyoming Natural Diversity Database (Fertig 1997) ranks Z. h. preblei as S1: critically imperiled in the state because of extreme rarity (five or fewer extant occurrences or very few remaining individuals) or because of some factor of the subspecies life history that makes it vulnerable to extinction. The Wyoming Department of Game and Fish (1998) ranks the subspecies as nongame through their Nongame and Wildlife Habitat Protection Programs. GOAL The goal of this conservation strategy for Preble's meadow jumping mouse should work towards the sustainability, protection, and restoration of Z. h. preblei populations and habitats on both private and public lands to provide the spatial, genetic, and demographic structure needed to promote long-term species viability and provide species management flexibility. Specific objectives include the following. OBJECTIVES 1. 2. Document the present distribution of Z. h. preblei. This will require the following: • Conduct trapping surveys to better define the boundaries of the range of Z. h. preblei.; conduct trapping surveys in areas of potential sympatry of Z. h. . preblei with Z. princeps; evaluate effect of small mammal species richness and composition on the distribution and/or abundance of Z. h. preblei. • Further explore genetic relationships among different Z. h. preblei populations, among different subspecies of Z. hudsonius, and among different species of Zapus. Identify populations of Z. h. preblei where the rate of population growth ~ 1. This will require population studies to estimate the following parameters: • Survival (annual, over-summer, over-hibernation), and identity offactors affecting survival [e.g., weight, sex, age, abundance (i.e., density dependent Z. h. preblei response), habitat features (e.g., stream reach, vegetation composition), weather, predation, disease]. • Reproductive and developmental parameters (i.e., number oflitters per year, number of young per litter, age at first reproduction, juvenile survival), and identity of factors affecting reproduction [e.g., habitat features (i.e., food quality, nest site availability, cover availability), abundance (i.e., density dependent response), weather]. • Recruitment, and identity of factors affecting recruitment (e.g., weather, habitat features). • Immigration and emigration rates, and identity of factors affecting immigration, emigration (e.g., age, habitat features, abundance). �36 3. 4. 5. • Population structure (sex and age ratios) • Dispersal parameters (rate, who disperses, time of dispersal) • Abundance, and identity of factors affecting abundance (e.g., habitat features). • Rate of population change. Protect populations of Z. h. preblei where the rate of population growthis ;:::1. Maintain current range of natural variability of Z. h. preblei. Identify ecological requirements for sustaining viable populations of Z. h. preblei throughout its range of natural variability. This will require research to address the following: • Estimate habitat use [distances traveled (daily, seasonally), landscape features (connectivity with other potential sites, geology), seasonal use, hibemacula, nest sites, distance to nearest open water, hydrology (water quality, flow), abundance (i.e., density dependent responses), and cover]. • Dispersal habitat [end point descriptions (disperses from what to what), landscape features (connectivity with other riparian strips, corridor use, overland use]. • Physiology (e.g., estimate dependency of Z. h. preblei on open water, food habits, effects of varying water quality on Z. h. preblei physiology). • 6. 7. 8. 9. 10. 11. 12. 13. Protect habitats to sustain or restore populations of Z. h. preblei with high where the rate of population growth ;:::1. Promote protection, management, and possible restoration of habitat for conservation of Z. h. preblei in all currently or recently occupied and suitable habitat. Monitor the status of Z. h. preblei populations throughout its range to detect changes in local distribution. Identify threats to the conservation of Z. h. preblei. Eliminate or minimize threats to Z. h. preblei conservation. Integrate Preble's meadow jumping mouse conservation strategy objectives with management and habitat objectives of other Front Range riparian species. Promote scientific management of Preble's meadow jumping mouse. Promote public support for Z. h. preblei conservation efforts and scientific management of Preble's meadow jumping mouse through public education. RESEARCH NEEDS There are currently four research projects being conducted on Z. h. preblei, with a fifth planned to begin spring of 1998. Each of these projects is designed to provide further information on the ecology or demography of Preble's meadow jumping mouse. However, if research methodologies were standardized and coordinated across all the projects, comparability and quality of the data would be greatly enhanced. Such a coordinated effort would maximize data quantity, quality, and comparability over varying conditions throughout the range of Z. h. preblei in the most effective and efficient way possible. Comparable information gained across the ecological range of Z. h. preblei would then provide more useful information for use in developing sound management strategies for conservation of the mouse. To achieve such a coordinated research effort all project leaders must agree to follow data collection protocols, establish 'ownership' of data and future publications, and work cooperatively on the possible sharing of equipment and technical assistance during peak data collection periods. To facilitate such a mutual effort will require the participation of all project leaders conducting field studies of Z. h. preblei as well as cooperative investigators, and specialized data analysts with expertise in the study design, analysis, and interpretation of mark-recapture and telemetry data A proposed agenda including tasks, task leaders, and a timetable to initiate a cooperative research effort include: �37 Task Task Leader, Affiliation Date Identify all potential participants: project leaders, cooperating investigators, data analysts. T. Shenk, Colorado Division of Wildlife January 1998 Prioritize research questions. All project leaders* January 1998 Identify sites to best address prioritized research needs. All project leaders January 1998 Design studies specific to research question to be addressed at each site. All project leaders, cooperating investigators, data analysts January-February 1998 Evaluate needs at each site to conduct specified research. All project leaders March 1998 Identify discrepancies between needs and available funds, equipment, and personnel currently allocated for each site. All project leaders March 1998 Attempt to balance discrepancies. All project leaders, cooperating investigators March 1998 * To date: M. Bakeman, C. Meaney, T. Ryon, R. Schorr, T. Shenk The following outline lists research needs to provide information to develop sound management strategies for conservation of Z. h. preblei. Research needs are not listed in order of priority, but listed in a logical manner to provide completion of needs. There are three components of Z. h. preblei ecology that are currently unknown and yet key to any sound conservation strategy for the subspecies. These are (1) detailed demographic studies estimating survival and reproduction and determining the factors influencing each of the parameters, (2) detailed studies evaluating movements and dispersal habitat of individuals within and among populations, and (3) detailed studies to define hibernation needs, primarily descriptions of suitable hibernacula criteria and food requirements for sufficient fat storage prior to immergence. Demographic studies: Information on the population dynamics of Preble's meadow jumping mouse is necessary to determine which areas support populations where the rate of population growth is ~ 1. Key parameters to estimate include: Survival: • estimates of survival, including •.. annual •.. over-summer •.. over-hibernation • investigate possible factors affecting survival, including •.. weight •.. sex •.. age •.. abundance (i.e., density dependent response) •.. habitat features: stream reach, vegetation composition •.. weather •.. predation •.. disease Approach: mark-recapture techniques �38 Recruitment: • • estimate recruitment (as an alternative to reproductive parameters listed below) investigate possible factors affecting recruitment, including •.. weather •.. habitat features Approach: mark-recapture techniques Population structure: • estimate sex ratios • estimate age ratios Approach: mark-recapture techniques Abundance: • • estimate abundance investigate factors affecting abundance, including •.. habitat features (see under habitat use) Approach: mark-recapture techniques for closed populations Immigration, emigration: • • estimate rates of immigration and emigration investigate possible factors affecting immigration, emigration •.. habitat features (see under habitat use) •.. abundance (i.e., density-dependent response) Approach: estimate rates from mark-recapture data, identify existence with radio telemetry Reproduction: • • • • • estimate number of litters per year estimate number of young per litter estimate age at first reproduction estimate juvenile survival investigate possible factors affecting reproduction, including •.. habitat features: food availability, nest site availability, cover availability •.. abundance (i.e., density-dependent response) •.. weather Approach: from telemetIy work Dispersal Studies: Dispersal is a key process in metapopulation theory and to maintain genetic diversity between isolated populations. Key parameters to evaluate include: Population parameters • who disperses • time of dispersal • estimate rate Approach: estimate rate with mark-recapture, and mark-recapture data document who and when from both telemetry Habitat parameters • through what habitat • end point descriptions (disperses from what to what) • landscape features (connectivity with other riparian strips, corridor use, overland use) Approach: document where from both telemetry and mark-recapture data �39 Distribution Studies: Conservation of Z. h. preblei should maintain populations throughout the range of its natural variation and to try to identify ecological limits for the subspecies. Key concerns include: Range-wide distribution: • conduct trapping surveys to better define the eastern boundary of the range of Z. h. preblei • conduct trapping surveys in areas of potential sympatry of Z. h. preblei with Z. princeps Approach: determine from trapping surveys Habitat Studies: To identify and define habitat requirements of Z. h. preblei studies should be conducted to address the following: Habitat use: • estimate distances traveled: daily, seasonally • describe landscape features (connectivity with other potential sites, geology) • determine seasonal use • describe hibernacula • describe nest sites • estimate distance to nearest open water from other habitats used • describe hydrology (water quality, flow) in areas of use • evaluate effects of abundance on habitat used (i.e., density dependent responses) Approach: estimate from telemetry work Physiological Studies: Physiological studies will provide information on the mechanisms driving habitat selection. Physiological requirements: • estimate dependency of Z. h. preblei on open water • determine energetic requirements to survive hibernation Approach: estimate from laboratory studies Systematic Studies: To better define the relationship of Z. h. preblei to other subspecies of Z. hudsonius and other species of Zapus the following studies should continue. Molecular systematic relationships: • further explore genetic relationships among different Z. h. preblei populations • further explore genetic relationships among different subspecies of Z. hudsonius • further explore genetic relationships among different species of Zapus. Approach: explore through laboratory studies Systematic relationships: • link genetic relationships to systematic studies of Z. hudsonius. Approach: explore through museum studies Community Studies: Composition of the community where Z. h. preblei occur could help explain ecological tolerances of the subspecies, providing insight to the mechanisms determining its distribution. Small mammal assemblages: • comparison of small mammal assemblages in areas where populations of Z. h. preblei occur and areas where they do not II> species composition II> relative abundance Approach: estimate from trapping surveys �40 LITERATURE CITED Armstrong, D. M. 1972. Distribution of mammals in Colorado. University of Kansas, Museum of Natural History Monograph 3:1-415. Armstrong, D. M., M. E. Bakeman, A Deans, C. A Meaney, and T. R Ryon. 1997. Conclusions and recommendations in: Report on habitat findings on the Preble's meadow jumping mouse. Edited by M. E. Bakeman. Report to USFWS and Colorado Division of Wildlife. Bailey, B. 1929. Mammals of Sherburne County, Minnesota. Journal ofMammalogy 10:153-164. Bailey, V. 1923. Mammals of the District of Columbia Proceedings of the Biological Society. Washington 36:103-138. Bailey, V. 1926. A biological survey of North Dakota. U. S. Department of Agriculture North American Fauna No. 49:117-119. Colorado Division of Wildlife. 1997. Colorado's endangered, threatened, special concern, undetermined status and candidate species - Terrestrial species. Draft Report for the Colorado Division of Wildlife. Colorado Natural Heritage Program. 1997. Colorado's Natural Heritage: Rare and imperiled animals, plants, and plant communities. Volume III, No.1. Unpublished Report for the Colorado Natural Heritage Program. Compton, S. A, and RD. Hugie. 1993. Status report on zapus hudsonius preblei, a candidate endangered species. Pioneer Environmental Services, Inc. Report for the USFWS. Logan, Utah Logan, Utah. Cranford, J. A 1983. Ecological strategies of a small hibernator, the western jumping mouse Zapus princeps. Canadian Journal of Zoology 61:232-240. EG&G. 1993. Report of Findings: 2nd Year Survey for the Preble's meadow jumping mouse. Prepared by Stoecker Environmental Consultants for ESCO Associates, Inc., Rocky Flats Environmental Technology Site, Jefferson County, Colorado ERO Resources. 1995. Environmental review of South Boulder Creek Management Area. Prepared for City of Boulder Real Estate/Open Space Department. Prepared by ERO Resources Crp., Denver, Colorado in association with Stoecker Ecological Consultants, Boulder, Colorado. Fertig, W. 1997. Wyoming plant and animal species of special concern. Wyoming Natural Diversity Database. Laramie, Wyoming. Fitzgerald, I P., C. A Meaney, and D. M. Armstrong. 1994. Mammals of Colorado. Denver Museum of Natural History, University Press of Colorado. Niwot, Colorado. Hafuer, D. I, K E. Petersen, and T. L. Yates. 1981. Evolutionary relationships of jumping mice (genus Zapus) of the southwestern United States. Journal of Mammalogy 62:501-512. Hall, E. R 1981. The mammals of North America John Wiley and Sons, Inc., New York, New York, 2 volumes. Hamilton, W. J., Jr. 1935. Habits of jumping mice. American Midland Naturalist 16:187-200. Harrington, F. A, A Deans, M. E. Bakeman, and B. J. Bevirt. 1996. Recent studies of Preble's meadow jumping mouse at the Rocky Flats Site, Colorado. Journal of the Colorado-Wyoming Academy of Science 28(1): Abstract 18. Krutzsch, P. H. 1954. North American jumping mice (genus Zapus). University of Kansas Publications, Museum of Natural History 7:349-472. Levins, R 1970. Extinction. Lectures in Mathematical Life Sciences 2:75-107. Long, C. A 1965. The mammals of Wyoming. University of Kansas Publications, Museum of Natural History, 14:493-758. Meaney, C. A and N. W. Clippinger. 1995. A survey of preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. Report prepared for the Colorado division of Wildlife. Meaney, C. A, N. W. Clippinger, A Deans, and M. OShea-Stone. 1996. Second year survey for Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. Report prepared for the Colorado Division of Wildlife. �41 Meaney, C. A, A Deans, N. W. Clippinger, M. Rider, N. Daly, and M. O'Shea-Stone. 1997. Third year survey for Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. Report prepared for the Colorado Division of Wildlife. Olson, T. E., and F. L. Knopf 1988. Patterns of relative diversity within riparian small mammal communities, Platte River watershed, Colorado. Pg. 379-386 in: Proceedings of the symposium: Management of amphibians, reptiles and small mammals in North America Flagstaff, Arizona U. S. Forest Service, General Technical Report RM-166. Poly, W. 1., and C. E. Boucher. 1997. Record ofa creek chub preying on ajumping mouse in Bruffey Creek, West Virginia Brimleyana 24: 29-32. PTI Environmental Services. 1996a Preble's Meadow Jumping Mouse Study at Rocky Flats Environmental Technology Site, Annual Report 1996. Final. Rocky Flats Environmental Technology Site, Golden, Colorado. PTI Environmental Services. 1996b. Preble's Meadow Jumping Mouse Study at Rocky Flats Environmental Technology Site, Spring 1996. Final. Rocky Flats Environmental Technology Site, Golden, Colorado. Quimby, D. C. 1951. The life history and ecology of the jumping mouse, Zapus hudsonius. Ecological Monographs 21:61-95. Riggs, L. A, 1. M. Dempey, and C. Orrego. 1997. Evaluating distinctness and evolutionary significance of Preble's meadow jumping mouse: Phylogeography of mitochondrial DNA noncoding region variation. Final Report for the Colorado Division of Wildlife. Denver, Colorado. Ryon, T. R 1996. Evaluation of historical capture sites of the Preble's meadow jumping mouse in Colorado, final report. MS Thesis. University of Denver, Denver, Colorado. Ryon, T. R 1997. The evaluation of historic capture sites of the Preble's meadow jumping mouse in Colorado. in Report on habitat findings of the Preble's meadow jumping mouse. Edited by M. E. Bakeman. Report to USFWS and Colorado Division of Wildlife. Sheldon, C. 1934. Studies on the life histories of Zapus and Napaeozapus in Nova Scotia Journal of Mammalogy 15:290-300. Sheldon, C. 1938. Vermont jumping mice of the genus Zapus. Journal of Mammalogy 19:324-332. Svihla, A and R D. Svihla 1933. Notes on the jumping mouse, Zapus trinotatus trinotatus Rhoads. Journal ofMammalogy 14:131-134. USFWS. 1997a Proposal to list the Preble's meadow jumping mouse as an endangered species. USFWS 50 CFR part 17. USFWS. 1997b. Interim survey guidelines for Preble's meadow jumping mouse. USFWS. Denver, Colorado. Whitaker,1. 0., Jr. 1963. A study of the meadow jumping mouse, Zapus hudsonius (Zimmerman), in cental New York. Ecological Monographs 33:3. Whitaker, J. 0., Jr. 1972. Zapus hudsonius. Mammalian Species 11:1-7. Wyoming Game and Fish Department. 1998. Native species status classification system. Cheyenne, Wyoming. ��43 APPENDIX A Evaluating distinctness and evolutionary significance of Preble's meadow jumping mouse: Phylogeography of mitochondrial DNA non-coding region variation. Final Report to the Colorado Division of Wildlife Reply to: Box 9528, Berkeley, CA 94709 Tel: 5101531-4848 Fax: 5101530-0580 Email: b&i3igc.org EVALUATING DISTINCTNESS & EVOLUTIONARY SIGNIFICANCE OF PREBLE'S MEADow JUMPING MOUSE: PHYLOGEOGRAPHY OF MITOCHONDRIAL DNA NON-CODING FINAL REPORT DECEMBER 8, 1997 Submitted by Lawrence A. Riggs, Ph.D. John M. Dempc:y, M.A. Cristian Orrego, Ph.D. Biosphere Genetics, Ine, Berkeley, CA Submitted to Judy Sheppard ·Terrestrial Section Colorado Division ofWddIife Denver,CO REGION VARIATION �I· 44 Evaluating Distinctness & Evolutionary Significance of Preble's Meadow Jumping Mouse: Phylogeography of Mitochondrial DNA Non-Coding Region Variation Lawrence A Riggs, John M. Dempcy, and Cristian Orrego SUMMARY We have generated molecular genetic data fO[Jl:!otal.Qf92.jwnping mice including 71 sampled from 20 populations in Colorado and four populations in Wyoming, and 21 specimens representing three reference groups of closely related taxa, DNA sequences obtained for a,433 basepair (bp)-long portion of the mitochondrial DNA non-coding region (sometimes referred to as the D-Ioop) in all samples indicate that a group of populations ranging from southeastern Albany County, Wyoming, south along the Front Range of the Rocky Mountains to western Las Animas County, Colorado, fonn a coherent group, both genetically and geographically. This group, which we refer to as the "Preble's group" is distinct from four oftive other populations sampled by live trapping or from museum specimens and may be somewhat differentiated from the fifth of these. The Preble's group is also clearly distinct from two of the three reference groups and less markedly differentiated from the third. Phylogenetic analyses based on these data are still at a preliminary stage and indicate that the Preble's group of populations is most closely allied with, and not strongly differentiated from, samples of Z h. intermedius from Minnesota. Of the four populations that appear to be distinct from the Preble's group, two sampled near the Colorado-New Mexico border are closely allied with museum specimens identified as Z. h. luteus from New Mexico. Two others from the vicinity of Cheyenne, Wyoming, are most similar to representative samples of Z princeps from western Larimer Co., Colorado and are next most closely allied with a Z. princeps specimen from northern New Mexico. A suggestion from our analysis is that Z. hudsonius from Indiana (possibly either Z. h. americanus or Z. h. intermedius based on distribution) may be the most ancestral of the populations and taxa sampled and may have shared a common ancestor with progenitors of forms presently known as Z. princeps princeps and Z. h. luteus. BACKGROUND ' , Recent applications of molecular genetic methods in systematics, population genetics, and conservation biology have demonstrated that variation in DNA markers can , be highly infonnative in the evaluation of genetic distinctness and evolutionary significance, two important detenninants in decisions regarding application of the Federal Endangered Species Act of 1973 as amended and interpreted by the U. S. Congress. The study described here sought to discover whether and how molecular data might support and help objectify the view, based on other more traditional criteria, of the Preble's mouse as an evolutionary unit distinct from other species and subspecies in the genus Zapus. \ t �45 Final Reportl12-8-97 Preble's DNA study Work funded by the Colorado Division of Wildlife (hereafter CDW) and performed by Biosphere Genetics, Inc. has evaluated the ability of three different approaches to quantifying variation in the DNA molecule to provide reliable and informative data addressing issues of key importance in the decision of whether or not to list the Preble's mouse. Previous reports to CDW have summarized our finding that RAPD (randomly amplified polymorphic DNA) markers are not sufficiently repeatable for use in the context of this study. Methods to assay variation in another class of genetic markers found in nuclear DNA called microsatellites or simple sequence repeats, although -potentially promising, are not yet sufficiently well developed and validated for zapodids to apply at this time. A third approach, DNA sequencing of the mitochondrial DNA noncoding region which includes the D-loop, emerged from preliminary work as the method. most appropriate and informative, in combination with other more traditional criteria, for the determination of whether populations of the Preble's meadow jumping mouse (Zapus hudsonius preblel) constitute one or more distinct evolutionary units with significance warranting protection under the Federal Endangered Species Act. Results based on Sequence variation in the mitochondrial DNA non-coding region assayed in a subsample of the 54 mice presumed to be Preble's obtained during the summer of 1996 indicated: (1) that 7 populations sampled at sites ranging from Longmont to Colorado Springs in Colorado varied relatively little from one another, (2) that two other populations sampled in Las Animas County near Lake Dorothey were similar to one another but differed quite markedly from the first group, and (3) that none of the individuals sampled in Laramie '{ ~ County in Wyoming could be assayed using the techniques successfully applied to the other samples, suggesting that this population may also differ to some degree from those in Colorado. Field surveys of the occurrence of the Preble's meadow jumping mouse conducted by several individuals and organizations during the summer ofl996 were extended to additional sites in the summer of 1997. CDW sponsored some of these efforts and coordinated the collection, recording and delivery of tissue samples for DNA analyses performed by Biosphere Genetics, Inc., also under contract With the Division. In addition, the U. S. Air Force supported DNA analysis of samples. obtained from trapping effOrts during the' summer of 1996 on the Warren Air Force Base and the inclusion of reference material representing two additional subspecies of Zapus hudsonius, Z h. pa/ustris, and Z h.~pesUi~ . METHODS Tissue Sampling and Storage A sampling protocol and data forms for using in obtaining samples for.DNA analysis were developed for CDW and distributed to all parties participating in the coordinated trapping effort. The disseminated protocol, adapted and updated from the .U.S. FIShand Wddlife Service standardized protocol, described handling and instrument cleansing procedures to minimize contamination of the sample with human or other mammalian'DNA Very small samples of tissue were obtained from live-trapped �I . 46 Preble's DNA study Final Reportl12-8-97 specimens of Zapus by using a tagging punch (National Band and Tag Co., Newport, KY) to remove one or more plugs of tissue from the outer portion of each mouse's ear. In some cases, ear tissue was obtained by clipping or notching the ear with scissors. Tissue plugs or pieces from a given animal were placed in a cryogenic screw-cap vial filled 113112full with 95% ethanol and labeled with an alpha-numeric code intended to uniquely specifYtrapping location and the individual animal caught. Hair samples were collected by some participants and placed either in a separate labeled vial or in the same vial containing the ear tissue. Samples were conveyed to CDWand accumulated into lots for delivery to BGl When'samples were transported from the field or shipped by common carrier, they were at ambient temperature. Upon receipt at BGY,samples were kept refiigerated at 4°C until the time of DNA extraction. Specimens obtained from field swveys and miscellaneous museum specimens used to expand coverage of the potential range of the taxa examined in this study are listed in Table 1 of the Appendix to this report. Reference specimen tissues used for comparison with field-sampledjumping mice were provided from four separate museum collections (see Appendix Y,Table 2) as toes clipped from museum specimens or as small pieces of internal organs maintained in the museum's frozen collection, usually at -70°C. Toe clips were sbipped at ambient temperature and refiigerated at 4°C upon receipt. Frozen specimens were sbipped on dry ice and extracted upon receipt with any remaining tissue being transferred to ethanol for further storage. Reference specimens are listed in Table 2 of the Appendix to this report. DNA Extraction & Purification We extracted DNA from plugs or pieces of ear tissue using the Hair Lysis Buffer (HLB) method similar to that of Greer et ale(1995). The tissue was rinsed, either with a jet of triple-distilled water or by agitation in a drop of distilled water on Parafilm®, then. placed in a volume ofHLB and incubated at 56° C with periodic agitation for 10-12 hours. In most cases, this process completely digested the sample leaving only a trace of particulate matter suspended or settled in the bottom of the tube. With the particulates centrifuged to the bottom, aliquots of the extract were withdrawn for fractionation and for purification prior to use in DNA amplifications. Raw DNA extracts were run out on 1.5% agarose gels and stained with ethidium bromide to quantify the DNA by comparison with staining intensity of Lambda Hind m fragments run on the same gel. We used Prep-AGenenl (Bio-Rad, Hercules, CA) to purifYthe DNA preparations before sequencing. Amplification of Mitochondrial DNA Non-coding Region Sequences We began our examination of the mitochondrial non-coding region using the primers designated as ECO-THR. (L15996) and BAM-TDKD (H16401)to amplify an approximately 550 bp-long segment between the threonine tRNA gene near the . cytochrome b end of the non-coding region and the TOKD site approximately halfway through the non-coding region. 1 This primer combination, first used to study human Inc BOO- and BAM- portions ofthesc primer designations refer to the addition of restriction cuzyme scqucnccs at the S' cud of the primer sc:qucncc. These "'tails" are not essential for the present application and we will usc primer designations with and without this notation interchangeably in this report. Biosphere Genetics, Inc. "page 3 . �47 Preble's DNA study Final ReportlI2-8-97 populations (Vigilant et aI. 1989) had worked well in studies of other mammals and typically revealed amounts of sequence variation appropriate to differentiating subspecies and geographically isolated populations. Because the non-coding region includes a structure described as a displacement or ~1)" loop known from work on human and cattle DNA, we will occasionally refer to the region of DNA studied here as the D-Ioop. Amplifications with the above primers were performed in 12.5 J.dreaction volumes using 1 J.dof template DNA, 1 unit of the DNA polymerase, Klentaq-I (Ab Peptides, St. Louis, MO), and other reaction components in standard proportions. A T~~e PHC-2 .. thetmocylcer was used to implement a program using an initial denaturation at 94°C for 3 minutes followed by 42 cycles with denaturing at 94° for 4S seconds., annealing at 55° for 1 minutes, and extension at 72° for 1 minutes fonowed by a final extension at 72° for 5 minutes. Products were fractionated on 1.5% agarose gels. ~plification of the 5S0 bp D-loop fragment was successful, but primer dimer and secondary products also appeared, sometimes in considerable amounts. We experimented with a variety of modifications to obtain cleaner product (primer reduction, additions ofMgCh at various levels, alternative forms of DNA polymerase, etc) and were ultimately able to obtain product with a . minimum'of these additional elements in many, but not all, samples of mice presumed to be Z h. preblei. Following the optimization effort, revised reaction conditions were used to assess D-Ioop fragment amplification success with all new DNA extracts in 12.5 J.dreactions, then applied with slight modifications to the thermocyler program to amplify targeted products in 50 J.dreaction volumes to obtain sufficient quantities for automated sequencing. Two or three 50 J.dreactions were run for each sample, depending on prior estimates of the amount of template available in individual DNA extractions. The quantity and quality of product generated by each reaction was assessed by agarose gel fractionation. Reaction volumes were combined when necessary, then cleaned with the solid-phase reversible immobilization method (DeAngelis et al. 1995) using carboxyl coated magnetic particles (perSeptive BioSystems Inc, Cambridge, MA). Cleaned, resuspended DNA products were then dried down to a pellet in the bottom of a 1.5 ml eppie tube for shipping by overnight service at ambient temperature to an automated sequencing service, Resolution of Amplification Problems Some DNA extracts, most notably those of Z princeps and of putative Z h: . preblei pOpulations sampled at the Warren Air Force Base in Wyoming, did not amplify well or at all with the ECO- TIIRlBAM- TDKD primer combination. We first suspected that this OCCUlTed because the TDKD primer designed from sequence information on the human D-loop was not finding sufficient complementarity on the heterologous primer site in at least some Zapus populations or taxa. Conventional wisdom suggested that the ECO-1HR primer site should be·highly conserved in vertebrates. Based on this reasoning, we designed two new primers for the middle of the D-loop. We examined D-Ioop sequences available from Genbank for rat (Rattus norwegicus), mouse (Mus musculus), vole (Clethrionomysg/areolus), gopher (Thomomys bottae), and kangaroo rat (Dipodomys ordii) as well as our own sequence data for Zapus. Sequences for the Pa~e4 �t . 48 Final Reportl12-8-97 . relevant portion of the D-Ioop for Mus, Clethrionomys and Zapus were used to identify a primer site most likely to be reasonably conserved in muroid and dipodoid rodents. The primers designed for this new site are designated PRDL L-15738 (forward primer) and PRDL H-15720 (reverse primer). We had these primers synthesized (Operon Technologies, Alameda, CA) for testing and possible use in resolving the difficulties mentioned above. The relative positions and priming directions of all primers are shown in Figure 1. . We used PRD~!!-15~~0 iq combination with ECO-THR. in ail attempt to amplify those samples that had notamplltied with the ECO- THRIBAM- TOKD primer combination. We also attempted to .amplifythe entire D-Ioop by using ECO-THR.in combination with PIlE-rev H29~ a primer previously designed for general use in amplifYingportions of the D-Ioop in mammals (A Gavazzi, unpublished). Fmally we ..tested PRDL L-15738 in combination with PHE-rev H29 for the purpose of generatingD. loop fragments spanning the TOKD site for automated sequencing of this site in Zapus. Testing of these primers on Zapus DNA extracts revealed that: I) the ECO-THRIPRDL H-lS720 primer combination amplified in only two of20 samples and did nothing to solve the previous difficulties encountered with the ECO- THRIBAM- TOKD combination; 2) the ECQ-THRIPHE-rev H29 combination produced a fragment ca. 1500 bp long, apparently spanning the D-Ioop successfully in some cases, but not with the DNA extracts that had been previously problematic, and 3) the PRDL L-15738IPHE-rev H29 combination produced multiple bands in some cases and no amplification in others. We inferred from these results that our original problem in amplifying some Zapus samples, most notably those assigned to.Z. princeps and the samples from the Warren Air Force Base, was not with the TDKD site, but rather with annealing of the ECO-THR primer designed for a conserved section of the human threonine tRNA gene to the heterologous sequence in Zapus. Reexamination of the sequences successfully obtained in preliminarywork on Zapus suggested that a primer site that would be more highly.conserved between Zapus and other mammals might be present internal to the one primed by ECO..THR. Further testing showed that an available primer designated PROC, for priming of a sequence on the proline tRNA gene, when used in combination with the BAM-TOKD primer, was uniformly very successful in amplifying a ca. 433 bp fragment (excluding primer sequences) of the D-loop. While we have been reluctant to give up on efforts to access .information about variation existing in and around the THR site, the PROClBAM-TOKD primer combination was clearly the tool of choice for compiling the dataset required for this study within the time frame required. The results presented in this report are for the 433 bp segment of the D-loop falling between the proximal ends of these primer sites, Automated Sequencing DNA sequencing of both the 550 bp and 433 bp fragments of the D-Ioop generated as described above was performed by The University of Maine DNA Sequencing Core Facility using an ABI model 373A Stretch DNA sequencer. Results of . one sequencing pass in each direction were received bye-mail as attached files ABI . format. Pherograms in these files were viewed with the program Chromas (v. lA, Conor m Biosphere Genetics. Inc. PageS �Final Reportl12-8-97 Preble's DNA study 1HR PROC ---+ ---+ IdUlAlDlltRNAnDl PRDLLlS738 N_-_COdiac_·_1qICIIl_. 1-1 ~ PRO H-lS720 ____.lllRNAnEl ~ rosn PHEA . Fig. I. Rclativc positions of the primers used in this study for amplification of the mitochondrial DNA non-coding region, which includes the D-loop. Primers TIm. and roKD were from Vigilant et a1. (1995), PHE-rev H29 was from Anita Gavazti and primers prdl L-15738 and prdl H-IS720 arc two new primers designed for this study to work with Zapus and other rodents. Primers are used in pair-wise combinations to amplify segments of DNA lying between their positions on this simplified map. The relative positions of coding and non-coding regions are indicated by labeled areas. McCarthey, Griffith University, Brisbane, Queensland, Australia) and machine-assigned sequences were exported as text files for use in subsequent analysis. Data Analysis Text files containing the two sequencing reads generated from opposite ends of the amplified fragment were entered into the program MacDNASIS (v. 1, Hitachi Software Engineering Co., Ltd., San Bruno, CA) and combined to obtain a consensus sequence for the amplified region of each individual mouse sampled. Sequences obtained from both THR-TOKD and PROC- TDKD amplifications were used as available. However, since the more inclusive THR-TDKD sequences were not available for individuals assigned to·z. princeps and samples ultimately found to be more closely related to Z. princeps than to the Preble's samples, and since the longer sequences were of variable quality at the beginning of the S' to 3' read in a number of the Preble's samples, we 1:15edonly the 433 bp sequence available in all samples in all subsequent data analysis. Consensus sequences were aligned manually using the multiple sequence option of MacDNASIS. The program MacClade (v. 3, Maddison, 1992) was used to create' a data matrix of character states for S8 variable sites in the aligned sequences for phylogenetic analyses. Phylogenetic trees were generated using the parsimony algorithm in PAUP (v. 3.1.1, Swofford, 1993). To expedite review ofVarlous options for the rooting of trees; one individual was selected as most representative of each population (fewest changes from the most commonly shared state at the 58 variable sites treated as c~cters in the DNA sequence) and used in the depiction of .most-parsimonious trees and in bootstrap analyses. Bootstrap analyses used the 50% majority rule with 100 repHcations. Biosphere Genetics. Inc. Page 6 49 �5~rcble's DNA study Final ReportlI2-8-97 Sirici .- .. ....:.. I I , I Figure '2. Strict consensus tree, identical to the single most parsimonious tree generated using the "branch and bound" option ofPAUP v. 3.1.1 (Swofford 1993) onsequences for all samples analyzed. Length of this shortest tree is 61 evolutioruuy steps; consistency index (Cl) = 0.869. Although not supported by bootstrap analysis in the context of phylogenetic analysis, SUb-groupings of'populations considered here to be part of the Preble's group suggest that population subdivision may be discemable by more appropriate population-analytie methods. Biosphere Genetics. Inc. 1. �51 Final Reportl12.S-97 Preble's DNA study RESULTS Figure 2 illustrates phylogenetic relationships among all samples included in the study to date, inferred from maximum parsimony analysis (length = 61 steps, consistency index [CI] =.0.869). Bootstrap analysis for the entire dataset was not practical for this report. However, bootstrap analyses were performed for various outgroup scenarios using one individual from each population as described above. No outgroup was designated for the analysis shown in Figure 2. _ -- .. The important relationships in Figure 2 are perhaps more easily seen in the unrooted, circular format tree generated using one representative individual from each population (Figure 3).. No meaning is attached to the length of branches hi this tree. The 19 branches that join the baseline in the center directly (11 o'clock to S o'clock on the diagram) and four more samples that "areshown grouped together and joining the baseline with a bootstrap value of S3 (indicating little support for treating this group of samples as distinct from the other 19) form a relatively homogenous group. Also not strongly differentiated is the group of three representing reference samples of Z. h. intermedius from Minnesota and Z. h. campestris from South Dakota as well as an individual museum . specimen designated as only Z. hudsonius from Johnson County, Wyoming. Bootstrap values indicate strong support for recognizing samples of Z. princeps, Z. h. luteus, and populations affiliated with them in the diagram as distinct from the Preble's group. Further observations following from the analysis are discussed in the following section." DISCUSSION Preliminary data analyses conducted before all new populations sampled during the summer of 1997 and reference material had been analyzed examined three options for the rooting of the phylogenetic tree. When the Indiana population of Z. hudsonius was assigned as the outgroup (one most parsimonious tree, length"= 63, CI = 0.921), the strong relationships in the tree were the same ones seen in the tree generated when no outgroup was assigned (Figures 2 and 3). Essentially, populations of jumping mice ranging from southeastern Albany County in Wyoming south along the Front Range of the Rocky Mountains to western Las Animas County in Colorado were seen to form a coherent group within which separate populations or groups of populations were not strongly differentiated. Designating Z. princeps as the outgroup resulted in Indiana Z. hudsonius appearing as the sister group to the assemblage of presumed Preble's populations with the populations from the Colorado-New Mexico border"and the Warren Air Force Base joining the tree at an intermediate level. Bootstrap values for the resulting tree were uniformly high, but we have not yet resolved doubts about whether Z. princeps can properly be treated as ancestral to Z. hudsonius. Finally, designating Z. hudsonius intermedius from Minnesota as the outgroup in earlier analyses resulted in a tree in which all the branches to the Preble's populations samples joined the baseline as undifferentiated from the outgroup. Biosphere Genetics. Inc. PageS �52 Preble's DNA study Final Reportl12-8-97 MRD97o:lMNH9716 ROX9701 CGA 1 80S9706 ._._ EPC970t ZHCS0C2 _ LOSt --- RSSt076~ RF96100 Figure 3. Bootstrap consensus tree (50% majority rule, 100 replications), based on analysis of single most-representative individual from each population or reference group, obtained with no outgroup designated. Bootstrap values are shown next to relevant branches in the tree. Bootstrap values greater than 90% indicate those relationships which are strongly supported. . . Biosphere Genetics. Inc. - Paec 9 . ., . �Preble's DNA study Final Reportl12-8-97 53 There are several notable observations for individual population samples among our findings based-on analyses completed in late November, 1997. We summarize these here in list form with population/sample codes in parentheses referring to designations in the tables of the Appendix: 1. The sixjumping mice caught at the Warren Air Force Base (CCF) in the summer of 1996 (identified by field workers as Preble's).and an individual jumping mouse caught in Weld County, Colorado just south of Cheyenne, Wyoming (LTC), are indistinguishable from reference samples of Z. princeps (CZP) provided by Dr. Bruce Wunder from Larimer County, Colorado. Bootstrap values indicate that this group is, in tum, most closely affiliated with, but distinct from, a sample of Z princeps from TaosCounty, New Mexico. . 2. The eastern-most sample identified in the field as Z. h. preblei, a single individual from Elbert County, Colorado (HAY), is confirmed to belong in the Preble's group by DNA sequence obtained from a sample of three hairs. 3. Jumping mice caught at two sites in the Lake Dorothey area on the Colorado-New Mexico border (CCR and WFS), suspected on the basis of morphological traits to be similar 10 Z h. luteus (Cheryl A Jones, personal communication), are confirmed to be so on the basis of the non-coding region sequence data. There is also a reasonably strong indication (bootstrap value = 88) that these two populations and the Z. h. luteus samples are, together, likely to have shared a common ancestor with progenitors of Z princeps reference samples and the populations sampled north (CCF) and south (LTC) of Cheyenne, Wyoming. This finding is in contrast to the conclusions ofHafher et al. (1981) (but see next comment). 4. A suggestion from our analysis (not strongly supported by bootstrap analysis) is that Z hudsonius from Indiana (possibly either Z. h. americanus or Z h. intermedius based on distribution) may be the most ancestral of the populations and taxa sampled and may have shared a common ancestor with progenitors offomis presently known as Z. princeps princeps and Z. h. luteus. We speculate that the forms recognized as subspecies of Z princeps and Z hudsonius present today in the Rocky Mountain and Northern Great Plains region may be the derivatives of two separate range expansions occuning at different times during the Pleistocene. 5. Two 'reference samples obtained from the Denver Museum of Natural Histocy, originally identified as Z princeps from the San Isabel National Forest ·in.western Las Animas County, appear on the basis of non-coding region sequence data to belong to the Preble's group. 6. A singlejumping mouse specimen obtained by Chris Garber from the Medicine Bow National Forest in Wyoming, identified as a Preble's mouse by Dr. David Armstrong on the basis of his analysis ofpoody preserved skeletal material, is confirmed to be a Preble's by DNA sequencing from a piece of preserved skin. 7. One reference collection sample identified as Z h. pallidus from Garden County, Nebraska (ZHPNEG), and two samples identified as z: h. campestris from ~eston Biosphere Genetics. Inc. Page 10 �I· 54 Preble's DNA study Final~nV12-8-97 County, Wyoming (ZHPWYW), are indistinguishable from other samples in the Preble's group in the present analysis. 8. Two reference collection samples identified as Z. h. campestris from Custer County, South Dakota, are found to be most similar to a museum specimen identified only to species level as Z. hudsonius from the vicinity of Buffalo in Johnson County, Wyoming. - ._._ Further Research Needs Additional sample -materials and analyses may be required to achieve a rigorous view of the relationships of the group referred to here as Preble's to other subspecies and species of the genus Zapus in North America The choice ofan appropriate outgroup remains unresolved, and could benefit from further input from those knowledgeable about evolution and morphology of the Zapodids in North America and elsewhere. Based on extensive work on limb myology of the Dipodoidea (Stein, 1990), the superfamily to which the genus and the rest of the family Zapodidae belong, the genus Sictsta (the birch mice), represented by species in Europe and Asia, would appear to be an appropriate outgroup, However, divergence between Zapus and Sicista may be too great to be effectively measured by variation in the D-Ioop. Another North American genus, Napeozapus, has been suggested as a possible outgroup for this analysis. However, slightly greater adaptation for saltatorial ability and a higher tooth count in Napeozapus than in Zapus would suggest that the former is derived rather than ancestral to the latter. Sequence information for the cytochrome b gene potentially forthcoming from other researchers (T. Yates, personal communication) may be helpful in sorting out relationships at the species and higher taxonomic levels and may be a useful complement to the noncoding region sequence data in elucidating relationships potentially traceable to - Pleistocene glaciation events. We have begun to examine a portion of the cytochrome b gene and the neighboring threonine gene in a separate study. . The DNA sequence results reported here indicate the need for a reexamination of external and skeletal morphology in Zapus. _In at least a few cases (DMNH, LDS), the sequence data suggest that _even species level identifications cannot always be made unambiguously in the field. Patterns of morphological variation across the range of the species and subspecies of interest may be such that more careful examination of museum accessions is to be recommended as well. One promising dental character that may distinguish Z. hudSonius-from Z. princeps is the presence of a deep anteromedian fold in the anteroconid of the molar ml in the former as described by Klingener (1963) and applied in an analysis of cave floor faunal remains by Hafner (1993). It will be interesting to learn what this character may say about the taxonomic identity of the samples referenced above and about specimens available in museum collections for the areas near the northern and southern boundaries of the Preble's mouse range. . Further analysis of molecular markers at the species, subspecies, and population levels is needed to confirm the findings suggested by this study and to further delineate patterns of variation bearing on conservation and management ofZ. h. preblei. We have material in hand to continue to expand sample sizes for a number of the populations already surveyed and our initial contacts with several museums in the context of the Biosphere Genetics, Inc. Page 11 �Preble's DNA study Final Reportl12-8-97 present study indicate that sample sizes of comparative taxa can also be increased for at least some parts of the relevant subspecis ranges. Because there has been relatively little recent survey activity in Wyoming and only a few reference specimens from museum collections were included in this study, the northern extent of populations assignable to the Preble's group is poorly defined. New trapping survey initiatives and DNA analysis of more material available from museums would address this situation. We also recommend that a nuclear DNA dataset be generated to complement the mitochondrial DNA dataset. A portion of our work not reported here has evaluated two classes ofDN~ markers potentially appropriate to ~~ task and begun applications of microsatellite methods to development of markers likely to be applicable both to phylogenetic analysis of relationships at the species and subspecies level and to population genetic analyses supporting assessment and monitoring of population differentiation and gene flow, genetic .parameters of population viability, and mating system analysis.. We expect that further examination of the results obtained in this study, both by ourselves and others, will lead to a more comprehensive definition of research needs and priorities along two possibly divergent paths, one inclined to address fundamental issues of systematics and biology, and the other focused more sharply on the near-term requirements that may be imposed by the outcome of the pending decision on the status of the Preble's mouse by the U.S. Fish and WIldlife·Service. We urge that planning attention be devoted to providing for both ongoing research and for integration of assessment and monitoring activities into any prospective recovery effort...Molecular ecology has much to offer in both areas of endeavor. . Conclusion Based on resultsto date, we conclude that mitochondrial DNA non-coding region (D-loop) sequence data appear consistent with the view that a geographically contiguous set of populations previously recognized as the Preble's meadow jumping mouse (2: h.. preb/et) form a homogenous group recognizably distinct from other nearby populations and from another geographically-adjacent species of the genus. Biosphere Genetics. Inc. Page 12 55 �56 Preble's DNA study REFERENCES Final Reportl12-8-97 CITED DeAngelis, M.M., D. G. Wang, and T. L. Hawkins. 1995. Solid-phase reversible immobilization for the isolation of peR products. Nucleic Acids Research 23(22):4742-4743. Greer, C.E., C.M. Wheeler, and M.M. Manos. 1995. PCR amplification from Paraffinembedded tissues: Sample preparation and the effects of fixation. pp .99-112, in: PCR Primer, A Laboratory Manual, C.W. Dieffenbach and G.S. Dveksler (eds.) CHSL Press, New York, 1995. Hafuer,D.l. 1993. Reinterpretation of the Wiconsinan mammalian fauna and paleoenvironment of the Edwards Plateau, Texas. 1. Mamm. 74(1): 162-167. Hafner; 0.1., K.E. Petersen, and T.L. Yates. 1981. Evolutionary relationships of jumping mice (genus Zapus) of the Southwestern United States. J. Mamm. 62(3):501-512. Klingener. D. 1963. Dental evolution of ZaPlIS. J. Mamm. 44:248-260. Maddison, W.P. and D.R Maddison. 1992. MacClade v. 3: analysis of phylogeny and character evolution. Sinauer Associates, Inc. Sunderland, Massachusetts. Stein, B. R 1990. Limb myology and phylogenetic relationships in the superfamily Dipodoidea (birch mice, jumping mice, and jerboas). Z. zool. Syst. Evolut.-forsch. 28:299-314. Swofford, D.L. 1993. PAUP - Phylogenetic Analysis Using Parsimony - version 3.1.1. Computer program distributed by the lllinois Natural History Survey. Campaign, Illino~. . Vigilant, L., R Pennington, H. Harpending, T.D. Kocher, and A.c. Wilson. 1989. Mitochondrial DNA sequences in single hairs from a southern African population. Proc. Natl. Acad. Sci. USA, 86:9350-9354. Walsh, P.S., D.A Metzger, and R Higushi. 1991. CheleX® 100 as amedium for simple extraction of DNA for PCR-based typing from forensic material. Biotechniques. 10(4):506-513. Biosphere Genetics. Inc. Page 1J �57 Appendix Tables in this appendix contain information on the populations and species samples included in . this study. All samples received 'for which DNA was extracted are listed.' Those for which DNA"sequence data was successfully obtained for the analyses reported in"the main document are indicated as explained in each table. . Additions and corrections were still being received as the final draft of this report was going to press. Questions about incomplete or inaccurate data should be referred to the Colorado Division of WIldlife, Terrestrial Section, Attn: Judy Sheppard. �Table 1. Sampling locations for jumping mice analyzed as putative Preble's mice as (Zapus hudsonius preblei). This table includes both samples obtained by 1ivc-trapping during the summers of 1996 and 1997, and samples from museum specimens not identified to subspecies or otherwise designated as reference material (included in Table 2). Numbers of samples for which DNA sequence data were successfully obtained in time for the analyses presented in the main body of the report are shown in parentheses under the number of samples processed. The individuals included in the analyses are indicated· by an.asterix in the column headed "Original sample #s." Samplo Codo BAD BOS Loc:elity/Jitonamo & desaiption or otha' source information Baclwatcr Creek. Natrona Co., Wyoming releY. and cruldnlnldo Nt At Soarb Boulder Creek. tleer intcnectjon with Eat Boulder DitdI. on City of Boulder Opat Space. Boulder CO. 5300 ft. LousMllo Quaclnmglo, TIS, R 70 W cs, No. 1 (1) 13 (1) BVC Bem.r Creek. 2 112mils SW of Monument, EI PlIo Co., CO. 7000 ft. Pa1mcr Lake quaclnmgto, T 11 S, R 68 W 4 (2) CCF Crow Creek, Warren Air Forco BaM. Laramio Co., WY. [elCMltfon and quaclnmgto N/A] TI4N,R67W 6 (6) CCR Clicorica Creek. Lake Dorothey State Wildlifo AreI, Las Animas Co., CO [E1eY.-?] 6 (4) N Brancb, Middlo Forie, Lodgepolo Creek. Polo Mt. Unit, Medicine Bow National ForeSt, ce. 2 mL S. 15 mL B Laramie, Albany eo;, WY, 7600 ft. (Quadranglo - NtAl Deadman Creek. U. s. Air Forco Academy, EI Paso Co., CO, [elCMltfonand quadrangle N/A] 1 (1) COA DC 5 (2) Original samplo'l . CU 13469B089702 . S089703 BOS9704 BOS9705 BOS9706B089707 B089708 B089709 B089710 B089711 B089712 B089715 B089716 1/+114 Section N/A Lat-Long coordinates N/A UI'M Coords. N/A 3, W 112 3, WII2 3, WII2 3, WII2 3, W 112 3, WII2 3, WII2 3, WII2 3. WII2 3, WII2 3, W 112 3, W 112 3, W 112 3~59' 30"N . 105°12'55" W 4818450.00,4427090.00 4818450.00, 4427090.00 4818450.00,4427090.00 4818450.00,4427090.00 4818450.00,4427090.00 4818450.00,4427090.00 4818450.00.4427090.00 4818450.00,4427090.00 4818450.00,4427090.00 4818450.00,4427090.00 4818450.00,4427090.00 4818450.00,4427090.00 4818450.00.4427090.00 1/4 1/4 1/4 1/4 3~03'27.2" 104°53'49.8" Collettor(l) P. Robinson & E. Andenon C. Meaney & N. Clippinger Sponsorin&'Participating organization(. ) Univ.ofColorado Museum. Boulder, CO Colorado Division of Wildlife Clrron Meaney. Consultant i i BVC9701· BVC9702 BVC9703BVC9704CCF9601CCF96O:ZCCF9ti03CCF9604CCF9605·CCF9606482483503 504 .• 505506[Specimen tag] 29,NE 29,m 29,m 29,NE 27 27 27 27 27 27 N/A T15N R71 W sec 11 N/A N/A C.Oarbec (collector) D. Aimstrong University Museum. Univ. of Colorado 97DC02N 97DC03N . 97DC02S97DC05S97DC07S N/A N/A 4318360.00,513700.00 4318360.00,513700.00 4318360.00,513700.00 4318380.00,513650.00 4318300.00.513650.00 P. Sdluerman, S. AsIc, I. Hobert Colorado Natural Heritage Program NlA 3~00'02" 104°21'39" 5088970.00,4323168.00 5088970.00, 4323168.00 5088970.00,4323168.00 5088970.00.4323168.00 NlA N. Clippinger NlA C. Alones C.F1emming C.Paguo Colorado Division of Wildlife Carron Meaney. Consultant Colorado Natural Heritage Program U. S. Air Force Dmva' Museum of Natural History , Page A-2 U1 co �Table 1 continued (2nd page). £PC Eat Plum Creek. 6500 ft. Doaglu Co••CO. DawIcn Butte quadrangle. T 9 S, R 67 W 3 (1) EPC9701EPC9702 EPC9703 9,E 112 9,E 112 9,E 112 39"16'46.3" N 104°53'44.2" W 5090040.00,43478021 Hay Gulch, tn'butatyto Running Creek near Pmcer, Elbert Co., CO. 6,225 ft. Clbin Gulch quldrangle. T 7 S R 64 W. . Colt Creek. Rmsom'Edwm!a Homestead Ranch, 1e.ff'cnonCo., CO, (Elev. -1) iEtdondo- Springs, CO) T 2 S, R 70 W lib De Smctneer Bu1Jiro,1oIJnsonCo., WY. (efeYlltfonand quadrangle N/A1 1 (1) HAY·l- E 112 N/A 540660.00, 4368470.00 (?) ADien, 9601· 7,8,17,18 [Information necdedftom collcctora1 [Informationneeded) C.F1emming C.Pague N/A N/A NlA 4.SW 114 9NW,NW 4,SW 1/4 9NW,NW 4;SW 114 9NW,NW 4. SW 1/4 9NW,NW 40046'11.0" 105~1'36.7" 4696010.00,45132560.00 4696010.00,45132560.00 46915010.00,45132560.00 41596010.00,45132560.00 4696010.00,45132560.00 4696010.00,45132560.00 46915010.00.45132560.00 4696010.00.45132560.00 4696010.00,45132560.00 46915010.00 45132560.00 ACrodcdt J.M. Merritt D. Armstrong A Deans A Deans 5090040.00, 43478021 5090040.00, 43478021 509.0040.00 43478021 HAY 1CC IDS Late Pine Creek. Lower Clcrokeo SlIte WildlffOArea, Larimer Co., CO. 6200 ft. Uvermoro Mountain quadrangle. T 10 N, R 71 W LPC LTC MRD RBC .. RF Late Tree Creek, It 1·25,nOl1han portion Werd Co., CO. 5,900 ft. Carr Southwest quadrangle. T. 11·12 N. R. 67 W. Mmh.n Road, Dry Creek Ditch '2. between Marsh.n road and South Boulder Creek, Boulder Co., CO. 5700 ft. Louisville ..L _t. T 1_S,R 70W . Rabbit Creek, Lower Ctcrokeo SWA CI. 9 mi. NW Uvcrmore. Lat:imerCo.'CO. 6300 ft. Uvermoro Mountain quadranglo ,T 10 N. R 71 W Roc:ty flits Emiloninentll TocIIiloloz.ySite, 1oIftnon eo. CO. 5780ft. Louisville. CO, T 2 S; R 70 W Annendix A 4 (3) I (I) 10 (4) 1 (1) 9602- 9603 9604CUU132· LPC9701· LPC9702· LPC9703· LPC9704LPC9705 LPC9706 LPC9707 LPC9708 LPC9709 LPC9710 LTC-I· 1 (1) MRD9701· 8 (4) RBC9701· RBC9702· RBC9703· RBC9704· RBC9705 RBC9706 RBC9707 RBC9708 4,SW1/4 9NWNW NENE,SE SE 16,NW 1/4 40057' 5.8" N 10 (6) 1/4 Colorado Division of Wildlife Carron Meaney, Consuhant 5063300.00, 45333900.00 A Dean. 39"58'1.6" 105°13'59.7" 4798400.00, 44242200.00 A Deans 40049'25.6" 105°21"8.6" 4715010.00,45212170.00 4715010.00.45212170.00 4715010.00.45212170.00 4702850.00,45194120.00 4715010.00,45212170.00 4715010.00,45212170.00 4702850.00,45212170.00 4702850.00.45212170.00 A. DeanS Colorado Division of Wildlife Carron Meaney. Consultant T.Ryon PTI Environmental Services 10~S5'29.3" 18. SE 1/4 15,NW 1/4 10,SW 114 10,SW 1/4 10, SW 1/4 10,SW 1/4 10 SW 1/4 W N/A ZAHU96-62· ZAHU96-63. 1.1niv.of Colorado M~ Boulder. CO I . 21,NW Colorado Division of Wildlife Catron Meaney. Consultant Colorado Division of Wildlife Carron Meaney. Consultant Colorado Natural Heritage Program Colorado Division of Wildlife Catron Meaney. Consuhant Colorado Division of Wildlife Clrron Meaney, Consuhant ZAHU96-64- ZAHU96-65· HU96-100· HU96-101RF·97·102 RF·97.103 RP-97-104 RP-97-105 1l,5W 1I,5W 11.5W 11.5 W U1 4832800.00,44148500.00 4832800.00.44148500.00 4832800.00.441485.00.00 4832800.00.44148500.00 \D Pace A-J �0\ Table 1, continued (3rd page). ROX SCR Roxborou~ State Patknear juMtiClQ ofWiUow Creek Iftd IJItJe Winow Creek. Dougln Co .• CO. 7300 ft. Kusler quadrangle, T 7 S, R 69 W Smith Credc. U. S. Air Forco Acadany, EI Paso Co .• CO. 6,860 ft. Monument quadrangle, T12S,R67W a 2 (2) 6 (6) STV St. Vrain R., Boulder County Open Spice, HyzlCI2O,Boulder Co ••CO. 5,060 ft. Hyzlene quadrangle, T 3 N. R 70 W 6 (5) WFS West.Fort SdlldJheim Credc, Lab Domhey State Wildlife Area. Lal Animaa Co .• CO [Elev.-1) . (QuadranRle - ?1. f'l'wrnbI)JRngJSeo - 11 Woodhouse R.andl, Douglal Co., CO. 5,760 ft. Kassler qullldranglo, T 7 S, R68 W 4. (4) WCIIl Plum Credc. Dougln Co., CO (Detailed desaiption needed from coDec:tor1 [BIllY. -1) [Quadrangle - 1), T 7 S, R 68 W 8 (8) WHR WPC 7 (5) 24,E 112 39" 25'44.$" 105°03'48.9" 39" 26'10.1" 105°04'24.0" NfA ROX9702' 24,E 112 SCRI' SCR2' SCRl' SCRS' SCR6' SCR,. STVI STV2' STV3' STY4' STY5' STY6' 530' 6NE,SE If4 36, SE 1/4 A Deans Colorado Division of Wildlife Carron Meaney. Consult.ant NfA C. Meaney Colorado Division ofWidlife NfA N/A C. Meaney Colorado Division of Wildlife NlA 3?OOO'25" 104~2'32" N/A C.A]ones Denver Museum of Natural History 36,NE 114, SWI/4 NfA N/A C. Meaney IIId N. Clippmger Colorado Division of Wildlife 36,SW 114, EII2 [Need info. from collector) [Infonnation needed) ? ? '31' 532' 533' WHR1' WHR2" WHR3 WHR4' WHRS' WHR6' WHR7 WPCl' WPC2' WPC3' 4945290.00, 43643880.00 4936!80.00. 43651170.00 I I ~, WRN Monument Creek near ccafluencowitb Pine Credc, N.lldo of Woodman Road, Et Paso Co., CO. 5 (1) WRP Wbito RIIIch pm, Ralston Credc. off'Hwy 43 nOl1h ofOoldeu, Jeft"erson Co., Colondo, 5740 ft. Ratstm Buttes quadrangle, T 2 S, R.70 W 3 (2) Appendix A ROX9701' WPC5' WPC6' WPC7' WPC8' 97WRN130 97WRN138' 97WRN142 97WRN143 97WRN75 WRP9725 WRP9741' WRP9749' I I , N/A 31 NfA 4309600.00.5139400.00 4309580.00.5139400.00 4309550.00,5139400.00 4309550.00,5139400.00 P.SdluennI!I S.Ask J. Hobert 04774210.00, 4401f4950.00 04774210.0Q, 4401f4950.00 04774210.00, 4401f49S0.00 Jill P&:tcrson Colorado Natural Heritage Program Page A-4 �Table 2. RcfereDce specimens of jumping mice provided by museums and other investigators. used as representatives of recognized species and subspecies of the genus Zapus for comparison with specimens listed in Table 1. SamploCodo CZP DMNH RSS ZAHU ZHCSDe ZHCWYW ZHLNMO Spec:Ima1information provided by IOUrCO individual or inttJtutJm NOIU Creek. FR 1$6 oft"Col~Hwy 14, LarImer Co. CO. 10,000 ft. AssIgncdto ZopIII prll'lCflpll No. Purptoiro Campground, 2625 m. San babel NItfClllIIIForat, IA. Anima. eo.. Colorado. AssImcdto Zarnll prlnc.pl at DMNH ZopIII hrldlltmhll aamplcs provided by Bruce Wunder. Anoka County, MInnesota. Blaine, NE 1/4 of'NE. Sec. I, T31N, R23W Premmcd to be Z. h. Intmn,dtti,l ZopIlIlmdlltmhlllllll1'lcs provided by Bruce Wunder, origin in Indiana; may be Z. b. Immnldttll or Z. h. amlrlctmflll Ztrprtl Imdlltmfllil camputrll from Custer Co., SO, provided bytbo Museum ofNlltural ~. 'Univ.ofK.lntat ZopIII hrldlltmhll ~1Itrl1L from Weston Co••WY, provided bytbo MWICUID ofN1ltura1 m.Ocrv. Urriv. ofKmsa. Ztiprlllhrldlltmhlll (ltltetlll) from Otero Co., NM 2 (1) Original.amplo nOlo CUB' CZP9' CZP 12' CZP 13' 7916' 7917 2 (2) 4 (4) 1/4-114 Lat-Lcng Secilon N/A N/A UTM Coordinates NlA CollcctorlSource Cooperating Institution B. Wunder Colorado State Univenity I N/A 37"15'00" N 105°6'30"W N/A C. A Jones Denver Muso:umof Natural History RSS 1076' RSS 1077* N/A N/A N/A B. Wunder Robert Sikes ElmerBumey Colorado Stat" University University ofMinnescu 2 (2) ZAHU-253' ZAHU-300' N/A NlA N/A Colorado State University 2 (2) 109994' 109995' N/A N/A NlA B. Wunder cotlcdcd by John Whittaker R.Timm T.Hotme. 2 (2) 42469' 42470' N/A N/A N/A R.Timm T.HotmCs Natural History Muso:um, Univ. of Kansas. Lawrence 2 (2) NK878' NK879' N/A N/A N/A T. Yates B.GannClll. Museum ofSouthwestc:m Biology. Univ. of New Mexioo. Albuquerque, NM Museum ofSoutbwestc:m Biology, Univ. of New Mexioo. Albuquerque, NM Natural History Museum, Univ. of Kansas. Lawrmce Natural History Museum, Univ. of Kansas. Lawrmce Univ. of Colorado Museum. Boulder, CO Museum ofSoutbwestc:m Biology. Univ. of New Mexioo. Albuquerque, NM ZHLNMS ZopIIIIhrldllonhll (ltltetlll) ftom SandoVIIICo., Fcntm lAke, NM 2 (2) NK3835' NK3837' N/A N/A N/A T.Y~ B.qannon ZHPNEC ZopIIilhrldilonillil palllcltlil ftom Cherry Co., NE ZopIIil hrldlomllil pallfcltlll ftom Garden Co., NE ZopIIilprll'lCflPILprlnc.plI from Ptarmigan ciinp:OrandCo CO . ZopIIIIprlnc.PI prll'lCflP' ftom Tlos Co., NM 2•. (0) 2 (1) 2 -87043 87044 N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A N/A R.Timm T.Hotmes R.Timm T.Holmes AdeQueiroz D.ArmsIronIt T. Yates B.Gannon ZHPNEO ZPPCOO ZPPNMI' (0) 2 (1) 115762' 115763 CU14912 CU14913 NK811 NK814' I .' Natural History Museum. Univ. afKansas. Lawrmce N/A - not available for 'this report. In some cases the indicated information may be obtained from the person or institution supplying the material analyzed or can be inferred from other available information. 0'1 ~ Appendix A r.II·!' ':,."1 ��63 I APPENDIXB STUDY State of __ -'C""-o""'l"""o"""'ra=d=o"-_ Project No. W-1S3-R-11 Work Plan NO. __ O=6=6=2 _ Task No. ....::3'-_ II PLAN Cost Center 3430 Mammals Research Conservation of Preble's meadow jumping mouse Temporal and spatial variation in the demography of Preble's meadow jumping mouse (Zapus hudsonius preblei) TEMPORAL PREBLE'S A. AND SPATIAL VARIATION IN THE DEMOGRAPHY MEADOW JUMPING MOUSE OF (Zapus hudsonius preblei) NEED On May 12, 1998 the U.S. Fish and Wildlife Service (USFWS) published a final rule in the Federal Register (63 FR 26517) to list Preble's meadow jumping mouse (Zapus hudsonius preblei) as 'threatened' under the Federal Endangered Species Act (ESA) of 1973, as amended. Recovery goals for Preble's meadow jumping mouse should work towards the sustainability, protection, and restoration of Z. h. preblei populations and habitats on both private and public lands to provide the spatial, genetic, and demographic structure needed to promote long-term species viability and provide species management flexibility. Recovery efforts for the subspecies will be most effective if reliable information is available on the basic ecology of the subspecies and this information used to design recovery efforts such as Habitat Conservation Plans. A review of studies conducted on Preble's meadow jumping mouse shows that there is insufficient information to fully address defining rangewide ecological requirements, limiting factors, limits of species tolerance, or population status (Shenk 1998). Most work to date has focused on geographic distribution (presence or absence of Z. h. preblei), taxonomy, and habitat descriptions of sites where mice have and have not been captured. For Preble's meadow jumping mouse in particular, information on dispersal, habitat use, and population dynamics is most needed to identify minimal ecological requirements of the subspecies. Monitoring, by way of estimating demographic parameters (such as survival, abundance, and reproduction), of a single population over time provides an opportunity to document demography of a species, estimate temporal fluctuations in demography, and gain insights into the temporal variation inherent in demographic parameters. Monitoring of multiple populations over time and over multiple geographic locations provides the opportunity to gain further understanding of population processes and detaches time effects from spatial effects. Population monitoring activities do not constitute scientific experiments, in the spirit of manipulation of salient ecological variables, however, replication of monitoring activities for natural populations over long periods of time and in diverse geographic locations can lead to insights into population processes (Cook and Campbell 1979). These insights can then be translated into hypotheses useful for predicting changes in population demography resulting from either natural perturbations (e.g., flooding events) or anthropogenic modifications (e.g., gravel mining). Experimentation would then be required to test these hypotheses and establish cause and effect. There are three components of Z. h. preblei ecology that are currently unknown and yet key to any sound conservation strategy for the subspecies. These are (1) detailed demographic studies estimating survival, reproduction, immigration, emigration, and abundance and determining the factors influencing each of the parameters, (2) detailed studies evaluating movements and dispersal habitat of individuals within and among populations, and (3) detailed studies to defme hibernation needs, primarily descriptions of suitable hibernacula criteria and food requirements for sufficient fat storage prior to immergence. The overall objective of this study is to provide estimates for all three of these �64 needs from three different populations of Preble's meadow jumping mouse. Thus, providing the opportunity to estimate spatial variation in the demography of the species. The three study populations occur in areas of different habitat matrices. The first study area limits animals within the population to a single drainage. The second population occurs in an area that combines a tributary and main drainage. The third population occurs in an area with both a tributary and main drainage as well as a series of ponds and irrigation ditches available to the animals. Continuing the study for multiple years will provide the opportunity to estimate temporal variation in the demography of the subspecies. B. OBJECfWES Specific objectives for this study on Preble's meadow jumping mouse include: 1. 2. 3. 4. 5. 6. 7. c. Evaluate temporal and spatial variation in daily and seasonal movements of Preble's meadow jumping mouse. Estimate and evaluate temporal and spatial variation of over-summer, over-winter, and annual survival rates for Preble's meadow jumping mouse. Estimate and evaluate temporal and spatial variation of reproduction, immigration, and emigration for Preble's meadow jumping mouse .. Estimate and evaluate spatial variation in abundance and density for Preble's meadow jumping mice. Evaluate temporal and spatial variation of habitat use in Preble's meadow jumping mouse. Document temporal and spatial differences in composition of foods consumed by Preble's meadow jumping mice. Collect genetic tissue samples for future analyses to document and compare variation within and among populations of Preble's meadow jumping mouse. EXPECfED RESULTS There are three components of Z. h. preble; ecology that are currently unknown and yet key to any sound conservation strategy for the subspecies. These are (1) detailed demographic studies estimating survival, reproduction, immigration, emigration, and abundance and determining the factors influencing each of the parameters, (2) detailed studies evaluating movements and dispersal habitat of individuals within and among populations, and (3) detailed studies to define hibernation needs, primarily descriptions of suitable hibernacula criteria and food requirements for sufficient fat storage prior to immergence. The overall objective of this study is to provide estimates for all three of these needs from three different populations of Preble's meadow jumping mouse providing the opportunity to estimation spatial variation in the demography of the species. Continuing the study for multiple years will provide the opportunity to estimate temporal variation in the demography of the subspecies Performance Indicators 1. Provide descriptions of movements of mice for three Preble's meadow jumping mouse populations. 2. Provide estimates of survival and reproduction for three Preble's meadow jumping mouse populations. 3. Provide estimates of immigration, emigration, and abundance for three Preble's meadow jumping mouse populations. 4. Describe over-summer foods consumed by three Preble's meadow jumping mouse populations. 5. Quantify habitat characteristics of three sites where Preble's meadow jumping mouse are known to occur. 6. Analyze and publish results of research �65 D. ApPROACH Study Site Selection All three study sites selected are areas where Preble's meadow jumping mouse (PMJM) has been found in the last two years. To evaluate spatial differences in movement ofPMJM, sites were selected that provided a variety of habitat matrices available to the mouse. The first site selected, CDOW Maytag property, has one primary water source available to the mouse. This water source is East Plum Creek. Therefore we predict mice will restrict their movements to up and down this single drainage. The second site, Colorado Open Lands Pine Cliff Ranch, provides both a tributary (Garber Creek) and a main stem drainage (West Plum Creek). This provides the opportunity to investigate whether PMJM will move overland to move from one drainage to another or if they are restricted to moving only along riparian corridors. The third study site, CDOW Woodhouse Property and Dupont Property, provides an area containing a tributary (Indian Creek),a main stem drainage (West Plum Creek), and a series of ponds and irrigation ditches scattered throughout the property. This provides an even greater opportunity to investigate how much the mice will use upland areas or if they restrict their movements strictly to riparian corridors. Each of these unique habitat matrices provides the opportunity to estimate spatial variation in nightly and seasonal movements of PMJM. Movement Study Background Movement and dispersal pattern information will be key to any conservation strategy designed for Preble's meadow jumping mouse. Key factors include (1) which segment of the population disperses, (2) when.do they disperse, (3) through what habitat do they disperse, (4) how far will individuals disperse (i.e., what is the maximum distance that separates adjacent populations) and (5) how critical is dispersal (both into and out of a population) to the persistence of a given population. The only currently available data on dispersal and/or movement for Z. h. preble; are from marked mice at Rocky Flats Environmental Technology Site (T. Ryon unpublished data). Two mice, an adult female and an adult male, were observed approximately 1.6 kilometers from previous locations (incidences occurred separately). Each of the locations were in the same drainage (Woman Creek). Objectives The PMJM movement study is designed to describe nightly and seasonal movement patterns of PMJM and to describe habitats used by PMJM. Specific objectives include: 1. 2. 3. 4. 5. Describe nightly movements of PMJM. Evaluate difference in nightly movements as they relate to sex, age, and habitat available to the animals. Describe 30-day (or life of the radio transmitter) interval movements ofPMJM. Evaluate difference in 30-day (or life of the radio transmitter) interval movements as they relate to sex, age, and habitat available to the animals. Describe seasonal movements ofPMJM. Evaluate difference in seasonal movements as they relate to sex, age, and habitat available to the animals. Describe habitats where mice occur: movement corridors, end point descriptions (disperses from what to what), and landscape features (connectivity with other riparian strips). Estimate the mean amount of time PMJM spend in each available habitat. Approach Movement data will be documented primarily from locations of radio-tagged mice from each of the three study populations. To place radio transmitters on the animals, mice will be captured in Sherman live traps. Mice weighing> 18 grams will be fitted with MD-2C, I-gram radio transmitters supplied by Holohill Systems Ltd. (used successfully on PMJM by R Schorr, personal �66 communication). A maximum of 30 mice at each study site will be radio-tagged. The following procedures will be used to capture and attach the transmitters to the mice. Trapping session details Timing of surveys 1. Four 7-day trapping sessions will be conducted during the following weeks: June 2-9, 1998; July 21-28, 1998; September 8-15, 1998; and June 2-9, 1999. 2. Due to the nocturnal nature ofPMJM, traps will be set between 19:00hrs and 21:00hrs and checked as early as possible in the morning beginning at 5:00hrs (to reduce stress and the potential for predation on trapped animals). Time required to complete the trap lines will vary depending on how many animals are caught. Trapping protocols 1. Trappers will be advised to follow the Center for Disease Control's Hantavirus instructions and recommendations when dealing with rodents which include: a Baseline blood serum samples will be collected and stored at Poudre Valley . Hospital for all surveyors. b. Respirators are available for use by surveyors if they request their use. c. Surgical gloves will be used at all times when handling rodents. d. All equipment will be rinsed first with a 10% bleach solution, then water after use. e. Traps and any equipment that may have rodent feces or urine on it will not be transported in the interior of a vehicle. f Traps will be cleaned after each use by a mouse. g. If a surveyor becomes ill with hantavirus symptoms they will report immediately to a medical facility. 2. Trappers must be in possession of both a federal and state collecting permit. Trapping methodology 1. Small mammal Sherman live traps (folding and non-folding) will be used to conduct the trapping sessions. 2. Traps will be set in two parallel lines of trap stations (1 trap per station) on either side of the drainage. Trap stations will be 5 meters apart for a total of 250 meters; the parallel transects will be 10 meters apart unless extent of habitat, terrain topography, or stream hydrology do not allow. 3. Location of transects will be recorded on field data sheets and identified on 7Yz minute topographic maps. Locations will also be recorded to the nearest 5 meters in UTM coordinates using a Trimble Geo-Explorer GPS. 4. In case of windy conditions or large number of trap-tampering predators (i.e., raccoons, foxes, coyotes, etc.), traps will be secured to the ground with hoops of heavy-gauge malleable wire, stakes, or with other materials that can effectively secure/immobilize the traps. 5. A small (-1 inch) ball of polyester quilt or wool (fleece) will be placed in each trap as nesting/bedding material. 6. Baiting materials will be Manna Pro Sweet 3-way Livestock feed which contains no animal matter. Ingredients include flaked barley, flaked corn, flaked oats, and cane molasses. Peanut butter will be used to stick the bait to the trap. 7. Checking of the traps will be conducted by two surveyors; one person will handle the traps and animals captured, the other person will record the data Handling of captured animals 1. All animal captures will be recorded. �67 2. 3. 4. 5. If an animal has been captured in a trap, a ziplock plastic bag will be placed over the end of the trap. The trap will be opened allowing the animal to fall into the plastic bag. The animal will be identified to species while in the plastic bag. If the animal is not a PMJM, identification of the animal will be recorded and the animal set free. Each captured PMJM will be scanned to detect the presence/absence of a PIT tag and/or radio-collar. If a PIT tag or radio-collar is detected and the mouse has been captured at that same site within the 7-day trapping effort, identification of the mouse will be recorded and the mouse will be released. If the animal is a PMJM and no PIT tag or radio collar is 'detected the PMJM will be anesthetized for further processing, as follows. All PMJM will be PIT-tagged. If the PMJM weighs> 18 g a radio-collar will also be put on the animal Anesthesia The following protocol has been used successfully on PMJM by R Schorr (personal communication). l. Measure 1 ml of Metofane (methoxyflurane) onto one cotton ball. 2. Place ball into ziplock bag and seal (to keep Metofane fumes in bag). 3. Lay bag still and move away. This should minimize stress for the mouse and the time the mouse struggles in the bag, decreasing the amount of time required for the metofane to work. 4. After the animal stops movement, wait 1 minute before removing it from the baggie. The animal should remain anesthetized for 2-3 minutes. 5. Time animal is exposed to Metofane and reaction to the anesthesia will be recorded. PIT tagging the animal The following protocol has been used successfully on PMJM (C. Meaney, personal communication). 1. Each PMJM will have a PIT tag inserted above the shoulder blades by lifting the skin on its back and inserting the needle with,the PIT tag under their skin and injecting the tag. Verification of the PIT tag identification number will be made before insertion into the mouse by running the PIT tag scanner across it. 2. Skin behind the opening will then be pinched to prevent emergence of the tag. 3. Surgical glue will then be applied to the opening to prevent the PIT tag from exiting the body and prevent infection of the mouse. 4. A PIT tag scanner will be held over the mouse to again verity PIT tag identification number and that it still works after the insertion procedure (note: PIT tag will be scanned twice during application to mouse, or once if mouse was previously PIT tagged). 5. PIT tag identification number will be recorded on the field data form and the mouse released. 6. If, at any time during the handling of the PMJM the animal appears to be severely stressed (dramatic changes in heartbeat, respiration and responsiveness or gums turning blue) the animal will be released immediately. Collaring the animal The following protocol has been used successfully on PMJM (R Schorr, personal communication). 1. Make sure transmitter is functioning before collaring the animal with the transmitter by tuning the receiver to the transmitter frequency to make sure there is a signal. 2. Slide crimp and then tubing onto the antenna �68 3. 4. 5. 6. 7. 8. Guide antenna through the channel of the transmitter. As antenna exits the transmitter, guide it through the crimp. Keep loop made by antenna large enough to fit over the mouse's head. Once over the neck of the animal, pull antenna to reduce collar size, making sure rubber tubing is on the dorsal side of the neck. The purpose of the rubber tubing is to prevent damage to the collar and prevent the person attaching the collar from cinching the collar too tight. Make sure collar can rotate freely around the animals neck, but not loose enough to be removed. Pinch metal crimp to hold the antenna in the desired collar position. Approximately 2 inches of antenna will be pointing out behind the animal. Measurements 1. Each PMJM will be weighed while in the bag, recording the weight in grams using a Pesola spring balance. 2. Sex, age (juvenile, adult), and reproductive condition (pregnant, lactating or nonbreeding if it is female; for males the position of the testes will indicate if in breeding status or non-breeding status) will be noted. 3. Each PMJM will be measured (total length, length of body, length of hind foot - heel to distal end of claws) in millimeters. Measurements of total body and tail length will be taken with the mouse on a firm surface to minimize error. 4. Capture of every PMJM will be documented by taking a photograph of the mouse on first capture. If the mouse has been captured previously as noted by the detection of a PIT tag, no photograph needs to be taken. 5. If there are feces in the trap where a PMJM was captured, the feces will be collected in a plastic bag labeled with date and location of the site. Fecal samples will be kept cool until returned to the CDOW office where they will be frozen. Fecal samples will be analyzed by the Composition Analysis Laboratory, Inc, 622 liz Whedbee, Fort Collins, CO for content. Handling of trap mortalities and injuries 1. All trap mortalities will be recorded on a 'Trap Mortality' data form which includes information on species, potential duration of time spent in the trap, any information available on cause of death, etc. 2. All animals found dead in a trap will be double-bagged in a plastic bags and placed in a cooler with ice. Specimens should be frozen as soon as possible and deposited the CDOW freezer at the office in Fort Collins. All specimens will be sent to the Colorado State University Diagnostic Laboratory for detection of any disease and/or other conditions that may have contributed to cause of death. All PMJM specimens will be returned to the CDOW. 3. A museum card will be completed and attached to each PMJM specimen and given to Cheri Jones, Curator of Mammalogy at the Denver Museum of Natural History, for study skins and tissue storage once the Diagnostic Laboratory has completed their evaluation. 4. If an animal is severely injured (e.g., severed limb, large lacerations) it will be euthanized by soaking a cotton ball in Metofane (methoxyflurane) and placing the cotton ball and the mouse in a ziplock baggie until the animal stops breathing. All specimens will then go through steps 1-3 above. 5. If an animal appears to be only slightly injured (e.g., broken tail, small laceration) the animal will be released to the wild. If an animal appears to be cold stressed attempts will be made to warm it by holding it in the surveyors hands and/or against their body. If the animal appears to be heat stressed, isopropyl alcohol will be applied with a cotton swab to the ears, arm pits, and feet to cool it down. �69 Training of field crews 1. All field technicians will be required to spend a full day at the Denver Museum of Natural History Zoology Department viewing and handling specimens of all small mammal species likely to be captured in Douglas County. 2. All field technicians will be provided with a small mammal field guide and a key specifically designed to list the species most likely to be captured on the trapping sites. 3. All field technicians will participate in 4 days of practice trapping at the Frank State Wildlife Area from May 26-29. During these trapping sessions all technicians will learn how to place, bait and set traps. Field identification will be practiced on all animals caught each day and verified by crew leaders. All field technicians will also practice handling, measuring, implanting PIT tags, and taking genetic tissue samples from deer mice (Peromyscus maniculatus) until they are competent in this procedure. Data collection Once transmitters are in place, locations of individual mice will be made nightly. Because very little is known about the movements of PMJM the pilot protocols will be established. As we learn more about PMJM movements, protocols will be adjusted to maximize efficiency and data collection. To describe nightly movements the following parameters will be estimated: 1. Minimum and maximum distances moved each night animal is followed. 2. Mean minimum and maximum distances moved each night for all mice. 3. Minimum and maximum distances moved away from the stream center each night by individual mice. 4. Mean minimum and maximum distances moved away from the stream center each night for all mice. To describe 30-day (or life of the radio transmitter) movements the following parameters will be estimated: Minimum and maximum distances moved each 30-day (or life of the radio transmitter) 1. interval animal is followed. Mean minimum and maximum distances moved each 30-day (or life of the radio 2. transmitter) for all mice. Minimum and maximum distances moved away from the stream center each 30-day (or 3. life of the radio transmitter) interval by individual mice. Mean minimum and maximum distances moved away from the stream center each 304. day (or life of the radio transmitter) interval for all mice. Locations: 1. A minimum of 6 locations per individual mouse per night will be made. As many mice as possible will be tracked on a given night as is feasible to assure the minimum 6 locations per individual. Each individual mouse should be tracked a minimum of 3 nights per week. 2. The 6 locations per night per individual will be scattered throughout the night to maximize information learned about nightly movements (i.e., 8 locations taken within a single hour contains less information than 8 locations taken one each hour). 3. Three bearings will be taken for each location from pre-determined way stations to estimate location of the mouse. 4. Daytime locations will be taken as time permits. �70 Demography Study Background Information on the population dynamics of Preble's meadow jumping mouse is necessary to determine which areas support viable populations. To begin to evaluate the viability of a population information on key demographic parameters must be obtained. Below is a summary of what is know to date on the demography of Preble's meadow jumping mouse. Abundance: There is no information on range-wide abundance of Preble's meadow jumping mouse. However, from trapping surveys it appears that where the subspecies does occur it exists in low densities and thus one of the rarer components of the small mammal assemblage present. Reproduction: Meadow jumping mice have been observed to produce up to three litters per season (Whitaker 1963). Breeding peaks appear to occur in early to mid-June and August with a possible third litter in September (Whitaker 1963). Juvenile Z. h. preblei have been observed in June, August, and September (Meaney et al. 1996, 1997, PTI 1996a, M. Bakeman unpublished data, T. Ryon unpublished data), suggesting two litters per year. Z. hudsonius typically have litters of 5-6 young per litter (Quimby 1951). Age of first reproduction is unknown for Z. h. preblei, however, females of Z. hudsonius have been observed to give birth at 3 months (i.e., females born in June have been observed to give birth in August of the same year). Gestation period is approximately 18 days (Quimby 1951). Young remain dependent on the female for approximately 18 days (Quimby 1951). No evidence of male parental care exists for Z. hudsonius (Whitaker 1963). Survival: No information exists on survival rates for populations of Z. h. preblei. Whitaker (1963) reported a 67% loss of individuals over hibernation and that average body mass of individuals emerging from hibernation was greater than the average for mice entering hibernation. Because no mice are mown to store food in their hibernacula, this indicates that the lighter individuals died during hibernation and only those entering with higher masses survived. All the energy they use during hibernation and the periodic arousals (the energetically most expensive part of hibernation) must be the fat they carry into hibernation (B. Wunder, personal communication). Thus, the ability to put on sufficient fat for overwinter survival during hibernation is a critical factor in the life history of these mice. Besides insufficient fat storage prior to hibernation, other observed mortality factors in Z. hudsonius include predation (Whitaker 1963, Poly and Boucher 1997) and cannibalism (Sheldon 1934). Other assumed mortality factors for Z. h. preblei include starvation, exposure, and disease. Population Structure: Armstrong et al. (1997) reported an overall sex ratio for all captured Preble's meadow jumping mice of 5l.6 males: 48.4 females; approximately 86.0% of captures were identified as adults. However, Armstrong et al. (1997) suggested that these data be interpreted with caution because of possible differences in field techniques. Longevity: Very few individuals of Z. h. preblei have been permanently marked. Therefore, recapture information, necessary to determine longevity, is minimal. Several recaptures have yielded adults surviving through two years, indicating a longevity of at least three years (T. Ryon, unpublished data). One recapture history from Rocky Flats Environmental Technology Site recorded an adult male in 1996, two years after it was first captured as an adult in 1994, indicating survival of at least four years (PTI 1996b). Quimby (1951) found that only a low percentage of Z. hudsonius lived two years or more, but gave two records of mice that lived for at least two years under natural conditions. Whitaker (1963) reported a female living at least two years. Objectives Objectives of the demography study are to: 1. Estimate abundance of PMJM at each of three study populations. 2. Estimate over-summer, over-winter, and annual survival ofPMJM at each of three study populations. 3. Estimate temporary emigration ofPMJM at each of three study populations. 4. Estimate immigration of marked PMJM back into each of three study populations. 5. Evaluate the affect of weight, sex, age, abundance (i.e., density dependent response), �71 6. and habitat features such as stream reach, vegetation composition and density on survival, reproduction, abundance, temporary emigration, and immigration of marked animals back into three study populations of PMJM. Estimate age and sex ratios of PMJM at each of three study populations. Approach Mark-recapture estimation techniques will be used to estimate abundance, over-summer survival, over-winter survival, temporary emigration and immigration of marked animals back in to the three study populations of PMJM. Abundance: Abundance will be estimated using Pollock's Robust Design (see Kendall et al. 1997, 1995 and Kendall and Nichols 1995 for detail). The robust design is a combination of the CormackJolly-Seber (CJS)(Cormack 1964, Jolly 1965, Seber 1965) live recapture model and the closed capture models. The key difference from the CJS model is that instead of just one capture occasion between survival intervals multiple capture occasions are used. These occasions are close together in time allowing the assumption that no mortality or emigration occurs during theses short time intervals. The closely spaced encounter occasions are termed "trapping sessions" and each trapping session can be viewed as a closed capture survey. Four 7-day trapping sessions will be conducted during the following weeks: June 2-9, 1998; July 21-28, 1998; September 8-15, 1998; and June 2-9, 1999. Survival: By using the estimate of the probability that an animal is captured at least once from the trapping sessions designed to estimate abundance, survival between the longer intervals can be estimated. Four 7-day trapping sessions will be conducted during the following weeks: June 2-9, 1998; July 21-28, 1998; September 8-15, 1998; and June 2-9, 1999. Thus, estimates of over-summer survival, over-hibernation survival, annual survival and survival from June-July, July-August, and August-September can be made. Temporary emigration and immigration: The longer intervals between trapping sessions also allows estimation of temporary emigration from the trapping area, and immigration of marked animals back to the trapping area Reproduction: Reproductive parameters will be estimated by following radio-tagged females to nest sites. At each nest found the following will be recorded: date, number of young in each nest and number of litters observed for individual females throughout the year Habitat Use Study Background The habitat matrix within the range of Z. h. preblei is mixed grasslands adjacent to the Colorado Front Range along the Piedmont and along the base of the Laramie Mountains in Wyoming and extends to the Colorado plains. Within this matrix, Preble's meadow jumping mice occur along stream drainages that contain patches of suitable vegetation. Suitable habitat appears to have at least two major components. The first component is a supply of open water, at least in part of the active season (M. Bakeman, C. Meaney, personal communication). Secondly, areas where Preble's meadow jumping mouse has been found have dense cover (M. Bakeman, C. Meaney, personal communication). If Preble's meadow jumping mouse behaves as a metapopulation in the classical sense of a set of local populations linked by infrequent dispersal then habitat includes not just one area of suitable habitation but also areas suitable for nearby mouse populations. These suitable areas must also be linked by dispersal habitat. If the mice are dependent on dense riparian habitat for dispersal as well as for areas to reproduce, persistence of discrete populations would require a mosaic of suitable discrete riparian patches interconnected with dispersal corridors of similarly dense riparian vegetation. If mouse populations function in a source (populations where growth rate ~ 1) and sink (those populations where �72 growth rate < 1, maintained through immigration) system, it will be critical to identity and protect those populations serving as sources. Thus, for a source-sink population critical habitat will include those areas that support source population, dispersal habitat to sink areas will be less critical. If local mouse populations are functionally discrete, such a mosaic of interconnected areas of suitable habitat would provide a buffer for local, and source, populations against deleterious stochastic events by providing the opportunity for local population failures to be 'rescued' by immigration from other populations. Areas of suitable habitat must also provide requirements to survive throughout the life cycle. These requirements must provide necessities for both the active period and hibernation periods. During the active period suitable habitat must provide requirements for daily survival, reproductive activities (breeding, nesting, and rearing of young to independence), and dispersal. The hibernation period requires sufficient food supplies to assure fat storage prior to hibernation and suitable hibernacula, Habitat providing all seasonal and life cycle requirements mayor may not occur in a single contiguous area If not in a contiguous area, habitat patches must occur in a mosaic of usable areas where suitable corridors exist for seasonal movement among sites. Based on studies of Z. h. preblei and Z. hudsonius elsewhere, Z. h. preblei apparently occurs mostly in undergrowth consisting of grasses, forbs, or both in open wet meadows and riparian corridors, or where tall shrubs and low trees form an overstory and provide adequate cover (Armstrong et al. 1997). Meadow jumping mice are widespread in abandoned grassy fields, but is often more abundant in thick vegetation along ponds, streams, and marshes or in rank herbaceous vegetation of wooded areas (Whitaker 1963). Preble's meadow jumping mice have been trapped in natural riparian areas as well as areas altered by anthropogenic influence including ditches and wetlands adjacent to interstate highways, cement-lined ditches with tall cover, ditches along driveways and moderate road use, and moderate cattle grazing (M. Bakeman, personal communication). The majority of sites where Z. h. preblei have been found consist of multistoried cover but the species composing the cover vary greatly (Armstrong et al. 1997, Meaner et al..1997). Vegetation composition of the dense cover varies considerably and includes both native and non-native species (Meaner et al. 1996, 1997, Armstrong et al. 1997, M. Bakeman, personal communication). The herbaceous understory can be primarily grasses or forbs or a mixture of the two. Few of the sites, however, are dominated by fewer than two understory species. The tall shrub canopy at most sites is willow (several species), although scrub oak, birch, and alder occur in sites south of the Palmer Divide (Armstrong et al. 1997). Ponderosa pine is the most common tree at higher elevations. The mouse appears to tolerate weedy or exotic species in areas that are structurally diverse and species rich; nearly every successful site contained Canada thistle (Armstrong et al. 1997). Thus, the mouse does not appear to have an affinity toward any single plant species but instead favors sites that are structurally diverse and provide adequate cover and food throughout its life cycle. Preble's meadow jumping mouse mice are typically not found in upland areas away from riparian habitats but are most often captured where either ground water daylights to seep springs or on main water channels (M. Bakeman, T. Ryon, personal communication) suggesting a dependence on open water, at least during their active periods. Movement of Z. h. prebleion Woman Creek at Rocky Flats Technology Site suggests mice move along corridors of shrub cover, generally Salix exigua (PTI 1996a, T. Ryon, unpublished data), suggesting dispersal habitat is similar to habitats used for other activities. Jumping mice of the genus Zapus are true hibernators, spending much of their lives in hibernation. Meadow jumping mice spend approximately 7 months (~210 days) per year in hibernation (Quimby 1951) whereas estimates for Z. princeps indicate that some populations (e.g., in the western mountains of Utah) spend up to 300 days per year in hibernation (Cranford 1983). Males emerge prior to females (Bailey 1923, Bailey 1929, Hamilton 1935, Quimby 1951, Whitaker 1963) with the earliest annual recorded dates for Z. hudsonius males being April 25-May 16 and females May 4-26. Latest fall capture date for an adult male was October lOin 1994 at South Boulder Creek (ERO Resources 1995) and September 28 in 1995 at the VanFleet Parcel on South Boulder Creek (Armstrong et al. 1997). A juvenile male was captured as late as October 26 and a female juvenile on �73 October 27 both in 1995 at Rocky Flats Environmental Technology Site (M. Bakeman, unpublished data). Jumping mice hibernate in underground burrows (Quimby 1951, Whitaker 1963). They are excellent burrowers and create their own hibernacula, Meadow jumping mice are generally solitary hibernators, however, there have been occurrences of more than one mouse found in a single hibernaculum. One hibernaculum, located on Rocky Flats Environmental Technology Site, used by Z. h. preblei has been located (Armstrong et al. 1997). This site was 9m above a creek bed (Walnut Creek); it had a thick cover of chokecherry (Prunus virginianay and snowberry (Symphoricarpos spp.), the mouse was found in a leaf litter nest 30cm beneath the ground in coarse textured soil (Armstrong et al. 1997). Four possible hibernacula were located by tracking radio-telemetered mice at the U. S. Air Force Academy in fall 1997. These sites are located 7, 12, 29, and 31m from a creek bed (R Schorr, personal communication). There was no consistency among sites in aspect (N/NW, S/SE, E, and none [level ground)). Three sites were in vegetation dominated by coyote willow (Salix exigua), one site was in vegetation dominated by snowberry and mullein (Verbascum thapsus). However, all four hibernacula appear to be below coyote willows. These four U. S. Air Force Academy sites have not been disturbed to protect any hibernating mice and therefore are only possible hibernacula because there is no confirmation a mouse is actually hibernating there. Confirmation of a true hibernaculum cannot be made until a chamber, or nest is located. These sites may also possibly be locations of radios discarded by the mice or dead mice. Objective The objective of the habitat study is to identify and refine habitat requirements of Z. h. preblei, including hibernation sites, and to determine if they influenced any of the demographic parameters that will be estimated in this study. Approach The general approach is to measure both site specific and landscape features at each of the three population study sites to document the extent of spatial variation. These quantitative measures of spatial variation will be used in the analyses to determine if they influenced any of the demographic parameters that will be estimated in this study. The following habitat characteristics will be measured and recorded for each site. Micro-site habitat characteristics 1. Cover will be measured at 20 random locations along the 250 meter sampled stream stretch. Mean cover and standard error of the mean will be estimated. Cover will be estimated using a vegetation profile board (Nudds 1977) that allows for an assessment of visual obstruction in 0.5 meter vertical intervals above ground. The board will be 1.5 meters high and 30.48 cm wide. The board is marked in alternate black and white colors at 0.5 meter intervals. Horizontal cover is assessed in each interval by viewing the board from 15 meters away in a randomly chosen direction. The percentage of each interval concealed by vegetation is recorded as 0-20, 21-40, 41-60, 61-80, and 81-100% estimated concealment. 2. Vegetation species composition and richness will be estimated using the Modified-Whittaker nested vegetation sampling method (Stohlgren et al. 1995). A modified Whittaker plot is 20meters x 50-meters with 101m2 and 210m2 subplots arranged systematically around the perimeter and 1 100m2 subplot centered in the inside of the 20 meter by 50 meter plot. Species composition and percent cover of each species is recorded for each subplot. 3. Soil samples will be taken at each site for a hydrometer textural analysis (percent sand, silt and clay). Samples will be collected using a soil probe. Samples will be taken from 0-12 inches, and 12+ to 24 inches. The 0-12 inch sample will be taken first, removing the probe and soil. Sample will be placed in a labeled ziplock bag. The probe will then be placed in the same hole for the 12+ - 24 inch probe. With that soil sample being placed in a separately labeled ziplock bag. The hydrometer textural analysis will be conducted at the CSU Soils Laboratory. �74 4. 5. 6. 7. Mean stream width will be estimated by measuring stream width at 30 locations along the entire site sampled. The 30 locations will be stratified equidistant from each other to cover the entire stream stretch in increments of site stream length/20. Describe potential hibernacula: upland vegetation, take soil sample at site. Describe nest sites (what they are made of, placement). Compare parameter estimates from 1-6 for the three population study sites. Landscape habitat characteristics The following landscape habitat characteristics will be measured using either GIS, topographic maps, or aerial infrared photography. 1. Estimate distance to nearest human habitation and human disturbance. 2. Estimate connectivity of sites to other streams. 3. Estimate total length of stream stretch at each site and total length of stream stretch with suitable habitat at each site. 4. Compare parameter estimates from 1-3 for the three population study sites. Hibernation site measurements Hibernation sites will be identified by following radio-collared mice in late September and early October. Once a radio-signal remains stationary for 7-10 nights we will assume we have located a hibernaculum. The following measurements will be taken at each hibernation site: 1. 2. 3. 4. Distance ofhibernaculum to stream. Vertical elevation from hibernaculum to stream. Soil sample (as described above for the Habitat Use Study). Vegetation species richness within a 5-m radius. Density of vegetation (measured as described above for the Habitat Use Study). Composition of Seasonal Food Consumption Study Background Specific food habits are unknown for Preble's meadow jumping mouse. Armstrong et al. (1997) summarized what is currently known about food habits of meadow jumping mice in general as follows: Studies of food habits in central and eastern United States indicate they are governed by availabilitymorethan preference(Whitaker1963). Grass seeds of several species are probably the most important component of the diet, and mice will shift to those species that have available seed. Invertebrates and fungi are also readily eaten. Mice feed on both adult and larvalinvertebrates, especially Coleoptera (beetles). Invertebratefeeding is very important in the spring as mice emerge from hibernation, and may consist of half of the diet at that time. Mice also feed on various species of fungi, which are often encourtteredduring burrowing activity. As the growing season progresses, graminoidseeds dominatethe diet. There is no reason to believe food habitats of one subspecies should differ greatly from those of other subspecies. This belief is further supported by the indirect evidence available that food habits of Z. h. preblei are similar to that described above (i.e., the observation of the piles of grass sterns observed in areas where the subspecies is known to occur). As true hibernators, meadow jumping mice do not cache food in their hibernacula, Therefore, it is assumed they require high quality food for high fat accumulation prior to immergence. Preble's meadow jumping mice have been observed to increase body weight by 10% in the 2-3 weeks prior to immergence. Laboratory studies show the mouse can gain mass at rates up to 1.0 grams per day (B. Wunder, personal communication). This ability to increase fat reserves at such a rate is used by the �75 mice for preparation to enter hibernation. However, for the mice to successfully gain enough fat prior to hibernation to ensure a high probability of survival throughout hibernation, food of sufficient quality must be available. No specific information is available on what foods Preble's meadow jumping mice eat to meet these ecological requirements. However, from late August through early September, when adults begin to gain weight, there are many species of grarninoids that have available seed. In most years, grasshoppers are also readily available and probably dominate invertebrate biomass in most habitats. Objective The objective of this study is to document seasonal changes in foods consumed by Preble's meadow jumping mice. Approach Fecal samples from traps where PMJM are captured during the four trapping sessions will be collected and analyzed for composition and percentage of each discrete food type identified in the total sample. Changes in foods composition by either food type and or percent occurrence will be analyzed for spatial and temporal differences. Spatial differences will be defined as differences among the three populations studied. Temporal differences will be defined as differences among the three trapping sessions (June, July, September). Molecular Systematic Study Background The family Zapodidae (jumping mice) consists of small to medium-sized mice with enlarged hind feet and exceptionally long tails. Four living genera are recognized in this family, two of which, Zapus and Napaeozapus, are found in North America (Hall 1981). There are three living species of the genus Zapus: Z. trinotatus (Pacific jumping mouse), z. princeps (western jumping mouse), and Z. hudsonius (meadow jumping mouse). Z. h. preblei is one of twelve living subspecies of the species Z. hudsonius (Krutzsch 1954, Hafner et al. 1981). Z. h. preblei was first described by Krutzsch (1954) from a specimen collected by E. A. Preble in 1895 near Loveland, Colorado. Quimby (1951) described the species Z. hudsonius as follows: 'A mouse-like rodent with greatly enlarged hind feet and an exceptionally long tail. The forelegs are relatively short. The ears are somewhat conspicuous. The body is clothed in moderately long, somewhat dense hair of a rather coarse texture and several colors. The dorsal portions are marked by a broad stripe of brownish hairs many of which are tipped with black giving the region a grayish-black appearance. The sides are bright yellowish-orange, whereas the underparts and feet are white. The tail is bicolor, dark above and light below, and sparsely covered with hair which is longer on the terminal part. The mammae are eight, and quite prominent in lactating females. The male genitalia are inconspicuous except during the breeding season when the scrotal sac becomes enlarged. The testes enlarge and may be either abdominal, inguinal, or scrotal during this period.' The skull of Z. hudsonius is small and light with a narrower braincase and smaller molars than in Z. princeps (from Fitzgerald et al. 1994 p. 18). However, Fitzgerald et al. (1994) urge caution In distinguishing the two species of Zapus in Colorado. Such caution is further warranted by the recent conflicting identifications of mice based on genetic characteristics. Analysis of mitochondrial DNA sequence data from 92 individual mice indicates that mice sampled from southeastern Albany County, Wyoming, south along the Front Range of the Rocky Mountains to western Las Animas County, Colorado (Purgatoire Campground, San Isabel National Forest), form a coherent genetic group (Riggs et al. 1997). This group of samples are distinct from samples obtained from mice from three other populations. Genetic samples from mice captured in the Dorothey Lakes area of southern Colorado (Las Animas County) group together and are most closely allied with Z. h. luteus, a subspecies described from New Mexico (Riggs et al. 1997). The single genetic sample collected from Weld County, Colorado (Lone Tree Creek) and six samples obtained �76 from Warren Air Force Base in Laramie County, Wyoming are most similar to reference samples of Z. princeps from Colorado (Larimer County) and New Mexico (Taos County) (Riggs et al. 1997). The sequence data indicate that the samples from specimens identified as Z. h. lute us and samples from specimens of Z. princeps are more closely allied with each other than either is with the samples defining the Z. h. preblei group. This closer alliance of the samples of Z. h. luteus with Z. princeps rather than Z. h. preblei conflicts with results ofHafuer et al. (1981), based on a combination of pelage, morphologic, and genetic data, which support a closer alliance with Z. hudsonius. A more complete biosystematic evaluation of jumping mice is needed to clarify and further refine relationships among populations of the group referred to as Z. h. preblei as well as to other subspecies and species of the genus Zapus. Such an evaluation requires detailed analyses of pelage, morphometric, and genetic data from sufficient numbers of individuals to adequately represent the populations of interest. However, the mitochondrial DNA non-coding (D-loop) sequence data available at this time are consistent with the view that a geographically contiguous set of populations previously recognized as Preble's meadow jumping mouse form a homogenous group recognizably distinct from other nearby populations and from another geographically-adjacent species of the genus (Riggs et al. 1997). Therefore, given the genetic data available, Riggs et al. (1997) conclude Preble's meadow jumping mouse is a distinct population of Z. hudsonius. Objective The objective of the genetic component of this study is to document genetic variation within and among the three study site populations. Efforts are underway to secure funding for this study. Approach Genetic tissue samples will be collected from at least 10 and not more than 30 PMJM captured at each study site. A sample of at least 10 individuals from a population will provide enough information to document the genetic variation within the population, samples sizes in excess of 30 contribute little to documenting the genetic variation within the population (T. Quinn, personal communication). Genetic tissue sampling: collection of ear tissue samples using ear punch tool The following protocol for collection of genetic tissue samples has been used successfully on PMJM (M. Bakeman, C. Meaney, T. Ryon, R Schorr, personal communication). Before checking traps, ear punch tools will be cleaned by immersing them in a 10% bleach solution for a few minutes, then rinsing thoroughly with clean water. Tools will be dried thoroughly. A small screw-cap container will be useful to contain the bleach solution and to receive used ear punch tools. A sports bottle or other container with a squirt or pop-up top may be used for rinsing. Any previously used containers will be washed thoroughly by hand or in the dishwasher and not used for other purposes (e.g., drinking) while in the field. 1. A fresh pair of clean latex gloves will be used when handling each mouse. 2. With a clean ear punch tool, 2 tissue plugs will be obtained from the mouse's ear. If there is excessive bleeding, gentle pressure will be applied to the site with a small, dry piece of cotton for 1 minute. Punch may be applied to deliver the plug as a small circlet of tissue (about the size of the head of a pin) onto a finger of your gloved hand. Placement of the punch may be done in any way that is convenient. 3. Place the finger tip tightly over the end of an opened sample vial and shake to wash the tissue plug into the ethanol. 4. Repeat until all three tissue plugs have been collected into the same sample vial. Inspect the tube to see that all plugs have made it into the ethanol. 5. Close the sample vial--screw the cap on firmly and check that there is no leakage of ethanol when the tube is inverted. Label both the vial and the corresponding record on the data sheet (see item 6 below) with a.unique identifier as follows. Use a seven-place alpha-numeric code composed of the following: �77 - A three-letter designator for the survey location (e.g., WPC = West Plum Creek, GCR = Garber Creek). A number beginning with two digits indicating the year (e.g., 98) followed by 2 digits specifying the individual trapped, numbered sequentially. Example: WPC9801 6. = first animal sampled at West Plum Creek in 1998. Note: The combined seven-place alpha-numeric code needs to be a unique identifier for an individual mouse across alllocations and sites being trapped in studies coordinated by the Colorado Division of Wildlife. Record information for the individual sampled on the data sheet provided. Repetitive entries may be completed before or after the field session but do not allow much time to elapse before doing this--what is obvious to you at the time may not be so obvious later on. Information for each sample needs to be complete and unambiguous if the sample itself is to be useful. After returning from-the field, samples will be kept in a cool place or refrigerated until delivery to the CDOW office at 317 West Prospect, Fort Collins, CO where they will be kept in the freezer for analyses. E. LOCATION CDOW Maytag Property is located seven miles south of Castle Rock, Colorado along East Plum Creek. CDOW Woodhouse Property is located on Indian Creek, a tributary of Plum Creek near Louviers, Colorado. The Dupont Property is located west and south of the town of Louviers and contains the eastern stretch of Indian Creek and its confluence with Plum Creek. Pine Cliff Ranch, owned by Colorado Open Lands, is located on Garber Creek and extends eastward to its confluence with West Plum Creek, 5 miles west of Sedalia Training sessions for all field technicians and crew leaders will take place at the Frank State Wildlife Area and Kodak State Wildlife Area located in Weld County, Colorado. Data analyses and office work will be conducted at the CDOW Research Center, 317 West Prospect, Fort Collins, Colorado. F. SCHEDULE April-May April April-May May June 2-9 June-July July July 21-28 July-Aug Sep 8 -15 Sept-Oct, Sept.-Oct. December April 1999 Complete Study Plan for a demography study of Preble's meadow jumping mouse Select study sites Purchase equipment for summer 1998 field season Advertize for, hire, and train 4 technicians Conduct first trapping session Collect movement data Collect site and landscape scale habitat data Conduct second trapping session Collect movement data Conduct third trapping survey Collect movement data Collect data on possible hibernation sites Complete preliminary report for Preble's meadow jumping mouse technical working group Complete Final report for 1998 summer field season �78 F. PERSONNEL Tanya Shenk Gary C. White 2 Crew Leaders 2 Field Technicians G. Principal Investigator Statistical Consultant CD OW temporary technicians CNHP temporary technicians BUDGET Operating Expenses radio transmitters (150 @$150) radio receiver (3 @ $2565 ) pit tag scanner (6 @$600) pit tags (200 @$10.00) 3 portable communication radios & battery recharger maps computerso~are, upgrades office supplies photographic equipment, expenses field equipment misc. hardware $ 22,500 . $ 7,695 $ 3,600 $ 2,000 $ 2,100 $ 500 $ 1,500 $ 300 $ 500 $ 800 $ 500 Personnel 2 field technician for 4 months each 2 crew leader for 6 months $ 12,000 $ 30,000 30,000 miles @0.12 per mile $ 3,600 vehicle rental: 3 vehicles, 4 mo. @$120 per month overnight travel 30 days @$75 per day $ 1,440 Travel TOTAL $ 2,250 $ 75,365 �79 I. LITERATURE CITED Armstrong, D. M., M. E. Bakeman, A Deans, C. A Meaney, and T. R Ryon. 1997. Conclusions and recommendations in: Report on habitat findings on the Preble's meadow jumping mouse. Edited by M. E. Bakeman. Report to USFWS and Colorado Division of Wildlife. Bailey, B. 1929. Mammals of Sherburne County, Minnesota Journal ofMammalogy 10:153-164. Bailey, V. 1923. Mammals of the District of Columbia Proceedings of the Biological Society. Washington 36:103-138. Cook, T. D. and D. C. Campbell. 1979. Quasi-experimentation: design and analysis issues for field settings. Houghton-Mifllin, Boston. Cormack, R M. 1964. Estimates of survival from the sightings of marked animals. Biometrika 51:429-438. ERO Resources. 1995. Environmental review of South Boulder Creek Management Area Prepared for City of Boulder Real Estate/Open Space Department. Prepared by ERO Resources Crp., Denver, Colorado in association with Stoecker Ecological Consultants, Boulder, Colorado. Fitzgerald,1. P., C. A Meaney, and D. M. Armstrong. 1994. Mammals of Colorado. Denver Museum of Natural History, University Press of Colorado. Niwot, Colorado. Hafner, D. 1., K. E. Petersen, and T. L. Yates. 1981. Evolutionary relationships of jumping mice (genus Zapus) of the southwestern United States. Journal of Mammalogy 62:501-512. Hall, E. R 1981. The mammals of North America John Wiley and Sons, Inc., New York, New York, 2 volumes. Hamilton, W. J., Jr. 1935. Habits of jumping mice. American Midland Naturalist 16:187-200. Jolly, G. M. 1965. Explicit estimates from capture-recapture data with both death and immigration stochastic model. Biometrika 52:225-247. Jones, C. A 1996. Mammals of the James John and Lake Dorothey State Wildlife Areas. Final Report, submitted to the Colorado Division of Wildlife and Colorado Natural Areas Program. Kendall, W. L., and 1. D. Nichols. 1995. On the use of secondary capture-recapture samples to estimate temporary emigration and breeding proportions. Journal of Applied Statistics.22:751762. Kendall, W. L., K. H. Pollock, and C. Brownie. 1995. A likelihood-based approach to capturerecapture estimation of demographic parameters under the robust design. Biometrics 51 :293308. Kendall, W. L., J. D. Nichols, and J. E. Hines. 1997. Estimating temporary emigration using capturerecapture data with Pollock's robust design. Ecology 78:563-578. Krutzsch, P. H. 1954. North Americanjumping mice (genus Zapus). University of Kansas Publications, Museum of Natural History 7:349-472. Meaney, C. A, N. W. Clippinger, A Deans, and M. OShea-Stone. 1996. Second year survey for Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. Report prepared for the Colorado Division of Wildlife. Meaney, C. A, A Deans, N. W. Clippinger, M. Rider, N. Daly, and M. O'Shea-Stone. 1997. Third year survey for Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. Report prepared for the Colorado Division of Wildlife. Nudds, T. D. 1977. Quantifying the vegetative structure of wildlife cover. Wildlife Society Bulletin 5:113-117. Poly, W. 1., and C. E. Boucher. 1997. Record ofa creek chub preying on ajumping mouse in Bruffey Creek, West Virginia Brirnleyana 24: 29-32. PTI Environmental Services. 1996a Preble's Meadow Jumping Mouse Study at Rocky Flats Environmental Technology Site, Annual Report 1996. Final. Rocky Flats Environmental Technology Site, Golden, Colorado. �80 PTI Environmental Services. 1996b. Preble's Meadow Jumping Mouse Study at Rocky Flats Environmental Technology Site, Spring 1996. Final. Rocky Flats Environmental Technology Site, Golden, Colorado. Quimby, D. C. 1951. The life history and ecology of the jumping mouse, Zapus hudsonius. Ecological Monographs 21 :61-95. Riggs, L. A., J. M. Dempey, and C. Orrego. 1997. Evaluating distinctness and evolutionary significance of Preble's meadow jumping mouse: Phylogeography of mitochondrial DNA noncoding region variation. Final Report for the Colorado Division of Wildlife. Denver, Colorado. Seber, G. A. F. 1965. A note on the multiple recapture census. Biometrika 52:249-259. Sheldon, C. 1934. Studies on the life histories of Zapus and Napaeozapus in Nova Scotia Journal of Mammalogy 15:290-300. Shenk, T. M. 1998. Conservation assessment and preliminary conservation strategy for Preble's meadow jumping mouse (Zapus hudsonius preb/ei). Special Report. Colorado Division of Wildlife (in review). Stohlgren,T. 1., M. B. Falkner, and L. D. Schell. 1995. A Modified-Whittaker nested vegetation sampling method. Vegetatio 117:113-121. USFWS. 1997. Interim survey guidelines for Preble's meadow jumping mouse. USFWS. Denver, Colorado. Whitaker, 1. 0., Jr. 1963. A study of the meadow jumping mouse, Zapus hudsonius (Zimmerman), in cental New York. Ecological Monographs 33:3. �81 APPENDIXC STUDY PLAN State of Colorado Project No. W-153-R-ll Work Plan No._-"'0=66""'2"-Task No. _.",,2'-- _ _ Cost Center 3430 Mammals Research Conservation of Preble's meadow jumping mouse (ZaQUShudsonius preble;) Habitat use and distribution of Preble's meadow jumping mouse (Zapus hudsonius preblei) in Larimer and Weld Counties, Colorado HABITAT USE AND DISTRIBUTION OF PREBLE'S MEADOW JUMPING MOUSE preble,) IN LARIMER AND WELD COUNTIES, COLORADO (Zapus hudsonius A. NEED On May 12, 1998 the U. S. Fish and Wildlife Service (USFWS) published a final rule in the Federal Register (63 FR 26517) to list Preble's meadow jumping mouse (Zapus hudsonius preblei) as 'threatened' under the Federal Endangered Species Act (ESA) of 1973, as amended. Scarcity of suitable habitat presumably limits current distribution of Preble's meadow jumping mouse and thus, maintenance of quality habitat has been identified by the USFWS (63 FR 26517) as the principal conservation goal. Although Meaney et al. (1997) reported an improved ability to recognize suitable habitat for Preble's meadow jumping mouse, a more refined and complete definition of potentially suitable habitat for the mouse does not exist. Because the protection of potentially suitable habitat for Preble's meadow jumping mouse may occur under the ESA, as well as protection of known locations where the subspecies occurs, the definition of potentially suitable habitat, as determined by the USFWS, will directly influence site specific regulatory procedures. Ideally, a definition of potentially suitable habitat for the subspecies would identify areas where the Preble's meadow jumping mouse could survive and reproduce in sufficient numbers to sustain populations throughout its range of natural variability over an extended length of time. During the active period of the life cycle of the mouse, suitable habitat must provide requirements for daily survival, reproductive activities (breeding, nesting, and rearing of young to independence), and dispersal. The hibernation period requires habitat with sufficient food supplies to assure fat storage prior to hibernation and hibernacula sites. Habitat providing all seasonal and life cycle requirements mayor may not occur in a single contiguous area If not in a contiguous area, habitat patches must occur in a mosaic of usable areas where suitable corridors exist for seasonal movement among sites. Because very little is known about the ecological requirements of the subspecies, potentially suitable habitat is currently defined only as areas of well-developed, dense herbaceous vegetation consisting of a variety of grasses, forbs and thick shrubs in close proximity to open water. This definition is vague and possibly incomplete. Very few surveys for determining the presence or absence of Preble's meadow jumping mouse have been conducted in Larimer or Weld Counties, Colorado. Of the surveys nine conducted since 1990, only three sites yielded captures of Preble's meadow jumping mouse. Therefore, very little information is known about habitat use or current distribution of the subspecies in these two counties. Quantifying and comparing both landscape and site specific characteristics of habitats where mice are found and not found in these two counties would further our understanding of habitat use by the subspecies, particularly in the northern half of its probable range. Any new locations where Preble's �82 meadow jumping mouse are found during this study would contribute to the range-wide distribution map currently available for the subspecies. Repeated annual visits to these same randomly selected sites would also provide information on population persistence at a given site. Such a monitoring scheme would be necessary for evaluating the continued status of the mouse at each site, providing further information as to the suitability of the habitat for long-term persistence of the population. Both Larimer and Weld Counties include habitat that is currently perceived to be outside the ecological limitations of Preble's meadow jumping mouse. Larimer County provides the opportunity to explore elevationallimitations, currently thought to be 7400 feet (2260m) (USFWS 1997); Weld County extends beyond the currently believed eastern boundary of the subspecies. Both of these boundaries will be challenged by selecting survey sites within the two counties beyond the perceived ecological limits of the subspecies. Genetic tissue samples will be collected on all Preble's meadow jumping mice captured to assist identification through DNA analysis, particularly for those animals captured outside the currently perceived ecological limits, and to provide information for future studies to better define the relationships among different populations of Z. h. preblei, other subspecies of Z. hudsonius and other species of Zapus. Thus, the primary objective of this study is to quantify and define habitats used and identify ecological limitations of the subspecies in Larimer and Weld Counties, Colorado. Such information could be used by the USFWS to further refine the current definition of potentially suitable habitat for Preble's meadow jumping mouse. B. OBJECTNES Specific objectives for this study on Preble's meadow jumping mouse (PMJM) include: 1. Defme and quantify habitat characteristics at sites where PMJM was found and sites where PMJM was not found. 2. Identify elevational and eastern boundary limitations for PMJM in Larimer and Weld Counties, Colorado. 3. Describe the distribution of the PMJM in Larimer and Weld Counties, Colorado. 4. Select sites and establish monitoring protocols for determining long-term trends in populations of PMJM. 5. Collect genetic tissue samples to assure identification ofPMJM captured during this study and for future genetic analyses to better define the relationship of populations of Z. h. preblei to each other, other subspecies of Z. hudsonius and other species of Zapus. C. EXPECTED RESULTS Conservation of Z. h. preblei should include maintaining populations of the subspecies throughout the range of its natural variation and to try to identify ecological limits for the subspecies. Key concerns include range-wide distribution, habitat use, and population persistence at a given site. This study will address these concerns. Comparisons of habitat variables from both successful and unsuccessful sites will be used to better define suitable habitat for the subspecies. Habitat variables recorded will include both site level characteristics such as vegetational composition, cover, and composition of other small mammal species as well as landscape level characteristics including connectivity with other potential sites, geology, hydrology, and distance to development. Repeated annual visits to the randomly selected sites will provide information on population persistence and site fidelity of individuals at the given sites. Such a monitoring scheme will be necessary for evaluating the continued status of the mouse at given sites and provide further information on the suitability of the �83 habitat to sustain populations of PMJM over the long term. Genetic tissue samples collected from PMJM captured will be used to confirm identification of the mice and for future analyses to better define the relationship among (1) different populations of Z. h. preblei, (2) other subspecies of Z. hudsonius and (3) other species of Zapus. Performance Indicators 1. Provide a definition of and quantify habitat characteristics at sites where PMJM was found and sites where PMJM was not found in Larimer and Weld Counties, Colorado. 2. Update the known distribution ofPMJM in Larimer and Weld Counties, Colorado. 3. Provide monitoring sites and protocols for determining persistence trends ofPMJM at a given site. 4. Genetically identify all PMJM captured. 5. Complete a genetic tissue sample collection for future genetic analyses to better define the relationship of populations of Z. h. preble; to each other, other subspecies of Z. hudsonius and other species of Zapus. 6. Analyze and publish results of research. 7. Update and modify the CDOW conservation plan for PMJM. D. ApPROACH Distribution Study Background The meadow jumping mouse (Z. hudsoniusy is broadly distributed across North America from the Atlantic to Pacific coasts, extending south into the United States to Alabama and Georgia and west across the Great Plains to the base of the Rocky Mountains. In general, it is a common inhabitant of moist, grassy and herbaceous fields. Eleven living subspecies have been described (Whitaker 1972). Hafner et al. (1981) describe a twelfth subspecies, Z. h. luteus. Z. h. preble; occurs only in eastern Colorado and southeastern Wyoming (Krutzsch 1954, Long 1965, Armstrong 1972). From its limited ecological and geographic distribution, Fitzgerald et al. (1994) suggest it is an Ice Age relict, once widespread in tallgrass prairie across the eastern plains of Colorado but now restricted to scattered localities on the Colorado Piedmont. Similar relict populations of meadow jumping mice in the White Mountains of Arizona and the Sacramento Mountains and Rio Grande Valley of New Mexico are described as the subspecies Z. h. luteus (Hafner et al. 1981). Numerous surveys have been conducted since 1990 to establish the current distribution of Preble's meadow jumping mouse. Surveys have been funded by the U. S. Fish and Wildlife Service, Rocky Flats Environmental Technology Site, Colorado Division of Wildlife, U. S. Air Force Academy, Warren Air Force Base, Colorado Department of Transportation, City of Boulder Open Space, and City of Boulder Greenways Program (Armstrong et al. 1997, P. Plage, personal communication). From 1990 to 1997 such surveys have yielded captures of Preble's meadow jumping mouse, based on field identification and supported by the genetic analyses, at the following sites: 1. 2. 3. 4. Lone Pine Creek, Larimer County, Colorado Rabbit Creek, Larimer County, Colorado St. Vrain Creek and associated tributaries in Boulder County, Colorado City of Boulder Open Space, along South Boulder Creek and its tributaries, Boulder County, Colorado �84 5. 6. 7. 8. 9. 10. 11. 12. Coal Creek, Jefferson County, Colorado Rocky Flats Environmental Technology Site, Jefferson County, Colorado White Ranch Park, Ralston Creek, Jefferson County, Colorado Plum Creek drainages including Indian Creek, West Plum Creek, and East Plum Creek, Douglas County, Colorado Roxborough State Park, Douglas County, Colorado Hay Creek, Elbert County, Colorado Monument Creek on the U. S. Air Force Academy (AFAC) and tributaries of Monument Creek off the AFAC including Smith Creek, Pine Creek, and Jackson Creek, El Paso County, Colorado Medicine Bow National Forest, Albany County, Wyoming Two more sites yielded mice that were identified as Z. h. preblei in the field but were genetically found to be more closely allied with the western jumping mouse, Z. princeps. These sites include: l. 2. Warren Air Force Base, Laramie County, Wyoming Lone Tree Creek, Weld County, Colorado Similarity of the habitat at these two sites compared to those where Z. h. preblei (as determined genetically) were found suggests the possibility of areas of sympatry or parapatry between the two species of Zapus. According to Fitzgerald et al. (1994) the distributions of Z. princeps and Z. hudsonius do overlap. The area of overlap occurs in eastern Wyoming. The distribution of Z. hudson ius in Colorado is now known to be larger than shown in Fitzgerald et al. (1994). The boundaries are currently as far south as Las Animas County (based on genetically identified specimens of Z. h. prebleii. Captures of Z. princeps and Z. hudsonius (as identified in the museum) have occurred as close as eight miles of one another within the same drainage (Armstrong 1972). Z. princeps were reported captured in 1981 (Olson and Knopf 1988) at the Lone Pine site in Larimer County, Colorado, where Z. h. preblei were captured this year. Because neither specimens or genetic samples were taken in the 1981 study, identification of those mice will remain in question. The discrepancy may be explained by field misidentification or Meaney et al. (1997) also suggest this discrepancy might be explained by displacement of Z. princeps with Z. h. preblei sometime in the sixteen intervening years between trapping efforts. Although assumed to have different ecological requirements, genetic evidence presented here suggests further investigation of possible distributional overlap between Z. h. preblei and Z. princeps. Several notable results from the genetic analysis of specimens acquired from the Denver Museum of Natural History may affect the location of the southern distribution of Z. h. preblei. One site yielded mice identified as Z. p. princeps in the field but were found to be genetically more closely allied with Z. h. preblei. This site is the most southern location to date of Z. h. preblei and is located at l. Purgatoire Campground, San Isabel National Forest, Las Animas County, Colorado Also from Las Animas County were mice collected and identified as Z. h. luteus (Jones 1996), an identification also supported by the genetic analysis. These mice were from: 1. 2. Lake Dorothey State Wildlife Area, Chicorica Creek, Las Animas County, Colorado Lake Dorothey State Wildlife Area, West Fork Schwacheim Creek, Las Animas County, Colorado These sites suggest a more northerly distribution of Z. h. luteus, currently known only from limited areas in New Mexico and Arizona (Hafner et al. 1981). Identifying the southern boundary of Z. h. preblei clearly needs further study. �85 Objective Conservation of Z. h. preblei should require maintaining populations of the subspecies throughout the range of its natural variation and to try to identify ecological limits for the subspecies. Specific objectives for the distributional component of this study are to (1) clarify the distribution of the subspecies in Larimer and Weld Counties, Colorado, and (2) define and quantify the amount of suitable habitat within Larimer and Weld Counties. Approach To meet these objectives, this study will include (1) constructing a sampling frame of potentially suitable habitat within the area of interest (Larimer and Weld Counties, Colorado) and (2) conducting trapping surveys to determine presence or absence at 30 sites randomly selected from the sampling frame. Specific approaches to each of these tasks follow. Sampling frame A sampling frame will be developed to delineate potentially suitable habitat for PMJM from unsuitable habitat. To produce a preliminary map displaying existing potential habitat for Preble's meadow jumping mouse would require at least three primary layers of ecological information. Two initial layers, hydrology and vegetative cover would provide a map displaying all areas where these two ecological requirements of Preble's meadow jumping mouse co-occur. The hydrology layer must be in enough detail to provide locations of intermittent and small order streams as well as identify water source (e.g., irrigation ditches, stream, seep). Degree of density of vegetative cover is sufficient for initial mapping efforts. Demarcation of shrub, trees and grasses will suffice in these initial efforts rather than species identification. However, not all combinations of vegetative cover and water provide sufficient habitat to support Preble's meadow jumping mouse. Thus, mapping potential habitat in such a way would produce a map displaying far more potential habitat than exists. For example, an extremely critical ecological requirement for the survival of the mouse, that to date we have very little information for, is potential hibernacula sites. As a possible index to hibernacula habitat, a third GIS layer, indicating the presence of alluvial deposits may provide some insight in to hibernation requirements. Areas where alluvial deposits co-occur with the presence of water and vegetative cover may provide a more realistic map of potentially suitable habitat for the mouse. Our ability to survey a site selected at random will be restricted to those areas where we are able to obtain permission to access the land. We should be able to survey any randomly selected site on public lands and thus our inference should be unrestricted. Such readily available access will probably not occur on privately owned or leased lands, thus, restricting inferences made on private lands. By stratifying the sampling frame, this lack of access on private lands will not restrict inferences that can be made on public lands. Inference on private lands will depend on the percent of access we get from private landowners. The sampling frame will be developed from the 1:24,000 scale Hydrographic GIS layer developed by the CDOW (Reese Tietje, unpublished data). From this sampling frame a three-stage sampling design will be used to select the 30 random sites for doing PMJM surveys. Primary sampling units will be 3rd order streams. Of the 150 3rd order streams that exist within the sampling frame, 30 will be selected on public lands, 30 on private lands. The secondary sampling unit will be the stream stretch selected within the primary (3rd order stream) unit. The tertiary sampling unit will be the actual sites along the stream stretch selected at the secondary sampling stage that will be sampled. Each of the primary, secondary, and tertiary sampling units will be selected using simple random sampling. If the secondary sampling unit does not yield potentially suitable habitat (i.e., no dense vegetation) secondary sampling units will continue to be selected at random until one yields potentially suitable habitat. Determination of suitable habitat will be made by field site visits. Total stream length of all the secondary sampling units will be recorded as well as total stream length containing potentially suitable habitat. From these measurements an estimate of the probability �86 of potentially suitable habitat occurring within the primary sampling unit will be made for each area surveyed. A mean estimate, over all sites surveyed, of the probability of a primary sampling unit having potentially suitable habitat will then be made. Because of the random selection of primary sampling units, inference can be made as to the probability of potentially suitable habitat existing on all 3rd order streams within the sampling frame. By recording presence or absence of PMlM within the tertiary sampling units we can estimate the probability of PMJM occurring within potentially suitable habitat. Because of the random selection of sampling units in each of the two strata (public and private lands) we can then estimate the probability of PMJM occurring in potentially suitable habitat within both public and private lands. Trapping surveys Trapping surveys to determine presence/absence ofPMJM will primarily be conducted following protocols defined by the USFWS (1997). Additional guidelines have also been developed to accommodate the needs of this project. A brief summary of these guidelines follow. Landowner contact 1. To request permission to conduct surveys on private property, landowners will first be notified by letter. The letter will briefly describe the project and outline specifically how they might be affected if PMJM were found on their property. The letter will be followed by a phone call to determine if permission will be granted by the landowner for access to the property or not. 2. No private property will be surveyed without prior consent of the landowner. 3. To request permission to conduct surveys on public land, the agency (e.g., USFS, State Forest, etc.) responsible will be contacted. Timing of surveys 1. Trapping surveys will be conducted within the period from June 1 to September 15, 1998. These dates correspond to dates when the mouse is known to be out of hibernation. 2. Due to the nocturnal nature of PMJM, traps will be set between 1900hrs and 21OOhrs and checked as early as possible in the morning beginning at 500hrs (to reduce stress and the potential for predation on trapped animals). Time required to complete the traplines will vary depending on how many animals are caught. 3. Absence ofPMJM at a site will be defined as no captures ofPMJM from a minimum of three consecutive nights and 750 trapnights (a trap night is defined as the sum total number of traps available each night). 4. Presence ofPMJM at a site will be defined as at least one capture ofPMJM at the site. However, to accommodate sample size requirements for the monitoring and genetics study (see below) trapping efforts will continue until at least 10 individuals are tagged with Passive Integrated Transponders (PIT) tags. Survey protocols 1. Surveyors will be advised to follow the Center for Disease Control's Hantavirus instructions and recommendations when dealing with rodents which include: a Baseline blood serum samples will be collected and stored at Poudre Valley Hospital for all surveyors. b. Respirators are available for use by surveyors if they request their use. c. Surgical gloves will be used at all times when handling rodents. d. All equipment will be rinsed first with a 10% bleach solution, then water after use. e. Traps and any equipment that may have rodent feces or urine on it will not be transported in the interior of a vehicle. �87 2. 3. f. Traps will be cleaned after each use by a mouse. g. If a surveyor becomes ill with hantavirus symptoms they will report immediately to a medical facility. Surveyors must be in possession of both a federal and state collecting permit. If PMJM is found at previously undocumented sites these locations must be reported within one day to the USFWS. Survey methodology 1. Small mammal Sherman live traps (folding and non-folding) will be used to conduct the surveys. 2. Traps will be set in two parallel lines of trap stations (1 trap per station) on either side of the drainage. Trap stations will be 5 meters apart for a total of 250 meters; the parallel transects will be 10 meters apart unless extent of habitat, terrain topography, or stream hydrology do not allow. 3. Location of transects will be recorded on field data sheets (see Appendix B) and identified on 7Yz minute topographic maps. Locations will also be recorded to the nearest 5 meters in UTM coordinates using a Trimble Geo-Explorer GPS. 4. In case of windy conditions or large number of trap-tampering predators (i.e., raccoons, foxes, coyotes, etc.), traps will be secured to the ground with hoops of heavy-gauge malleable wire, stakes, or with other materials that can effectively secure/immobilize the traps. 5. A small (-1 inch) ball of polyester quilt or wool (fleece) will be placed in each trap as nestinglbedding material. 6. Baiting materials will be Manna Pro Sweet 3-way Livestock feed which contains no animal matter. Ingredients include flaked barley, flaked com, flaked oats, and cane molasses. Peanut butter will be used to stick the bait to the trap. 7. Checking of the traps will be conducted by two surveyors; one person will handle the traps and animals captured, the other person will record the data Handling of captured animals l. All animal captures will be recorded (see Appendix A, Data Forms). 1. If an animal has been captured in a trap, a ziplock plastic bag will be placed over the end of the trap. The trap will be opened allowing the animal to fall into the plastic bag. The animal will be identified to species while in the plastic bag. 2. If the animal is not a PMJM, identification of the animal will be recorded and the animal set free. 3. Each captured PMJM will be scanned to detect the presence/absence of a PIT tag. If a PIT tag is detected and the mouse has been captured at that same site within the 4-day trapping effort, identification of the mouse will be recorded and the mouse will be released. 4. If the animal is a PMJM and no PIT tag is detected the PMJM will be weighed while in the bag, recording the weight in grams using a Pesola spring balance. 5. Sex, age (juvenile, adult), and reproductive condition (pregnant, lactating or nonlactating, inactive if it is female; for males the position of the testes - scrotal, inguinal, or abdominal) will be noted. 6. Each PMJM will be measured (total length, length of body, length of hind foot - heel to distal end of claws) in millimeters. Measurements of total body and tail length will be taken with the mouse on a firm surface to minimize error. 7. Capture of every PMJM will be documented by taking two photographs of the mouse on first capture. If the mouse has been captured previously as noted by the detection of a PIT tag, no photograph needs to be taken. �88 8. 9. 10. 11. 12. 13. 14. Each PMJM will have a PIT tag inserted above the shoulder blades by lifting the skin on its back and inserting the needle with the PIT tag under their skin and injecting the tag. Verification of the PIT tag identification number will be made before insertion into the mouse by running the PIT tag scanner across it. Skin behind the opening will then be pinched to prevent emergence of the tag. Surgical glue will then be applied to the opening to prevent the PIT tag from exiting the body and prevent infection of the mouse. A PIT tag scanner will be held over the mouse to again verify PIT tag identification number and that it still works after the insertion procedure (note: PIT tag will be scanned twice during application to mouse, or once if mouse was previously PIT tagged) .. PIT tag identification number will be recorded on the field data form and the mouse released. If there are feces in the trap where a PMJM was captured, the feces will be collected in a plastic bag labeled with date and location of the site. Fecal samples will be kept cool until returned to the CDOW office where they will be frozen. Fecal samples will be analyzed by the Composition Analysis Laboratory, Inc, 622 Y2 Whedbee, Fort Collins, CO for content. If, at any time during the handling of the PMJM the animal appears to be severely stressed (dramatic changes in respiration, heartbeat or responsiveness or gums turning blue) the animal will be released immediately. Handling of trap mortalities and injuries 1. All trap mortalities will be recorded on a 'Trap Mortality' data form which includes information on species, potential duration of time spent in the trap, any information available on cause of death, etc. (see Appendix A, Data Forms). 2. All animals found dead in a trap will be double-bagged in a plastic bags and placed in a cooler with ice. Specimens should be frozen as soon as possible and deposited the CDOW freezer at the office in Fort Collins. All specimens will be sent to the Colorado State University Diagnostic Laboratory for detection of any disease and/or other conditions that may have contributed to cause of death. All PMJM specimens will be returned to the CDOW. 3. A museum card will be completed and attached to each PMJM specimen and given to Cheri Jones, Curator of Mammalogy at the Denver Museum of Natural History, for study skins and tissue storage once the Diagnostic Laboratory has completed their evaluation. 4. If an animal is severely injured it will be euthanized by soaking a cotton ball in Metofane (methoxyflurane) and placing the cotton ball and the mouse in a ziplock bag until the animal stops breathing. All specimens will then go through steps 1-3 above. 5. If an animal appears to be only slightly injured (e.g., broken tail, small laceration) the animal will be released to the wild. If an animal appears to be cold stressed attempts will be made to warm it by holding it in the surveyors hands and/or against their body. If the animal appears to be heat stressed, isopropyl alcohol will be applied with a cotton swab to the ears, arm pits, and feet to cool it down. Training of field crews 1. 2. All field technicians will be required to spend a full day at the Denver Museum of Natural History Zoology Department viewing and handling specimens of all small mammal species likely to be captured in Larimer and Weld Counties. All field technicians will be provided with a small mammal field guide and a key specifically designed to list the species most likely to be captured on the trapping sites. �89 3. All field technicians will participate in 4 days of practice trapping at the Frank State Wildlife Area and/or the Air Force Academy from May 26-29. During these trapping sessions all technicians will learn how to place, bait and set traps. Field identification will be practiced on all animals caught each day and verified by crew leaders. All field technicians will also practice handling, measuring, implanting PIT tags, and taking genetic tissue samples from deer mice (Peromyscus maniculatus) until they are competent in this procedure. Habitat Use Study Background Site scale: Based on studies of Z. h. preblei and Z. hudsonius elsewhere, Z. h. preblei apparently occurs mostly in undergrowth consisting of grasses, forbs, or both in open wet meadows and riparian corridors, or where tall shrubs and low trees form an overstory and provide adequate cover (Armstrong et al. 1997). Meadow jumping mice are widespread in abandoned grassy fields, but is often more abundant in thick vegetation along ponds, streams, and marshes or in rank herbaceous vegetation of wooded areas (Whitaker 1963). Preble's meadow jumping mice have been trapped in natural riparian areas as well as areas altered by anthropogenic influence including ditches and wetlands adjacent to interstate highways, cement-lined ditches with tall cover, ditches along driveways and moderate road use, and moderate cattle grazing (M. Bakeman, personal communication). The majority of sites where Z. h. preblei have been found consist of multistoried cover but the species composing the cover vary greatly (Armstrong et al. 1997, Meaney et al. 1997). Vegetation composition of the dense cover varies considerably and includes both native and non-native species (Meaney et al. 1996, 1997, Armstrong et al. 1997, M. Bakeman, personal communication). The herbaceous understory can be primarily grasses or forbs or a mixture of the two. Few of the sites, however, are dominated by fewer than two understory species. The tall shrub canopy at most sites is willow (Salix spp.), although scrub oak (Quercus gambelli), birch (Betula spp.), and alder (Alnus spp.) occur in sites south of the Palmer Divide (Armstrong et al. 1997). Ponderosa pine (Pinus ponderosa) is the most common tree at higher elevations. The mouse appears to tolerate weedy or exotic species in areas that are structurally diverse and species rich; nearly every successful site contained Canada thistle (Cirsium arvense) (Armstrong et al. 1997). Thus, the mouse does not appear to have an affinity toward any single plant species but instead favors sites that are structurally diverse and provide adequate cover and food throughout its life cycle. Preble's meadow jumping mouse mice are typically not found in upland areas away from riparian habitats but are most often captured where either ground water surfaces to seep springs or on main water channels (M. Bakeman, T. Ryon, personal communication) suggesting a dependence on open water, at least during their active periods. Movement of Z. h. preblei on Woman Creek at Rocky Flats Technology Site suggests mice move along corridors of shrub cover, generally Salix exigua (PTI 1996a, T. Ryon, unpublished data), suggesting dispersal habitat is similar to habitats used for other activities. Small mammal assemblage: Preble's meadow jumping mice are only one component of the small mammal community inhabiting areas where they were captured. These mice were more often found at sites with high species richness and small mammal abundance (Armstrong et al. 1997, Meaney et al. 1997). The variety and relative abundance of other small mammalian species trapped during surveys for Preble's meadow jumping mouse provide some framework for comparison of small mammal assemblages at sites where Z. h. preblei occurred and sites where none were captured. All trapped sites were either historical sites of known occurrence or apparently suitable habitat for Z. h. preblei, providing a common basis for comparison. Three small mammal species (Spermophilus variegatus, Peromyscus nasutus, and Rattus norvegicus) were not captured where Z. h. preblei occurred but were captured in unsuccessful sites. However, captures of these three species only occurred at one or two sites and the species comprised an extremely small percentage of the species �90 caught in those areas providing too little information for speculating on possible interactions between these species and Z. h. preblei. The higher occurrence and percent composition of capture of Mus musculus, the house mouse, in areas where Z. h. preblei was not caught might suggest degradation of habitat for Z. h. preblei in those areas or possibly competition between the two species (Ryon 1996). The house mouse, thrives in areas of human habitation and croplands, but also occurs in abandoned fields and ditch banks where they may displace native rodents (Fitzgerald et al. 1994). Ryon (1996) also reported the presence of domestic cats (Felis catus) at sites where Z. h. preblei historically occurred but were not found during his study. Contrarily, C. Miller (personal communication) reported house cats along South Boulder Creek where Preble's meadow jumping mice are known to occur. Landscape scale: Areas of suitable habitat must provide requirements for PMlM to survive throughout its life cycle. These requirements must provide necessities for both the active period and hibernation periods. During the active period suitable habitat must provide requirements for daily survival, reproductive activities (breeding, nesting, and rearing of young to independence), and dispersal. The hibernation period requires sufficient food supplies to assure fat storage prior to hibernation and suitable hibemacula, Habitat providing all seasonal and life cycle requirements mayor may not occur in a single contiguous area If not in a contiguous area, habitat patches must occur in a mosaic of usable areas where suitable corridors exist for seasonal movement among sites. The habitat matrix within the range of Z. h. preblei is mixed grasslands adjacent to the Colorado Front Range along the Piedmont and along the base of the Laramie Mountains in Wyoming and extends to the Colorado plains. Within this matrix, Preble's meadow jumping mice occur along stream drainages that contain patches of suitable vegetation. Suitable habitat appears to have at least two major components. The first component is a supply of open water, at least in part of the active season (M. Bakeman, C. Meaney, personal communication). Secondly, areas where Preble's meadow jumping mouse has been found have dense cover (M. Bakeman, C. Meaney, personal communication). If Preble's meadow jumping mouse behaves as a metapopulation in the classical sense [a set of local populations which interact via dispersal of individuals moving among populations and where local extinctions and recolonizations occur (Levins 1970)] then habitat includes not just one area of suitable habitation but also areas suitable for nearby mouse populations. These suitable areas must also be linked by dispersal habitat. If the mice are dependent ort dense riparian habitat for dispersal, as well as for areas to reproduce, persistence of discrete populations would require a mosaic of suitable discrete riparian patches interconnected with dispersal corridors of similarly dense riparian vegetation. If mouse populations function in a source (populations where growth rate <!: 1) and sink (those populations where growth rate < 1, maintained through immigration) system, it will be critical to identify and protect those populations serving as sources. Thus, for a source-sink population critical habitat will include those areas that support source population, dispersal habitat to sink areas will be less critical. If local mouse populations are functionally discrete, such a mosaic of interconnected areas of suitable habitat would provide a buffer for local, and source, populations against deleterious stochastic events by providing the opportunity for local population failures to be 'rescued' by immigration from other populations. Objective The objective of the habitat study is to identify and refine ecological requirements preblei in Larimer and Weld Counties, Colorado. of Z. h. Approach The general approach is to measure both site specific and landscape features of 30 sites selected at random to be surveyed for the presence of PMJM. A comparison of sites where PMJM are found and are not found will then be made in an attempt to refine our current understanding of the ecological requirements of the subspecies. The following habitat characteristics will be measured and recorded for each site. �91 Micro-site habitat characteristics 1. Cover will be measured at 20 random locations along the 250 meter sampled stream stretch. Mean cover and standard error of the mean will be estimated. Cover will be estimated using a vegetation profile board (Nudds 1977) that allows for an assessment of visual obstruction in 0.5 meter vertical intervals above ground. The board will be 1.5 meters high and 30.48 cm wide. The board is marked in alternate black and white colors at 0.5 meter intervals. Horizontal cover is assessed in each interval by viewing the board from 15 meters away in a randomly chosen direction. The percentage of each interval concealed by vegetation is recorded as 0-20, 21-40, 41-60, 61-80, and 81-100% estimated concealment. 2. Vegetation species composition and richness will be estimated using the Modified-Whittaker nested vegetation sampling method (Stohlgren et al. 1995). A modified Whittaker plot is 20meters x 50-meters with 101m2 and 210m2 subplots arranged systematically around the perimeter and 1 100m2 subplot centered in the inside of the 20 meter by 50 meter plot. Species composition and percent cover (optical estimate) of each species is recorded for each subplot. 3. Soil samples will be taken at each site for a hydrometer textural analysis (percent sand, silt and clay). Samples will be collected using a soil probe. Samples will be taken from 0-12 inches, and 12+ to 24 inches. The 0-12 inch sample will be taken first, removing the probe and soil. Sample will be placed in a labeled ziplock bag. The probe will then be placed in the same hole for the 12+ - 24 inch probe, with that soil sample being placed in a separately labeled ziplock bag. The hydrometer textural analysis will be conducted at the CSU Soils Laboratory. 4. Mean stream width will be estimated by measuring stream width at 30 locations along the entire site sampled. The 30 locations will be stratified equidistant from each other to cover the entire stream stretch in increments of site stream length/30. 5. Compare parameter estimates from 1-4 for sites where PMJM were captured and sites where PMJM were not captured. Landscape habitat characteristics The following landscape habitat characteristics will be measured using either GIS, topographic maps, or aerial infrared photography. 1. Estimate distance to nearest human habitation and human disturbance. 2. Estimate connectivity of sites to other streams from 1:24,000 topographic maps. 3. Estimate total length of stream stretch at each site and total length of stream stretch with suitable habitat at each site. 4. Compare parameter estimates from 1-3 for sites where PMJM were captured and sites where PMJM were not captured. Monitoring Study Background For purposes here, population persistence is defined as the presence of Z. h. preblei at the same site for multiple years. The majority of recent locations of meadow jumping mice have been the result of survey efforts that focused only on determining presence of Z. h. preblei. Once survey protocols (USFWS 1997) are met [i.e., a minimum of 400 trapnights (one trap set for one night = 1 trapnight) conducted] sites are typically not revisited eliminating the possibility of determining population persistence at these sites. Thus, these and other sites not yet surveyed mayor may not have persistent populations. Areas where trapping was conducted over multiple years as part of further research, yielded three sites in Colorado that support persistent populations: Rocky Flats Environmental Technology Site, the U. S. Air Force Academy near Colorado Springs, and Boulder Open Space on South Boulder Creek. Besides establishing the presence of Z. h. preblei over multiple years, other considerations of persistence include (I) presence in consecutive years versus a 'blinking' in and out of a population as would be expected in a metapopulation, (2) how populations are sustained in a given area (e.g., source or sink populations), (3) maximum abundance supportable by a given area, and (4) fluctuations in �92 abundance of a population over years. If Z. h. preblei exist in areas as part of a metapopulation or as a series of source-sink populations it would be critical to conserve and protect all key sub-population areas as well as critical habitat for dispersal. Detailed population studies, including estimating and determining factors affecting survival, reproduction, and dispersal rates, must be conducted to determine if Z. h. preblei occurs as either metapopulations or in source-sink populations. There have been no such studies conducted on Z. h. preblei or Z. hudsonius elsewhere to provide any supporting or refuting evidence for either metapopulation or source-sink structure. Consistently low abundance in a given area due to limiting ecological requirements (e.g., size of area of suitable habitat) or fluctuations in population abundance, including years of low abundance, must be considered in any conservation strategies for Z. h. preblei because of the threat of complete loss of the population due to catastrophic or extreme environmental conditions. Some evidence exists for such fluctuations in population abundance at Rocky Flats Environmental Technology Site. Trapping surveys on the Woman Creek drainage yielded the following captures of Z. h. preblei: seven in 1993 (EG&G 1993), zero in 1994 (T. Ryon, unpublished data), one in 1995 (T. Ryon, unpublished data), two in 1996 (PTI 1996a), and 33 in 1997 (T. Ryon, unpublished data). Trapping effort was consistent. from 1995-1997. Repeated trapping surveys conducted over different years intermittently yielded successful captures of Z. h. preblei along the St. Vrain Creek and lower Coal Creek (three in 1989, zero in 1992, zero in 1994, zero in 1996, one in 1997, M. Bakeman, unpublished data). If protocols in each year were the same, these data also suggest populations may undergo fluctuations in abundance. Objective The objective of the monitoring study is to establish a monitoring protocol to evaluate the status (i.e., presence or absence) of Z. h. preblei populations at specific sites and to detect changes in local distribution from year to year. Detection of PIT-tagged individuals from year to year will also provide evidence for site fidelity and/or movements of individual mice from one area to another. Approach Trapping surveys will be repeated each year at 20 randomly selected sites of those surveyed. Of the sites surveyed, monitoring sites will be established from a random selection of ten sites where no PMJM are captured and 10 sites where PMJM are captured in 1998. These 20 sites will be surveyed every year thereafter. 1. 3. 4. 5. 6. 7. Trapping surveys will be conducted as described in the Distribution Study. Habitat will be evaluated at all sites as described in the Habitat Use Study and comparisons made to results from previous years. Each PMJM captured will be PIT-tagged as per the protocols outlined in the Distribution Study. Population persistence at a given site will be evaluated as the number of years when a given site hadPMJM. Recaptures of PIT-tagged individuals will be mapped from year to year to provide evidence for site fidelity and/or movements of individual mice from one area to another. Detailed protocols for long-term data analyses will be designed once a state-wide, long-term monitoring program is fully developed. Molecular Systematic Study Background The family Zapodidae (jumping mice) consists of small to medium-sized mice with enlarged hind feet and exceptionally long tails. Four living genera are recognized in this family, two of which, Zapus and Napaeozapus, are found in North America (Hall 1981). There are three living species of the genus Zapus: Z. trinotatus (Pacific jumping mouse), Z. princeps (western jumping mouse), and Z. hudsonius (meadow jumping mouse). Z. h. preblei is one of twelve living subspecies of the species Z. �93 hudsonius (Krutzsch 1954, Hafner et al. 1981). Z. h. preblei was first described by Krutzsch (1954) from a specimen collected by E. A. Preble in 1895 near Loveland, Colorado. Quimby (1951) described the species Z. hudson ius as follows: 'A mouse-like rodent with greatly enlarged hind feet and an exceptionally long tail. The forelegs are relatively short. The ears are somewhat conspicuous. The body is clothed in moderately long, somewhat dense hair of a rather coarse texture and several colors. The dorsal portions are marked by a broad stripe of brownish hairs many of which are tipped with black giving the region a grayish-black appearance. The sides are bright yellowish-orange, whereas the underparts and feet are white. The tail is bicolor, dark above and light below, and sparsely covered with hair which is longer on the terminal part. The mammae are eight, and quite prominent in lactating females. The male genitalia are inconspicuous except during the breeding season when the scrotal sac becomes enlarged. The testes enlarge and may be either abdominal, inguinal, or scrotal during this period.' The skull of Z. hudsonius is small and light with a narrower braincase and smaller molars than in Z. princeps (from Fitzgerald et al. 1994 p. 18). However, Fitzgerald et al. (1994) urge caution in distinguishing the two species of Zapus in Colorado. Such caution is further warranted by the recent conflicting identifications of mice based on genetic characteristics. Analysis of mitochondrial DNA sequence data from 92 individual mice indicates that mice sampled from southeastern Albany County, Wyoming, south along the Front Range of the Rocky Mountains to western Las Animas County, Colorado (Purgatoire Campground, San Isabel National Forest), form a coherent genetic group (Riggs et al. 1997). This group of samples are distinct from samples obtained from mice from three other populations. Genetic samples from mice captured in the Dorothey Lakes area of southern Colorado (Las Animas County) group together and are most closely allied with Z. h. luteus, a subspecies described from New Mexico (Riggs et al. 1997). The single genetic sample collected from Weld County, Colorado (Lone Tree Creek) and six samples obtained from Warren Air Force Base in Laramie County, Wyoming are most similar to reference samples of Z. princeps from Colorado (Larimer County) and New Mexico (Taos County) (Riggs et al. 1997). The sequence data indicate that the samples from specimens identified as Z. h. luteus and samples from specimens of Z. princeps are more closely allied with each other than either is with the samples defining the Z. h. preblei group. This closer alliance of the samples of Z. h. luteus with Z. princeps rather than Z. h. preblei conflicts with results ofHafuer et al. (1981), based on a combination of pelage, morphologic, and genetic data, which support a closer alliance with Z. hudsonius. A more complete biosystematic evaluation of jumping mice is needed to clarify and further refine relationships among populations of the group referred to as Z. h. preblei as well as to other subspecies and species of the genus Zapus. Such an evaluation requires detailed analyses of pelage, morphometric, and genetic data from sufficient numbers of individuals to adequately represent the populations of interest. However, the mitochondrial DNA non-coding (D-Ioop) sequence data available at this time are consistent with the view that a geographically contiguous set of populations previously recognized as Preble's meadow jumping mouse form a homogenous group recognizably distinct from other nearby populations and from another geographically-adjacent species of the genus (Riggs et al. 1997). Therefore, given the genetic data available, Riggs et al. (1997) conclude Preble's meadow jumping mouse is a distinct population of Z. hudsonius. Objective Because of the uncertainty of the identification of the mice captured in Weld County, Colorado, and because currently perceived ecological boundaries of PMJM will be challenged in the survey effort, the objective of the genetic component of this study is to genetically identify the PMJM captured during this study. Therefore, genetic tissue samples will be collected from each PMJM captured to confirm identification of the animal. These genetic tissue samples could also provide further data to better define the relationships of different populations of Z. h. preblei as well as to other subspecies of Z. hudsonius and other species of Zapus through molecular systematic relationships and to link these genetic relationships to systematic studies of Z. hudsonius. Efforts are underway to secure funding for these studies. �94 Approach Genetic tissue samples will be collected from every PMJM captured during the study. To date, positive identification of the individual to subspecies cannot be made using only genetic information. However, results of the DNA analysis combined with the morphometric data that will also be collected (e.g., body length, tail length) and the photograph of the individual should provide sufficient information to support the field identification to subspecies. As established by the USFWS (1997) Survey Guidelines, presence of PMJM is established when only one individual is captured. Thus, we could stop our trapping efforts after the first PMJM capture. However, because we would also like to provide sufficient genetic data to more fully explore the relationships of different populations of Z. h. preblei we will attempt to collect genetic tissue samples from a minimum of 10 individuals per successful survey site. A sample of at least 10 individuals from a population will provide enough information to document the genetic variation within the population (T. Quinn, personal communication). Genetic tissue sampling: collection of ear tissue samples using ear punch tool The following protocol has been used on PMJM (M. Bakeman, C. Meaney, T. Ryon, R Schorr, personal communication). Before checking traps, clean the ear punch tools by immersing them in a 10% bleach solution for a few minutes, then rinsing thoroughly with clean water. Dry tools thoroughly. A small screw-cap container will be useful to contain the bleach solution and to receive used ear punch tools. A sports bottle or other container with a squirt or pop-up top may be used for rinsing. Wash any previously used containers thoroughly by hand or in the dishwasher and do not use for other purposes (e.g., drinking) while in the field. 1. Use a fresh pair of clean latex gloves when handling each mouse. 2. With a clean ear punch tool, obtain 2 tissue plugs from the mouse's ear. If there is excessive bleeding, gentle pressure will be applied to the injured area for approximately 1 minute. Punch may be applied to deliver the plug as a small circlet of tissue (about the size of the head of a pin) onto a finger of your gloved hand. [Placement of the punch may be done in any way that is convenient or useful as an ear mark in the context of your own work. A consistent location on one ear can help identify recaptured mice that have already been sampled.] 3. Place the finger tip tightly over the end of an opened sample vial and shake to wash the tissue plug into the ethanol. 4. Repeat until all three tissue plugs have been collected into the same sample vial. Inspect the tube to see that all plugs have made it into the ethanol. 5. Close the sample vial--screw the cap on firmly and check that there is no leakage of ethanol when the tube is inverted. Label both the vial and the corresponding record on the data sheet (see item 6 below) with a unique identifier as follows. Use a seven-place alpha-numeric code composed of the following: - A three-letter designator for the survey location (e.g., STY = St. Vrain, SCR = Smith Creek). A number beginning with two digits indicating the year (e.g., 98) followed by 2 digits specifying the individual trapped, numbered sequentially. Example: STY9801 = first animal sampled at St. Vrain in 1998. 6. Note: The combined seven-place alpha-numeric code needs to be a unique identifier for an individual mouse across all locations and sites being trapped in studies coordinated by the Colorado Division of Wildlife. Record information for the individual sampled on the data sheet provided. Be sure to include all data requested. Repetitive entries may be completed before or after the field session but we �95 recommend not allowing much time to elapse before doing this-what is obvious to you at the time may not be so obvious to us, or to you, later on. Information for each sample needs to be complete and unambiguous if the sample itself is to be useful. After returning from the field, samples will be kept in a cool place or refrigerated until delivery to the CDOW office at 317 West Prospect, Fort Collins, CO where they will be kept in the freezer for analyses. E. LOCATION Trapping surveys will be conducted at 30 sites selected at random from the sampling frame developed for Larimer and Weld Counties, Colorado. No survey site will occur at elevations greater than 7600 feet and not east of the UTM Easting Coordinate of 602000. The molecular systematic studies will be conducted in genetic laboratories using the genetic tissue samples collected in the field (ear punches). Training sessions for all field technicians and crew leaders will take place at the Frank State Wildlife Area located in Larimer County, Colorado 2.3 miles east ofI-25 on Highway 392 and 0.5 miles south on County Road 13. Data analyses and office work will be conducted at the CDOW Research Center, 317 West Prospect, Fort Collins, Colorado. F. SCHEDULE March 1998 April 1998 May 1998 May 1998 May 1998 June-September 1998 October 1998 Oct 1998-March 1999 December 1998 April 1999 April 1999 G. Purchase equipment for summer 1998 field season Complete sampling frame and select trapping sites Complete Study Plan for a distribution study of Preble's meadow jumping mouse Advertize for, hire, and train technicians Train field crews Conduct trapping surveys, collect site habitat data Conduct GIS analyses of the sites surveyed Conduct analyses Provide USFWS with a preliminary report of the results of the study Complete analyses for 1998 season. Submit Final Report for the 1998 season to the CDOW and USFWS· PERSONNEL Tanya Shenk Gary C. White 1 CDOWTFTE 4 CNHP temporary employees Principal Investigator Statistical Consultant Field Crew leader Field Technicians �96 H. BUDGET The following is a budget to conduct habitat use and distributional information for 30 sites to be completed in summer 1998. Operating Expenses PIT Tags PIT tag scanners PIT tag software genetic sampling kit expenses traps topographic maps photographic equipment, expenses laboratory analyses (fecal, soil) camera trap supplies misc. equipment (gloves, cotton, bait, etc.) $ $ $ $ $ $ $ $ $ $ Personnel 4 field technician for 3.5 months each 1 crew leader for 5 months GIS contract work Statistical consultant $ 24,000 $ 24,000 $ 4,000 $ 3,000 Travel 30,000 miles @0.12 per mile vehicle rental: 3 vehicles, 4 mo. @$120 per month overnight travel 30 days @$75 per day $ $ $ TOTAL $ 80,290 I. 3,000 1,800 300 500 2,000 500 3,500 1,200 5,000 200 3,600 1,440 2,250 LITERATURE CITED Armstrong, D. M. 1972. Distribution of mammals in Colorado. University ofK.ansas, Museum of Natural History Monograph 3:1-415. Armstrong, D. M., M. E. Bakeman, A. Deans, C. A. Meaney, and T. R Ryon. 1997. Conclusions and recommendations in: Report on habitat findings on the Preble's meadow jumping mouse. Edited by M. E. Bakeman. Report to USFWS and Colorado Division of Wildlife. EG&G. 1993. Report of Findings: 2nd Year Survey for the Preble's meadow jumping mouse. Prepared by Stoecker Environmental Consultants for ESCO Associates, Inc., Rocky Flats Environmental Technology Site, Jefferson County, Colorado Fitzgerald, J. P., C. A. Meaney, and D. M. Armstrong. 1994. Mammals of Colorado. Denver Museum of Natural History, University Press of Colorado. Niwot, Colorado. Hafuer, D. J., K. E. Petersen, and T. L. Yates. 1981. Evolutionary relationships of jumping mice (genus Zapus) of the southwestern United States. Journal of Mammalogy 62:501-512. �97 Hall, E. R 1981. The mammals of North America John Wiley and Sons, Inc., New York, New York, 2 volumes. Jones, C. A 1996. Mammals of the James John and Lake Dorothey State Wildlife Areas. Final Report, submitted to the Colorado Division of Wildlife and Colorado Natural Areas Program. Krutzsch, P. H. 1954. North American jumping mice (genus Zapus). University of Kansas Publications, Museum of Natural History 7:349-472. Levins, R 1970. Extinction. Lectures in Mathematical Life Sciences 2:75-107. Long, C. A 1965. The mammals of Wyoming. University of Kansas Publications, Museum of Natural History, 14:493-758. Meaney, C. A, N. W. Clippinger, A Deans, and M. OShea-Stone. 1996. Second year survey for Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. Report prepared for the Colorado Division of Wildlife. Meaney, C. A, A Deans, N. W. Clippinger, M. Rider, N. Daly, and M. O'Shea-Stone. 1997. Third year survey for Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. Report prepared for the Colorado Division of Wildlife. Nudds, T. D. 1977. Quantifying the vegetative structure of wildlife cover. Wildlife Society Bulletin 5:113-117. Olson, T. E., and F. L. Knopf. 1988. Patterns of relative diversity within riparian small mammal communities, Platte River watershed, Colorado. Pg. 379-386 in: Proceedings of the symposium: Management of amphibians, reptiles and small mammals in North America. Flagstaff, Arizona U. S. Forest Service, General Technical Report RM-166. PTI Environmental Services. 1996a Preble's Meadow Jumping Mouse Study at Rocky Flats Environmental Technology Site, Annual Report 1996. Final. Rocky Flats Environmental Technology Site, Golden, Colorado. Quimby, D. C. 1951. The life history and ecology of the jumping mouse, Zapus hudsonius. Ecological Monographs 21:61-95. Riggs, L. A, 1. M. Dempey, and C. Orrego. 1997. Evaluating distinctness and evolutionary significance of Preble's meadow jumping mouse: Phylogeography of mitochondrial DNA noncoding region variation Final Report for the Colorado Division of Wildlife. Denver, Colorado. Ryon, T. R 1996. Evaluation of historical capture sites of the Preble's meadow jumping mouse in Colorado, final report. MS Thesis. University of Denver, Denver, Colorado. Stohlgren,T. J., M. B. Falkner, and L. D. Schell. 1995. A Modified-Whittaker nested vegetation sampling method. Vegetatio 117:113-121. USFWS. 1997a Proposal to list the Preble's meadow jumping mouse as an endangered species. USFWS 50 CFR part 17. �98 USFWS. 1997b. Interim survey guidelines for Preble's meadow jumping mouse. USFWS. Denver, Colorado. Whitaker, J. 0., Jr. 1963. A study of the meadow jumping mouse, Zapus hudsonius (Zimmerman), in cental New York. Ecological Monographs 33:3. Whitaker, J. 0., Jr. 1972. Zapus hudsonius. Mammalian Species 11:1-7. �o o Entered Checked SMALL MAMMAL TRAPPING FORM DATE '_' _ _f _ SITE I.D (DRAINAGE). TOPO QUAD. NAME HANDLER~ _ ~TypE _ TEMPERATURE 085 NVM Tn, _ NO. OF TRAPS ~------------- COMMON NAME 'lNJI MORT l.JThffi TOTAL WT(&) l.ITMN RECORDER. SAMPLE SITE CLOUD COVER (0-8) BAG WT(&). TAlL LENGTH (mm) PAGE OF _ TIME START _ Body lAneth HIND FOOT (mm) (mnt) PRECIPITATION EAR (mm) SEX TIME FINISH _ _ REP, COND liE •• PIUKI PIT Ta,II RADIO FREQ PJM· Photo Site PboCoI COMMENTS > O~ ~~ 'TlZ o ~ ~ ~ > IQ_ \0 COMM~: _ �100 Zapus hudsonius preblei Injury/Mortality Documentation Found dead Found severely injured, euthanized Slightly injured, returned to wild Died during handling Dat~Time: _ Location: _ Weather Conditions: ___;___; _ Approximate Time Trap Set: _ TimeTrapChecked:, _ Field Technician(s) Present: Information: Species: PIT TAG Number: Weight (g): Total Body Length (mm): TruILength(mm): HilldfuotLength(mm): Ear Length (mm): Sex: Reproductive Condition(s): Descriptionofmjury: Details of Probable Reasons for Injury or Mortality: Signature of Technician(s): _ _ _ _ _ _ _ ~ _ �101 Colorado Division Wildlife Research July 1998 of Wildlife Report SEGMENT state of Project No. Work Package Colorado W-153-R-ll No. 0663 Task No. Period Covered: Author: T.D.I. Personnel: July NARRATIVE Cost Center 3430 Mammals Program Species of Special Concern/Species at Risk Conservation Kit fox Conservation 1, 1997 - June 30, 1998 Beck T. Beck, G. Bock, D. Coven, J. Garner, P. Schnurr, L. Willmarth; CDOW R. Gill, M. McLain, ABSTRACT The Program Narrative was modified to more clearly outline the multiple stages of the development of a conservation strategy. The initial phase of biological assessment was begun this segment. Sixty-three cameras, triggered by active-infra-red sensors, were deployed throughout the Uncompahgre and Gunnison valleys. Seven photos of kit fox were obtained; all are believed to be different individuals. Four of the photos were at or near dens initially located by ground searches. In addition, 2 active kit fox dens were located; one of which had 2 individuals and the other only one. Of the 3 dens with pairs present, none appeared to have young-of-the-year. All but two of the photos and sightings were in the Peach Valley-Montrose East habitat area. This isolated area of habitat is only 90 km2 in area. The Gunnison Valley area comprises 316 km2 of habitat, of which approximately 50% is deemed marginally suitable because of lack of den sites. No efforts were made to trap and collar kit fox during the whelping season. ��103 KIT FOX (VULPES MACROTIS) Thomas STATUS IN COLORADO D. I. Beck P.N. Objective Develop and implement a conservation kit fox populations in Colorado. Segment 1. Develop western a conservation Colorado. 2. Survey for presence activated cameras. 3. Capture a.nd radio-collar Gunnison valley areas. strategy and conserve Objectives strategy to recover of fox in Gunnison METHODS to recover and conserve valley area with kit fox in infra-red adult kit fox in both the Uncompahgre AND and MATERIALS The earlier version of the Program Narrative was modified to more clearly outline the progressive stages of the conservation strategy development. modified document was submitted in May 1998. The Ground searches for kit fox sign and dens were conducted in the Uncompahgre and Gunnison valleys on 40 days during April-June 1998. Remote 3smm cameras, activated by active-infra-red sensors, were deployed in areas with fresh kit fox sign, areas with old fox dens, and likely travel areas. The camera system was developed by TrailMaster (Lenexa, KS) and was essentially the same system as used by Beck on black bear (Ursus americanus) studies (Beck 1995). The distance from the camera to the infra-red transmitter was 2 m; camera height above ground was 15-30 cm. A pipe was driven into the ground 10 cm in front of the infra-red transmitter and one of several baits (beaver meat, chicken flesh lure) were placed down in the pipe to minimize bird conflict. The recorder unit was programmed so that the infra-red beam had to be broken for 0.15 sec to be recorded ( value of P = 3) and only a 6 sec delay between successive pictures. Color print film, ASA 400, in rolls of 24 and 36 exposures were used. Cameras were left at locations for varying periods, depending on area, work schedule, and need for cameras in other areas. Observations of active kit fox dens were conducted at dusk on 9 evenings, in hopes of documenting presence of pups. Observations were made with binoculars or a night-vision scope at distances of 30 to 100 m. Maps of areal extent of salt desert shrub communities were obtained from CDOW Habitat Section. Also, IS-minute quad maps were used for plotting camera locations and all fresh fox sign. �104 RESULTS AND DISCUSSION Program Narratiye Modification The revised Program Narrative describes 5 phases in the development of a conservation strategy: Biological Assessment, Socio-Economic Assessment, Political Assessment, Conservation Planning and Implementation, and Conservation Evaluation. The development of a formal Conservation strategy will occur in Phase IV. It is believed that by doing the biological, social, and political assessments as precursors to the formal plan, support for the plan will be more wide-based and solid (Clark et al. 1994). Biological Assessment Analysis of the work reported by Fitzgerald (1996) suggested that summer was the period of lowest trap success, and the majority of search effort had been conducted in the summer. Thus it was deemed prudent to resurvey much of the available kit fox habitat during spring and fall seasons, utilizing a different technology than live-trapping. Ground searches for kit fox dens and spoor were conducted on 40 days during April-June, 1998 by either one or two researchers throughout the 406 km2 of habitat in the Gunnison and Uncompahgre valleys. Only 4 active dens were located, all in den complexes previously identified in the Uncompahgre Valley. Three of the dens supported kit fox pairs while the other had a single fox. Cameras were set near 3 of the 4 dens. The den in the Montrose land fill (kit fox pair) was not equipped with a camera because of the high human traffic flow by the den. The camera at the single den malfunctioned so no photos were made. We obtained 2 photos of kit fox at each of the other 2 dens. A total of 63 camera sets were made, for periods ranging from 7 to 28 days. Thirty-five of the sets were for periods of 10-15 days. Three kit fox photos were obtained in areas where we did not find active fox dens or spoor. Subsequent searches also resulted in no active dens found. Based on camera and ground searches, 8 kit fox were located in the Uncompahgre Valley and 2 in the Gunnison Valley just west of the Delta airport. Photos were obtained from all active den sites where cameras operated correctly; thus supporting the use of cameras as an effective survey tool. Based on 9 evening observations and 8 daytime examinations of den sites not appear that any of the 3 kit fox pairs produced any pups in Spring, Four natal dens of red fox (Vulpes vulpes) were located in the saltbush habitats in what appeared to be suitable kit fox habitat. it did 1998. The camera surveys also produced photos of the following: badger (3), house cat (3), dog (1), raccoon (7), cottontail rabbit (76), prairie dog (3), antelope ground squirrel (16), striped skunk (3), domestic cattle (6), magpies (44), horned lark (9), meadow lark (1), and a snake (1). Sixteen cameras produced no photos, 3 had mechanical malfunctions, and 2 were disturbed and made inoperable by people. The Uncompahgre Valley area is comprised of approximately 90 km2 of suitable kit fox habitat; arranged in an oblong pattern 30 km N-S with an average width of less than 4 km. Eight of the observed kit fox were in this area. This area is separated from the Gunnison Valley habitats by agricultural land, housing developments, a highway, and a river; resulting in a 7-9 km long obstacle course to inhibit dispersal. Based on the range of kit fox densities summarized by White and Garrott (1997) (0.16-0.7jkm2) this area of habitat �105 would likely only this lower bound. support 14-63 kit foxes. Current density appears to be near The Gunnison Valley area is approximately 316 km2 of apparently suitable kit fox habitat. Only 2 kit fox were located in this region; both at the same camera and likely a pair. They were at the southern tip of the habitat area. Approximately 50% of the habitat is characterized by a volcanic rock cap which severely limits den sites. No old kit fox dens were located in these formations and only a few badger and coyote dens. It appears that the lack of old dens limits the value of this region. The northernmost section of possible habitat in the Gunnison Valley lies between Kannah Creek and the Colorado River. It is approximately 74 km2 in area but separated from the upper valley by extensive irrigated farm lands and new housing developments. No fresh or old kit fox sign was discovered in this area. It was decided not to trap and collar adult foxes during the whelping season because of the added stress of this activity to the individual foxes (Cypher 1997). The small number of kit foxes present seems to justify minimum intrusion during this period. Literature Cited Beck, T.D.I. 1995. Development of black bear Wildlife, Fed. Aid. Rep. W-153-R-8. llpp. Clark, T.W., R.P. Reading, A.L. Clarke. Recovery:Finding the Lessons, Improving Washington, D.C. 450 pp. inventory techniques. colo. Eds. 1994. Endangered Species the Process. Island Press, Cypher, B.L. 1997. Effects of radiocollars Wildl. Manage. 61(4):1412-1423. on San Joaquin Kit Foxes. J. J.P. 1996. Status and distribution of the kit fox (Vulpes in Western Colorado. Colo. Div. Wildlife, Final Fed. Aid. Rep. W78 pp. Fitzgerald, macrotis) 153-R-7. White, P.J. and R.A. Garrott. Can. J. Zool. 75:1982-1988. Prepared Div. 1997. Factors by Thomas Beck Wildlife Researcher regulating kit fox populations. ��107 Colorado Division Wildlife Research July 1998 of Wildlife Report JOB PROGRESS REPORT state of Project Colorado No. Work Package Cost Center W-1S3-R-11 No. __~0~6~6~3,- Manunals Program _ Author: Species of Special Lynx Conservation Task No. Period 3430 Covered: July Concern Recoyery 1, 1997 - June 30, 1998 D. F. Reed ABSTRACT A conservation strategy/recovery plan for lynx in Colorado was prepared and the draft revised January 12, 1998 (Chapter 1 in Seidel et al. 1998). This conservation strategy to recover and conserve lynx populations in Colorado provided general background on the species and generally identified the habitat goals and objectives of identifying potential habitat, identifying linkage zones, selecting suitable habitats, refining habitat suitability models, and refining habitat protection, as well as, conservation actions to be taken. Subsequently, more specific protocols were developed to assess potential lynx habitat based on the species' primary prey, the snowshoe hare.· The first protocol was for the winter track survey which was conducted during the winter. The second was for the snowshoe hare pellet counts (Krebs plots) which were begun toward the end of the segment. The routes selected for the winter track surveys were based upon accessibility and transportation mode (snowmobile, cross-country skis, etc.), and hence, were not randomly selected. Furthermore, sample sizes (routes per forest types) were small. Conclusions about representativeness should be made with caution. ��109 LYNX CONSERVATION/RECOVERY Dale F. Reed P.N. OBJECTIVE Develop a conservation Colorado. strategy to recover SEGMENT 1. Prepare 2. Assess a conservation potential and conserve in OBJECTIVES strategy/recovery habitat lynx populations plan for lynx in Colorado. for lynx reintroduction. STUDY AREA The study area includes the forests (Aspen, Douglas Fir, Lodgepole Pine, mixed conifer, mixed forest, Ponderosa Pine, and Spruce-fir) and deciduous oak above 7,500 ft throughout the north-south western half of Colorado. METHODS AND MATERIALS The methods used in preparing the "Conservation Strategy/Recovery Plan" (Seidel et al. 1998) involved extensive literature review, writing, rewriting, editing, and coordination with the numerous authors, species experts, and agencies participating in its completion. The methods used in assessing potential habitat for lynx (Felis lynx) involved 1) a winter track survey where snowshoe hare (Lepus americanus) tracks (crossings and parallel movements) were counted in snow 24-28 hours after snowfalls (described by Byrne 1998) and 2) counts of snowshoe hare pellets (Kreb's plots; Krebs et al. 1987) via randomly selected points in the forest types and deciduous oak (see Study Area) as determined by statewide GAP data. RESULTS Results as in the "Conservation Strategy/Recovary Plan" are summarized following Executive Summary (Seidel et al. 1998:vii-x): EXECUTIVE in the SUMMARY Endangered Species Since life began on this planet many species have come and gone through natural changes in physical and biological conditions. Since these extinctions occur naturally why should we spend money and effort to conserve species that are nearing this end? How do they benefit society if restored? Congress addressed these questions in the preamble of the Endangered Species Act of 1973, "recognizing that endangered species of fish, wildlife, and plants are of aesthetic, ecological, educational, historical, recreational, and scientific value to the Nation and its people." Congress further stated its intent to protect and conserve the ecosystems and habitats. The State of Colorado �110 has adopted similar protections for species of special concern. The primary force driving the loss of these species is habitat destruction or human exploitation. While we do not know the causes for the decline in lynx (Lynx canadensis) in the Southern Rocky Mountains (SRM) human activity is at least partly responsible. We have the knowledge to conserve and reestablish these species. We also have the responsibility. History and Distribution Lynx historically occurred, at low densities, as forest carnivores in the SRM. There have been 12 investigations reported in Colorado since 1979 to document the presence of lynx or wolverine. Although the presence of both species has been suspected but not confirmed, and no viable populations have been found. There have been no field studies in the southern ranges in Wyoming or in New Mexico. Wyoming conducted a survey in 1986 of reported sightings but few of these reports were documented (Reeve et al. 1986). This draft does not include information on the Wyoming and New Mexico portions of the SRM. The final strategy will include a coverage of these areas. Any future decisions regarding lynx that affect neighboring states will require further investigation and coordination. Current Status At this time the lynx is classified as a federal "candidate" species by the Fish and Wildlife Service. In Colorado the lynx is classified as state endangered species and in New Mexico are protected species. Wyoming classifies lynx as Native Species Status Category II (Oakleaf et al. 1996). The Forest Service classifies lynx as a "sensitive species." Conservation strategy On July 8, 1997, representatives of the u.S. Forest Service, u.S. Fish and Wildlife Service and the Colorado Division of Wildlife (CDOW) met to discuss a cooperative program for the conservation and reestablishment of lynx and wolverine in Colorado. On August 4, 1997, those agencies plus the National Park Service signed a letter agreeing to jointly prepare "A Candidate Conservation Strategy for Lynx and Wolverine in Colorado." The Colorado Division of Wildlife is the lead agency for these state listed species. Species Goal The goal of this strategy is the conservation and reestablishment of lynx within their former ranges by establishing populations which reprodu~e sufficiently to allow emigration into unoccupied suitable habitats. If this is accomplished, it would permit the downlisting of these species by the states from endangered. Both species are considered in this document due to expected economies of scale that will be realized in the habitat assessment, acquisition, and monitoring portions of this strategy. This document could serve as the basis for future recovery efforts in Colorado. Risks to the Species We can only speculate on the history of lynx populations in Colorado. Some speculate that trapping, hunting, and poisoning played a significant role in population reductions. If so, Amendment 14 approved by the voters of Colorado in 1996 greatly reduced that threat to future populations because it strongly restricts the use of poisons, leghold, kill-type and snare trapping devices in the State of Colorado. Removal of this historical source of mortality would enhance the probability of success for conservation and/or potential future reintroductions of lynx. A second primary mortality factor for lynx populations reported from other sites within their distribution is mortality of kittens or kits either through starvation or disturbance during rearing. �111 These are factors that Most of the potential land with the majority lands in Colorado are not known how much of can be addressed by specific management practices. habitat for these species in the SRM occurs on public on National Forest System lands. Many National Forest designated and managed as wilderness. It is currently that area has potential to support lynx populations. Habitat Goals for Lynx The first goal is to describe and map potential habitats for lynx in the SRM. The second goal is to protect those habitats that may be important for whatever lynx may still exist in the state and potential habitats for future reintroduction or reestablishment efforts. Habitat Evaluation Actions for Lynx Lynx foraging habitat requirements are inextricably linked to habitat requirements and population distribution of snowshoe hares (Lepus americanus) because that species is a staple prey item. Therefore habitats with abundant snowshoe hare populations would be considered the primary criterion for selecting potential locations for lynx reintroductions. The first step in the strategy will be to assess the potential habitats both for foraging and denning potential using GIS vegetation mapping. Key areas will be identified that contain at least the minimum requirements for species survival. The criteria used for this selection will come from the significant body of knowledge that exists for lynx habitats in other regions (Table 1). These same criteria have been used to develop potential habitat management guidelines that will be recommended to land managers until information obtained from monitoring reintroduced individuals suggests changes to those recommendations. Once key blocks of potential habitat are identified, the CDOW intends to conduct two surveys to detect snowshoe hare occurrence and estimate densities to validate or adjust initial habitat assumptions. Data from both surveys will be used to identify the 2 habitats with the greatest potential to support Colorado lynx. The actual reintroduction sites will be identified from the field surveys and modified or confirmed from advice from a peer review panel of qualified scientists with recognized expertise in lynx ecology and management. Reestablishment of lynx Reestablishment of viable lynx populations in Colorado requires reintroductions. A reintroduction would involve assessing potential release sites for habitat suitability, radio-marking lynx prior to release, and intensive monitoring of radio-marked animals. Lynx populations in Canada and Alaska currently are increasing toward peak population levels (Fontana pers. commun. 1997). Lynx selected for reintroduction otherwise would be killed and sold as pelts to be traded as fur. By paying a compensatory price, the CDOW can obtain lynx for reintroduction at a reasonable price and increase their chances of survival. When lynx are at the lower end of their population cycle, population levels would be insufficient to provide sufficient animals for reintroduction. Since wild lynx populations cycle every 10 to 12 years, there is a narrow 4- to 5-year window of opportunity to reintroduce lynx in Colorado's most suitable habitats. It is estimated that lynx would be needed to be released each year ( in each of 2 habitats each year) for 2 successive years to establish several viable breeding populations. Species Monitoring Radio-marked lynx will be intensively monitored by frequent relocations both from the air and the ground. Recorded data would include habitat selection patterns, seasonal home ranges, reproductive success, survival, and probable cause of death when mortalities are detected. This information would be �112 evaluated critically to determine if additional releases of these species into unoccupied habitats is warranted before additional releases are attempted. Revision Any strategies for conserving lynx in the SRM must be regarded as tentative because very little is known about their specific ecology and habitat requirements in the SRM. As the body of knowledge increases and new information is gained from post-release monitoring of both lynx and wolverine, the Conservation strategy would be revised. Formal lynx and wolverine status reviews would be scheduled periodically (e.g., annually) by the Lynx and Wolverine Conservation strategy Team to revise the lynx/wolverine Conservation Strategy as needed. Budget A draft budget is included in Appendix B. It is estimated that the cost to complete the first 3 years of this Conservation strategy could approximate $2.5 million. The Division of Wildlife has committed to funding $700,000, leaving $1.8 million to be acquired from other sources. Habitat Protection Following the pre-release habitat suitability surveys, it will be necessary to develop interim guidelines to delineate, preserve and protect lynx habitats on public lands that are deemed critical to the conservation of these species. These interim habitat protection guidelines periodically would be revised as additional information surfaces from the post-release monitoring of radiomarked individuals. Guidelines will be based on the best scientific and economic information available; will conform to existing laws and regulations; and will be subjected to further peer and public review prior to implementation. These guidelines will be developed and amended into this document at a later date subject to concurrence by all signatories. Results of the winter track surveys were reported by Byrne (1998). Counts of snowshoe hare pellets (Kreb's plots; Krebs et al. 1987) were only begun toward the end of this segment - hence the results from this second method will not be available until the next segment. LITERATURE CITED Byrne, G. 1998. A Colorado winter track survey for snowshoe hares and other species. Colo. Div. Wildl. 35pp. Krebs, C. J., B. S. Gilbert, S. Boutin, and R. Boonstra 1987. Estimation of snowshoe hare density from turd transects. Can. J. zool. 65:565-567. Seidel, J., B. Andree, S. Berlinger, K. Buell, G. Byrne, B. Gill, D. Kenvin, and D. Reed. 1998. Draft strategy for the conservation and reestablishment of lynx and wolverine in the southern Rocky Mountains. U.s. Forest service, National Park Service, U.S. Fish and Wildlife Service, and Colorado Division of Wildlife. 115pp. prepar;~:;:Z)Ai0L~ Dale F. Reed Wildlife Researcher �113 Colorado Division Wildlife Research July 1998 of Wildlife Report JOB PROGRESS State of Project Work Colorado Cost Center No. -DW~-~1~5~3~-~R~-~1~1L- _ Package No. ~0~8~8~0~ Author: 3430 Mammals. Research _ Black-footed Task No. Period REPORT Ferret Recoyery Monitoring and Managing Black-footed Ferrets Covered: July M. A. Wild Personnel: Disease in 1, 1997 - June 30, 1998. and K. T. Castle E. Wheeler. ABSTRACT We prepared a Program Narrative describing research to support black-footed ferret (Mustela nigripes) recovery efforts in Colorado. The black-footed ferret is a federally listed endangered species in the United States. Blackfooted ferrets have been extirpated from Colorado, but are scheduled to be reintroduced to the wild at the Little Snake Management Area (LSMA) in Moffat County, Colorado in 1999. Our research can be sub-divided into two broad sections: disease monitoring and flea control as a tool to manage sylvatic plague in prairie dogs and black-footed ferrets. Disease monitoring will be performed using collection of carnivores (primarily coyotes) from the LSMA twice per year. Carnivores will be examined post-mortem for signs of active disease and serology will be performed to test for exposure to diseases potentially threatening black-footed ferrets. Results of testing in FY97/98 show that toxoplasmosis and tularemia are present at LSMA, but potential impact on black-footed ferrets is unknown. Two potentially devastating diseases, canine distemper and plague, are also present at LSMA. Although prevalence of titers to canine distemper virus (CDV) in coyotes appears low at LSMA, an increasing trend 'in prevalence is suggested., Titers to plague (Yersina pestis) were found in 65\ and 48\ of coyotes tested in the summer and winter collections, respectively. Titers were present in both adult and juvenile animals, suggesting ongoing plague activity in some areas. The second section of our Program Narrative describes experiments with captive white-tailed prairie dogs to evaluate insect growth regulators (IGRs) on control of prairie dog fleas (genus Oropsylla). Preliminary work to establish a captive colony of >30 white-tailed prairie dogs and to develop techniques to rear Oropsylla in an insectary have been initiated this fiscal year. ��115 MONITORING AND MANAGING Margaret DISEASE IN BLACK-FOOTED A. Wild and Kevin FERRETS T. Castle P. N. OBJECTIVES 1. Monitor ferrets enzootic disease activity threatening reintroduced into the LSMA. 2. Write a Final Program Narrative describing evaluate management techniques to minimize reintroduced black-footed ferrets. SEGMENT survival of black-footed an experiment to develop and disease-related mortality of OBJECTIVES 1. Determine the level of activity of canine distemper virus, Aleutian disease, toxoplasmosis, tularemia, and leptospirosis in carnivores and plague in prairie dogs (using coyotes as sentinels) in the LSMA by sampling ~20 coyotes in summer and winter. 2. Write a Final Program Narrative describing evaluate management techniques to minimize reintroduced black-footed ferrets. METHODS Carnivore Disease an experiment to develop and disease-related mortality of AND MATERIALS Survey Infectious diseases can severely impact the success of black-footed ferret (Mustela nigripes) reintroduction efforts. As part of the black-footed ferret reintroduction protocol, we monitored disease activity in carnivores in the Little Snake Management Area (LSMA), Colorado. Coyotes (Canis latrans) were collected during a summer and winter sampling period. Post-mortem examination and sample collection was performed as described in the Planning Program Narrative (Wild 1997). In addition to serologic assay for plague (Yersina pestis) using the passive hemagglutination inhibition test (PHA/PHI), sera collected in the winter sampling were also tested using the plague ELISA test. Proposed Research Experiments We compiled data and performed literature searches and interviews to prepare Program Narrative. We initiated work on Phase I--Pilot Study as per the Program Narrative and captured white-tailed prairie dogs using techniques described in the attached protocol (Attachment 1). Black-footed Ferret Reintroduction We assisted in preparation of the Black-footed ferret allocation request submitted to the US Fish and Wildlife Service by the Colorado-Utah blackfooted ferret recovery working team in 1997 and 1998. a �116 RESULTS Carnivore Disease AND DISCUSSION Survey with the assistance of USDA Wildlife Services and the Bureau of Land Management (BLM) we collected 20 coyotes from the LSMA between 21-25 July 1997 and 21 coyotes between 10-11 February 1998. Coyotes were collected using a combination of calling and aerial gunning. Death occurred rapidly and the collection technique was adequate, but much less efficient in summer than when used in the winter months. Coyotes were collected from various locations in the >4700 mi2 management area; however, we focused on the Powderwash area and near the site of the pre-conditioning pens. No lesions indicative of active disease were noted on gross examination of carcasses. We found no serologic evidence of exposure to leptospirosis (serovars canicola, grippo, hardjo, ictero, and pomona). Coyotes were not tested for Aleutian Disease (a mustelid disease). Serologic titers were positive to toxoplasmosis in two coyotes (one' adult, one juvenile) collected in the winter sampling. Fifty percent of the coyotes sampled during the summer had positive titers to tularemia, while <5% of those sampled in winter were positive. Positive titers could have been associated with predation on rabbits, prairie dogs, or other rodents or exposure to ectoparasites. Duration of titers to tularemia are likely of short duration «6 mo). The impact of tularemia or toxoplasmosis on blackfooted ferrets is not currently known; however, ferrets and humans are susceptible to these diseases. Although a relatively small proportion of coyotes had positive titers (~1:16) to canine distemper virus (CDV), data collected over the last 18 mo suggest a trend toward increasing prevalence of CDV at the LSMA (Fig. 1). Because CDV is a serious disease threat to blackfooted ferrets, a priority in the coming year will be to monitor the trend of CDV prevalence. Likewise, plague activity on the LSMA is of concern. In the summer collection, both juvenile (7/13) and adult (5/7) animals showed evidence of exposure to plague (titers >1:8; Fig. 2). Seropositive juveniles indicates that some active plague is continuing in the area. Samples collected in the winter were tested using the standard PHA/PHI test and also an ELISA test. The ELISA test may be more specific than the PHA/PHI test; however, it has not been fully validated and accepted (H. Edwards, pers. commun.). Results of the standard PHA/PHI test indicate about half (10/21) of the animals samples had positive plague titers (Fig. 2). Alternatively, results of the ELISA test suggest that 81% (17/21) of animals samples were positive. Anecdotal evidence suggests that localized outbreaks of plague may be occurring in some areas of the LSMA, while plague activity may be lesser in the other areas (M. Albee, pers. commun.). This localization of plague activity is not uncommon. Proposed Research Experiments Given the results of these initial surveys, plague appears to be the most significant disease threat currently to black-footed ferrets reintroduced into the LSMA. Further, it is now more apparent that research emphasis needs to be placed on management and control of plague in prairie dogs and black-footed ferrets. The Program Narrative describing this research is attached (Attachment 2). To support this research, we captured 34 white-tailed prairie dogs from the Arapaho National Wildlife Refuge (ANWR), Colorado. Initially, four adult males were captured in April to evaluate methodology. Thiry juveniles were then captured in June. Spring/summer capture is required because white-tailed prairie dogs hibernate. Further, juveniles can be easily differentiated from �117 adults in early summer (June). One prairie dog died from trauma at FWRF, but all prairie dogs remained healthy during the quarantine period. The housing and care protocol (see Program Narrative) appears sufficient for maintaining prairie dogs. The white-tailed prairie dogs are more fractious than anticipated; however, training continues in an attempt to habituate the animals to handling. We also initiated investigation of maintaining prairie dog fleas (Oropsylla spp.) in artificial insectaries. Flea mortality has been high immediately after capture from ANWR and transport to Fort Collins. We will continue to modify our protocols to increase survival. Fleas that survive transport have been successfully maintained in the rearing medium. pinky (neonatal) mice are provided to the fleas for a 24-hr period every 2-3 days as a blood source. Fleas have been observed to lay eggs, but the numbers (a few/day) are, as expected, low. Black-footed Ferret Reintroduction The U.S. Fish and Wildlife Service approved an allocation of black-footed ferrets for reintroduction into the LSMA in fall 1997; however, due to delay in publishing the final rule on experimental designation of the joint Colorado-Utah reintroduction sites, black-footed ferrets were not received this year. The Colorado-Utah site has received conditional approval for allocation of 20 black-footed ferrets later in 1998. LITERATURE CITED Wild, M. A. 1997. Monitoring and managing disease in black-footed ferrets. Colorado Div. Wildl. Res. Rep., 0880-1, Jul 1996 - Jun 1997, Fort Collins. Prepared by Magaret A. Wild Wildlife Researcher ��119 ATTACHMENT PROTOCOL Prepared 17 March by: Kevin 1998 T. castle, 1 FOR LIVE-TRAPPING PRAIRIE DOGS M.S. The following procedures are based on my personal experience with trapping a variety of mammals, and on the prairie dog trapping experiences of Hoogland (1995) and A. Schwartz (pers. commun.). Hoogland reported only 12 trap-related deaths in over 10,000 captures, and Schwartz observed only 5 deaths in approximately 1,700 captures; the average mortality rate is approximately 0.1%, so trapping mortality is unlikely if these procedures are followed. If an animal is severely injured (fracture, severe laceration), such that its ability to survive is markedly impaired, or it would suffer undue misery and stress, it will be euthanized. Euthanasia will be performed using an overdose of inhalant anesthetic, by placing the prairie dog in an induction chamber with a halothane-soaked gauze. If two trapping-related deaths occur, trapping will cease and the protocol will be reviewed and modified. Minor injuries or metabolic imbalances will be treated by, or under the supervision of, a veterinarian. One or two single or double-door Tomahawk traps (6" x 6" x 24") are placed near active burrows; single door traps seem to selectively capture juvenile prairie dogs, whereas double-door traps will capture both juveniles and adults. Traps are baited with rolled oats and/or "horse sweet mixture"; horse sweet mixture is sticky, and therefore tends to stay in place, rather than roll or blow away. During a 1-2 week "prebaiting" session, the traps are wired closed (to prevent unwanted, accidental entry), and bait is spread around the traps and burrows. Prebaiting allows the prairie dogs to become accustomed to having strange objects in their environment, and lets them associate the traps with the presence of food. In addition, the traps "weather" and take on a more natural scent; this is especially important for new, unused traps that may smell of oils used in the manufacturing process. After the prebaiting session, traps are set (opened and baited) just prior to sunset, and checked (minimally) at noon and at dusk the following day. It is better to set the traps at night rather than in the morning, to avoid disturbing any animals that become active early. Once disturbed, it may take from 15 minutes to several hours for an animal to re-emerge from its burrow, and any other animals may become more wary. Prairie dogs are diurnal, and are typically not seen above ground in inclement weather, so it is unlikely that any will be captured at night or when temperatures are very cold or very hot. Nonetheless, traps are not set during rain or snow, or when daytime ambient temperature is expected be below 20°F or above 90°F. Prairie dogs should be handled by experienced mammal trappers, wearing heavy leather or welder's gloves. Upon capture, each animal is placed into a coneshaped nylon or canvas bag for non-chemical immobilization. They are sexed and weighed, and age is determined (Hoogland 1995; Cox and Franklin 1990). A numbered monel ear tag is attached to each animal. Released animals should be tagged also, to minimize the handling of animals recaptured at a later date. Animals are then transferred to individual mesh-bottom holding cages (minimum size = 18" x 18" x 18"), and the cages are covered for transport. At least part of the holding cage should remain covered at all times, to provide a hiding place for the animal; if possible, a nest box containing bedding material (e.g. "Bed-o'-cobs", Care Fresh, hay) should be provided. Rodent chow, �120 alfalfa cubes, and water are provided ad lib. if transport/holding time will exceed 2 h. Animals held in the field should be kept indoors if possible, but minimally need to be protected from the elements and from potential predators. They should be checked every 3-4 h the first 1-2 days after capture, to assess health and acclimation to captivity, and at least once per day thereafter. Literature Cited: Hoogland, J. L. 1995. animal. University The black-tailed pra~r~e dog: social of Chicago Press, Chicago, 557 pp. life of a burrowing �121 ATTACHMENT PROGRAM state of Project Work No. Package NARRATIVE Colorado Cost Center W-153-R Mammals No. __~0~8~8~0~ Task No. A. 2 _ 3430 Program Black-footed Ferret Conservation Monitoring and Managing in Black-footed Ferrets Disease NEED The mission of the Colorado Division of Wildlife (CDOW) is to "perpetuate the wildlife resources of the state and provide people the .opportunity to enjoy them" (state of Colorado, 1994). Further, the CDOW Long Range Plan established a goal directing the Division to "cooperate with federal, state, county and local government agencies, private landowners and other organizations in evaluating the potential for restoration of extirpated species" (state of Colorado, 1994; Goal 8). The black-footed ferret (Mustela nigripes) is a federally listed endangered species in the United states. Black-footed ferrets have been extirpated from Colorado, but are scheduled to be reintroduced to the wild at the Little Snake Management Area (LSMA) in Moffat County, Colorado in 1999. Coyote Basin, Utah and White River Resource Area, Colorado are also under consideration for blackfooted ferret reintroduction efforts in the near future. The overall goal of ferret reintroduction is to contribute to the national black-footed ferret recovery plan objectives (U. S. Fish and Wildlife service, 1988). The specific reintroduction objective for LSMA is to sustain a natural breeding population of 30 adult black-footed ferrets. Success of ferret reintroduction programs in Wyoming, South Dakota, and Arizona have been hampered by multiple factors. Mortality rates, primarily from predation and disease, have been high (D. Biggins, pers. commun.; P. Marinari, pers. commun.). It has become apparent that improved methods for preconditioning ferrets prior to release, predator management, and disease management are prerequisite to success of future reintroduction efforts. Here, we will begin investigating the potential for impact of diseases on black-footed ferrets in the LSMA reintroduction site. Disease, primarily canine distemper and sylvatic plague, has been a significant factor limiting captive as well as free-ranging black-footed ferret populations. Mortality of black-footed ferrets from infection with canine distemper or sylvatic plague is extremely high (Williams et al., 1988; Williams et al., 1994). Further, mortality from sylvatic plague in prairie dogs can markedly impact prey base for black-footed ferrets. Before black-footed ferrets are reintroduced to the site, data on these potentially devastating diseases must be collected and unique plans for their management devised. B. OBJECTIVES 1. Monitor ferrets enzootic disease activity threatening reintroduced into the LSMA. survival of black-footed �122 2. C. Develop techniques to manage plague in the LSMA using insect growth regulators applied orally to prairie dogs. EXPECTED RESULTS AND BENEFITS Disease Monitoring Improved techniques for disease management will likely be required for the successful reintroduction of the extirpated black-footed ferret to Colorado. We must monitor enzootic disease activity to determine immediate risks to black-footed ferrets. Monitoring will also provide information on the epizootiology of diseases that will assist in formulating management plans for black-footed ferrets and may also aid in understanding population dynamics of other inhabitants. of the short grass prairie. Performance Indicator: Monitor disease status of ~40 carnivores in the LSMA annually and based on these findings make disease management recommendations the Colorado-Utah black-footed ferret recovery working team. to Flea Control We believe that the proposed research will provide a novel and effective means of controlling plague in prairie dogs and black-footed ferrets. Sylvatic plague is transmitted primarily via the bite of infected fleas. Control of flea populations can be used to break the cycle and reduce the occurrence of disease. Products, such as lufenuron and pyriproxyfen, administered orally to pet animals have effectively controlled fleas (Blagburn et al, 1994; Hink et al., 1994; Palma et al., 1993). If these products are efficacious and safe in prairie dogs, and to black-footed ferrets which consume treated prairie dogs, then they could be used to control fleas, and plague, at reintroduction sites. Further, these products could be used as an alternative method of wildlife management in areas where sylvatic plague is a threat to public health. Performance Indicator: Provide a novel technique for the management in the LSMA to protect the endangered black-footed ferret. D. of plague APPROACH Disease Monitoring Rationale: Infectious diseases can severely impact the success of black-footed ferret reintroduction efforts. Canine distemper and sylvatic plague pose the greatest threats to black-footed ferret survival (Williams et aI, 1988; Williams et aI, 1994). Wild canids and mustelids, as well as domestic dogs, are susceptible to canine distemper and can serve as reservoirs for infection (Williams, 1982). Plague is maintained primarily in rodent populations, such as prairie dogs (Quan, 1982). Therefore, prior to reintroduction, status of diseases such as canine distemper and plague in sympatric wildlife populations must be determined to assess the risk to black-footed ferrets. We will monitor activity of canine distemper in carnivores, primarily coyotes (Canis latrans), by determining serologic titers and performing postmortem examination. Titers are useful in ascertaining if carnivores have recently been exposed to the virus. Postmortem examination is required to determine if carnivores have active infection with canine distemper. We will monitor plague activity in prairie dogs using coyotes as sentinels. Although coyotes are rarely involved in plague transmission, serologic testing of coyotes for titers to plague can �123 help define the extent of plague activity in rodent prey populations (Thomas and Hughes, 1992). Moreover, by sampling juvenile animals in the summer, recent plague activity can be identified. This survey of carnivores, combined with collaborative work by the Bureau of Land Management investigating changes in prairie dog numbers, will provide important information on activity of diseases that are potentially devastating to black-footed ferrets. Disease management activities will be driven by these findings. Methods: We will use survey of carnivores as a tool to determine status of canine distemper and sylvatic plague in LSMA. Survey~ will be performed annually in January-March and July-September. Each survey will consist of a minimum sample of 20 coyotes. Aerial gunning will be used as the primary means of sample collection. Summer collections will focus on juvenile animals. Animals will be located by personnel in a fixed wing aircraft. Personnel from USDA Wildlife Services skilled in coyote control will fly over and shoot animals at close range (about 10-45 m) using a 12 g shotgun loaded with copper coated BB's or #4 buck shot. Multiple shots will be fired at the head of the animal. Ground crews will locate and recover the animal carcass. If the animal was not fatally wounded, ground crews will euthanatize animals with a shot to the neck or chest. Location of carcass collection will be recorded within 1/4 section. Surveys will be supplemented with additional samples collected opportunistically from carcasses of coyotes and other carnivores, including badger (Taxidea taxus), red fox (Vulpes vulpes), and striped skunk (Mephitis mephitis), killed as a result of hunting or by unintentional vehicular collision. Diagnostic techniques will include collection of blood samples for serology, collection of lower jaw for age determination, and postmortem examination to detect evidence of active disease. Serum will be harvested from blood samples, frozen, and submitted to Wyoming state Diagnostic Laboratory (WSVL, Laramie) for determination of titers to canine distemper using the serum neutralization test and to plague (Yersinia pestis) using the passive hemagglutination inhibition test. Titers to toxoplasmosis, tularemia, Aleutian disease (mustelids only), and leptospirosis will also be determined using samples collected during the initial carnivore survey. Gross necropsy will be performed by or under the direct supervision of a veterinarian. Routine tissue collection will include: samples of gross lesions, kidney, urinary bladder, lung, and brain placed in 10% buffered formalin; samples of lung, small intestine, and brain frozen; and samples of lesions stored fresh for immediate analysis. Samples of parasites will also be collected opportunistically; however, all badgers with dermatitis will be sampled for Filaria taxideae. Fixed tissues will be submitted to Dr. E. S. Williams, WSVL for histologic examination. Frozen samples will be stored by Colorado Division of Wildlife for analysis as needed (i.e., as indicated by histologic' or serologic findings). Fresh samples will be submitted to WSVL or Colorado State Veterinary Diagnostic Laboratory (CSVDL, Fort Collins). Parasites will be stored and submitted to WSVL or CSVDL as needed. Additional blood samples will be collected for serology as available from carcasses harvested by local hunters and from unintentional vehicular collision. Serum samples will be handled as previously described. Analysis: A titer of ~1:16 will be considered positive for antibodies against canine distemper virus or for antibodies against Yersinia pestis. We will use Fisher's exact test (or extensions) to statistically analyze the prevalence of antibodies among age classes and over time. �124 Flea Control Rationale: Mortality of black-footed ferrets from infection with sylvatic plague is extremely high (Williams et al., 1994). Further, mortality from sylvatic plague in prairie dogs can markedly impact prey base for black-footed ferrets. Plague has severely hampered reintroduction efforts in other states (P. Marinari, pers. commun.) and preliminary data suggest that plague activity is present in some areas of the LSMA (M. Albee, unpub. data). Successful reintroduction will likely require management techniques to control plague. Plague i~ maintained primarily in rodent populations, such as prairie dogs, where it is transmitted via the bite of an infected flea. Fleas of the genera Opisocrostis and Oropsylla have been associated with plague transmission in prairie dogs (Ubico et al., 1988). Insecticide dusts have been applied to prairie dogs burrows in an attempt to kill fleas and thus control plague. Carbaryl has been the most commonly used insecticide; however, application is labor intensive and activity is short-lived (Barnes, 1993). Permethrin has been shown effective up to 84 days after application, but the authors warn that application at the recommended dosage rate would be highly laborious (Beard et al., 1992). Recently developed compounds used to control fleas in pet animals offer a promising alternative to insecticide dusts. Although topically applied flea adulticides, such as imidacloprid, are not feasible for use in wildlife; insect growth regulators (IGRs) with ovicidal and/or larvicidal activity could be delivered orally to free-ranging animals via bait. The IGR lufenuron is a benzoylphenylurea derivative which inhibits formation of chitin in the exoskeleton of insects (Cohen, 1987). A single oral dose of lufenuron has been shown effective in controlling the cat flea (Ctenocephalides felis felis) for at least 30 days in treated dogs (Hink et al., 1994) and cats (Blagburn et al., 1994). Davis (1997) reported a significant reduction in fleas on ground squirrels (Spermophilus beecheyi) that had been treated with lufenuron. Another IGR, pyriproxifen, exerts activity by mimicking natural insect juvenile hormones. pyriproxifen has been shown effective in vitro in inhibiting normal egg productiop for 80 hr post-treatment (Palma et al., 1993) and may have other longer lasting means of control as well (T. Miller, pers. commun.). If these compounds proved efficacious over a period of time (~ 1 mo) when administered orally to prairie dogs, they could be applied over large areas of prairie dog habitat via a treated bait. However, prior to management application, the products should be evaluated in a controlled laboratory setting to determine an effective dose, duration of efficacy, product safety, and to formulate an acceptable bait carrier. If results of laboratory tests are positive, the products should be tested in a controlled field study to determine efficacy under natural conditions. Without this evaluation, the management potential of IGRs for controlling plague and protecting the health of the endangered blackfooted ferret, as well as human health, will be difficult to discern. Our work will be divided into three phases. Phase I will be pilot work to establish a colony of captive prairie dogs and to develop an environment suitable for flea survival and reproduction both on the prairie dogs (and in their bedding) and in an artificial medium. Before we can evaluate the efficacy of IGRs, we must be sure that fleas will survive and reproduce on our captive prairie dogs. Further, we will need to artificially rear prairie dog fleas for infestation of prairie dogs and assessment of egg viability. These techniques are not currently available (Barnes, 1993). Phase II will evaluate efficacy and safety of lufenuron and pyriproxifen fed to captive prairie dogs. Finally, in Phase III we will evaluate potentially efficacious IGRs (from phase II) in a limited field application. White-tailed prairie dogs (Cynomys leucurus) will be used as the study animal because this is the species of prairie dog present at the LSMA black-footed ferret reintroduction site. �125 Phase I--Pilot study Methods: Capture of prairie dogs. White-tailed prairie dogs will be captured at the Arapaho National Wildlife Refuge, Colorado. One or two single or double-door Tomahawk traps (15 cm x 15 cm x 60 cm) will be placed near active burrows. Traps will be baited with horse sweet mixture (sweetened grain mixture). During a 1-2 week prebaiting session, the traps will be wired closed (to prevent unwanted, accidental entry), and bait will be spread around the traps and burrows. After the prebaiting session, traps will be set (opened and baited) just prior to sunset, and checked (minimally) at noon and at dusk the following day. Prairie dogs are diurnal, and are typically not seen above ground in inclement weather, so it is unlikely that any will be captured at night or when temperatures are very cold or very hot. Nonetheless, traps will not be set during rain or snow, or when daytime ambient temperature is expected be below -6 C or above 32 C. Prairie dogs will be handled by experienced mammal trappers, wearing heavy leather or welder'S gloves. Upon capture, each animal will be physically restrained in a cone-shaped nylon or canvas bag. They will be sexed and weighed, and age determined (Hoogland, 1995; Cox and Franklin, 1990). A numbered monel ear tag will be attached to each animal. The animals will then be transferred to an individual mesh-bottom holding cage (minimum size = 45 cm x 45 cm x 45 cm) with nest box and covered for transport. Feed (e.g., rodent chow, alfalfa cubes) and water will be provided ad lib if transport/holding time will exceed 2 hr. Care and housing of prairie dogs. Prairie dogs will be maintained indoors in individual cages (45 cm x 90 cm x 45 cm) with nest boxes at the Foothills Wildlife Research Facility (FWRF). Animals will be quarantined for 14 days upon return to FWRF to assess disease (plague) status. Due to our study needs, prairie dogs will not be treated to control external parasites. During the quarantine period, all personnel will be required to wear room-specific gloves and coveralls when in the room, whether observing animals or cleaning cages. Personnel will be informed of the clinical signs of plague in prairie dogs and humans and instructed to monitor themselves and their clothing for fleas. If plague is diagnosed in a prairie dog, all animals in the building will be euthanized with an overdose of halothane inhalant anesthetic. Our custom cage design is based on specific study needs (providing the prairie dogs with sufficient living space, while allowing us to rear and collect fleas), discussions with M. Metzger, and the published work of Hilton (1971). Individual prairie dog cages will be constructed of wood and wire fencing, with "no-see-um" netting over the top and covering potential flea escape routes. Half of the cage serves as a nest box, and the other half serves as a feeding area. Two hinged doors allow access to each half of the cage for cleaning or animal handling. The nest box is a 40 x 28 x 23 cm plastic container fitted with a wooden lid. The nest box has a 10 cm diameter hole in one wall to provide access to the feeding area; a 10 cm diameter piece of PVC tubing joins the nest box to the feeding area. The access hole can be covered to confine the animal to either side of the cage as necessary. Previous work on ground squirrels (Hilton, 1971) suggests that the animals are unlikely to defecate/urinate in the nest box, and Hoogland (1995) noted that there was little fecal matter in the burrows and nests of black-tailed prairie dogs he studied. Our preliminary work shows that defecation in the nest box is minimal, and that urination is very rare; we therefore do not expect the nest boxes to become soiled during experiments. �126 The floor of the feeding area consists of 1 cm mesh wire fencing, to allow feces, urine, and spilled food to fall through to a pan below. This mesh size prevents the animal's feet from becoming wedged in the floor, but may encourage the animal to spend most of its time in the nest box when not feeding. It is important for us to keep the animal in the nest box as much as possible, in order to enhance flea reproduction; many fleas of the genus Oropsylla are "nest fleas" and are associated with the host's body only when feeding (Hilton, 1971; M. Metzger, pers. comm.). Confining the prairie dogs and fleas to the nest box will also facilitate collection of fleas and.their eggs. A food dish and water bottle will be wired to the side of the feeding area. A 2.5 cm deep, removable, galvanized sheet metal pan will be placed under the entire cage to catch food, feces, urine, and any fleas that escape the nest box. All prairie dogs will be checked at least once per day. Fresh feed (rodent chow, alfalfa cubes) and water will be provided daily, and cage bottoms will be cleaned of feces each day. Pans will be cleaned and bedding will be replaced as needed, but a minimum of 2 times per week. Rooms (or nest boxes) will be maintained at approximately 24 C and 75% relative humidity to ensure flea survival and reproduction (M. Metzger, pers. commun.; Metzger and Rust, 1997). Two south-facing windows in the animal room should provide sufficient light for these fossorial animals, and will allow us to maintain the animals on a natural photoperiod. We do not anticipate any prairie dog mortality due to capture or experimental procedures. If an animal becomes sick or injured, and recovery is not likely, it will be euthanized with an overdose of halothane inhalant anesthetic. Pilo~ s~udy. We will capture and house 3-5 prairie dogs for initial evaluation of our methods. Prairie dogs will be observed daily and weighed at least once per week, to help assess health and acclimation to captivity. If two or more animals do not adjust to our captive conditions (they refuse food and/or water or appear sick) we will review and modify our procedures. All prairie dogs that die in captivity will undergo a full post-mortem examination. Minor adjustments to husbandry may be made as needed to meet animal needs. One objective of the pilot study is to determine if prairie dogs can be trained to allow repeated grooming by handlers, without being chemically immobilized. Hoogland (1995) successfully groomed black-tailed prairie dogs without chemical immobilization, and one of us (KTC) repeatedly handled captive black-tailed prairie dogs that were distracted with a treat, such as a carrot. We will therefore attempt to handle each animal on a daily basis, in order to establish a routine whereby they become accustomed to being groomed. If we can train the animals to allow grooming, chemical immobilization will not be necessary. If chemical immobilization appears necessary, we will test the following methods to determine which to use in Phase II: 1) animals will be placed in an induction chamber and anesthetized with isoflurane. The animals will be removed from the chamber within 15 seconds after cessation of movement and anesthesia will be maintained via a mask; or 2) animals will be placed into a restraining cone and injected intramuscularly (IM) with a combination of ketamine (30 mg/kg) and diazepam (0.5 mg/kg) (Kreeger, 1997). During anesthesia, animals will be monitored visually for respiration, heartbeat, and signs of distress. Trained or immobilized prairie dogs will be groomed with a fine-toothed flea comb for 3 minutes; grooming will take place up to 3 times per week. After immobilization, each animal will be returned to its cage, and closely observed until it regains normal functions. �127 Concurrent to establishing the captive prairie dog colony, we will develop a technique to artificially rear prairie dog fleas of the genus Oropsylla. Although prairie dogs can harbor several genera of fleas, we have selected to maintain only Oropsylla because they are among the most common fleas of prairie dogs and they have been implicated in the transmission of plague (Ubico et al., 1988). Fleas will be collected by combing trapped prairie dogs and by swabbing of burrows (Barnes et al., 1972). Fleas will be placed in 500 ml glass jars each containing about 125 g of larva-rearing media consisting of wheast (Red star Biologicals), dried beef blood (Monfort Biologicals), powdered dog chow, and sand. In the laboratory, live fleas will be anesthetized with CO2 and identified by comparison to a reference collection of preserved fleas. Because fleas are sensitive to changes in temperature and humidity, they will be maintained in an incubator at about 23-24 C and ~70% relative humidity (Metzger, pers. commun.). Appropriate relative humidity will be maintained using a super-saturated solution of potassium chloride (Winston and Bates, 1960). A natural photoperiod will be approximated within the incubator through the use of fluorescent lights and a timer. Fleas will be monitored daily for survival and reproduction. Modifications will be made as necessary to optimize flea performance. To determine adequacy of the cage environment for flea survival and reproduction, we will infest prairie dogs with about 20-50 male and female fleas. We will collect and count fleas from prairie dogs and their bedding at weekly intervals. Fleas will be collected from prairie dogs by combing (while distracted with a treat or while under anesthesia) for 3 min and from bedding by sifting and visual inspection. Fleas will be returned to the prairie dog after counting. Collections, and if needed reinfestations, will be continued until successful colonization occurs. If a severe allergic reaction or other health problem associated with flea infestation occurs, the prairie dog will be removed from the study and provided veterinary care or euthanized. Analysis: Evaluation of husbandry techniques and artificial flea rearing procedures will be subjective. No statistical analyses will be performed. Phase II--Laboratory Evaluation of IGRs Methods: Prairie dogs will be captured and maintained in captivity as described in Phase I. Laboratory evaluation of pyriproxifen and lufenuron will be divided into four experiments--efficacy dose titration, efficacy of controlling flea infestations in a simulated burrow environment, product safety, and bait formulation. Detailed Study Plans will be written for each of these experiments based on results of the Pilot Study; however, general procedures will be as follows. Efficacy dose titration. In this experiment we will determine an effective dose of pyriproxifen and of lufenuron to interrupt reproduction in prairie dog fleas and monitor duration of the products' efficacy. Prairie dogs (n = 21) will be blocked by sex and randomly divided into two treatment groups and a control group. In the first experiment, lufenuron will be evaluated at two dose rates. Treatment group 1 (n = 7) will receive 15 mg/kg and treatment group 2 (n = 7) will receive 45 mg/kg lufenuron PO. The control group (n = 7) will receive excipient suspension without lufenuron po. After a sufficient washout period, the experiment will be repeated using pyriproxifen at anticipated dosages of 50 mg/kg and 100 mg/kg PO for treatment groups 1 and 2, respectively. Prairie dogs will be infested with fleas prior to treatment and periodically after treatment to maintain flea infestations. We will collect flea eggs prior to treatment and at 5 weekly intervals and determine egg �128 viability by rearing in artificial media. Number of eggs hatching in a specific time period (to be determined in Phase I) will be determined and compared between groups and over time. We may also collect blood from prairie dogs during the weekly samplings if an assay to determine serum levels of test product can be developed. Development of a successful artificial rearing system for prairie dog fleas is prerequisite to the performance of this experiment. If such a protocol is not developed, we will attempt to determine an effective dose using the methods described below for evaluation of efficacy in a simulated burrow environment. In this case, duration of efficacy will be more difficult to determine. Efficacy of con~rolling flea infes~a~ions in a simula~ed burrow environmen~. If results of the efficacy dose titration experiment suggest that a single oral dose of pyriproxifen and/or lufenuron is effective in controlling reproduction of prairie dog fleas for at least 4 wk, we will further evaluate the product(s) in a simulated burrow environment. The purpose of this experiment is to evaluate the ability of the test products to control prairie dog flea populations over an extended period. Prairie dogs (n = 20) will be blocked by sex and divided into treatment and control groups. They will be infested with fleas and treated monthly with pyriproxifen or lufenuron. Adult fleas on the animal and in the cage will be counted at 1-2 wk intervals for 12 wk. After counting, fleas will be returned to the animal. Number of fleas recovered will be compared between treatment groups and over time. Produc~ safe~y. We will determine safety of an overdose of pyriproxifen and/or lufenuron. Prairie dogs (n = 20) will be divided into a treatment and control group. Prairie dogs in the two groups will be maintained similarly except the treatment group will receive an amount of product equal to that contained in the maximum calculated daily intake of treated bait. Overdose feeding will continue for a 3 day period. Health, attitude, and weight change of prairie dogs in the treatment and control groups will be monitored during, and for 1 wk following, this treatment. Based on type of action, these products should be safe in mammals (Blagburn et al., 1995; T. Miller, pers. commun.); however, if animals become moribund, they will be euthanatized. All dead animals will receive a complete postmortem examination. If the overdose of test product is safe in prairie dogs, product safety will be tested in black-footed ferrets or black-footed ferret x Siberian polecat hybrids. Ferrets maintained at other research facilities will be used as test animals and will be divided into a treatment and control group. Prairie dogs treated with three times the maximum daily dosage of test product (as described above) will be fed to ferrets in the treatment group for 7 days. Health, attitude, and weight change of ferrets in the treatment and control groups will be monitored during, and for 1 wk following, this treatment. Bai~ formula~ion. When an effective dose of pyriproxifen and/or lufenuron is determined, we will begin investigating formulation of a bait carrier. Because of differences in the chemical properties of the test products, unique formulations will likely be required for each product. Due to its high palatability, we will initially attempt to incorporate the test product into horse sweet feed (sweetened grain mixture); however, incorporation into a pelleted feed may be required to adequately deliver the product. Palatability of treated bait will be evaluated in feed-deprived (hungry) prairie dogs as well as those fed ad libitum maintenance diets (satiated). Blood samples may be collected from prairie dogs to determine serum level of test product after bait consumption. �129 Analysis: Efficacy dose titration. Efficacy of test product will be determined by comparing developmental success of eggs produced by fleas exposed to treated prairie dogs vs. eggs produced by fleas exposed to control group prairie dogs. These formulas will be used in efficacy determination: Developmental Percentage success efficacy = number emergent adult fleas x 100 number eggs collected mean developmental success (control)mean developmental success (treated) mean developmental success (control) x 100 Mean developmental success values for eggs collected from prairie dogs of the treatment groups will be analyzed using analysis of variance. in each Efficacy in a simulated burrow environment. Efficacy of test product will be determined by comparing number of fleas recovered from treatment and control prairie dogs and their cages on each sampling day using analysis of variance. Significant reductions in numbers of fleas on treated prairie dogs would signify interruption of the flea life cycle attributable to treatment with the test product. Product safety and Bait formulation: Evaluation of product safety acceptability of bait will be subjective. No statistical analyses performed. Phase III-Field Evaluation and will be of IGRs Methods: Based on results of laboratory evaluations, we will evaluate pyriproxifen and/or lufenuron in a limited field application. Only product(s) that significantly reduced flea numbers on treated hosts for ~1 mo and that. showed no adverse effects on captive prairie dogs and ferrets will be field tested. We will identify similar, yet geographically distinct, colonies of healthy prairie dogs for evaluation of the test product(s). Treatment colony(s) will receive bait fortified with the test compound while the control colony will receive bait only. Treatments will be applied at monthly intervals during the summer. Observations will be made to determine prairie dog numbers and consumption of bait by prairie dogs and non-target species. Flea indices (number of fleas per individual and number of fleas per burrow) will be determined by counting and identifying fleas collected by combing live-trapped prairie dogs and by swabbing burrows (Barnes et al., 1972). Blood samples may also be collected from prairie dogs to determine serum level of test product. Collections will be made at monthly intervals beginning ~1 mo prior to initiation of treatment, continuing through the active season of prairie dogs (September), and resuming the following summer. Flea indices will be compared between treatment and control colonies and over time. Analysis: Efficacy of test product will be determined by comparing flea indices of control and treatment colonies at each sampling using analysis of variance. Significant reductions in numbers of fleas on prairie dogs at treated sites would signify interruption of the flea life cycle attributable to treatment with the test product. �130 Schedule: Objective 1. Disease survey 2. Phase I--Pilot study 3. Phase II--Laboratory study 4. Phase 111- Field study 5. Analyze data, report results E. Fiscal Year 1998-2002 1998-1999 1999-2000 2000-2001 2001-2002 LOCATION Planning, captive animal research, data analysis, and reporting will be carried out at the Colorado Division of Wildlife's Foothills Wildlife Research Facility, 4330 W. Laporte Ave., Fort Collins, Colorado. Carnivore disease survey Moffat County, Colorado. will be conducted at the Little Capture of prairie dogs to establish a captive colony National Wildlife Refuge, Jackson County, Colorado. Location reported F. Year Estimated Costs $13,000 20,500 21,400 21,400 13,000 Ell!: R~gyit:~m~n:tfZ PFTE TFTE 0.5 0.5 0.5 0.5 0.5 0.25 0 0.5 0.5 0 PERSONNEL Margaret A. Wild1 Kevin T. Castle1 Thomas A. Miller2 Craig Parks3 Co-principal investigator Co-principal investigator Co-investigator Co-investigator lColorado Division of Wildlife, Fort Collins, Colorado Virbac, Inc., Fort worth, Texas 3Novartis Animal Health U.S., Greensboro, North Carolina 2 but will be COST 1998 1999 2000 2001 2002 G. will be at Arapaho of Phase III (field study) is yet to be determined, in the detailed Study Plan for that experiment. ESTIMATED Fiscal Snake Management Area, �131 H. LITERATURE CITED Barnes, A. M. 1993. A review of plague and its relevance to prairie dog populations and the black-footed ferret. Proceedings of the symposium on the management of prairie dog complexes for the reintroduction of the blackfooted ferret. J. L. Oldemeyer, D. E. Biggins, B~ J. Miller, and R. Crete, eds. 96pp. Barnes, A. M., L. J. Ogden, and E. G. Campos. 1972. Control of the plague vector, Opisocrostis hirsutus, by treatment of prairie dog (Cynomys ludovicianus) burrows with 2% carbaryl dust. Journal of Medical Entomology 9: 330-333. Beard, M. L., S. T. Rose, A. M. Barnes, and J. A. Montenieri. 1992. Control of Oropsylla hirsuta, a plague vector, by treatment of prairie dog burrows with 0.5% permethrin dust. Journal of Medical Entomology 29: 25-29. Blagburn, B. L., J. L. Vaughan, D. S. Lindsay, and G. L. Tebbitt. 1994. Efficacy dosage titration of lufenuron against developmental stages of fleas (Ctenocephalides felis felis) in cats. American Journal of Veterinary Research 55: 98-101. Blagburn, B. L., C. Barnett. 1995. (Ctenocephalides American Journal M. Hendrix, J. L. Vaughan, D. S. Lindsay, and S. H. Efficacy of lufenuron against developmental stages of fleas felis felis) in dogs housed in simulated home environments. of Veterinary Research 56: 464-467. Cohen, E. 1987. Interference with chitin biosynthesis in insects. In Chitin and benzoylphenyl ureas, series entomologica, vol. 38, J. E. Wright and A. Retnakaran (eds), Dr. W. Junk, Publishers, Boston, pp. 33-42. Cox, M. K. and W. L. Franklin. black-tailed prairie dogs. 1990. Journal Premolar gap technique of Wildlife Management for aging live 54:143-146. Davis, R. M. 1997. Use of an orally administered insect development inhibitor (lufenuron) as a flea control agent in the California ground squirrel, Spermophilus beecheyi. Fourth International Symposium on Ectoparasites of Pets, pp 31-35. Hilton, D. F. J. 1971. A method for rearing fleas of ground squirrels. Transactions of the Royal Society of Tropical Medicine and Hygiene 66: 188189. Hink, W. F., M. Zakson, and S. Barnett. 1994. Evaluation dose of lufenuron to control flea infestations in dogs. Veterinary Research 55: 822-824. Hoogland, J. L. 1995. animal. University The black-tailed pra~r~e dog: social of Chicago Press, Chicago, 557 pp. of a single oral American Journal life of a burrowing Kreeger, T. J. 1997. Handbook of Wildlife Chemical Immobilization. International Wildlife Veterinary Services, Inc., Laramie, WY. Metzger, M. E. and M. K. Rust. 1997. Effect of temperature (Siphonaptera: Pulicidae) development and overwintering. Entomology 34: 173-178. of 342 pp. on cat flea Journal of Medical �132 Palma, K. G., S. M. Meola, and R. W. Meola. 1993. Mode of action of pyriproxifen and methoprene on eggs of ctenocephalides felis (Siphonaptera: Pulicidae). Journal of Medical Entomology 30: 421-426. Quan, T. J. 1982. Plague. In Diseases of wildlife in Wyoming, N. Kingston, W. R. Jolley, and R. C. Bergstrom (eds), Wyoming Department, Cheyenne, pp. 67-71. Rust, M. K. 1992. (Siphonaptera: 242-245. Influence Pulicidae) State of Colorado. 1994. Division of Wildlife. E. T. Thorne, Game and Fish of photoperiod on egg production of cat fleas infesting cats. Journal of Medical Entomology Long Range 32pp. Plan. Department of Natural Thomas, C. U., and P. E. Hughes. 1992. Plague surveillance testing of coyotes (Canis latrans) in Los Angeles County, Journal of Wildlife Diseases 28:610-613. Resources, by serological California. Ubico, S. R., G. o. Maupin, K. A. Fagerstone, and R. G. McLean. 1988. plague epizootic in the white-tailed prairie dogs (Cynomys leucurus) Meeteetse, Wyoming. Journal of Wildlife Diseases 24: 399-406. U. S. Fish and Wildlife 87pp. Service. 1988. Black-footed Ferret 29: Recovery A of Plan. Williams, E. S. 1982. Canine distemper. In Diseases of wildlife in Wyoming. E. T. Thorne, N. Kingston, W. R. Jolley, and R. C. Bergstrom (eds), Wyoming Game and Fish Department, Cheyenne, pp. 10-13. Williams, E. S., K. Mills, D. R. Kwiatkowski, E. T. Thorne, and A. BoergerFields. 1994. Plague in a black-footed ferret (Mustela nigripes). Journal of Wildlife Diseases 30:581-585. Williams, E. S., E. T. Thorne, M. J. G. Appel, and D.-W. Belitsky. 1988. Canine distemper in black-footed ferrets (Mustela nigripes) from Wyoming. Journal of Wildlife Diseases 24:385-398. Winston, P. W., and D. H. Bates. 1960. Saturated solutions humidity in biological research. Ecology 41: 232-237. for the control of �133 SEGMENT State of Project No. Work Package Task No. Colorado W-153-R No. __~0~8~8~0~ NARRATIVE _ Cost Center 3430 Mammals Program Black-footed Ferret Conservation Monitoring and Managing Disease in Black-footed Ferrets Planning Program Narrative Objectives: 1. Monitor enzootic disease activity threatening ferrets reintroduced into the LSMA. 2. Develop techniques regulators applied survival of black-footed to manage plague in the LSMA using orally to prairie dogs. Segment Objectives: 1. Determine the level of activity of canine distemper plague in prairie dogs (using coyotes as sentinels) sampling ~40 coyotes. insect in carnivores and in the LSMA by 2. Establish a colony of captive white-tailed prairie infestation with fleas of the genus Oropsylla. 3. Develop 4. Determine an effective dose of pyriproxifen and of lufenuron fleas of the genus Oropsylla on prairie dogs. Estimated techniques Segment Costs to artificially = Personal Services Permanent employees Temporary employees Contracts Total Personal Services Operating supplies Travel expenses Capital egyipment Total Operating Total Costs rear dogs that can sustain fleas of the genus $46,540 (0.5 FTE) and services growth $26,040 o 10rOOO $36,040 $ 8,300 2,200 o $10,500 $46,540 Oropsylla. to control �� https://cpw.cvlcollections.org/files/original/741451162f021db928ae9dd98ad7de82.pdf eebfa541860e03c7633ae6cd7558b2fc PDF Text Text 135 Colorado Division Wildlife Research July 1998 of Wildlife Report JOB PROGRESS State of project No. Work Package Colorado Cost Center W-153-R-11 Mammals No. __~3~0~0~1~ _ Task No. Period Personnel: 3430 Program Deer Conservation Experimental Covered: Author: REPORT Deer ~nyentory July 1, 1997 - June 30, 1998. R. M. Bartmann, T. M,. Pojar. R. Arant, L. Bennett, G. Bock, D. C. Bowden, D. Coven, M. Leslie, D. Masden, K. Miller, J. Olterman, M. Potter, G. Schoonveld, H. Spear, S. Steinert, J. Thomson, B. Watkins, G. C. White, and D. Younkin. Abstract An inventory system was developed to enable more precise monitoring of mule deer (Odocoileus hemionus) population status. It was implemented in 2 Data Analysis Units (DAU) in late 1997. A spreadsheet model was used to identify key parameters for model inputs. Allocation of sampling effort among the surveys required to obtain parameter estimates was based on annual variability associated with those estimates and sensitivity of the model to change in those estimates. Key parameters are annual doe survival~ overwinter fawn survival, fawn:doe ratio and proportion of does in the population obtained from age/sex surveys, population size, and harvest. In D4, the fawn:doe ratio estimate was 58.8:100. None of 30 radiocollared does died during the segment and 10 of 38 fawns died for a 74% survival. In D19, the fawn:doe ratio estimate was 34.2:100. Five of 31 does died for an 84% survival and 20 of 39 fawns died for a 49% survival. A quadrat census to estimate population size could not be done in either DAU. ��137 EXPERIMENTAL Richard DEER M. Bartmann INVENTORY and Thomas M. Pojar P. N. OBJECTIVES 1. Develop and test an experimental deer inventory system in 5 DAU's in western Colorado to determine efficacy in monitoring deer population status. 2. Publish results in a peer-reviewed scientific SEGMENT doe survival journal. OBJECTIVES 1. Estimate the annual (Uncompahgre). 2. Estimate the over-winter D-19 (Uncompahgre). 3. Estimate the December fawn:doe and buck:doe Feather) and D-19 (Uncompahgre). 4. Estimate winter (Uncompahgre). 5. Develop a spreadsheet model for predicting deer population DAU's D-4 (Red Feather) and D-19 (Uncompahgre). 6. Analyze fawn survival deer density data and prepare rate in DAU's in DAU's an annual rate D-4 Federal D-4 (Red Feather) in DAU's ratios D-4 (Red Feather) in DAU's (Red Feather) and D-19 D-4 and (Red and D-19 Aid Job Progress performance in report. INTRODUCTION During the early 1990's, mule deer populations in much of Colorado and the west began declining. The Colorado Division of Wildlife (CDOW) has collected data on deer populations for many years. These data consist primarily of harvest estimates collected state-wide every year, age/sex classifications made in most Game Management Units (GMU) every 1 or more years, and population estimates made in a few GMU's or DAU's infrequently. These data were assembled to try and determine if and when a deer decline occurred and its magnitude (Colo. Div. Wildl., unpubl. rep.). It was concluded the occurrence of a large-scale deer decline might be inferred but evidence was weak. This was partly because the data were inconsistent in the timing and manner in which they were collected, and also because additional key data on survival rates were not available. Thus, a new deer monitoring system was needed that would enable assessing deer population performance in a more timely and precise manner. STUDY AREAS It was originally around the state. suffice. Because planned to establish a new deer inventory system in 5 DAU's Later, it was decided that for evaluation purposes, 3 would of financial constraints, the system was implemented in 2 �138 DAU's the first year with the third DAU to be included the second year. The 2 DAU's selected for initial implementation of a new deer inventory system were D-4 ( Red Feather) in the northern front range and D-19 (Uncompahgre) in southwest Colorado. DAU D-4 consists of GMU's 7, 8, 9, 19, and 191. Winter range is a mountain shrub type with Ponderosa pine (Pinus ponderosa)- Douglas fir (Pseudotsuga menziesii) overstory in many places. DAU D-19 consists of GMU's 61 and 62. Winter range is primarily a pinyon (Pinus edulis)-juniper (Juniperus osteosperma)-sagebrush (Artemisia tridentata) type with considerable oakbrush (Quercus gambelli) interspersed. METHODS Survival Basic methods for both DAU's were the same, although timing of their implementation varied. Annual doe survival and over-winter fawn survival were estimated from a sample of radio-collared animals. Seventy radiocollars were available the first winter and allocated to 30 does and 40 fawns. Deer were captured in early winter by netgunning from a helicopter. Annual doe survival was estimated for the period 1 December to 30 November. Over-winter fawn survival estimated from 1 December to 15 June when they were considered adults. Capture locations were originally randomized throughout the winter range in each DAU, but problems developed with private land access and finding deer. Consequently, the helicopter crew was sent to accessible areas where deer were known present and instructed to put out a certain number of doe and fawn radiocollars. Each radiocollar had a mortality mode set for a 3-4-hour delay with frequencies in the 148-151 MHz range. Fawn collars were a drop-off type so they could be retrieved and reused. This precluded getting survival information on yearling females from 15 June to 30 November. Data from Piceance Basin indicates this is not a critical period from a survival standpoint, except during the hunting season, and hunting mortalities are censored anyway. Radiocollared deer were monitored from the air a minimum of once every 2 weeks from December through June. Each mortality was checked as soon as possible to try and determine cause of death. Age/Sex Classifications Age/sex classifications were used to estimate fawn recruitment to December and post-season buck:doe ratios. A random-route survey was already in place in D4 and was used without modification for this study. Fifteen existing census quadrats were randomly selected and a route passing through each quadrat devised. The route was flown with a Bell Soloy helicopter using a GPS unit for navigating from point to point. In D-19, a random-route survey was designed similar to that in D-4. Each GMU was divided from north to south into strata at 1,000-meter intervals along UTM coordinates. Two existing census quadrats were randomly selected in each strata within a GMU and, starting at the top of the GMU, a line was drawn to connect quadrats in a zig-zag pattern from north to south. The process was then repeated for the other GMU. �139 Deer in groups encountered along the flight path were classed as fawns, does, bucks, or unknown. Estimates of fawn:doe and buck:doe ratios were based on groups using the estimator developed by Bowden et al •. (1984).· Population Size Deer population size was estimated from counts of deer on 0.65-km2 quadrats using procedures described by Kufeld et ale (1980). A quadrat census system was established in D-4 in 1985 and has been flown 7 times, the last in January 1997. The current design includes 98 quadrats in 15 strata, but the original data on the total sample area and strata poundaries could not be found. There was not enough time to gather the needed data to define a new total sample area and re-stratify, so the quadrat census was to be flown as in the past. The exception was that a Bell Soloy helicopter was to be used instead of a Jet Ranger. Snow conditions along the front range are undependable. When adequate snow does occur to provide a good counting background, it usually lasts only a few days. Helicopters need to be scheduled in advance, so the chance of encountering good counting conditions was minuscule. Therefore, it was decided the D-4 survey would be flown regardless of snow cover. A quadrat census was established in GMU 62 in 1976. It was expanded to include GMU 61 in 1981. There are 346 quadrats in 13 strata which took -70 hours to survey. Only 30 hours were allocated for the census, so the number of quadrats was reduced to 140 by random selection. Population Model The POPII model is currently used to help manage deer populations. The model is complex and requires numerous parameter estimates with no supporting data. As a result, people are free to manipulate various parameters to try and make the model fit their preconceived ideas. A much simpler model is needed for which key inputs are derived from sample-based estimates. The new model will help determine the key types of data required and how to allocate costs and effort to obtain them. RESULTS Doe Survival ~: Thirty does were captured and radiocollared from 4-7 January 1998. The delay from the desired late November period was caused by contract problems. Monitoring was done from a fixed-wing aircraft about once every 2 weeks through June 1998. No does died during that period (Table 1). ~: Does were captured and radiocollared from 15-20 December 1997. Here again contract problems delayed the process. The first two does captured were given fawn radiocollars by mistake. Consequently, 31 does were finally radiocollared. Monitoring from a fixed-wing aircraft was done weekly the first month, but was increased to twice weekly the rest of the winter and spring to try and locate mortalities quicker. As of 30 June, 5 of the 31 does had died for a survival estimate of 84%. Annual doe survival is monitored until 30 November, so this rate could go lower. �140 Table 1. Fates of mule deer does and fawns radiocollared in OAU's Feather) and D-19 (Uncompahgre) during the 1997-98 winter. D-4 Fate 0-4 (Red D-19 Does Fawns Does Fawns 30 28 26 19 Coyote 3 3 10 Lion 1 1 1 Survived Bobcat 1 Unknown Predator 2 1 Starvation 1 1 Accident 1 1 Undetermined 3 5 Fawn Survival ~: Thirty-eight fawns were captured and radiocollared during the same period as for does. Two collars were temporarily misplaced and not put out. Ten fawns died for an over-winter survival rate of 74%. ~: A doe radiocollar was erroneously placed on a fawn. This, together with the 2 fawn collars placed on does, resulted in only 39 fawns being radiocollared. Twenty fawns died by 15 June for an over-winter survival rate of 49%. The fawn that received a doe collar was 1 of the mortalities which eliminated the need to recapture it to prevent choking when it got older. Age/Sex Classifications ~: The classification survey was flown 16-18 December. There were 950 deer classified for a fawn:doe ratio of 58.8 (95% CI = 52.9-65.2) and buck:doe ratio of 23.8 (95% CI = 19.4-28.6). ~: The classification survey was flown 11-13 December. The random routes only took about 5 hours to complete with 358 deer classified. To use more of the 15 hours allocated for the survey, a non-random route was flown through the longer north-south length of each GMU. This produced another 831 deer in about the same amount of time as . for the random routes. L For all data combined, the fawn:doe ratio estimate was 34.2 (95% CI = 30.937.7) and the buck:doe ratio estimate was 11.1 (95' CI = 8.9-13.4). Comparing results from 2 route types, the fawn:doe ratio estimate was marginally higher (35.3 vs. 31.8) while the buck:doe ratio was nearly twice as high (12.8 vs. 7.0) on the non-random routes. Thus, the observer's knowledge about where to find large numbers of deer could bias ratio estimates, particularly for bucks. Population Size ~: Contract problems prevented getting a helicopter before cutoff date for conducting the survey, so it was not flown. the mid-January �141 ~: Adequate snow conditions did not occur until late January. A population estimate has not been made since 1994, so it was decided to do the survey even though it was past the 15 January cutoff. Work was started 27 January, but weather and other problems prevented the survey from being completed in a reasonable time span, so it was canceled on 12 February after 63 quadrats had been flown. A valid population estimate cannot be made based on this incomplete data. Population Model Development of a population model to identify key inputs for refining the deer inventory system was begun with cooperators from Colorado state University. Deer population dynamics are much more complicated than the model portrays, but routine measurement of the wide array of inputs required for a more complicated model is unrealistic. Thus, it is a reasonable trade-off between what we can measure from a practical standpoint and what. is needed to predict deer population status for management purposes. The model concentrates on predicting changes in the adult doe portion of the population as this is the key to evaluating population status. The model has only 2 age classes (fawns and adults) assigned to 3 categories (fawns, does, and bucks). Fawns are initially recruited to the population on 1 December when the fawn:doe ratio is estimated from age/sex classifications. The model contains 4 parameters that are year-specific: annual buck survival, annual doe survival, over-winter fawn survival, and December fawn recruitment. Buck survival has no influence on population status unless their numbers drop low enough to affect breeding; a phenomenon yet to be documented in hunted populations. Therefore, there is no real need for an estimate of buck survival. An indication of status can be derived from the buck:doe ratio estimate obtained from age/sex surveys, and the number of new bucks recruited is estimated from male fawns that survive to become yearlings. Thus, only the last 3 parameters need to be considered for the model. The strategy for managing a mule deer population is to estimate the doe segment so harvest can be adjusted to maintain the December doe population at some desired level. To do this, 3 more parameters have to be estimated in addition to the 3 mentioned above: total population size, the proportion of the population that is does, and doe harvest. A harvest estimate is already available from another source and fawn recruitment and the proportion of does in the population are obtained from the same survey, so we only need to consider 4 surveys to obtain 5 parameter estimates. The problem now comes down to how to allocate costs and effort to these 4 surveys to derive estimates of the 5 parameters that will result in the most precise prediction of the December doe population or, alternatively, the annual rate of population increase. Either will indicate the number of does that must be harvested to maintain the desired doe population level. This process is ongoing and will be completed in the next segment. �142 LITERATURE CITED Bowden, D. C., A. E. Anderson, and D. E. Medin. 1984. sampling plans for mule deer sex and age ratios. Journal of Wildlife Management. 48:500-509. Kufeld, R. C., J. H. Olterman, and D. C. Bowden. 1980. A helicopter census for mule deer on Uncompahgre Plateau, Colorado. Journal Wildlife Management. 44:632-639. -"7 Prepared by / . .J./~~~'"·/""'ht.~:::i·~d~~;:.~~Bartm~~~a:::::n::~=-=·-=··.= .....:....=-._._. __ - ....:::===----WildLife Researche~ Wildlife Researcher quadrat of �143 Colorado Division Wildlife Research July 1998 of Wildlife Report JOB FINAL REPORT State of Project No. Work Package Colorado Cost Center W-1S3-R-11 Mammals No. __~3~0~0~1~ _ peer Reproduction Covered: Author: Personnel: Program peer Conservation Task No. Period 3430 July 1, 1997 - June 30, 1998. R. M. Bartmann, K. Larsen, Assessment T. M. Pojar. M. W. Miller, S. F. steinert, and G. C. White. Abstract Blood samples were taken from 29 mule deer (Odocoileus hemionus) does captured 4-7 January 1998 by helicopter netgunning in Data Analysis Unit (DAU) 0-4 (Red Feather) and tested for pregnancy via assay of pregnancy-specific protein B (PSPB). Twenty-seven does were deemed pregnant for a 93% pregnancy rate and all except possibly 1 were considered to have conceived during the first estrus. ��145 DEER REPRODUCTION Richard M. Bartmann ASSESSMENT and Thomas M. pojar P. N. OBJECTIVES 1. Estimate the variation in pregnancy-specific protein B (PSPB) levels mule deer does in Game Management Unit. (GMU) 19 (Poudre). 2. Develop a study plan to compare current mule deer reproductive rates to historic data to determine if differences exist that might contribute to the recent decline in deer populations in Colorado. SEGMENT the variation in GMU 19. OBJECTIVES 1. Estimate periods 2. Test for differences in pregnancy winter sample periods. 3. Develop a study plan to compare current and historic mule deer in at least 2 areas of Colorado. 4. Analyze data in PSPB levels and prepare an annual in in mule rates deer does during 2 winter of mule deer does between Federal reproductive Aid Job Progress the 2 rates of report. INTRODUCTION An assessment of mule deer population status during the early to mid-1990's indicated that many populations across western Colorado had declined to various extents. However, the cause(s) was not identified. One indication of a problem was detection of a significant decline in December fawn:doe ratios in some DAU's over the past 20+ years. This suggested a possible problem with reproduction or neonatal mortality. A logical starting point to try and identify a problem was the breeding season--were does being bred? This was also the easiest aspect to investigate because it did not require killing deer to obtain samples. STUDY AREAS GMU 19 was selected for study because there was historic data for comparison. Winter range along the northern front range is a mountain shrub type with Ponderosa pine (Pinus ponderosa)-Douglas fir (Pseudotsuga menziesii) overstory. METHODS Pregnancy detection was via blood analysis to determine levels of PSPB. PSPB levels in blood have been used to detect pregnancy with a high degree of accuracy in various species including mule deer (Wood et ale 1986). Detection times post-conception have varied among species, but is generally considered reliable after about 24 days with cattle (Sasser et ale 1986). Therefore, delaying blood sample collection until early January was considered to provide an adequate interval post-breeding for detecting PSPB. Delaying the second set of collections until early February would detect pregnancies from �146 conceptions during of late breeding. the second estrus and provide an indication of the extent From 15-20 does (~l-year-old) were to be captured with Clover traps in early January and again in early February. However, helicopter netgunning of deer for survival work in DAU D-4, of which GMU 19 is part, was delayed until early January, so it was decided to collect blood samples from does captured for that project. Blood samples were submitted to BioTracking in Moscow, Idaho to assess levels of PSPB to determine pregnancy status. RESULTS Blood samples were obtained from 29 does captured 4-7 January 1998. Twentyseven were considered pregnant and 2 not pregnant. One doe was at the cut-off point of 93% (93.1%) and may have been a late breeder. The range in values for the pregnant does was 84.3-93.1 with mean 88.5 and SE 0.36. Values for the 2 non-pregnant does were 102.6 and 102.8. The 27 pregnancies yielded a 93% pregnancy rate with all except possibly 1 apparently bred during the first estrus. Thus, late breeding did not appear a problem so no collections were done in February. The 93% pregnancy rate is barely significantly lower (P = 0.062) than the 100% rate derived from data for 49 does collected in GMU 19 from January-May 196165 (Anderson 1965a, 1965b, 1966). However, it is still considered a high rate with the conclusion that conception is apparently not a problem in DAU D-4. A next step would be to determine fetal rates, but this would involve euthanizing a large number of does during mid to late winter. One option is to have hunters collect does, but hunting seasons that late in winter are not well accepted. Therefore, no further work is planned to evaluate reproductive status until a satisfactory collection method is found for estimating fetal rates. LITERATURE CITED Anderson, A. E. 1965a. Reproductive studies. Colo. Dep. Game, Fish and Parks, Game Res. Rep. January: 165-195. Anderson, A. E. 1965b. Reproductive studies. Colo. Dep. Game, Fish and Parks, Game Res. Rep. January: 522-543. Anderson, A. E. 1966. Reproductive studies. Colo. Dep. Game, Fish and Parks, Game Res. Rep. January: 275-307. Basser, R. G., C. A. Ruder, K. A. Ivani, J. E. Butler, and W. C. Hamilton. 1986. Detection of pregnancy by radioimmunoassay of a novel pregnancyspecific protein in serum of cows and a profile of serum concentrations during gestation. Biol. Reprod. 35:936-942. Wood, A. K., R. E. Short, A. Darling, G. L. Dusek, R. G. Sasser, and C. A. Ruder. 1986. Serum assays for detecting pregnancy in mule and whitetailed deer. Journnl of i-lildlifeManagemeni:. 50:684-687 • -» Prepared by / I~(~£-WildLife ReBearche~ Wildlife Researcher : . --==- �147 Colorado Division Wildlife Research July 1998 of Wildlife Report JOB PROGRESS REPORT State of Project Work No. Package Colorado Cost Center W-153-R-ll Mammals No. ~3~0~0~1L_ Task No. Period _ 3430 Program Deer Conservation Monitoring and Managing Chronic Wasting Disease in Deer Covered: July 1, 1997 - June 30, 1998 Authors: M. W. Miller Personnel: S. Berry, K. Larsen, Wheeler, M. A. Wild, K. 1. O'Rourke, T. R. Spraker, and E. S. Williams S. Tracy, E. ABSTRACT Deer from throughout Colorado were examined for occurrence of chronic wasting disease using a combination of targeted surveys and harvest/road-kill surveys. We continued to develop and modify a statewide targeted surveillance program for acquiring, examining, and reporting on CWO suspects submitted from Colorado. Between June 1997 and May 1998, 6 chronic wasting disease (CWO)· cases were diagnosed among 16 "suspect" deer submitted from known endemic portions of northeastern Colorado; CWO was not diagnosed in any of 7 additional "suspect" deer submitted from elsewhere in Colorado. All confirmed CWO cases originated in game management units (GMUs) where the disease had been detected previously. Harvest and road-kill surveys were used to estimate CWO prevalence in enzootic management units. About 4.2% of deer harvested in Larimer County data analysis units (DAUs) (D4 or D10) and about 1.1% of deer harvested in South Platte River bottom units (DAU D44 plus GMUs 87, 90, 93, and 95) tested positive for CWO via immunostaining; none of the 171 deer harvested or culled in other select GMUs outside known enzootic areas tested positive for CWO. Prevalence estimates may be somewhat liberal because subclinical cases where either histopathological lesions or anti-PrP immunostaining reactions in brain tissue were included; of the 44 cases identified, ~none appeared to be suffering from clinical CWO and only 26 of 44 (59%) "positive" deer showed both histopathological lesions and immunostaining. In contrast to earlier observations derived from targeted surveillance data, CWO prevalence among male and female mule deer did not differ (P = 0.86). Age distribution of CWO-positive mule deer did not differ from sex-specific age distributions of negative animals. These data support the belief that CWO epidemiology is driven primarily by lateral transmission. �148 Requiring head submissions increased survey sample sizes for deer >4-fold compared to voluntary submissions in 1996. Based on harvest estimates from surveyed GMUs, compliance with mandatory submission regulations was about 75%. Unstaffed barrel sites still appear to be the most efficient method for sample collection~ Data from both targeted surveillance and surveys indicate that eastern Larimer County remains the most significant focus of CWO in Colorado, although some natural spread may be occurring both southward and eastward. Targeted ,surveillance of clinical suspects appears to be the most sensitive approach for initially detecting CWO in deer (and elk) populations throughout Colorado, and should be continued statewide. Once detected, combinations of harvest and road-kill surveys will be employed to estimate prevalence, monitor prevalence trends, and compare prevalence among DAUs. 'Because CWO appears to be more prevalent in deer than in elk in endemic areas, it follows that harvest surveys in high-risk areas should focus on deer rather than elk to be most effective in confirming absence of CWO in nonendemic areas. Cwo affected about 25% (5/25) of the adult (~1-yr-old) mule deer and about 46% (5/11) of the adult white-tailed deer in resident Foothills Wildlife Research Facility herds during the last biological year; CWO accounted for 83% of adult mule deer mortality and 100% of adult white-tailed deer mortality. Resident mule deer also represented the source of infection (treatment) in an ongoing 10-yr study of cattle susceptibility to CWO. No signs of neurological disease were observed in any of the 12 calves (subjects) or 12 mule deer fawns from the Rocky Mountain Arsenal National Wildlife Refuge (contact controls) housed with naturally~infected resident deer since July 1997; one calf died from gastrointestinal obstruction and bloat about 2 wk after entering the deer paddocks, and one fawn that apparently failed to adjust to captivity died about 1 mo after capture, presumably from hypothermia secondary to malnutrition. �149 MONITORING AND MANAGING CHRONIC WASTING DISEASE IN DEER M. W. Miller P. N. OBJECTIVES (1) Design, conduct, and report results of: (a) targeted surveillance to estimate and detect changes in distribution of chronic wasting disease (CWO) in free-ranging deer populations; and (b) harvest or road-kill surveys to estimate and detect prevalence of CWO in enzootic deer populations. (2) Design, captive changes in conduct, and report results of experimental studies using deer naturally or experimentally infected with cwo. SEGMENT OBJECTIVES (1) Conduct and report results of targeted surveillance to estimate and detect changes in distribution of CWO in free-ranging deer populations statewide. (2) Conduct and report results of harvest surveys to estimate prevalence of CWO in DAUs D4, D10( +GMU 29), and D44 (+ GMUs 87, 93, and 95). (3) Continue experimental evaluation natural contact exposure. (4) Observe deer. epizootiological features of cattle susceptibility of naturally-occurring to cwo via CWO in captive INTRODUCTION Chronic wasting disease (CWO) affects native deer and elk, causing behavioral changes and progressive loss of body condition that invariably lead to the death of affected animals (Williams and Young 1992). Neither the causative agent nor its mode of transmission have been identified. There are no tests currently available for diagnosing CWo in live animals, and postmortem tests require microscopic examination of brain tissue. There are no known treatments for cwo. Previous attempts to eradicate CWO from research facilities failed on at least 2 occasions (Williams and Young 1992; Miller et al., 1998). Although similar in some respects to other transmissible spongiform encephalopathies that affect domestic sheep (scrapie) and cattle (bovine spongiform encephalopathy; "mad cow disease"), there is no evidence suggesting cwo can be naturally transmitted to domestic livestock, or that scrapie or SSE can be transmitted to native cervids. Moreover, there is no evidence suggesting that cwo presents a threat to human health. "Chronic wasting disease" was first recognized by biologists in the 1960's as a disease syndrome of captive deer held in wildlife research facilities in Ft. Collins, CO, and was subsequently recognized in captive deer, and later in captive elk, in wildlife research facilities near Ft. Collins, Kremmling, and Meeker, CO and Wheatland, WY (Williams and Young 1980, 1982). Since 1981, CWO has also been diagnosed in free-ranging mule deer, white-tailed deer, and elk from northcentral Colorado; most of these diagnoses have been made since 1990 �150 (Spraker et al. 1997). Although CWO was first diagnosed in captive cervids, the original source of CWO is unknown; whether CWO in captive cervids really preceded CWO in wild cervids, or vice versa, is equally uncertain (Spraker et al. 1997). At present, the known world-wide distribution of CWO in wild cervids appears to be limited to northeastern Colorado and southeastern Wyoming. In Colorado, free-ranging CWO cases have primarily originated from along the Front Range near Estes Park and west of Ft. Collins-Loveland. Cases were diagnosed in 6 different game management units (GMUs)(191, 9, 19, 20, 94, and 96) prior to 1996, but two GMUs (19, 20) yielded about 85% of the documented cases; the affected GMUs comprise portions of 3 deer (04, 010, 044) and 2 elk (E4, E9) data analysis units (DAUs). There is no evidence that wild deer or elk outside northeastern Colorado are infected with CWO. The significance of CWO and its impacts on native deer and elk populations are unclear. Simulation models of CWO dynamics predict that this disease could cause significant declines in affected deer and elk populations (Miller and McCarty, unpublished data). In light of CWO's potential impacts on wildlife resources and the difficulties inherent in eliminating CWO from captive or wild cervid populations once established, it seems most prudent to assume CWO could adversely affect native deer and elk populations and manage to reduce its occurrence and prevent its further spread. A more complete understanding of CWO is fundamental to developing a comprehensive management program. In 1996, ongoing surveillance efforts were enhanced to provide a better tool for estimating and monitoring changes in CWO prevalence in enzootic areas and for potentially detecting emergence of CWO in new areas. Reliable estimates of CWO prevalence are particularly critical to detecting"trends, predicting potential impacts of disease on long-term population performance, and assessing efficacy of management interventions; moreover, such data are needed to guide policy decisions and to provide information to hunters and other publics. Ultimately, surveillance data will "be the foundation of an adaptive resource management plan for CWO in deer and elk; that plan will provide a mechanism for incorporating new knowledge gained through surveys, modeling, and experimental studies into a continuously evolving management program formulated to reduce the occurrence of CWO and minimize the risk of its spread to other native deer and elk populations in Colorado. MATERIALS AND METHODS Surveillance We monitored deer populations throughout Colorado for occurrence of CWO using a combination of targeted surveillance and harvest or road-kill surveys. These were organized and conducted as follows: Targeted (= clinical disease) surveillance: Deer showing clinical signs consistent with those seen in chronic wasting disease were collected by field personnel statewide and brain tissues examined for evidence of spongiform encephalopathy. The "suspect case" profile was defined as follows: • • Species: Age: mule deer white-tailed ~ 18 months deer �151 • Signs: emaciated and abnormal behavior &/or indifference to human activity &/or increased salivation &/or tremor, stumbling, incoordination &/or difficulty or inefficiency in chewing/swallowing increased drinking and urination &/or Where possible, submissions were subjected to complete necropsy; in some situations, only heads were available for examination and sampling. In all cases, histopathology of brain tissue (Williams and Young 1993) was used to diagnose CWO; in some cases, immunohistochemistry or other ancillary tests were used to confirm or support diagnoses. Harvest surveys: In order to obtain reliable estimates of CWO prevalence that will serve as a basis for monitoring responses to management interventions, we continued conducting harvest surveys on select deer populations. During the 1997-1998 hunting seasons, fresh brain and select lymphatic tissues were collected from deer harvested in enzootic GMUs; deer harvested or culled in other select GMUs throughout Colorado were also sampled as negative controls. Brain tissues were examined at the Colorado State University Diagnostic Laboratory for histopathological lesions (Williams and Young, 1993)or anti-PrP immunostaining reactions (O'Rourke et al., 1998) consistent with CWO infection. Because sample sizes for most individual GMUs were too small to provide reliable prevalence estimates by GMU, we pooled data by DAU for comparisons within and among species. We estimated specificity of imunohistochemistry using data from deer harvested outside known enzootic areas. Ages were estimated via replacement (4-6-mo-old, 16-18-mo-old, ~28-moold) and cementum annuli using first incisors from all positive deer and a random sample of negative deer (103 males and 100 females) harvested in four GMUs (9, 191, 19, 20); using these data, we compared sex-specific age frequency distributions for CWO-affected and unaffected male and female mule deer. Epizootiological Studies Epizootology of naturally-occurring CWO in captive mule deer and white-tailed ~ (Miller and Wild): Naturally-occurring CWO was a sporadic disease of resident FWRF mule deer prior to 1985 (Williams and Young, 1992), and also has occurred sporadically since 1994; no cases had been observed in resident white-tailed deer since their addition to FWRF in 1993. We maintained and observed 25 ~1-yr-old mule deer and 11 ~l-yr-old white-tailed deer in paddocks at CDOW's Foothills Wildlife Research Facility (FWRF) during July 1997-June 1998; 10 fawns were also born and recruited into the resident mule deer herd. Deer received natural forage, pelleted rations (high energy supplement and "browser" diet), and alfalfa hay; water and mineralized salt were available ad libitum. All deer were evaluated daily for clinical signs of CWO (and other health problems) in conjunction with routine feeding and handling activities. Resident mule deer also represented the source of infection (= "treatment") in an ongoing study of cattle susceptibility to CWO (see below). Cattle susceptibility to CWO (Williams and Miller): We continued monitoring cattle for clinical evidence of natural CWO transmission as part of a coordinated interagency effort to study cattle susceptibility to CWO. Twelve 4-mo-old calves purchased from a private ranch located near Sheridan, WY, were placed in paddocks at the FWRF with naturally-infected captive mule deer in July 1997 (see above for description of resident deer herd). Twelve 4-mo-old mule deer fawns were captured at the Rocky Mountain Arsenal National Wildlife �152 Refuge (RMANWR) in late September and placed in the same paddocks as contact controls for natural transmission (extensive ongoing surveillance of RMANWR deer has continued to confirm that this population is free of CWO). In a separate but related study, 20 additional 6-mo-old mule deer fawns were captured at the RMANWR in early December. Each fawn was given a single oral dose of about 5 g of fresh brain tissue homogenate from captive mule deer with clinical CWO. Fawns were then released into a separate paddock physically removed from the contact transmission study. These experimentally-infected fawns will serve two purposes: as controls for domestic calves orally inoculated with a single oral dose of about 50 g of this same CWO brain homogenate (Williams et al., 1998), and as subjects of a study on the pathogenesis of CWO in mule deer (Appendix A). RESULTS AND DISCUSSION Surveillance Targeted (= clinical disease) surveillance: Between June 1997 and May 1998, 6 chronic wasting disease (CWO) cases were diagnosed among 16 "suspect" deer submitted from known endemic portions of northeastern Colorado; CWO was not diagnosed in any of 7 additional "suspect" deer submitted from elsewhere in Colorado. All confirmed CWO cases originated in game management units (GMUs) where the disease had been detected previously. Males and females were represented equally. All CWO cases were observed and submitted during November-February, well within the October-April timeframe of most clinical case submissions (Spraker et al., 1997; Miller, 1997). Encephalitis, intoxication, neoplasia, and pneumonia were diagnosed among the 17 suspects not suffering from CWO; for most (9/17) non-CWO cases, no definitive explanation for clinical signs and condition could be determined from the samples submitted. Harvest surveys: We sampled and examined 1421 deer harvested in enzootic GMUs during 1997 archery, muzzleloader, and rifle seasons (Table 1). About 4.2% .(39/933) of deer harvested in Larimer bounty DAUs (04 and 010) tested. positive for CWO via immunostaining. Four GMUs (9, 191, 19, 20) yielded all of the positive deer detected in Larimer County units (04 and 010) (Table 1; Fig. 1); among these units, prevalence ranged from about 3-16%. Prevalence in sympatric elk populations (0-0.2%; Miller, 1998) was much lower than in deer (P < 0.009). Based on data from 1997 archery, muzzleloader, and rifle seasons, combined estimated mean CWO prevalence in 04 and 010 deer populations (4.2%) appeared to be essentially unchanged from 1995 (5.9%) and 1996 (5.7%); for the 3 GMUs (191, 19, 20) where sufficient samples sizes were available for 3 successive years (1995-1997), prevalence has not differed over time (P ~ 0.3). CWO was detected in about 1.1% (5/449) of deer harvested in South Platte River bottom units (DAU 044 plus GMUs 87, 90, 93, and 95·) (Table 1). Among river bottom and adjacent plains units, positive deer were harvested in GMUs 91, 95, 951, and 96 (Table 1; Fig. 1). None of the 171 deer harvested or culled in other select GMUs (mainly and 104) outside known enzootic areas tested positive for CWO. 66, 67, �153 rg 00 o o o o • MD PositiveMdpos.txt • ELK PositiveElkpos.txt .•. WTD PositiveWtdpos.txt WTD NegativeWtdneg.tx o MD NegativeMdneg.txt RiverColo riv CountiesCounties A . /\I o \1---' ___, s Figure 1. About 4.2% of deer harvested in larimer County data analysis units (OAUs) (04 or 010) and about 1.1% of deer harvested in South Platte River bottom units (OAU D44 plus GMUs 87,90, 93, and 95) tested positive for CWO via immunostaining. Most positive cases were from eastern larimer County. As reorted previously (Miller, 1997), the foregoing prevalence estimates may be somewhat liberal because the definition of "positive" included subclinical cases where either histopathological lesions or anti-PrP immunostaining reactions in brain tissue were observed. Of the 44 cases identified, none appeared to be suffering from clinical cwo. Twenty-six of the 44 (59%) positive deer showed both histopathological lesions and immunostaining; the other 18 were classified as positive solely on the basis of immunostaining reactions. Although no known "false positives" were identified among the 171 deer examined from outside known enzootic DAUs (specificity ~0.979), further evaluation of both sensitivity and specificity of existing diagnostic techniques still appears warranted. In contrast to observations derived from targeted surveillance data (Spraker et al., 1997; Miller, 1997), CWO prevalence among male (5.5%) and female (4.8%) mule deer did not differ (P = 0.86) for animals harvested in four Larimer County GMUs (9, 191, 19, 20). Antlerless permit numbers issued for 1997 failed to produce targeted sample sizes of 250 adult (~1 yr) does/DAU: we received 195 usable samples from 04 and 151 from 010, including samples from archery, muzzleloader, and rifle seasons. Additional does will be examined during 1998 to increase sample sizes for comparison of prevalence between sexes. Age distribution of CWO-positive mule deer did not differ from respective sexspecific age distributions of negative animals (Fig. 2). These data support the belief (Miller et al., 1998) that CWO epidemiology is driven primarily by lateral transmission. If maternal transmission were the primary mode of transmission (as is believed to be the case in scrapie of domestic sheep), then an age distribution skewed toward younger aged animals would be expected, particularly among does where older-aged animals are more abundant. In light of an estimated 18-24 mo incubation period for CWO in deer (Williams et al., 1998), the rarity of preclinical disease in yearlings (-16-17-mo-old deer) �154 appears to be the strongest argument against maternal transmission being the most common route for CWO transmission. Although survival rates for male mule deer are substantially lower than for females, CWO appears equally prevalent among males and females and uniformly distributed among respective age classes. These data may reflect the potential inefficacy of random culling in reducing CWO prevalence, although relatively high female densities and the potential for female-male transmission could confound such interpretations. Requiring head submissions increased survey sample sizes >4-fold compared to voluntary submissions in 1996: in 04/010, we collected 938 deer heads, as compared to -200 collected in 1996. Limited licenses also may have aided in increasing deer submissions. Based on harvest estimates from surveyed GMUs (COOW, unpubl. data), compliance with mandatory submission regulations was about 75%. Unstaffed barrel sites still appear to be the most efficient method for sample collection. Epizootiologieal Studies Epidemiology of naturally-occurring CWO in captive mule deer and whitetailed deer (Miller and Wild): Five mule deer and 5 white-tailed deer developed clinical cwo during the last biological year (May, 1997-April 1998) (Fig. 3); two additional mule deer cases and one additiona white-tailed deer case occurred during May-June 1998. In all, CWO affected about 20% (5/25) of the adult (~l-yr-old) mule deer and about 46% (5/11) of the adult white-tailed deer in resident FWRF herds during the last biological year; CWO accounted for 83% of adult mule deer mortality and 100% of adult white-tailed deer mortality. ~ CWO-poalilv. D N.D.1I •.• ~ ...•• r::: • ~• :::J 1:r 7-'D ••,D B. Female mule d.er Age cia •• Rgure 2. Age frequency distributions did not differ between CWO-positive and negative (A) male or (8) female mule deer sampled via harvest surveys. A.ilule du, · ." '0 -·,." z , •• , , •• 7 , •• 3 · ... '0 .! ,." z V •• r Since mule deer were reintroduced into Figure 3. Both mule deer and white-tailed deer herds FWRF in late 1990, 17 of 59 (29%) resident at CDOW's Foothills Wildlife Research Facility animals that survived to >1 yr of age are now infected with chronic wasting disease. have succumbed to CWO. The index case in the most recent epizootic occurred in October 1994; 4-5 clinical cases Annual clinical have occurred in each subsequent biological year (Fig. 3A). prevalence has ranged from 2-20% (Fig. 4). Generating comparable epizootic �155 behavior in a forecasting model (Miller and Mccarty, unpubl. data) requires a transmission coefficient (~) of about 3.5 infectious contacts/infectious animal/yr (Fig. 4); two assumptions of this spreadsheet model, no environmental source of infection and a l2-mo incubation period prior to onset of clinical disease, seem somewhat questionable in light of recent experiences and may need to be reevaluated. _a-wd,.."..DldMr 50 ~ fI) '0 [EJ~~oI" 25 40 20 30 15 ~ B r:::: '0 ~ fI) ..D E 20 ::::J Z 10 ~ l:. Based on observations made over the last 5 10 12 mo, CWO apparently can be an explosive disease 'in white-tailed deer populations. It follows that the relative rarity of o o 1995 1994 1996 1997 CWO in free-ranging white-tailed deer is Year probably more a function infrequent exposure rather than natural species resistance; white-tailed deer are relatively rare in the foothills of Figure 4. Modeled CWO dynamics eastern Larimer county where CWO is most simulated trends observed in FWRF deer prevalent. The source of infection for since 1994 when relatively high transmission our captive white-tailed deer herd is coefficients (13 = 3.5) were used. unclear. One possibility is introduction via two breeding males (C93, W93) that were intermittently housed with CWO-infected mule deer males; another possibility is transmission from infected mule deer via fenceline contact or environmental contamination. The essentially simultaneous occurrence of CWO in two does that had never been housed with mule deer seems to support the latter alternative. Cases in two additional animals 9 and 11 mo after the index case could be consistent with either a point source of exposure and variable incubation period or lateral transmission with a relatively short (-12 mo) incubation period. Data on pathogenesis, incubation periods, agent shedding, potential environmental sources of infection, and the relationship between infectiousness and onset of clinical signs for CWO in both white-tailed and mule deer are clearly needed to improve understanding of CWO epizootiology in both captive and free-ranging deer. Cattle susceptibility to CWO (Williams and Miller): One calf died from gastrointestinal obstruction and bloat about 2 wk after entering the deer paddocks, but the other 11 remained healthy throughout the first year of this 10-yr experiment. Similarly, 11 of the 12 control fawns from RMANWR remained healthy; one fawn that apparently failed to adjust to captivity died about 1 mo after capture, presumably from hypothermia secondary to malnutrition. Seven of the 25 >1-yr-old naturally-exposed resident FWRF deer developed clinical CWO and died or were euthanized during the first 12 mo of the study, thereby ensuring calves and control deer received some exposure to CWO. ACKNOWLEDGMENTS The statewide CWO monitoring and surveillance program described here relies heavily on efforts of dedicated field personnel throughout the Colorado Division of Wildlife, and truly represents a division-wide effort to improve our understanding and management of this important disease problems. In addition to those specifically listed, we collectively thank all of those �156 regional and area biologists, district and area wildlife managers, volunteers, deer and elk hunters, and others who assisted by submitting suspect cases, harvested animals, or road-killed animals throughout the year. LITERATURE CITED Miller, M. W. 1997. Monitoring. and managing wildlife chronic wasting disease in Colorado. in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-10, WP2, J17. Colorado Division of Wildlife, Fort Collins, Colorado, USA, pp. 37-46. 1998. Monitoring and managing wildlife chronic wasting disease in elk. in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-11, WP3002, T3. Colorado Division of Wildlife, Fort Collins, Colorado, USA, in press. , M. --wasting A. Wild, and E. S. Williams. 1998. Epizootiology disease in captive Rocky Mountain elk. J. Wildl. of chronic Dis. 34: 532-538. O'Rourke, K. I., T. V. Baszler, J. M. Miller, T. R. Spraker, I. SadlerRiggleman, and D. P. Knowles. 1998. Monoclonal antibody F89/160.1.5 defines a conserved epitope on the ruminant prion protein. J. Clin. Microbiol. 36: 1750-1755. Spraker, T. R., M. W. Miller, E. S. Williams, D. M. Getzy, W. J. Adrian, G. G. Schoonveld, R. A. Spowart, K. I. O'Rourke, J. M. Miller, and P. A. Merz. 1997. spongiform encephalopathy in free-ranging mule deer (Odocoileus hemionus), white-tailed deer (Odocoileus virginianus), and Rocky Mountain elk (Cervus elaphus nelsoni) in northcentral Colorado. J. Wildl. Dis. 33:1-6. Williams, E. S., and S. Young. 1980. Chronic Wasting disease of captive mule deer: A spongiform encephalopathy. Journal of Wildlife Diseases 16: 89-98. , and --Journal of __ ~' and Scientifique 1982. Spongiform encephalopathy Wildlife Diseases 18: 465-471. of Rocky Mountain elk. 1992. Spongiform encephalopathies in Cervidae. Revue et Technique Office International des Epizooties 11: 551-567. , --deer and 1993. Neuropathology of chronic wasting disease in mule (Odocoileus hemionus) and elk (Cervus elaphus nelsoni). Veterinary Pathology 30: 36-45. , M. W. Miller, --Susceptibility of T. J. Kreeger, H. Van Campen, and T. R. Spraker. 1998. cattle to cervid spongiform ~ncephalopathy. unpublished grant proposal, submitted to USDA, CSREES, National Research Initiative Competitive Grants Program, 88 pp. c Wildlife Research Veterinarian �157 Table 1. Results of 1997 CWD harvest surveys -- archery, muzzleloader, & rifle seasons. DEER DAU GMU D4 7 29 0 8 160 0 191 212 10 74 12 19 123 6 Total 598 28 20 335 11 Total 335 11 29 39 0 Total 39 0 87 37 0 90 1 0 91 54 1 . 92 41 0 93 53 0 94 55 0 95 97 2 951 56 1 96 55 1 449 5 9 DI0 Boulder Plains Total # Examined # Positive Prevalence (95% CI) 0.047 (0.031-0.067) 0.033 (0.016-0.058) 0 (0-0.09) 0.011 (0.004-0.025) �158 �159 AJ;lpendix A STUDY PLAN Pathogenesis Elizabeth of Chronic Wasting S. Williams Disease and Michael in Mule Deer W. Miller Need Chronic wasting disease (CWO) is a transmissible spongiform encephalopathy (TSE)of native North American deer and elk (Williams and Young, 1980, 1982, 1992). The neuropathology of clinical CWO is well-described (Williams and Young, 1980, 1982, 1992, 1993; Spraker et al., 1997), but CWO pathogenesis remains largely unstudied. Among clinical CWO cases, the parasympathetic nucleus of the vagus and the olfactory stria are most consistently and severely affected (Williams and Young, 1993). Typical but less abundant lesions occur in the parasympathetic nucleus of the vagus, and sometimes the olfactory stria, in apparently subclinical CWO cases detected during harvest surveys (E. S. Williams, unpubl. data; T. R. Spraker, pers. comm.). Brain and various lymphoid tissues often stain intensively with anti-prion protein (PrP) immunostaining in clinical cases (Williams and Young, 1992; Spraker et al., 1997; E. S. Williams, unpubl. data; T. R. Spraker, pers. comm.). Immunostaining of brain and select lymphoid tissues, particularly tonsil, are usually positive in subclinical cases. Positive staining of tonsil and brain tissue can occur in the absence of spongiform lesions. Lymphoid Prpsc propagation apparently precedes development of detectable neurological lesions in scrapie (Schreuder et al., 1996, van Keulen et al., 1995, 1996), but not bovine spongiform encephalopathy (Wells et al., 1996). The presence of Prpsc in central nervous system and peripheral tissues has lead to development of immunohistochemistry (Miller et al., 1993; van Keulen et al., 1995, 1996; Schreuder et al., 1996) and immunoblotting (Farquhar et al., 1989; Ikegami et al., 1991; Race et al., 1992) as preclinical and clinical diagnostic tests for scrapie in sheep. These antemortem tests are based on biopsy and examination of lymphoid tissues, typically tonsil, lymph node, and spleen for Prpsc• Positive staining of lymphoid and salivary gland tissues in the absence of either lesions or staining in brain tissue has also been observed in deer and elk, but interpretation of these observations remains equivocal. Improved understanding of CWO pathogenesis would clearly facilitate the interpretation of diagnostic findings in subclinical CWO suspects detected via harvest surveys. Such understanding would also enhance interpretation of data being gathered in studies designed to evaluate antemortem diagnostic tests for CWO. Moreover, reliable pathogenesis data may offer valuable insights into initiation, duration, and routes of agent shedding, as well as other important aspects of CWO epizootiology. Here, we propose to study the pathogenesis of CWO in mule deer after oral exposure to infectious brain material. Objectives The specific objectives of this study are to: (1) describe the pathogenesis of CWO in mule deer after oral exposure to infectious material using histopathology, immunohistochemistry, and Western blot analyses; and (2) compare susceptibility, pathogenesis, and incubation periods between male and female deer. Additionally, animals challenged in this study will serve as controls for ongoing cattle challenge trials (Williams et al., 1997). �160 Materials and Methods We will study the pathogenesis of CWO in mule deer after oral exposure to infectious brain material. Twenty 5- to 6-mo-old mule deer fawns (10 males, 10 females) will be captured from the Rocky Mountain Arsenal National Wildlife Refuge (RMANWR) and transported to the Colorado Division of Wildlife's Foothills Wildlife Research Facility (FWRF) in Fort Collins, Colorado. Previous surveillance work has revealed that deer residing at RMANWR are presently unaffected by CWO, thereby ensuring fawns will not be exposed to CWO until inoculated experimentally. For capture, fawns will be anesthetized with tiletamine HCl and zolazepam (Telazol~; 5 mg/kg) and xylazine HCl (2.5 mg/kg) or carfentanil HCl (0.03 mg/kg) and xylazine (1 mg/kg) delivered intramuscularly (1M) via projectile syringe; if carfentanil is used, anesthesia will be antagonized with naltrexone HCl (100 mg/mg carfentanil) delivered intravenously (IV)(25%) and subcutaneously (SC) (75%) after initial handling and processing. Additional xylazine (20-100 mg IV or 1M) will be administered as needed to keep fawns sedated until they arrive at FWRF. (See appendix for detailed capture protocols.) Upon arrival at FWRF, residual sedation will be antagonized with IV yohimbine HCl (0.25 mg/kg). Once fawns are able to swallow effectively, each will receive about 5 g of homogenized brain material pool collected from mule deer previously diagnosed with spongiform encephalopathy; presence of scrapieassociated fibrils in this homogenate was previously confirmed via negativestain electron microscopy (E. S. Williams, unpubl. data). This homogenate is the same being used for oral and intracranial cattle challenges (Williams et al., 1997). Homogenate will be deposited into the posterior oropharynx using a modified syringe. Once fawns have swallowed the homogenate, they will be released into a 3 ha paddock. Alfalfa hay, pelleted supplemental diets (highenergy and "browser" rations), mineralized salt blocks, and water will be provided ad libitum. All fawns will be observed daily by animal caretakers and evaluated at least monthly by an attending veterinarian for signs of CWO. To study CWO pathogenesis, two fawns' (one male, one female) will be randomly sacrificed at 3, 6, 12, 18, 24, 30, 36, or 42 mo after oral challenge. At the time of sacrifice, fawns will be anesthetized with tiletamine HCl and zolazepam (Telazol~; 5 mg/kg) and xylazine Hel (2.5 mg/kg) delivered via projectile syringe. We will collect blood, saliva, feces, and cerebrospinal fluid from each fawn, then administer about 400 mEq KCl intravenously to induce cardiac arrest. The incubation period of CWO after oral challenge is unknown; however, because clinical CWO developed in mule deer 18 to 24 mo after intracranial challenge (E. S. Williams, unpubl. data), we anticipate that most study animals surviving >24 mo will develop Cwo 24 to 36 mo after oral challenge. Deer developing severe clinical CWO (characteristic behavioral changes accompanied by estimated >20% weight loss) will be euthanized as described above. Two age-matched control deer (one male, one female) will be collected from the RMANWR on the same schedule as challenged deer are sacrificed. Control deer will be anesthetized in the field as described above, sampled, and euthanized with KCl; if field anesthesia becomes infeasible, control deer will be killed by shooting in the neck with a highpowered (~0.223 cal) rifle. Saliva, feces, blood, and cerebrospinal fluid will be stored at -70 C for potential evaluation via Western blot or infectivity studies. Carcasses will be transported to the Wyoming State Veterinary Laboratory (WSVL) for complete necropsy and tissue sampling. Samples collected will include brain, spinal �161 cord, and numerous other tissues (Table 1). These tissues will be divided and subsamples fixed in 10% neutral phosphate-buffered formalin or in PLP solution (periodate, lysine, paraformaldehyde) at a concentration of 2% paraformaldehyde (van Keulen et al., 1996); additional subsamples will be stored unfixed at -70 C for studies of Prpsc accumulation. Carcasses will be incinerated once sampling is completed. Tissues will be examined via histopathology, immunocytochemistry, Western blot, and/or negative-stain electron microscopy. Histopathology will be conducted as previously described (Williams and Young, 1993). Central nervous system tissues will be removed within 4 hr following euthanasia. Brains will be sagittally sectioned and half fixed in 10% neutral phosphate-buffered formalin. Samples of spinal cord and other tissues (Table 1) will be similarly fixed. The other half of the brain and portions of the spinal cord will be frozen at -70 C. Paraffin blocks will be prepared from olfactory tubercle and cortex; cerebral cortex (frontal, parietal, temporal, and occipital lobes); basal ganglia; three levels of thalamus; two levels of mesencephalon; three levels of pons and cerebellum; medulla oblongata at the obex; medulla caudal to the obex; and multiple levels of spinal cord to include cervical, thoracic, and lumbar regions. Tissues will be sectioned at 5-6 ~m and stained with hematoxylin and eosin, Bodian's silver stain, luxol fast blue stains, and glial fibrillary acidic protein (Dako, Carpinteria, California) immunocytochemistry for astrocytosis as appropriate. Lesions will be compared to those previously described for CWO in mule deer and elk and other natural TSEs of animals (Williams and Young, 1993; Hadlow, 1996). We will test for Prpsc in formalin- or PLP solution-fixed, paraffin embedded tissues using minor modifications of the technique of van Keulen et al. (1995). We will use a polyclonal rabbit antiserum against ME7 scrapie strain passaged in mice (Rubenstein et al., 1986) (Department of Virology, New York State Office of Mental Retardation and Developmental Disabilities, Staten Island, New York) or mouse monoclonal antibody against ovine Prpsc (F89/160.1.5; O'Rourke et al., 1997) (USDA/ARS, Pullman, Washington). Briefly, paraffin embedded tissue sections will be deparaffinized, treated with 99% formic acid for 30 minutes, rinsed with PBS, and autoclaved for 10 minutes at 122 C in 10 x Automation buffer. Sections will be incubated in 4% normal goat serum for 20 minutes, incubated overnight at 4 C in primary antibody at 1:1500 dilution, rinsed, and incubated for 30 minutes with biotinylated anti-rabbit or anti-mouse IgG (Vector Laboratories, Burlingame, California). Slides will be rinsed, and ABC reagent (Vectastain Elite ABC kit, Vector Laboratories) applied for 30 minutes, rinsed and incubated with AEC substrate solution (Dako Laboratories, Carpinteria, california) for 10 minutes. Slides will be counter stained with Harris hematoxylin, rinsed, "blued" with lithium carbonate, rinsed again, and coversli~d. Positive and negative brain sections, normal rabbit serum, and PBS will be used as controls. Adequate primary antibody is available for the entire study. L Western blot assays will be conducted with modifications of the techniques of Rubenstein et al. (1986) and Collinge et al. (1996). Brain or lymphoid tissues will be homogenized in Tris buffer saline (TBS) at 50 mg/500 ~l followed by a 5 minute low speed centrifugation. 100 ~l supernatant is combined with 6 ~l 22% sarcosyl in TBS, mixed, 4 ~l (5mg/ml) proteinase K is added, mixed again, then incubated at 37 C for 1-2 hours. Aliquots will be collected for protein concentration determination using Pierce's Micro BCA protein assay. Equal volume of concentrated sample buffer (120 roM Tris-HCl pH 6.8, 142 roM BME, 200 roM DTT, 4% SDS, 0.02% BPB, 20% glycerol) will be added and the sample boiled for 5 minutes. Hot samples will be loaded on Mini-Vertical 4%/12% SDS-PAGE �162 (Laemmli). The samples will be electrophoresed at 100 v for 1.5-2 hours with BioRad's Protein SOS-PAGE molecular weight markers. A semidry transfer to 0.45 ~m pure nitrocellulose will be conducted using Towbin buffer according to manufacturer's instructions. Post-transfer, the gel will be Coomassie Blue G250 stained while the membrane is floated on TBST20 (10 roM Tris pH 8.0, 150 roM NaCI, 0.1% Tween2o) until wet, then submerged and rinsed. The membrane will be incubated in 1% NGS -TBST2o for 30 - 60 minutes at room temperature, washed two-three times for 10 minutes each in TBST2o, and TBST20 1:5,000 diluted primary antibody (polyc1onal rabbit or monoclonal mouse anti-Prpsc) added to the blot and incubated 2 hours at room temperature or overnight at 4 C. The antibody will be removed, the membrane washed two-three times for 10 minutes each, and secondary antibody (Promega's goat anti-rabbit conjugated alkaline phosphatase) diluted in TBST20 (per manufacturer's instructions) added and incubated 2 hours at room temperature. The membrane will be again washed two to three times followed by three 2 minute washes in distilled water and then a 5 minute TBS wash done twice. Promega's BCIP/NBT color development solution (made according to manufacturer's instructions) will be added and development monitored. The reaction will be stopped by several distilled water rinses. The membrane and stained gel will be recorded via CCO camera, digitized, and quantitated (Bio-Rad Molecular Analysis System). Scrapie-associated fibrils will be purified using a modification of the techniques of Hilmert and Oiringer (1984). One to two g of brain will be homogenized in 10% sarcosyl (pH 7.4) and N-octano1 added for a 30 minute incubation at room temperature. The homogenate will be centrifuged for 10 minutes at 3,000 x G (JA-20 rotor, J-21 Beckman centrifuge) and supernatant collected and centrifuged for 30 minutes at 22,000 X G. The supernatant will then be transferred to Beckman quick seal tubes and centrifuged at 215,000 X G (75T rotor, L8-80 Beckman centrifuge) for 2 hours. The pellet will be resuspended in 1% sarcosyl and 10% NaCI, stirred for 60 minutes at 37 C, and microfuged for 15 minutes. The pellet will then be resuspended -in 1% sarcosyl, 10% NaCI, and 5 ~g proteinase Klml added and stirred for 2 hours at 37 C. The suspension will be microfuged for 10 minutes. Negative staining will be conducted according to Scott et ale (1987, 1990). The pellet will be resuspended in 50 ~l distilled water and a 300 mesh plastic coated, carbon stabilized grid floated on a drop of suspension for 10 seconds, blotted and negative stained with 2% potassium phosphotungstate, pH 6.6. Some additional modifications of the procedures for SAF detection (Stack et al., 1995; 1996) are currently being tested at WSVL. We will describe distribution of Prpsc and microscopic lesions as determined by each of the foregoing assays, and relate these to both the chronological progression of clinical CWO and the potential shedding of agent from infected deer. Because our study is largely descriptive, no statistical analyses of pathology data will be attempted. Among fawns surviving long enough to develop clinical CWO, mean incubation times for males and females will be compared using Student's t-test (a = 0.1), and the proportions of males and females developing clinical cwo will be compared using Fisher's exact probability test (a 0.1). = Literature Cited Collinge, J., K. C. L. Sidle, J. Meads, J. Ironside, and A. F. Hill. 1966. Molecular analysis of prion strain variation and the aetiology of 'new variant' CJO. Nature 383: 685-690. Farquhar, C. F., R. A. Somerville, and L. A. Ritchie. 1989. Post-mortem immunodiagnosis of scrapie and bovine spongiform encephalopathy. Journal of Virological Methods 24: 215-222. �163 Hadlow, W. J. 1996. Differing neurohistologic images of scrapie, transmissible mink encephalopathy, and chronic wasting disease of mule deer and elk. In Bovine spongiform encephalopathy The BSE dilemma, Vol. C. J. Gibbs, Jr. (ed.). Springer-Verlag, New York, New York, pp. 122-137. Hilmert, H., and H. Diringer. 1984. A rapid and efficient method to enrich SAF-protein from scrapie brains of hamsters. Biosciences Reports 4: 165170. Ikegami, Y., M. Ito, H. Isomura, E. Momotani, K. Sasaki, Y. Muramatsu, N. Ishiguro, and M. Shinagawa. 1991. Pre-clinical and clinical diagnosis of scrapie by detection of PrP protein in tissues of sheep. Veterinary Record 128: 271-275. Miller, J. M., A. L. Jenny, W. D. Taylor, R. F. Marsh, R. Rubenstein, and R. E. Race. 1993. Immunohistochemical detection of prion protein in sheep with scrapie. Journal of Veterinary Diagnostic Investigation 5: 309-316. Race, R. E., D. Ernst, A. L. Jenny, W. D. Taylor, D. Sotton, and B. Caughhey. 1992. Diagnostic implications of detection of proteinase K-resistant protein in spleen, lymph nodes, and brain of sheep. American Journal of Veterinary Research 53: 883-889. Rubenstein, R., R. J. Kascsak, P. A. Merz, M. C. Papini, R. I. Carp, N. K. Robakis, and H. M. Wisniewski. 1986. Detection of scrapie-associated fibril (SAF) proteins using anti-SAF antibodies in non-purified tissue preparations. Journal of General Virology 67: 671-681. Schreuder, B. E. C., L. M. J. van Keulen, M. E. W. Vromans, J. P. M. Langveld, and M. A. Smits. 1996. Preclinical test for prion diseases. Nature 381: 563. Scott, A. C., S. H. Done, C. Venables, and M. Dawson. 1987. Detection of scrapie-associated fibrils as an aid to the diagnosis of natural sheep scrapie. Veterinary Record 120: 280-281. Scott, A. C., G. A. H. Wells, M. J. Stack, H. White, and M. Dawson. 1990. Bovine spongiform encephalopathy: Detection and quantitation of fibrils, fibril protein (PrP) and vacuolation in brain. Veterinary Microbiology 23: 295-305. Spraker, T. R., M. W. Miller, E. S. Williams, D. M. Getzy, W. J. Adrian, G. G. Schoonveld, R. A. Spowart, K. I. O'Rourke, J. M. Miller, and P. A. Merz. 1997. Spongiform encephalopathy in free-ranging mule deer (Odocoileus hemionus), white-tailed deer (0. virgianus), and Rocky Mountain elk (Cervus elaphus nelsoni) in northcentral Colorado. Journal of Wildlife Diseases 33: 1-6. Stack, M. J., A. M. Aldrich, A. D. Kitching, and A. C. Scott. 1995. Comparative study of electron microscopal techniques for the detection of scrapie-associated fibrils. Research in Veterinary Science 59: 247-254. Stack, M. J., A. M. Aldrich, A. D. Kitching, and A. C. Scott. 1996. Comparison of biochemical extraction techniques for the detection of scrapieassociated fibrils in the central nervous system of sheep naturally affected with scrapie. Journal of comparative Pathology 115: 175-184. van Keulen, L. J. M., B. E. C. Schreuder, R. H. Meloen, M. Poe len-van den Berg, G. Mooij-Harkes, M. E. W. Vromans, and J. P. M. Langerveld. 1995. Immunohistochemical detection and localization of prion protein in bran tissue of sheep with natural scrapie. Veterinary Pathology 32: 299-308. van Keulen, L. M. J., B. E. C. Schreuder, R. H. Meloen, G. Mooij-Harkes; M. E. W. Vromans, and J. P. M. Langveld. 1996. Immunohistochemical detection of prion protein in lymphoid tissues of sheep with natural scrapie. Journal of Clinical Microbiology 34:1288-1231. Wells, G. A. H., M. Dawson, S. A. C. Hawkins, A. R. Austin, R. B. Green, I. Dexter, M. W. Horigan, and M. M. Simmons. 1996. Preliminary observations on the pathogenesis of experimental bovine spongiform encephalopathy. In �164 Bovine spongiform encephalopathy The BSE Dilemma, Vol. C. J. Gibbs, Jr. (ed.). Springer-Verlag, New York, New York, pp. 28-44. Williams, E. S. and S. Young. 1980. Chronic wasting disease of captive mule deer: A spongiform encephalopathy. Journal of Wildlife Diseases 16: 89-98. Williams, E. S. and S. Young. 1982. Spongiform encephalopathy of Rocky Mountain elk. Journal of Wildlife Diseases 18: 463-471. Williams, E. S., and S. Young. 1992. Spongiform encephalopathies of Cervidae. Scientific and Technical Review Office of International Epizootics 11: 551-567. Williams, E. S. and S. Young. 1993. Neuropathology of chronic wasting disease of mule deer (Odocoileus hemionus) and elk (Cervus elaphus nelsoni). Veterinary Pathology 30: 36-45. Williams, E. S., M. W. Miller, T. J. Kreeger, and H. Van Campen. 1997. Susceptibility of cattle to cervid spongiform encephalopathy. Unpublished study plan, 66 pp. Table 1. Tissues to be collected and detection of Prpsc. Nervous Tissue brain (multiple levels) pituitary CSF dura spinal cord (cervical, thoracic, lumbar) dorsal root ganglia trigeminal ganglia stellate ganglia sciatic nerve radial nerve Muscle Tissue diaphragm semitendinosus muscle triceps muscle longissimus dorsi muscle Alimentary Tissue tongue submandibular lymph node parotid salivary gland esophagus rumen omasum abomasum duodenum distal ileum and Peyer's patches spiral colon feces pancreas liver from deer exposed to CWO for histopathology Lymphoid Tissue spleen thymus tonsil submandibular lymph node retropharyngeal lymph node bronchial lymph node mediastinal lymph node hepatic lymph node mesenteric lymph nodes ileocecal lymph node hepatic lymph node superficial cervical lymph node popliteal lymph node Other Tissues kidney urine adrenal gland .lung nasal mucosa left ventricle blood (serum, buffy bone marrow skin (head) bone (rib) coat, clot) �165 Colorado Division Wildlife Research July 1998 of Wildlife Report JOB PROGRESS State of Colorado Project Cost Center W-153-R-11 Work Package Mammals 3001 3430 Research Deer Conservation Task No. Regulation of Mule Deer Population Growth by Fertility Control .Period Covered: Author: REPORT July 1, 1997 - June 30, 1998 Dan L. Baker Personnel: T. M •.Nett, J. Griess, M. A. Wild ABSTRACT Controlling the abundance of animals is fundamental to contemporary wildlife management. This is particularly true for wild ungulates. The most compelling motivation for regulating ungulate numbers is that overabundance causes problems that can be biological, economical, or political in scope. Resolving these problems requires controlling population growth. Achieving such control by traditional methods such as hunting or culling may not always be feasible. In these situations fertility control offers a promising alternative. Here, we conduct research to develop and test a practical and acceptable method of contraception in mammalian wildlife. We propose to use GnRH-toxin conjugates to selectively destroy gonadotroph cells in the anterior pituitary thereby preventing fertility in males or females. Using captive mule deer, we designed an experiment to evaluate the effective duration of GnRH-toxin conjugates. Together with knowledge of the population dynamics of the Rocky Mountain Arsenal mule deer population we began development of an interactive simulation model that will allow wildlife managers to choose specific tactics for applying contraceptive treatments. ��167 REGULATION OF MULE DEER POPULATION GROWTH BY FERTILITY CONTROL Dan L. Baker P. N. OBJECTIVES 1. To develop a practical in mammalian species which nuisance. 2. To demonstrate and acceptable technology to inhibit reproduction cause damage or constitute a significant public the feasibility 3. To predict population simulation modeling. impacts of such technology of alternative SEGMEHT 1. To develop and test GnRH-toxin in a field contraceptive application. regimes using OBJECTIVES conjugate 2. To develop a simulation model to evaluate regulate mule deer populations at the RMA. in captive mule the ability deer. of contraceptives to INTRODUCTION controlling the abundance of animals is fundamental to contemporary wildlife management. This is particularly true for wild ungulates. The most compelling motivation for regulating ungulate numbers is that overabundance causes problems that can be biological, economical, or political in scope (Jewel and Holt, 1981). Resolving these problems requires controlling population growth. Wild ungulate populations have traditionally been regulated by influencing death rates using controlled harvest or culling. However, there are an increasing number of circumstances where these traditional methods are not feasible. As a result, there is growing interest in controlling the growth of animal populations by influencing·fertility (Kirkpatrick and Turner 1985; Bomford, 1990; Garrott, 1995). Several techniques have been successful in controlling fertility of individuals (Plotka and Seal, 1989; Kirkpatrick and Turner, 1991; Plotka e~ al., 1992; Turner e~ al., 1996), but it remains unclear whether they are effective in regulating the growth of populations. Simulation models of the dynamics of ungulate populations regulated by contraception suggest that permanent sterilization of females offers one of the most efficacious and efficient approaches to population control (Boone and Wiegert, 1994; Garrott et al., 1992, Garrott, 1995f. One of the most promising new techniques to permanent sterilization involves linking analogs of gonadotropin releasing hormone (GnRH) to cytotoxins to form a GnRH-toxin conjugate. GnRH is a brain peptide that binds to receptors on gonadotrophs and stimulates synthesis and secretion of luteinizing hormone (LH) and follicle stimulating hormone (FSH). These two hormones, known as gonadotropins, control the proper functioning of the ovaries in the female and the testes in the male. By chemically linking a superactive analog of GnRH to a cytotoxin, it should be possible to specifically target that toxin to gonadotroph cells in the anterior pituitary gland. Since the gonadotrophs will internalize the GnRH-toxin complex as part of the normal process of de- �168 activation, the internalized toxin will cause death of the gonadotroph. Once the gonadotrophs are selectively destroyed, LH and FSH are unavailable for gamete production by the ovaries and testes and the animal becomes permanently sterile. Since GnRH is highly conserved across species, a single GnRH-toxin conjugate has the potential to induce sterility in both sexes and numerous vertebrate species. Many unanswered questions must be addressed before this potential contraceptive can be considered an effective and acceptable method of population control in free-ranging wild ungulates. Research is needed to evaluate the effective duration of this technique in deer, the effect on deer social structure, genetic variability of breeding populations, and the practicality of application in wild herds. In this study, we address unresolved questions in using GnRH-toxin conjugates as contraceptives in deer. Our objectives were: 1) to determine the most effective dose of GnRH-toxin and to evaluate the effective duration of treatment. 2) to develop an interactive simulation METHODS Study 1: Evaluate effective AND duration model conjugate to evaluate in mule deer the MATERIALS of GnRH-toxin conjugate Previously, we conducted controlled experiments with tame, penned mule deer to determine the most effective dose of GnRH analog and the season of year when treatments would be most effective in preventing fertility (Baker 1997). Here, we evaluate the effectiveness of GnRH-toxin conjugate in preventing pregnancy, the duration of effectiveness, and physiological side effects (if any) of treatments. We will evaluate the effectiveness of GnRH-toxin conjugate in 4 ovariectomized captive mule deer. Based on previous studies with domestic sheep, this is the minimum sample size for meaningful results. This experiment has never been conducted with any other wild species, thus results from this study will be used to estimate sample sizes for future investigations. The deer selected for this study will be hand-reared adult females maintained in captivity at the Foothills Wildlife· Research Facility, Fort Collins, Colorado. Protocol for Ovariectomy in Mule Deer Tonic secretion of pituitary LH is the result of an interplay between a stimulatory input from the brain and an inhibitory feedback from the gonads. In the intact female, estradiol secreted from the 90nads is a potent negative feedback hormone on LH secretion during anestrus (Goodman and Karsch 1980). Since measurement of LH secretion is the primary indicator of GnRH-toxin conjugate effectiveness, it is imperative that female mule deer in this experiment be ovariectomized. Ovariectomies of mule deer will be conducted at FWRF during the week of May 11, 1998. Deer will be isolated and fasted for about 24 hr prior to surgery to alleviate regurgitation and aspiration of rumen contents. On the day of surgery, anesthesia will be induced with intramuscular (1M) administration of 100 mg xylazine wlo 500 mg ketamine depending on response of the animal. Deer �169 will then be intubated and surgical anesthesia will be maintained using a rebreathing circuit with isoflurane. Anesthetized deer will be prepared for surgery by clipping hair in the abdominal area. They will then be carried to a designated surgery area, placed in right lateral recumbency, and the surgical site prepared using standard surgical scrub. The surgical area will be prepared with sterile towels and a surgical drape. Ovariectomy will be performed via mid-ventral laparotomy and will require 20 - 30 min per animal. Surgeons will be attired in a sterile gown, mask, cap, disposable shoe covers and sterile gloves. Surgical assistants will wear cap, mask, and disposable shoe covers. The ovarian artery will be ligated prior to removal of the ovary. The incision will be closed using a continuous .suture and the skin closed with interrupted mattress stitches. We will reverse anesthesia with yohimbine at a dose of 0.2 mg/kg (IV). To minimize infection, deer will be given ceftiofur sodium (1.1 mg/kg IV) administered perioperatively. Phenylbutazone (4 mg/kg) will be administered orally when deer have partially recovered from anesthesia and every 48 hr for up to one week, if needed. Animals will be placed in isolation pens for 24 hr and will be observed every 5 min during recovery from anesthesia. This procedure follows ARBL Standard Operating Procedure #1 Protocol for Ovariectomy/Luteectomy in Ewes and was modified for use in mule deer at FWRF. GnRH-toxin Conjugate Proiocol Approximately 6 weeks following ovariectomy, GnRH-toxin will be administered. Four ovariectomized deer will be moved from 5 ha pastures to individual isolation pens, sedated with xylazine (100 mg 1M), and administered IVan optimum dose of GnRH-toxin (3 ~g/50 kgBW). Deer will then be placed in individual isolation pens and fitted nonsurgically with indwelling jugular catheters. Indwelling catheters will remain in deer for up to 6 weeks and will be checked and flushed daily. Deer will remain in individual isolation pens for the first 6 weeks of the trial. This will minimize tranquilization and general handling of deer and allow daily evaluation of intake and general health of experimental animals. After 6 weeks, catheters will be removed and deer will be returned to 5 ha pastures. For subsequent trials, deer will be removed from 5 ha pastures one day prior to GnRH trials and fitted with indwelling jugular catheters. Sampling will be conducted the next day, catheters removed and deer returned to 5 ha pastures. One week after GnRH-toxin treatment, GnRH analog challenge trials will be conducted. Experimental animals will be moved to individual isolation pens and 3 ~g/50 kg BW GnRH analog (Baker 1997) will be administered through the indwelling cannula. Blood samples (5ml) will be collected at 0, 30, 60, 90, 120, lBO, 240, 300, and 360 min postinjection. After collections, blood will be held at 4 C for 24 hours and serum obtained by centrifugation. Serum will be stored at -20 C until analyzed. GnRH analog challenge trials will be repeated each week for 6 weeks, then twice monthly for 3 months, then once monthly for 1 year. Serum concentrations of LH will be quantified by means of ovine LH RIA (Niswender et al. 1969). The limit of sensitivity of the assay is 0.4 ng/ml. Response of the pituitary to GnRH-toxin conjugate will be assessed by 1) maximum LH response achieved postinjection, 2) total amount of LH secreted (ng/ml/min); estimated by calculating the area under the LH curve, and 3) an LH response of < 2.5 ng/ml following GnRH challenge will be considered sufficiently low to prevent ovulation. Data will be analyzed using least squares ANOVA for General Linear Models and the SAS Interactive Matrix Language. Response to treatment will be analyzed with two-way factorial �170 analysis of variance for a randomized complete block design with repeated measures structure. Factors will be GnRH-toxin conjugate and time. We will use a priori orthogonal contrast to test for differences among individual means. Schedule Pate April 23 Submit Actiyity research proposal is approved, to CPOW Animal perform Care and Use Committee May 12 If proposal deer June 23 Collect pre-treatment toxin conjugate June 30 Conduct 1 week GnRH analog challenge trial July 7 Conduct 2 week GnRH analog challenge trial July 14 Conduct 3 week GnRH analog challenge trial July 21 Conduct 4 week GnRH analog challenge trial LH samples, ovariectomies and treat August 4 Conduct 6 week GnRH analog challenge trial August 18 Conduct 8 week GnRH analog challenge trial of these trials Future dates will be dependent being of experimental animals. on results on captive mule deer with GnRH- and health and well- Study 2: Develop a simulation model to evaluate the ability of contraceptives to regulate mule deer populations at the Rocky Mountain Arsenal National Wildlife Refuge. Previously, we developed a general mathematical model to evaluate the ability of contraception to regulate animal populations (Baker and Hobbs 1996). However, in order for wildlife managers at the (RMA) to make decisions on the best tactics for treating this specific population of mule deer, we developed an interactive simulation model that will combine knowledge of mule deer population dynamics with an understanding of the constraints intrinsic in the GnRH-toxin conjugate technique. Using this model, managers will be able to evaluate the probable consequences of their decisions on implementing contraceptive regimes. Critical to model performance is knowledge of the sex and age composition the RMA deer herd before and after fertility control treatments are implemented. We provided this information by conducting herd composition surveys during 1996 and 1997. of Peer populations were sampled by establishing three permanent routes within RMA boundaries. Each route followed established roadways and was chosen to observe the maximum number of deer within each representative area of RMA. The length of each route was determined by the driving distance that could be covered by one vehicle during the period 0700 - 1200. One pair of observers �171 sampled one of three routes each day; the same pair of observers were consistent across all days. The deer population was sampled near the peak of the breeding season each year (approx. Nov. 22). Classification of deer were made from a vehicle using 8 x 50 binoculars and 40x spotting scopes. No classifications were attempted for deer that were further than 200 m from the observer or for deer that were bedded. Deer were classified by species as white-tailed or mule deer and by sex and age as mature buck, immature buck, doe, and fawn (Dasman and Taber 1956). We also recorded the number of abnormal deer observed. An individual group was determined by observing the behavior and spatial distribution of deer. To increase consistency among observers, we chose 25 m to be the maximum distance between individual groups. Prior to conducting sex and age classifications, all observes spent at least one day together independently classifying the same deer from as many groups as necessary to consistently obtain identical classifications. RESULTS AND DISCUSSIQH Study 1: Evaluate effective duration of GnRH-toxin conjugates Four female mule deer were successfully ovariectomized on May 11, 1998. Following surgery blood samples were collected from all deer to determine levels of LH (Table 1). After 8 weeks, LH concentrations had not increased sufficiently above baseline to conduct GnRH-toxin conjugate experiments. Blood samples are currently being collected each month to monitor LH levels. Previous studies indicate that for wild ungulates that breed seasonally, levels of circulating LH are significantly greater during the breeding season (fall) than during anestrus (May - Oct) (Curlewise et al. 1991, McCloud et al. 1991, Baker et al. 1995). Thus, we anticipate conducting these trials in late October when LH concentrations are increasing. Study 2: Develop to regulate mule Wildlife Refuge. a simulation model to evaluate the ability of contraceptives deer populations at the Rocky Mountain Arsenal National During Nov 20 - 22, 1997 we classified an average of 581 (se ± 7.0) mule deer (Table 2) and 197 (se ± 17.3) white-tailed deer each day (Table 3). Compared to 1996, this represented an increase of 30.1 % for mule deer and 33.2 % increase in white-tailed deer. We observed an increase in the total number of individuals observed in all age classes and for both species of deer. For both mule deer and white-tailed deer, the greatest increase was observed in young bucks (58.4 % in mule deer; 71.0 % in white-tailed deer). The smallest increase was shown for mature bucks for both species (7.3 % mule deer; 15.1 % white-tailed deer). The estimate for the mean buck/doe ratio for mule 'deer declined by 15 % (1.41 vs 1.20) from 1996 to 1997, however, the 95% confidence interval for the 1997 estimate was substantially improved over that in 1996. Statistically, the estimates for 1996 and 1997 are probably not different and are very close to the management objective of 1.25 - 1.5 bucks/doe. The mean buck/doe ratio for white-tailed deer increased slightly in 1997 (8.5%) but again this is probably not statistically different from 1996. The estimate for the mean fawn/doe ratio for mule deer declined from 1996 to 1997, however, this decline was slight (12.3 %) and given the large confidence interval for this ratio in 1996, it is unlikely that these two estimates are �172 statistica"ily different. For white-tailed deer, the mean fawn/doe ratio increased substantially in 1997 (25.8%), however, the precision of our estimates in 1996 was poor for this ratio (0.66-0.26) and much improved in 1997 (0.69-0.55), thus their may not be a statistically significant difference in the means for these two years. We are in the process of completing the statistical analysis of these data and we will provide a thorough analysis soon. For 1997, the precision of the estimates for all ratios were much improved. This can be attributed primarily to the 55 % increase in sample size (groups) for both mule and white-tailed deer. Weather conditions for conducting the classification surveys in 1997 were excellent each day and similar to conditions recorded for this same time period in 1996. The behavior of mule deer to our vehicles and the activities associated with these classifications appeared to be similar to that of 1996. Generally, mule deer seemed to be calm, undisturbed and well- habituated to our vehicles; we rarely had to classify moving or frightened mule deer. Similar behavior but to a lesser degree was observed for white-tailed deer in 1997. However, the behavior of white-tailed deer in 1997 was in contrast to previous years. In general, white-tailed deer appeared much less nervous and flighty, and more habituated to vehicles. This could account, in part, for the increase in numbers of white-tailed deer classified in 1997. In summary, it is important to point out that sex and age classification surveys are not intended to provide a census of the population size of deer at RMA. They are conducted to provide reliable information on recruitment rates into the populations and an estimate of the proportion of males to females. These data together with other biological information will be used in a simulation model that will assist resource biologist in managing deer populations during 1998. LITERATURE CITED Baker, D. L. 1997. Regulation of mule deer population growth by fertility control: laboratory, field, and simulation experiments. Pages 81- 87 in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-4, SP1, J1. Colorado Division of 'Wildlife, Fort Collins, Colorado, USA. Baker, D. L., and N. T. Hobbs. 1996. Regulation of mule deer population growth by fertility control: laboratory, field, and simulation experiments. Pages 113 -148 in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-4, SP1, J1. Colorado Division of Wildlife, Fort Collins, Colorado, USA. Baker, D. L, M. W. Miller, and T. M. Nett. 1995. Gonadotropin-releasing hormone analog-induced patterns of luteinizing hormone secretion in female wapiti (Cervus elaphus nelsoni) during the breeding season, anestrus, and pregnancy. Bio. Reprod. 52:1193-1197. Bomford, M. 1990. A role for fertility control in wildlife management? Bureau of Rural Resources Bulletin No.7, Department of Primary Industries and Energy, Canberra, 50pp. Boone, J. L., and R. G. Wiegert. 1994. Modeling deer herd management:sterilization is a viable option. Ecol. Model. 72:175-186. �173 Curlewis, J. D., B. J. McLeod, and A. S. I. Loudon. 1991. LH secretion and response to GnRH during seasonal anoestrus of the Pere David's deer hind (Elaphurus davidianus) J. Reprod. Fertil. 91:131-138. Dasman, R. F., and R. D. Taber. 1956. Determining structure in Columbian black-tailed deer populations. J. Wildl. Manage. 20:78-83. Garrot, R. A., D. B. Siniff, J. R. Tester, T. E. Eagle, 1992. A comparison of contraceptive technologies management. Wildl. Soc. Bull. 20:318-326. 1995. Effective using contraception. and E. D. Plotka. for feral horse management of free-ranging ungulate Wildl. Soc. Bull. 23:445-452. populations Goodman, R •.L., and F. J. Karsch. 1980. Pulsatile secretion of luteinizing hormone: differential suppression by ovarian steroids. Endocrin. 107:1286-1290. Jewell, P. A., and S. Holt. 1981. Problems in management wild animals.· Academic Press, New York, 360pp. of locally Kirkpatrick, J. F., and J. W. Turner, Jr. 1985. Chemical and wildlife management. Bioscien. 35:485-491. fertility and 1991. J. Zoo and Wildl. Med. Reversible contraception 22: 392-408. in nondomestic abundant control animals. McLleod, B. J., B. R. Brinklow, J. D. Curlewis, and A. S. I. Loudon. 1991. Efficacy of intermittent or continuous administration of GnRH in inducing ovulation in early and late seasonal anoestrus in Pere David's deer hind (Elaphurus davidianus). J. Reprod. Fertil. 52:229-238. Niswender, G.D., L. E. Reichert, Jr., A. R. Midgley, and A. V. Nalbandov. 1969. Radioimmunoassay for bovine and ovine luteinizing hormone. Endocrin. 84:1166-1173. Plotka, E. D., and U. S. Seal. 1989. Fertility tailed deer. J. Wildl. Dis. 25:643-646. ____ , D. N. Vevea, T. C. Eagle, J. R. Tester, Hormonal contraception of feral mares with Dis. 28:255-262. control in female white- and D. B. Siniff. 1992. silastic rods. J. Wildl. Turner, J. W. Jr., J. F. Kirkpatrick, and M. L. Irwin. 1996. Effectiveness, reversibility, and serum antibody titers associated with immunocontraception in captive white-tailed deer. J. Wildl. Manage. 60:45-51. Prepared by L_/?@L_ Dan L. Baker Wildlife Researcher _ �174 Table 1. Concentrations June - July, 1998. Date of LH (ng/ml) for ovariectomized mule deer during M-92 G92 S93 J93 June 9 1.8 3.3 2.9 5.4 July 7 3.3 1.7 1.9 2.8 Table 2. Summary November 1996-97. of mule deer classification Year surveys conducted 1997 1996 Mean in RMA during S.E. Mean S.E. Class: 36.3 1.7 87.3 6.8 Mature bucks 139.0 10.6 150.0 1.7 Adult does 124.3 11.8 197.0 2.0 Fawns 98.3 4.7 140.3 5.8 3.3 0.9 6.3 2.2 401.3 25.7 581.0 7.0 Young bucks Abnormal bucks Total Total animals groups (n) Ratios: . Mean buck/doe ratio: 95% CI Upper Lower 1.41 1.20 1.69 1.13 1.30 1.12 Mean fawn/doe ratio: 95% CI Upper Lower 0.8l 0.71 1.02 0.57 0.76 0.65 �175 Table 3. Summary of white-tailed RMA during November 1996 - 1997. Year deer classification surveys 1996 Day conducted at the 1997 1 2 3 1 2 3 31 35 43 84 78 100 119 143 155 153 147 150 104 124 145 200 193 198 bucks 102 89 104 150 141 130 animals groups (n) 361 93 394 104 449 131 589 238 567 234 587 271 1.44 1.44 1.37 1.18 1. 75 1.13 1. 73 1.13 1.60 1.12 1.22 1.14 1.20 1.12 1.30 1.22 0.98 0.72 0.72 0.75 0.73 0.65 1.28 0.67 0.93 0.49 0.87 0.56 0.79 0.71 0.79 0.67 0.71 0.59 Class: Young bucks Mature bucks Fawns Abnormal Total Total Ratios: Mean buck/doe ratio: 95% CI Upper Lower Mean fawn/doe ratio: 95% CI Upper Lower 1.16 1.26 ��177 Colorado Division Wildlife Research July 1998 of Wildlife Report JOB PROGRESS Colorado state of Project Work No. Package W-153-R-11 Mammals 3002 Elk Inyestigations Task No. Period Author: REPORT Research Estimating Developing PQpulation Covered:i July Survival Rates of Elk and Techniques to Estimate Size 1, 1997 - June 30, 1998 D. J. Freddy Personnel: R. Adams, T. Beck, J. Broderick, G. Byrne, D. Crane, R. Del Piccolo, J. Ellenberger, J. Frothingham, V. Graham, D. Homan, R. Kahn, K. Madariaga, D. Masden, C. Mehaffy, P. Neil, J. olterman, J. Thompson, P. Will, K. Wright, S. Yamashita, D. Younkin, CDOW; W. Andelt, D. Bowden, G. White, CSU; Diagnostic Laboratory, CSU; D. Ouren, USGS-BRO, BLM Glenwood springs, CO, USFS Rifle, CO, Rocky Mountain Elk Foundation, cooperating. Abstract During 1997-98 we monitored survival of 210 adult elk radio-collared in previous years and we conducted an applied sighting bias trial to further test methods of estimating elk density. We summarized average survival rates (± 95% CI) for elk between 1993-94 and 1997-98. Survival during summer-fall for females age ~12 months averaged 0.90 ± 0.03 inclusive of hunting deaths (n=524 elk-years) and 0.99 ± 0.01 with hunting deaths censored (n=476 elk-years). Survival during winter-spring for females age ~18 months averaged 0.97 ± 0.01 inclusive of hunting deaths (n=547 elk-years) and 0.99 ± 0.01 with hunting deaths censored (n=536 elk-years). Survival during summer-fall for males age ~24 months was 0.20 ± 0.09 inclusive of hunting deaths (n=82 elk-years) and 1.00 with hunting deaths censored (n=16 elk-years). 'Survival during winterspring for males age ~30 months was 1.00 (n=13 elk-years). Hunting related deaths were the primary cause of mortality for adult elk and annually removed 80% of the adult males, 11% of the yearling males, and 9% of the adult females. For adult females, deaths due to wounding and illegal kills exceeded the absolute number of females dying from natural 'causes during 5 years. High adult and calf natural survival rates allow this population to grow an estimated 7% annually with current levels of hunting mortality. Further tests of sighting bias revealed that failure to count all, elk in groups, as opposed to failure to detect groups, is likely the primary cause of negatively biased estimates of elk density based on quadrat sampling. Counting error may approach 25%. Therefore, sighting bias models developed to account for errors in detecting elk on quadrats do not adequately increase estimates of elk numbers. We did not find evidence indicating mark-resight estimates of elk numberS were positively biased. ��179 ESTIMATING SURVIVAL JOB PROGRESS REPORT RATES OF ELK AND DEVELOPING ESTIMATE POPULATION SIZE David TECHNIQUES TO J. Freddy P. H. OBJECTIVE Estimate survival rates of adult ,female, adult male, techniques to estimate population size. SEGMENT and calf elk and develop OBJECTIVES 1. Estimate winter, summer, and annual survival rates of adult female male elk from known fates of previously radio-collared elk. 2. Estimate density of elk in a portion of GMU-42 using a helicopter and employing mark-resight density estimators and random quadrat density estimators to evaluate potential errors in meeting assumptions of markresight theory and sighting bias correction theory. 3. Analyze data and summarize annually in Federal Aid Job Progress and Report. INTRODUCTION Our objectives are to provide reliable estimates of survival rates for calves during winter and for adult females and males throughout the year and to develop and test a system to estimate elk densities on winter ranges. Estimates of calf survival were obtained during winters 1993-94 through 199697 and summarized in Freddy (1997). We are continuing to obtain estimates of survival for adult males and females. For adults, we are interested in survival rates inclusive of hunting mortalities to document human-induced rates of survival and exclusive of hunting mortalities to estimate natural rates of survival. We are evaluating a system for estimating population size or density that incorporates estimates of sighting bias in conjunction with random sampling of search quadrats as sample units. OUr winter study area encompasses about 839 km2 (324 mi2) in the eastern half of Game Management Unit 42 south and east of Rifle, Colorado. This area is part of the Grand Mesa Elk Data Analysis Unit E-14. Elk winter habitats include juniper-pinyon woodland (Juniperus osteosperma-Pinus edulis), oakbrush-mountain shrub (Quercus gambelii-Amelanchier alnifolia), aspen (Populus tremuloides), sagebrush (Artemisia tridentata), and agricultural fields below elevations of 9,800 ft. In summer, elk use oakbrush-mountain shrub, aspen, and subalpine fir-Engelmann spruce (Abies lasiocarpa-Picea engelmannii) meadow systems throughout the Grand Mesa region up to elevations of 12,000 ft. (Freddy 1993). METHODS We placed radio collars (172-176MHz) having mortality sensors on elk calve.s age 6 months and adults age ~18 months during December 1993-1996 (Freddy 1997). Elk were captured using helicopter net-gunning and portable corral traps (Freddy 1997). During 1997-98, we monitored 210 adult elk having functioning radios as of July 1997. Collars were white and had black identification symbols on dorsal surfaces of collars to allow observers in helicopters to individually identify elk. �180 Estimating Elk Survival Rates We monitored life or death status of radioed elk with aerial surveys using a Cessna 185 at 2-4 week intervals from December 1993 to June 1998 and with daily ground surveys from December-April each year from 1993-94 to 1996-97. Survival rates (5) of radioed elk were calculated using the binomial estimator with a variance, VAR(S) = S(l-S)/n collars (White and Garrott 1990). Survival rates are expressed as the mean estimate ± the 95% confidence interval. We used x2-contingency tests to compare survival rates between sexes and among time intervals (PROC FREQ, SAS 1988). Sample sizes refer to numbers of individual radioed elk except when referenced as elk-years which denotes that individual radioed elk, because they survived for several years, were used repeatedly in successive yearly time intervals to calculate pooled average survival rates among years. Elk-years represents the numbers of times elk were at risk during time intervals. We defined 4 major time intervals for survival analyses: winter-spring was 1 December to 14 June, summer-fall was 15 June to 30 November, annual was 1 December to 30 November to coincide with timing of capture and collaring, and yearly was 15 June to subsequent 14 June used only for yearling elk aged 12-23 months. For adult elk during these time intervals, we calculated survival rates inclusive of natural and hunting-related mortalities, exclusive of hunting mortalities, and exclusive of natural mortalities. Excluding, or censoring hunting mortalities, provided estimates of natural survival rates. Excluding natural mortalities but including hunting mortalities provided estimates of hunting removal rates. Censoring elk associated with categories of mortalities reduced sample sizes used to estimate survival rates. Elk were also censored if radios failed or if their life/death status was unknown after an extended period of time. Archery, muzzleloading, and rifle hunting seasons occurred from about 1 September to 15 November each year during the summer-fall time interval. Laterifle seasons restricted hunters to taking only antlerless elk and occurred yearly from about 25 November-3l January which included portions of the summer-fall and winter-spring time intervals. Yearling spike-antlered males were generally not legal quarry in most areas frequented by radioed elk. Male elk become legal quarry when branch-antlered usually at age 2 years. Cause of death of radioed elk recovered in the field was estimated from presence or absence of gunshot wounds or bite wounds on carcass, predator tracks or scat at carcass site, physical positioning of carcass remains whether buried, covered, scattered, or consolidated (Wade and Browns 1982, Halfpenny and Biesiot 1986), relative amount of internal fat and marrow fat if present with carcass, and results of histopathology and marrow fat analyses. Fat content (percent dry matter) of bone marrow and estimates of age based on dental cementum were obtained for dead elk by the Colorado Division of Wildlife Laboratory while histopathology analyses were provided by the Colorado State University Veterinary Diagnostic Laboratory. Photographs were taken of nearly all mortalities so that physical evidence could be reviewed and judged by outside experts (pers. comm. T. Beck, W. Ahdelt). Estimating Elk Densities During 3 applied surveys in winters 199~ and 1997, we estimated numbers of elk on 132-137 mi2 of winter range with helicopter counts of marked (radioed) and unmarked elk on random quadrats and along nonrandom flight paths. Estimated numbers of elk for the 3 surveys based on actual counts of elk on quadrats were 51, 63, and 93% of the estimated numbers of elk obtained from markresight formulas using counts of marked and unmarked elk. Adjusting actual �181 quadrat counts with previously developed sighting bias models did not meaningfully increase quadrat estimates in comparison to mark-resight estimates (Freddy 1996, 1997). We had 3 important concerns: 1) sighting bias corrections developed during previous testing trials were biased high and provided inadequate corrections for applied surveys, 2) observers failed to see marks on elk in groups that were detected and counted accurately causing inflated mark-resight estimates of elk numbers, and 3) observers failed to count all elk within groups that were detected on quadrats with elk not counted inclusive of both marked and unmarked elk causing deflated counts of elk on quadrats. This last concern represented counting error that would not be accounted for by sighting bias corrections. In 1998, we conducted an applied sighting bias trial on quadrats along with mark-resight surveys to again compare estimates of elk numbers based on quadrat counts and mark-resight surveys. We selected 44 mi2 in West Divide and Alkali creeks that were a portion of the winter range sampled with quadrats and mark-resight surveys in 1996 and 1997 and a portion of the area used to conduct sighting bias trials in 1994 and 1995. This area was dominated by oakbrush and pinyon-juniper habitats. For this test, all marked elk were age ~18 months unlike previous sighting tests and winter range surveys that also included marked calves age 6 months. Locations of all radioed elk within this 44 mi2 area were obtained by telemetry using a Cessna 185 and ARNAV Star5000 GPS system on 26 February, 1998. Radioed elk represented a population of known size which we used to evaluate the magnitude of sighting detection bias. During 3 days from 27 February to 1 March, elk were counted on all 44, 1-mi2 quadrats using a Bell-Soloy helicopter flown at 40-50 mph. A flight crew was a pilot, navigator, and observer and crews changed each day among 3 observers, 2 navigators, and 1 pilot all of whom were involved in previous sighting bias tests and winter range surveys. Members of flight crews had no knowledge of the number or locations of marked elk within the survey area. Flight crews flew nearly 7 hours each day to complete quadrats which was similar to daily flying time and fatigue commonly associated with applied surveys. Quadrats were flown using the same helicopter and counting procedures that were used in previous sighting tests and surveys (Freddy 1994-1997). Observers recorded the standard sighting bias variables of group size, elk behavior, habitat type, percent screening cover, and percent snow cover for all groups seen. As flight crews completed quadrats, they maintained a list of marked elk seen and their GPS locations (Garmin Pilot III) and whether or not marked elk were or could have been individually identified. Those marked elk not seen or those seen but not individually identified during counts were then found by a second independent flight crew using a Bell-Soloy helicopter and telemetry. These missed elk were found on the same day an average 3.6 hours after quadrats were flown. For every missed' or unidentified marked elk group, the second flight crew recorded the GPS location (Garmin Pilot III) of the group and standard sighting bias variables. Those elk seen but not individually identified by quadrat crews were assigned as a specific individual elk based upon proximity of locations of where such elk were seen by quadrat crews compared to locations of these elk when found by the second flight crew. Locations and status of all marked elk were therefore known during the time quadrats were flown. A sighting trial occurred when marked elk were detected during quadrat counts and when marked elk were missed and later found by the second flight crew. To �182 maintain independent observations of marked elk, only 1 sighting trial occurred if there was more than 1 marked elk in a group. Otherwise, detecting or missing a marked elk constituted a sighting trial. Two mark-resight flights using the same helicopters were conducted on 1 and 2 March after quadrat counts were completed. These flights followed a nonrandom path through the same 44-mi2 with the intent of counting as many elk as possible within allotted flight time. The navigator for both flights had not participated in quadrat counts. The observer for the first flight was an observer during part of the quadrat counts while the observer for the second flight had not participated in quadrat counts. These 2 observers were the same persons who conducted mark-resight flights during winter range surveys in 1996 and 1997. At the conclusion of the second mark-resight flight, locations of collared elk not seen were confirmed to be within the sample area using the helicopter and telemetry. Weather conditions during flights were variable but similar to conditions experienced during previous winter range surveys. Quadrats were flown with generally acceptable visibility in hazy to bright sunshine and light to moderate wind speeds although brief snow squalls occurred on 27 and 28 February. Nonrandom flights were flown during hazy to bright sunshine and low wind speeds. Snow cover was nearly 100% in all areas with depths estimated to be 0.5 to 2.5 ft. Fresh snow was received on 24 and 25 February. Minimum night temperatures were 2-13 OF and were the coldest measured during the 14 days previous to flights. Maximum day temperatures were 27-35. OF. Number of elk on the search area was estimated using 3 approaches: from actual counts of elk on quadrats, from adjusting quadrat counts with sighting bias correction models, and from using the Bowden (BE) and lmmigration-EmigrationJHE (lMJHE) mark-resight estimators in program NOREMARK (White 1996). Elk counted on quadrats were potentially a total count because 100% of the search area was flown and because all 44 sample units were flown, there was no sampling variance associated with estimates based on quadrats. To adjust for elk groups likely missed on quadrats, we applied sighting bias corrections to elk groups counted on each quadrat which adjusted counts per quadrat and provided a sighting bias corrected total population estimate. We used 3 sighting correction models based on all sighting trials (n = 224) to .correct for probabilities of detecting elk groups, where probability of detection is [n = eU / 1 + eU] with u representing the linear regression of variables affecting the probability of detecting a group: 3. u = 1. u = 1.342(LogGroupSize(countedd - 0.131 2. u = 1.230(LogGroupSize(co=ted» - 2.127(Bedded) + 0.291 1.135(LogGroupSize(co=ted» - 1.822(Bedded) - 0.031(%Cover) + 1.517 probability of detecting an elk group was adjusted for group size, for bedded or not bedded behavior when detected, and for percent vegetation screening cover (Samuel et al. 1987, Freddy'1995). Relative AlC model performance values of 172, 160, and 154 for models 1-3, respectively, indicated model 3 was the most parsimonious model. We pooled counts of marked and unmarked elk seen during quadrat counts and mark-resight flights. Estimated numbers of elk based on all 3 flights pooled was considered to be the best estimate of population size and the benchmark against which quadrat counts were compared. Both the BE and lEJHE estimators account for movement of elk on and off the search area but the BE better accounts for heterogenous sighting probabilities among individual elk (Bowden 1995, White 1996). �183 Moyements We continued to precisely locate selected radioed elk at least once ~r month since their capture to document seasonal movements using a Cessna 185. These elk were originally selected at random from within trap zones and equalized by age class in January 1994. Elk from the original sample that died were replaced each subsequent January with randomly selected elk that were usually of the same sex, age class, and trap zone as elk that died. All replacement elk in January 1998 were yearling or older because additional calves were not marked in December 1997. As of 1 January 1998, these 44 elk were classified as 27 adult females, 5 yearling females, 4 adult males, and 8 yearling males. During June 1998, we again located an additional 28 adult females that were originally selected at random in 1995 to document their locations during the calving period. Females from the original 1995 sample that died were replaced each subsequent June with adult females randomly selected from the same trap zones of elk that died. As needed, we located other elk to document unusual movements. RESULTS AND DISCUSSIQH Survival Estimates Between 1 December 1993 and 14 June 1998, 181 radioed elk died of which 31 were calves 6-11 months old and 150 were adults ~12 months old (Table 1, Appendix I). Calves died of natural causes (Freddy 1997) while for adults ~12 months old, hunting accounted for 92% of 150 deaths. Yearlings Survival during summer-fall for yearling elk age 12-17 months was 0.89 ± 0.06 for males (n = 120) and 0.95 ± 0.04 for females (n = 128) when averaged among 4 years and inclusive of hunting deaths, and 1.00 for both males (n = 107) and females (n = 122) with hunting deaths censored. All deaths for males and females during summer-fall were due to hunting. Survival among years for males ranged from 0.83 to 0.97 and for females from 0.87 to 1.00 inclusive of hunting deaths. We failed to detect differences in survival between sexes within years (P ~ 0.13) and among years for both males (P ~ 0.39) and females (P ~ 0.09) inclusive of hunting deaths. Survival of females (0.95) tended to be higher than males (0.89) when averaged among years and inclusive of hunting deaths (P = 0.07) Cl'ables 2-5). Survival during winter-spring for yearling elk age 18-23 months was 0.98 ± 0.02 for males (n = 106) and 0.97 ± 0.0.03 for females (n = 121) when averaged among 4 years and inclusive of hunting deaths, and 0.98 ± 0.02 for males (n = 106) and 0.98 ± 0.02 for females (n = 120) with hunting deaths censored. Survival among years for males ranged from 0.96 to 1.00, inclusive or exclusive of hunting deaths. Survival for females ranged from 0.93 to 1.00 and 0.96 to 1.00 inclusive and exclusive of hunting deaths, respectively. We failed to detect differences in survival among ye~rs for both males (P ~ 0.51) and females (P ~ 0.22) inclusive or exclusive of hunting deaths. We also failed to detect differences in survival between sexes within years (P ~ 0.62) and among years (P ~ 0.76) inclusive or exclusive of hunting deaths (Tables 25). During winter-spring, there were 2 nonhunting deaths each for males and females and 1 illegal hunting death for a female (Table 1). Yearly survival for elk age 12-23 months was 0.87 ± 0.06 for males (n = 119) and 0.93 ± 0.05 for females (n = 127) when averaged among 4 years with hunting deaths included. Survival among years for males ranged from 0.79 to 0.97 and for females from 0.81 to 1.00. We failed to detect differences in yearly survival among years for males (P = 0.27) but survival was lower (0.81) for �184 females in 1996-97 (P = 0.02). Yearling males were subjected to about the same rate of illegal hunting during all years which accounted for most of the male mortality, while in 1996-97, females had a lower survival rate because 5 of 6 deaths were hunting related (Tables 2-5). With hunting deaths included, we failed to detect differences in survival between sexes within (P ~ 0.13) and among years (P = 0.15). Of 24 deaths involving yearling elk, 20 (83%) were hunting related (Table 1). With hunting deaths censored, yearly survival for elk age 12-23 months was 0.98 ± 0.02 for males (n = 106) and 0.98 ± 0.02 for females (n = 120) when averaged among 4 years. Survival among years for males and females ranged from 0.96 to 1.00. We failed to detect differences in yearly survival among years for males (P = 0.51) and for females (P = 0.50), and between sexes within and among years (P ~ 0.90) (Tables 2-5). Yearling spike-antlered males were generally not legal quarry during hunting seasons. Each year from 1994-1997, 29-32 yearling males radioed as calves entered the fall hunting seasons presumably as spike-antlered males. OVer 4 years, illegal harvest removed 10% of the yearling males with yearly rates ranging from 3 to 17%. Of 12 elk illegally killed, 11 were taken during rifle seasons and 1 during archery seasons. In addition to these 12 elk, 1 male was fatally wounded during archery season in an area where the elk was legal quarry. Overall, 11% of the yearling males were annually removed by hunting (Tables 1, 2-5). Adult Females Survival during summer-fall for all adult females age ~12 months was 0.90 ± 0.03 (n = 524 elk-years) when averaged among 4 years and inclusive of hunting deaths and 0.99 ± 0.01 (n = 476 elk-years) with hunting deaths censored. Survival among years ranged from 0.87 to 0.94 and from 0.99 to 1.00, inclusive and exclusive of hunting mortalities, respectively. We failed to detect differences in survival among years inclusive (P = 0.30) or exclusive (P = 0.46) of hunting mortalities (Table 6). Hunting was involved in 48 (96%) of 50 summer-fall deaths: 32 were killed, 12 were wounded, and 4 disappeared during hunting seasons and presumed legally killed. One natural death was attributed each to suspected predation and complications while calving. At the beginning of fall hunting seasons, there were 99, 129, 142, and 152 radioed adult females age ~12 months available to hunters in 1994, 1995, 1996 and 1997, respectively, representing 522 elk-years (Table 6, nonhunting deaths censored). During summer-fall, 9% of these radioeq elk were removed annually by all types of hunting mortality (range = 6-12%). Survival during winter-spring for all adult females age ~18 months was 0.97 ± 0.01 (n = 547 elk-years) when averaged among 5 years and inclusive of hunting deaths and 0.99 ± 0.01 (n = 536 elk-years) with hunting deaths censored. Survival among years ranged from 0.94 to 0.99 and from 0.97 to 0.99, inclusive and exclusive of hunting mortalities, respectively. We failed to detect differences in survival among years inclusive (P = 0.34) or exclusive (P = 0.40) of hunting mortalities (Table 6). There were 19 winter-spring deaths, of which 11 (58%) involved hunting with 5 killed and 3 wounded during rifle late-seasons and 3 illegally killed out of hunting seasons. Natural deaths were attributed to mountain lion predation (1), suspected predation (2), complications while calving (1), accidental fall (1), and unknown cause (3) (Table 1). During winter-spring, 2% of these radioed elk were removed annually by all types of hunting mortality. �185 Estimates of adult female survival using all females age ~12 months could be biased because of the constant yearly recruitment of yearling females into this radioed population of adults afforded by many radioed calves surviving to yearling age and no similar yearly recruitment of additional older adult females into the radioed population. We therefore estimated survival rates for adult females age ~24 months by excluding recruitment of yearlings age 1223 months and for 68 adult females initially collared as a cohort in December 1993 whose age at capture ranged from 18 months (n=6) to 10+ years. Survival during summer-fall for adult females age ~24 months was 0.89 ± 0.03 (n = 396 elk-years) when averaged among 4 years and inclusive of hunting deaths and 0.99 ± 0.01 (n = 354 elk-years) with hunting deaths censored. Survival among years ranged from 0.82 to 0.93 and from 0.99 to 1.00, inclusive and exclusive of hunting mortalities, respectively. We failed to detect differences in survival among years inclusive (P = 0.17) or exclusive (P = 0.38) of hunting mortalities (Table 7). Hunting was involved in 42 (95%) of 44 summer-fall deaths. During summer-fall, 11% of these radioed elk were removed annually by all types of hunting mortality (range 7-17%). Survival during winter-spring for adult females age ~30 months was 0.96 ± 0.02 (n = 420 elk-years) when averaged among 5 years and inclusive of hunting deaths and 0.98 ± 0.01 (n = 410 elk-years) with hunting deaths censored. Survival among years ranged from 0.93 to 0'.99 and from 0.97 to 0.99, inclusive and exclusive of hunting mortalities, respectively. We failed to detect differences in survival among years inclusive (P = 0.15) or exclusive (P = 0.39) of hunting deaths (Table 7). There were 16 winter-spring deaths with 10 (63%) attributed to hunting. During winter-spring, 2% of these radioed elk were removed annually by all types of hunting mortality (range 1-5%). Using the 1993 cohort of adult females, survival during summer-fall was 0.88 ± 0.05 (n = 189 elk-years) when averaged among 4 years and inclusive of hunting deaths and 0.99 ± 0.01 (n = 167 elk-years) with hunting deaths censored. Survival among years ranged from 0.82 to 0.94 and from 0.98 to 1.00, inclusive and exclusive of hunting deaths, respectively. We failed to detect differences in survival among years inclusive (P = 0.20) or exclusive (P = 0.55) of hunting mortalities (Table 8). Hunting was involved in 22 (96%) of 23 summer-fall deaths. Percent of these radioed elk removed during summerfall by all types of hunting-related mortalities averaged 12 annually (range = 6-17%). Interestingly, as this cohort collectively increased in age from 1994 to 1997, the annual percentage of elk removed by hunting during summer-fall steadily declined from 17 to 6%. For the 1993 cohort, survival during winter-spring age was 0.95 ± 0.03 (n = 233 elk-years) when averaged among 5.years and inclusive of hunting deaths and 0.98 ± 0.02 (n = 227 elk-years) with hunting deaths censored. Survival among years ranged from 0.92 to 1.00 and from 0.93 to 1.00, inclusive and exclusive of hunting mortalities, respectively. We failed detect differences in survival among years inclusive (P = 0.47) or exclusive (P = 0.17) of hunting deaths (Table 8). There were 11 winter-spring deaths, of which 6 (55%) involved hunting. During winter-spring, 3% of these radioed elk were removed annually by all types of hunting mortality. to Overall, survival rates for adult females during summer-fall and winter-spring were the same among age groups of adult females. Natural survival rates exceeded 0.98 for all time intervals and groupings (Table 9). �186 Annual survival rate using the 1993 cohort was 0.83 ± 0.07 (n = 199 elk-years) when averaged among 4 years and inclusive of hunting deaths and 0.97 ± 0.03 (n = 170 elk-years) with hunting deaths censored. Survival among years ranged from 0.78 to 0.94 and from 0.92 to 1.00, inclusive and exclusive of hunting mortalities, respectively. We failed to detect differences in survival among years inclusive (P = 0.17) or exclusive (P = 0.34) of hunting mortalities (Table 10). Annually, 14% of these radioed elk were removed by all types of hunting mortality (range 8-20%). Removal rates of females by hunting during summer-fall were compared between yearling females age 12-17 months and adult females age ~24 months. Removal rates were higher for adult females (0.11) than yearling females (0.05) when averaged among 4 years (P = 0.04). This difference was caused by extremes in removal rates in 1994 when 17% of adults and 3% of yearlings were removed (P =0.05) and in 1997 when 11% of adults and 0% yearlings were removed (P = 0.06). In 1995 and 1996, removal rates were not different (P > 0.41) but rates for yearlings (3%) tended to be lower than for adults (7-9%). These rates suggest that vulnerability of yearling and adult females to hunting is different. Adult Males Survival during summer-fall for adult males age 24-29 months was 0 •. 17 ± 0.09 (n = 75) when averaged among 3 years and inclusive of hunting deaths and 1.00 (n = 13) with hunting deaths censored. Survival among years ranged from 0.08 to 0.23 inclusive of hunting mortalities. We failed to detect differences in survival among years (P > 0.32) (Tables 2-4). All deaths were attributed to hunting resulting in removal rates averaging 83% collectively for all fall hunting seasons each year. Through fall 1997, 7 males had survived at least 1 year of hunting seasons and were available during at least 1 additional year of hunting seasons as 3 or 4 year-olds. Survival during summer-fall for adult males age ~24 months was 0.20 ± 0.09 (n = 82 elk-years) when averaged among 3 years and inclusive of hunting deaths and 1.00 (n = 16 elk-years) with hunting deaths censored. Survival among years ranged from 0.14 to 0.24 inclusive of hunting mortalities. We failed to detect differences in survival among years (P > 0.61) (Tables 2-4). All deaths were attributed to hunting resulting in removal rates averaging 80% collectively for all legal males during fall hunting seasons each year. Survival during winter-spring for adult males age ~30 months was 1.00 (n = 13 elk-years) during all years (Tables 2-5). There were no hunting or nonhunting deaths of adult males during winter-spring. Hunting was the cause of mortality among male elk age ~24 months. From the 1993-94 cohort of 36 male calves, 28 lived to age 24-months with 22 (79%) harvested in 1995 inclusive of 2 that disappeared 'and 2 wounding losses during rifle seasons. From the 1994-95 cohort of 33 male calves, 25 lived to 24months and 23 (92%) were harvested in 1996 including 2 that disappeared during seasons, 1 illegally taken during rifle seasons, and 1 wounding loss during archery/muzzleloading season. From the 1995-96 cohort of 33 male calves, 22 lived to 24-months and 17 (77%) were harvested in 1997 including 1 that disappeared, 1 illegally taken, and 4 wounding losses during rifle seasons. Comparative Survival of Adult Males and Females The differential impact of hunting on survival and recruitment of males and females to young adult age classes is demonstrated by net survival rates of �187 calf cohorts. For 1993-94, 1994-95, and 1995-96 calf cohorts, net survival from age 6 to 35 months averaged 0.10 for males and 0.73 for females (Tables 2-4). Only 3 (4%) of a potential 69 males survived from age 6 months to 48 months (Tables 2,3). Hunting Mortality Per Season Distribution of all types of hunting mortalities for male elk age >12 months during hunting seasons 1994-1997 was: Archery 15%, Muzzleloading 5%, RifleFirst 42%, Rifle-Second 25%, Rifle-Third 13%, Rifle-Late 0%, Illegal out-ofSeason 0%. Distribution of all hunting mortalities for female elk age >12 months was: Archery 14%, Muzzleloading 7%, Rifle-First 19%, Rifle-Second 31%, Rifle-Third 10%, Rifle-Late 15%, Illegal out-of-Season 4%. All female and nearly all male elk killed by hunters died within the Grand Mesa DAU E-14 or adjacent GMU 43. Exceptions were 6 adult males that dispersed long distances and were killed near Ruedi Reservoir, Black Mesa, Tomichi Dome, Gothic, and Anthracite Creek in GMUs 47, 63, 54, 55, 551, and 521 up to 100 mi south or east of their capture sites. These 6 males represented 9% of adult males age ~24 months that were taken by hunters. Adjusting Hunting Harvest Estimates Surveys used to estimate legal hunter harvest do not account for wounding losses or illegal kills. Estimates of the proportion of the legal kill that these additional deaths represent were made based on fates of radioed elk during hunting seasons 1994-97. We defined legal kill to be the number of radioed elk known to be harvested plus those whose radio signals disappeared during hunting seasons and were subsequently assumed to be legally killed. This legal kill would most likely represent the kill estimated by harvest surveys. Wounding losses and illegal kills represented deaths not accounted for by surveys. Total kill which was known for radioed elk could thus be expressed as (legal kill)*(X)= total kill where (X) represents a correction factor. correction factors by sex/age class were 1.16 for adult males, 1.11 for yearling males, and ~ 1.4 for all categories of adult females (Table 11). Rifle and late-rifle seasons involving adult female elk had high correction factors of 1.35-2.00. Losses of adult females not reported or estimated can be important in modeling populations. Total unreported losses of adult females during hunting seasons 1994-1997 was 18 which was 1.8x the 10 natural deaths of adult females in the comparative time interval of 1 December 1993-14 June 1998 (Tables 6, 11). An argument could be made that estimating unreported losses is as important as estimating natural mortality rates. Survival Rates and Population Modeling Survival and removal rates were incorporated into a simple spreadsheet model to assess trends in population growth (Table 12). Natural survival rates (hunting deaths censored) were used for summer-farl and winter-spring time intervals for adult males, yearling males, adult females, yearling females, and calves. Hunting deaths of all types were incorporated as total harvest removal rates for each sex/age class. Using total removal rates is a simpler method than using correction factors to account for unreported .harvest. This model assumed survival of calves 0-5 months of age during summer was 1.00 because the model uses post-season calf:cow ratios measured in January as an estimate of net calf recruitment to December. Recruited sex ratio of calves to December was assumed to be 1:1. �188 Modeling suggests that if harvest rates of antlered elk remained as measured and harvest rates of antler less elk were reduced to 0, the population could grow at an annual rate of 18%. Current rates of antlerless harvest allow annual growth rates of about 7.5%. To stabilize population growth, harvest rates on adult and yearling female and calf age classes must increase nearly 2X. population Estimates Movements of marked elk on and off the survey area during days when flights were conducted resulted in 34 marked elk (27 female, 7.male) and 38 marked elk (31 female, 7 male) within the area during quadrat and mark-resight flights, respectively. Elk movements were usually <600 yards and near sample area boundaries. Ratios of marked/unmarked elk counted were similar among quadrat and nonrandom flights ranging from 0.029 to 0.036 (Table 13). These ratios were generally lower than ratios of 0.031-0.042 observed during extensive survey flights in 1996 and 1997 when marked calves were available in addition to marked adults. Observers used 19.7±2.1 (95% CI) minutes to count 17.5 elk per quadrat (range=0-68). Observers 1B and 2E encountered similar densities of elk and had similar search times in oakbrush habitat while observer 3M had lower elk densities and higher search times in pinyon-juniper habitat (Table 14). During 3 extensive surveys in 1996 and 1997, these same observers used 17.6, 16.9, and 19.8 minutes to count 17.0, 25.9, and 17.3 elk, respectively, per quadrat (n=177). Searching effort during the 1998 sighting trial was therefore representative of searching effort during previous extensive surveys. During nonrandom flights, detection rates of marked elk were 0.32 and 0.45 with effective searching rates of 4.9 and 6.00 minutes per mi2 of search area during flights 1 and 2, respectively (Table 13). During nonrandom flights for extensive surveys in 1996 and 1997, detection rates of marked elk were 0.260.58 with effective searching rates of 3.3-3.9 minutes per mi2 of search area. Nonrandom flights in 1998 were therefore representative of flights during extensive surveys. Observers completed 32 sighting trials while counting elk on quadrats. Average sighting bias or detection rate for marked elk groups among observers was 0.78±0.15 (95% CI) (Table 14). This sighting bias rate was similar to the average sighting bias rate of 0.82+ 0.06 (95% CI) for these same 3 observers during sighting bias test trials (n=192) conducted in 1994 and 1995. Therefore, sighting bias rate estimated from test trials appears to be representative of sighting bias during applied surveys. Observers failed to detect 7 marked elk during sighting trials. When found after quadrats were completed, 6 of these elk were in groups of ~2 elk, 4 were bedded and appeared to have been bedded for several hours, 5 were in screening cover densities ~60%, and 5 were reluctant to move even when disturbed at close distances with the helicopter (Table 15). Importantly, the white collars were easily visible and thus, failure to detect marked elk was not likely due to failure to see marks. Of these 7 missed elk, 3 were males and 4 were females resulting in detection rates of 0.57 for adult males and 0.85 for adult females. Of the 7 missed elk, 3 were found in close proximity to other groups that were likely counted by quadrat crews. This lends credence to the possibility that some marked elk were not counted within detected groups because not all elk, �189 both marked and unmarked, were seen when groups were detected. This type of counting error is plausible as groups detected in dense pinyon-juniper or oakbrush often begin to move and observers concentrate on counting the moving elk and fail to see elk in the group that remain bedded or move in a different direction. Estimated number of elk on the search area was 1,179 based on the BE markresight estimator (Table 17). The 771 elk counted on quadrats represented 65% of the BE while sighting bias adjusted quadrat counts represented 73-75% of the BE. We propose that the 25% difference between sighting adjusted estimates and the BE represents counting error. This degree of counting error would equate to adding 2.2 elk per group to an average group size of 5.2 elk for the 148 groups detected (Table 18). Counting error, as opposed to detection error, appears to be the primary cause of negatively biased estimates of elk density based on quadrat sampling. Elk Moyements Yearling elk originally trapped as calves and surv1v1ng to age 18 months dispersed to winter ranges not affiliated with their natal capture winter range at rates of 43, 42, and 33% in winters 1995-96, 1996-97, 1997-98, respectively. Each year, 40-52% of the yearling females dispersed while only 25-33% of the yearling males dispersed. Yearling elk moved primarily west or south to winter ranges near the towns of Rulison, Parachute, Collbran, Hotchkiss, Paonia, and Paonia Reservoir. Specifically for the 1996 cohort of radioed calves, 28 males and 30 females survived to age 18 months in December 1997 (Table 5). By end of winter 199798, 19 (7M, 12F) or 33% had dispersed to other winter ranges (Table 19). Calves trapped in Alkali, Dry Hollow, and the Mammm creeks (zones E, F, G) comprised 79% of the dispersing yearlings and these elk moved primarily west to winter ranges near Collbran, Rulison, and Parachute. Calves trapped in West Divide Creek (zone D) comprised 16% of the dispersing yearlings and these elk moved to winter ranges near Collbran and also south towards Paonia Reservoir. We continue to cooperate with the USGS-BRO, USFS, and BLM with the support of the Rocky Mountain Elk Foundation to develop a GIS database allowing analyses relating elk locations, movements, and home ranges to habitat components. CONCLUSIQNS Hunting related mortalities were the primary cause of death for adult female and male elk. For adult males, all deaths to age 48 months were related to hunting. For adult females, deaths due to wounding and illegal kills exceeded the absolute number of females dying from natural causes during 5 years. Natural survival rates for adult females and males were ~0.98 during winterspring and summer-fall. High adult and calf survival rates allow this population to potentially increase 18% annually in the absence of hunting antlerless elk. Current hunting removal rates of 9% for adult females age ~12 months allows the modeled population to grow 7% annually. Failure to count all elk in groups, as opposed to failure to detect groups, appears to be the primary cause of negatively biased estimates of elk density based on quadrat sampling. Counting error may approach 25%. Sighting bias models developed to account for errors in detecting elk on quadrats inadequately increased estimates of elk numbers when compared to numbers estimated by mark-resight formulas. We did not find evidence indicating markresight estimates of elk numbers were positively biased. �190 LITERATURE CITED Bowden, D. C., and R. C. Kufeld. 1995. Generalized mark-sight population size estimation applied to Colorado moose. J. Wildl. Manage. 59: 840851. Freddy, D. J. 1993. Estimating survival rates of elk and developing techniques to estimate population size. Colo. Div. Wild1. Game Res. Rep. July: 83-117. Freddy, D. J. 1994. Estimating survival rates of elk and developing techniques to estimate population size. colo. Div. Wildl. Game Res. Rep. July: 27-42. Freddy, D. J. 1995. Estimating survival rates of elk and developing techniques to estimate population size. colo. Div. Wildl. Game Res. Rep. July: 63-79. Freddy, D. J. 1996. Estimating survival rates of elk and developing techniques to estimate population size. colo. Div. Wildl. Game Res. Rep. July: 87-108. Freddy, D. J. 1997. Estimating survival rates of elk and developing techniques to estimate population size. Colo. Div. Wildl. Game Res. Rep. July: 47-73. Hal.fpenny, J. C., and E. A. Biesiot. 1986. A field guide to mammal in North America. Johnson Books, Boulder, CO. 161pp. tracking Samuel, M. D., E. O. Garton, M. w. Schlegel, and R. G. Carson. 1987. Visibility bias during aerial surveys of elk in northcentral Idaho. Wildl. Manage. 51:622-630. SAS Institute Inc. 1988. SAS/STAT Cary, NC. 1028pp. User's Guide, 1982. Procedures Texas Agric. Expt. 6.03. SAS Institute, J. Inc., for evaluating predation on Sta. Publ. B-1429. 42pp. Wade, D. A., and J. E. Browns. livestock and wildlife. White, G. C. 1996. NOREMARK: Population estimation surveys. Wildl. Soc. Bull. 24:50-52. White, G. C., and R. A. Garrott. 1990. Analysis of wildlife data. Academic Press, Inc., San Diego. 383pp. from mark-resighting radio-tracking �191 Table 1. Causes of deaths in radio-collared elk between 1 December 1993 and 14 June 1998. Calves (M=male, F=female) were age 6-11 months and collared at age 6 months. Yearling males and females were age 12-17 months and juvenile males and females were age 18-23 months and all were collared at age 6 months. Adult males and females were ase >24 months at time of death. "Elk Ase!Sex Class at Death Yearling Juyenile Adult Total Cause of Death Calves All M F M F M F M F and -Code M F Natural causes o o 4 o o o 2 2 Malnutrition-6 2 2 o Unknown-Suspect o o o 3 1 4 Malnutrition-31 3 1 o o o o 1 5 13 Predation-Lion-3 o o o o 8 8 4 o o o o o Predation-Bear-35 o o o o o 0 o o o 1 1 Unknown Predator-5 o 1 o o o o Unknown-Suspect o 2 3 8 11 Predation-30 2 5 o o 1 1 2 o 0 o o 2 o 2 Accident-Birthing-32 o o o o 1 1 2 Accident-Fell-10 o 0 o o 1 o 1 1 o 2 2 4 6 Unknown Cause-11 2 1 o o o Subtotals .Q .Q .Q .a. .l.2. II sa 2. 111l 2. Legal Bunting 8 2 12 o 0 2 8 4 o Archery-33 o o 4 2 4 4 8 Muzzleloading-34 o 2 o o o 0 o 1 o 1 1 o Archery/Muzzle-27 o 0 o o o 29 22 7 22 7 Rifle-First-46 o 0 o o o o 20 o 0 10 10 10 10 o Rifle-Second-47 o o o 8 5 8 5 13 o 0 o Rifle-Third-48 o o o o 6 o 6 6 Rifle-Late-29 o 0 o o o o Subtotals .Q .Q .6..2. .Q .Q .Q sa aa sa II .i Wounding Loss 1 o 0 2 o 1 1 1 Archery/Muzzle-24 o o o 3 o 0 o o 1 1 2 Archery-52 1 1 o 3 2 o 0 o 1 1 2 Rifle-First-43 o o o 9 o 0 o 3 6 3 6 Rifle-Second-44 o o o 2 1 1 1 1 Rifle-Third-45 o 0 o o o o 1 1 o 1 o Rifle-Unk.Reg.-25 o 0 o o o o 3 o 3 o 3 Rifle-Late-26 o 0 o o o o Subtotals .Q .Q .Q .Q aa 1. II .a. .15. ~ ~ Illegal Bunting 5 o o o 5 Rifle-First-49 o 0 o o o 5 6 1 o o 6 Rifle-Second-50 o 0 5 o o o 1 o o o o 0 o o 1 Rifle-Third-51 1 o o 0 1 1 o o 1 Rifle-Unk.Reg.-9 o o o o 1 o o o 1 Archery Season-8 o 0 1 o o o 3 o 2 3 o Out-of-Season-7 o 0 o o o 1 Subtotals .Q .Q .Q 11 2. 2. .3. II .u: .Q ~ Presumed Buntin~ 1 1 Missing-ArchMuzz-21 o 0 o o o o o o 1 o 0 5 o 2 3 2 3 Missing-Rifle 1st-40 o o o 3 1 Missing-Rifle 2nd-41 o 0 o 1 o 1 2 1 o o 0 o o o o o o Missing-Rifle 3rd-42 o o o Subtotals .Q .Q .Q .Q .Q .2. .3. .5. .i .5. ~ Totals 17 14 13 6 2 3 66 60 98 83 181 • These elk were illegally shot as spike-antlered yearling males: (173.190/93) (173.232/93) (173.309/93) (173.320/93) (173.919/94) (174.059/94) (174.140/95) (174.200/95) (174.679/95) (174.729/95) (174.861/95) (173.210/96). (173.309/93) disappeared in 1994 during first rifle season when spike-antlered yearling males were not legal and was assumed to have been taken illegally. All other illegal deaths were confirmed. • These elk disappeared during hunting seasons and remained missing for several months and are assumed dead and legally harvested: (172.207/93) (172.649/93) (172.800/93) (172.961/93) (173.390/93) (173.439/93) (174.001/94) (174.181/94) (174.770/95). �Table 2. Survival rates, inclusive of nonhunting and hunting mortalities, for winter-spring (WS, 1 December-14 June) and summer-fall (SF, 15 June-30 November) time periods from 1 December 1993-14 June 1998 for the cohort of calves age 6 months radio-collared in December 1993. Survival rate for male and female calves pooled when age 6-11 mohths was 0.92 (95% CI 0.86-0.98, n=73). Survival rates (S) calculated as a mean estimate of (alive)/(alive + dead) and variance S(1-S)/n collars. Elk Elk Age (months) and Time Period (WS or SF dates) 18-23 24-29 30-35 36-41 42-47 6-59 6-11 (mos) l2.:.ll ~ ~ .WS WS SF WS SF WS SF WS SF ALL 1994-95 1995 1995-96 1996 1996-97 1997 1997-98 1994 1993-98 1993-94 MALES Survival L 95% CI U 95% CI n collars censo.redDied Nonhunt Hunt 0.89 0.78 0.99 36 0.88 0.76 0.99 32 0 o b 1.00 28 o o o o 0.21 0.06 0.37 28° 1. 00 0.00 0.00 0.50 0.00 1. 00 1. 00 0.00 0.00 4 4 2f o 22d o o o 2 o o o o 1 19 33 o 2e 1 33 4 4 0 4 0.95 0.87 0.97 0.92 1. 00 1. 00 1. 00 0.97 0.91 1. 00 37 35 34 34 33 0.84 0.71 0.97 32 censo red- o o o o o Died Nonhunt. Hunt 2 2 1h 1 1 5 o o o o 1 o o o o 1 1 5 FEMALES Survival L 95% CI U 95% CI n collars o 4 o 1. 00 o 22 0.97 0.91 o 2 1. 00 27 o o o o 3e,9 o 4 1 29 0.89 0.76 1. 00 27 0.96 0.87 1. 00 24 0.62 0.46 0.78 37 o o o 31 o 1 1 3 3 o 11 • Censored denotes collar failure and/or animal life/death status not known. Collar (173.309/93) disappeared 1994 hunting seasons and assumed dead. • Includes (173.241/93) collar failure August 1995 but seen January 1996. d Two collars (173.390/93). (173.439/93) disappeared 1995 hunting seasons and assumed dead. • Two collars censored; (173.241/93). collar failed August 1995. (173.381/93) slipped off December Includes (173.340/93) alive May 1997. 9 Censored elk was (173.249/93) slipped collar September 1997. • Collar (172.961/93) disappeared 1997 hunting seasons and assumed dead. h Collar (172.800/93) disappeared 1994 hunting seasons and assumed dead. b 1995. 14 ,_. It) (IJ �Table 3. Survival rates, inclusive of nonhunting and hunting mortalities, for winter-spring (WS, 1 December-14 June) and summer-fall (SF, 15 June-30 November) time periods from 1 December 1994-14 June 1998 for the cohort of calves age 6 months radio-collared in December 1994. Survival rate for male and female calves pooled when age 6-11 months was 0.90 (95% CI 0.83-0.97, n=69). Survival rates (S) calculated as a mean estimate of (alive)/(alive + dead) and variance S(l-S)/n collars. Elk Age (months) and Time Period (WS or SF dates) 18-23 24-29 30-35 36-41 42-47 ~ 6-11 (mos) l2...:.l1 WS SF WS SF WS. ALL SF WS 1996-97 1997 1995-96 1996 1997-98 1994-98 1995 1994-~5 MALES Survival L 95% CI U 95% CI n collars censozedDied Nonhunt Hunt 0.91 0.81 1. 00 33b 0 3 3 0 0.90 0.79 1. 00 30b 0.96 0.88 1. 00 26 0.08 0.00 0.19 25 o o 23 o 1c 1 1 3 o 23 o o o o FEMALES Survival L 95% CI U 95% CI n collars 0.89 0.78 0.99 36 0.97 0.91 1. 00 32 0.97 0.90 1.00 30 0.93 0.83 1. 00 29 0.96 0.89 1. 00 27 0.85 0.70 0.99 26 censo.red= o o o o o Died Nonhunt Hunt 4 4 1 2 1 4 o 1e 1 1 o o o o 1 o 2 1 4 3 o 1. 00 2d 0.50 0.00 1. 00 2 1. 00 o o o o o 1 o 1 1 C 31 4 27 1.00 22 o o o o Censored denotes collar failure and/or animal life/death status not known. b Includes (173.949/94) collar failure December 1994 but seen January 1996 Censored elk was (173.949/94) collar failure, which was killed in first rifle season d Includes (174.030/94) alive June 1997 . • Censored elk was (173.719/94) slipped collar May 1996. 0.03 0.00 0.09 32 lC 0.63 0.46 0.79 35 1d 13 5 8 1996. I-' 10 W �Table 4. Survival rates, inclusive of nonhunting and hunting mortalities, for winter-spring (WS, 1 December-14 June) and summer-fall (SF, 15 June-30 November) time periods from 1 December 1995-14 June 1998 for the cohort of calves age 6 months radio-collared in December 1995. Survival rate for male and female calves pooled when age 6-11 months was 0.88 in 1995 (95% CI 0.81-0.96, n=69). Survival rates (S) calculated as a mean estimate of (alive)/(alive+dead) and variance of S(l-S)/n collars. Elk Age (months) and Time Period (WS or SF dates) 6-11(mos) 12-17 18-23 24-29 30-35 6-35 WS SF WS SF WS ALL 1997 1997-98 1995-98 1996-97 1996 1995-96 MALES Survival L 95% CI U 95% CI n collars 0.86 0.74 0.98 35 censored- 2h 0.83 0.68 0.97 2,9. F 5 0.96 0.87 1. 00 24 o 0.23 0.04 0.41 22 1d 17 o 1 1 5 o 17 0.93 0.82 1. 00 4 1e 28 1.00 0.58 0.40 0.76 31 5 5 0 FEMALES Survival L 95% CI U 95% CI n collars 0.91 0.81 1. 00 34 0.87 0.75 0.99 31 27 0.83 0.66 0.99 23 0 o o 2f .19 3 3 0 4 2 4 o 1 1 o o o o cenaoredDied Nonhunt Hunt • Censored b Censored Censored d Censored • Censored , Censored 9 Censored C 4 1. 00 4 sh,e,d,e o o o Died Nonhunt' Hunt o 0.13 0.01 0.24 32 18 6 22 3f,9 13 4 9 denotes collar failure and/or animal life/death status not known. elk were (174.619/95) slipped collar May 1996, (174.800/95) capture mortality. elk was (174.660/95) slipped collar July 1996. elk was (174.671/95) likely collar failure July 1997 . elk was (174.190/95) slipped collar May 1998. elk· were (174.441/95) slipped collar August 1997. and (174.580/95) likely collar elk was (174.470/95) likely collar failure December 1997. failure July 1997. •.... 10 ,j:o �Table 5: Survival rates, inclusive of nonhunting and hunting mortalities, for winter-spring (WS, 1 December-14 June) and summer-fall (SF, 15 June-30 November) time periods from 1 December 1993-14 June 1998 for the cohort of calves age 6 months radio-collared in December 1996. Survival rate for male and female calves pooled when age 6-11 months was 0.86 (95% CI 0.81-0.96, n=69). Survival rates (S) calculated as a mean estimate of (alive)/(alive+dead) and variance of S(l-S)/n collars. Elk Age (months) and Time Period (WS or SF dates) 6-11(mos) 12-17 18-23 6-23 ALL WS SF WS 1996-98 1997-98 1997 1996-97 MALES Survival L 95% CI U 95% CI n collars cenaor-ed- 0.85 0.73 0.98 34 1b Died Nonhunt Hunt 5 5 0 FEMALES Survival L 95% CI U 95% CI n collars censo.redDied Nonhunt Hunt 0.86 0.74 0.98 35 0 5 5 0 • Censored b Censored 1. 00 0.97 0.90 1. 00 29 0 1 0 1 28 0 0 0 0 1. 00 1. 00 30 0 0 0 0 30 0 0 0 0 0.82 0.69 0.96 34 1b 6 5 1 0.86 0.74 0.98 35 0 5 5 0 denotes collar failure and/or animal life/death elk was (174.619/96) capture mortality. status not known. I-' 1.0 til �Table 6. Survival rates, inclusive of nonhunting and hunting mortalities, for winter-spring (WS 1 December14 June) and summer-fall (SF, 15 June-30 November) time periods from 1 December 1993 - 14 June 1998 for all Survival rates (S) calculated as a mean estimate of radio-collared adult female elk age ~12 months. (alive)/(alive + dead) and variance S(l-S)/n collars. Elk Age and Time Period (WS or SF dates) Adult Agylt Adult Adult Adult Adyl!;; Adylt AQylt AQJ.!.lt WS SF WS SF SF WS WS SF WS 1996 1995 1995-96 1996-97 1997 1997-98 1994-95 1994 1993-94 0.94 0.89 0.98 0.94 0.91 0.96 0 .. 99 0.'87 0.96 Survival 0.90 0.84 0.95 0.90 0.87 0.97 0.92 0.80 0.91 L 95% CI 0.94 0.98 1.00 0.98 0.96 1.00 1.00 0.94 1.00 U 95% CI i k m b 129b h 1271 100b f 68b e 95 9 119 143 152 138 n collars j n 1P 2 0 0 2 0 0 0 0 censoxed" 7 16 8d 3 13 2 4 13c 3 Died 4 1 1 0 0 1 1 1 1 Nonhunt 3 15 2 8 13 1 3 12 2 Hunt ---- 0 b Includes (172.011/93) collar failure April 1994, seen January 1996 • Censored denotes collar failure or life/death status unknown. d Collar (172.207/93) disappeared 1995 hunt seasons, assumed dead. • Collars (172.800/93,172.649/93) disappeared 1994 hunt seasons. f Composed of 35-12+, 6-24+, and 59-36+ months old females. • Composed of 6-18+ and 62-30+ months old females. h Composed of 32-12+, 34-24+, and 63-36+ months old females. 9 Composed of 34-18+ and ·61-30+ months old females. j Censored (172.011/93) failure and (173.719/94) slipped collar. 1 Composed of 30-18+ ~d 89-30+ months old females. I Composed of 27-18+ and 100-30+ month old females. k composed ot 31-12+, 29-24+, and 83-36+ months old females. m Composed of 30-12+, 23-24+, and 99-36+ months old females. n Censored (174.4-41/95) slipped collar August 1997 (174.580/95) likely failure June 1997 p Censored (174.470/95) likely collar failure August 1997. o Composed of 30-18+ and 108-30+ months old females. I-' 10 0'1 �Table 7. Survival rates, inclusive of nonhunting and hunting mortalities, for winter-spring (WS 1 December14 June) and summer-fall (SF, 15 June-30 November) time periods from 1 December 1993 - 14 June 1998 for radio-collared adult female elk age ~24 months. Survival rates (S) calculated as a mean estimate of (alive)/(alive + dead) and variance S(l-S)/n collars. Elk Age and Time Period (WS or SF dates) Adult Adyl!;; Adult 8,gult Agylt; 8,gylt; Agylt 8,gylt; Agyl!;; WS SF WS SF SF WS WS SF WS 1995 1995-96 1996· 1996-97 1997 1997-98 1994-95 1994 1993-94 0.93 0.89 0.93 0.99 0.89 0.98 0.82 0.93 0.95 Survival 0.84 0.88 0.88 0.97 0.84 0.96 0.87 0.72 0.90 L 95% CI 0.98 0.99 0.95 1. 00 0.95 1.00 0.91 0.99 1.00 U 95% CI 112 97 89 100 122 108 61 65 62 n collars 1b 1d 0 0 0 2c 0 0 cenaoxed0 7 6 12 1 13 2 4 12 3 Died 3 1 0 0 0 1 1 1 1 Nonhunt ·3 11 7 1 13 1 3 11 2 Hunt • Censored o Censored denotes collars collar failure or life/death (174.441/95) (174.580/95) status unknown. S d Censored Censored collar collar (172.011/93) 174.470/95) Table 8. Survival rates, inclusive of nonhuntng and hunting mortalities, for winter-spring (WS, 1 December14 June) and summer-fall (SF, 15 June-30 November) time periods from 1 December 1993 - 14 June 1998 for the group of adult female elk age ~18 months when radio-collared in December 1993. Survival rates (S) calculated as a mean estimate of (alive)/(alive + dead) and variance S(l-S)/n collars. Elk Age and Time Period (dates) Adult Adult 8,dylt Agylt; 8,dult Agyl!;; Adyl!;; 8,gl.l.lt My,lt WS SF SF WS WS SF WS SF WS 1995 1995-96 1996 1994-95 1996-97 1997 1994 1997-98 1993-94 0.93 0.92 0.88 1. 00 0.93 0.94 0.82 0.97 0.96 Survival 0.85 0.78 0.84 0.85 0.87 0.72 0.92 0.91 L 95% CI 0.97 1.00 1.00 0.99 1.00 0.91 1.00 1.00 U 95% CI 49b h 53b f 42h 65b f 68b e 391 361 36j 34j n collars 19 0 0 0 0 0 0 0 censor-ed- 0 d c 3 3 6 0 4 2 12 1 3 Died 0 3 0 0 1 0 0 1 1 Nonhunt 0 3 6 0 3 2 11 1 2 Hunt • Censored denotes collar failure or life/death status unknown o Collar (172.649/93) disappeared 1994 hunt seasons. • Composed of 6-·18+ and 62-30+ month old fe!llales. • Censored (172.011/93) collar failure April 1994. , Composed of 100% females 48+ months old. Includes (172.011/93) collar failure April 1994 but seen Jan. 1996. Collar (172.207/93) disappeared 1995 hunt seasons, assumed dead. , Composed of 100% females 24+ months old, h Composed of 100% females 36+ months old. l Composed of 100% females 60+ months old. b d I-' 10 -.J �Table 9. Summary of summer-fall (15 June-30 November) and winter-spring (1 December-14 June) survival rates inclusive and exclusive of hunting deaths among different age groupings of adult females averaged among years, 1993-94 - 1997-98. Survival rates (S) calculated as a mean estimate of (alive)/(alive + dead) and variance S(l-S)/n elk-years. Winter-Spring Average 5 Years Summer-Fall Average 4 Year's Age Grouping survival ±95% CI Elk-years Age Grouping Survival ±95% eI Elk-years Includes Hunting Age ~12 monthsb Age ~24 monthsC 1993 Cohortd Deathsa 0.90 0.89 0.88 0.03 0.03 0.05 524 396 189 Age ~18 months Age ~30 months 1993 Cohort 0.97 0.96 0.95 0.01 0.02 0.03 547 420 233 Excludes Hunting Age ~12 monthsb Age ~24 monthsC 1993 cohortd Deathse 0.99 0.99 0.99 0.01 0.01 0.01 476 354 167 Age ~18 months Age ~30 months 1993 Cohort 0.99 0.98 0.98 0.01 0.01 0.02 536 410 227 "Includes "Includes dIncludes "Includes hunting and natural deaths. bIncludes all females age ~12 months. only females age ~24 months by censoring yearling females recruited from calves. only females in 1993 trapped cohort having age 18 months to 10+ years. only natural deaths and represents natural survival rate. Table 10. Annual survival rates from 1 December-30 November each year 1993-94 - 1996-97 and overall survival through 14 June 1998 for the group of adult female elk age ~18 months when radio-collared in December 1993. Survival rates (S) calculated as a mean estimate of (alive)/(alive+dead) and variance S(1S)/n collars. Survival L 95% CI U 95% CI n collars censor-ed" Died Nonhunt Hunt Adult 19931994 0.78 0.68 0.88 68b•e 0 15C 2 13 Elk Age and Time Period Agult Adult Adult 19961995 1994 1997 1996 1995 0.94 0.86 0.84 0.87 0.75 0.75 1. 00 0.97 0.93 53b•f 36i 429 1h 0 0 d 2 6 10 0 3 1 2 3 9 " Censored denotes collar failure or life/death status unknown. " Collar (172.649/93) disappeared 1994 hunt seasons. • Composed of 6-18+ and 62-30+ month old females. 9 Composed of 36+ month old females. I Composed of 48+ month old females. (dates) AgyH 19931998 0.51 0.39 0.63 67 1h 33 6 27 Includes collar (i72.011/93) failed 4/1994 but seen alive 1/1996. Collar (172.207/93) disappeared 1995 hunt seasons, assumed dead . ' Composed of 100% females 24+ months old. h Censored (172.011/93) failure 4/1996. b d •.... 10 co �Table 11. Correction factors calculated to estimate total numbers of elk removed by all sources of hunting mortality from total numbers of elk 'legally removed' by hunting during 1994-1997 hunting seasons. Wounding Illegal Legal Total Correction Elk Sex/Age Class Or Loss Killb .xi u> xu iKilledc Factord Hunting Season fQ~ ~~~L8S~ ~lg~~~~ Males Age ~ 24 months Males Age 12-23 months Females Age ~ 12 months Females Age ~ 12 months Females Age ~ 24 months Females Age 12-23 months 57 7 2 66 0 1 12 13 1.16 1.11 e 41 15 3 59 1.44 35 12 3 50 1. 43f 37 14 2 53 5 1 1 7 1.40 44 6 2 52 1.18 26 9 0 35 1.35 Rifle Late Season for Females Age ~ 12 months 6 3 3 12 2.00 Archery for Males Age ~ 99 0 0 9 1. 00 4 3h 0 7 1. 75 4 1 0 5 1.25 EQ~ Hynting ~easQn~ Rifle pt, 2nd, & 3rd for Males Age ~ 24 months Rifle pt, 2nd, & 3rd for Females Age ~ 12 months 24 Archery for Females Age ~ months 12 mbnths Muzzleloading for Males Age ~ 24 months 1. 43 Muzzleloading for Females Age ~ 12 months 0 5 0 5 1. 00 • Legal kill equals known-killed plus dl.sappeared during hunting seasons and assumed legally killed. These eLk represent kill that could be estimated by hunter harvest surveys. b Wounding loss and illegal kills would not be estimated by hunter harvest surveys. Total elk removed by all sources of hunting. d Multiply legal kill by correction factor to estimate total elk removed by hunting. For yearling males, mUltiply modeled numbers of yearlings by 1.11 if yearling males are not legal quarry . • Removal rate measured from known illegal hunting. ! Correction factor calculated without including mortalities associated with late rifle seasons. 9 Includes one legal kill that may have been from either archery or muzzleloading seasons. h Includes one wounding loss that may have been from either archery or muzzleloading seasons. the legal C estimates of I-' 10 10 �Table 12. Data matrix for simple population model using spreadsheet software. We used a post-hunt (December) population composition of 50 calves:100 females age L12 months and 4 adult males:16 yearling males:100 adult females and an estimated population of 3,600 elk. Fall Hunting Natural Survival Natural Survival Removal Rate Elk Sex/Age Class Rate Winter-Spring Rate Summer-Fall Elk Sex/Age Class 0.80 Adult Males 1. 00 1. 00 Adult Males Age L30 months Age 2.24 months Yearling Males Age 12-17 months 1. 00 0.11 Yearling Males Age 18-23 months 0.98 Adult Females Age L 24 months 0.99 0.11 Adult Females Age 2.30 months 0.98 Yearling Females Age 12-17 months 1. 00 0.05 Yearling Females Age 18-23 months 0.98 Calves Age 0-5 months 1. 00 0.02a Calves Age 6-11 months 0.89 • Estrmated from harvest surveys. IIJ o o �201 Table 13. Elk counted during quadrat and nonrandom flights on 44 mi2 in West Divide and Alkali creeks, GMU 42, 27 February-2 March,1998. Survey Marked Elk Unmarked Total Marked/ Flight Flight Available Seen Elk Seen Elk Seen Unmarked Hours Quadrats 34A 27 744 771 0.036 17.2 NonRandom-1 38 12 420 432 0.029 3.6 NonRandom-2 38 17 534 551 0.032 4.4 Totals 38 56 1698 1754 0.033 25.2 a Four elk off sample area during quadrat counts were on sample area during nonrandom flights. Table 14. Summary of quadrat flights for each observer, 27 February - 1 March, 1998. Total Minutes Marked Elk/ Sighting Trial Seen/ Flown/. Observer Quadrats Total Unmarked Quadrat Quadrat Marked Elk (Date) Flown (Range) (Range) Seen(%) Missed Elk Seen Seen 20.6 1B 18 371 0.039 18.5 14 1 (0-68) (2/28) (0.93) (10-27) 2E (2/27) 17 3M (3/01) 9 loa 5 314 0.040 18.5 (0-55) 17.8 (10-35) 86 0.012 9.6 (0-29) 25.4 (18-39) (0.67) 1 1 (0.50) 17.5 19.7 771 0.036 25 7 (0-68) (10-39) . (0.78) aTwo elk groups each contained 2 marked elk that were seen but each group represented only 1 sighting trial. Observer 2E saw a total of12 marked elk. Totals 44 Table 15. Status of marked elk missed on quadrats during sighting trials and subsequently found using telemetry. Marked Statul2When Marked Elk Foynd %Snow %Screening Group Elk Cover Cover Habitat Observer Missed Size Behavior Sex 100 Juniper 50 Standing 1 1 IB M 2E 5 F F M F M 1 sa 1 1A 1 Bedded Bedded Bedded Moving Bedded Oakbrush Oakbrush Oakbrush Oakbrush Oakbrush 70 70 70 70 60 100 100 100 100 100 2A 50 30 Juniper Standing 3M 1 F AThese 3 elk may represent groups or portions of groups of elk that were detected during quadrat counts but marked elk and other elk in the group were not seen due to elk behavior or vegetation. For all 3 groups, additional marked or unmarked elk were seen in close proximity. �202 Table 16. Frequencies at which individual marked elk were seen during quadrat and nonrandom flights, 27 February-2 March, 1998. Maximum frequency equals 3. Frequencies represent heterogeneity of sighting probabilities among elk Total Marks Frequency Seen During Flightsa Frequency Seenb Seen 0 1 2 3 1 56 5 20 8 5 5 a Of the 56 marked elk seen, 46 or 82% were individually identified by numbers/symbols on the collars or telemetry frequency. Elk not identified had lost the numbers/symbols on collars and were seen during nonrandom flights. b Includes 5 observations of elk seen but not individually identified. C Of the 5 elk not seen, 3 were females and 2 were males. Table 17. Estimated numbers of elk in survey area based on quadrats, sighting bias adjusted quadrats, and Bowden and Immigration-Emigration Joint Hypergeometric mark-resight estimators calculated using program NOREMARK. Both mark-resight estimators account for populations not closed to movement during time of surveys. Conf. Int. Width 25% Conf. Limits Total Elk Upper Percenta Lower Elk Estimator Estimate 771 Quadrats 944 14 124 882 820 Quadrats Adj. Model 1b c 952 814 16 138 Quadrats Adj. Model 2 883 14 116 915 857 799 Quadrats Adj. Model 3d 1,430 457 973 39 Bowden Estimator 1,179 1,337 370 33 967 Imm/EmmJHE Estimator l,115e a Width as percent of total estimate. b Quadrat counts adjusted; (LOG GROUP SIZE)+(INTERCEPT); AIC=172. C Quadrat counts adjusted; (LOG GROUP SIZE)+(BEDDED)+(INTERCEPT); AIC=160. d Quadrat counts adjusted; (LOG GROUP SIZE)+(BEDDED)+(%COVER)+(INTERCEPT); AIC=154. e Represents daily population estimate on sample area. Extant population estimate was 1,162 elk. Table 18. February-1 Total Groups 148 Number and size of elk groups seen during quadrat March 1998. FI:~gyen!;;;:l!: of Grou:Q Sizes Seen 25-30 10-24 7-9 4-6 1 2-3 2 18 21 34 35 38 counts, 27 Average Group Size 5.2 �203 Table 19. Radio-collared elk captured as calves on winter range in GMU 42 during December 1996 that dispersed out of GMU 42 to a new winter range as of March 1998. Locations (GMUs) determined by aerial telemetry. Trap New Frequency Zone Agea Sex GMU Location Description porcupine Creek-Holms Mesa 172.899/96 G 21 F 42 West 173.450/96 Wallace Creek-Samson Mesa G 21 M 42 West East Fork Wallace Creek 173.789/96 42 West G 21 F Low Beaver Creek 173.870/96 G 21 F 42 West Plateau Creek-Hayes Mesa 174.059/96 G 21 M 421 Taughenbauh-Grass Mesas 174.220/96 F 21 42 West M Porcupine Creek-Holms Mesa 174.789/96LE G 21 M 42 West 174.929/96 D 421 Pole Gulch-Collier Creek 21 F East Muddy-Little Henderson Creek 174.949/96LE D 21 521 F Hawxhurst Creek~Grassy Gulch 174.969/96 D 421 F 21 East Muddy-Little Henderson Creek 174.998/96 E 521 21 F Hawxhurst Creek 175.020/96 F 421 21 F Low Beaver Creek 175.039/96 G F 42 West 21 West Muddy-Ault Creek 175.050/96LE E 521 F 21 42 West Wallace Creek-Sugarloaf Mtn. 175.070/96 E F 21 Low Fourmile Creek 175.089/96 A F 43 21 Wallace Creek-Sugarloaf Mtn. 175.180/96 E M 42 West 21 Plateau Creek-Red Mountain 175.189/96LE E M 421 21 Low Wallace Creek-High Mesa 175.199/96 G M 42 West 21 a b Age of elk in months as of March 1998. LE = Location elk routinely located. �204 Appendix I. Mortalities of 181 radio-collared elk from 1 December 1993 through 14 June 1998. Age is aBeroximate age in years of elk at death: C=calf age 6-11 months, Y=yearling age 12-23 months. Frequency 101 Q~ath Ira!2 Age Year Ca~tured Site Zone Sex Date Cause of Death & Code Number 172.030/93 BR A F 5 27-0ct-95 Legal kill second rifle season-47 172.039/93 GR B F 4 14-Jun-95 Unknown-suspect predation-30 172.070/93 BR A F 11 12-Feb-96 Unknown-11 172.080/93 GM A F 6 05-Nov-94 Legal kill third rifle season-48 172.090/93 GR B F 11 16-Jan-94 Wounding loss late rifle season-26 172.101/93 SR B F 8 14-0ct-96 Legal kill first rifle season-46 172.128/93 GC B F 8 31-0ct-96 Wounding loss second rifle season-44 172.139/93 F BC C 17 19-0ct-95 Wounding loss first rifle season-43 172.160/93 CC C F 3 23-0ct-94 Legal kill second rifle season-47 172.181/93 MC C F 3 03-Nov-94 Wounding loss second rifle season-44 172.201/93 GS C F 3 15-Nov-94 Wounding loss second rifle season-44 172.207193 SG 0 F 4 30-Nov-95 Disappear first rifle season-40 172.258/93 HY F C 3 04-0ct-94 Legal kill archery/muzzle season-27 1n.277/93 GS C F 3 04-0ct-94 Wounding loss archery/muzzle-24 172.290/93 SG 0 F 6 23-0ct-94 Legal kill second rifle season-47 172.290/94 SM 0 F 4 16-Sep-96 Legal kill muzzleloading season-34 172.300/93 AC E F 8 30-0ct-97 Wounding loss second rifle season-44 172.369/93 AC E F 3 18-Sep-94 Legal kill archery season-33 172.369/94 FM B F 4 14-Jan-96 Legal kill late rifle season-29 172.409/93 AC F E 8 29-0ec-94 Wounding loss late rifle season-26 172.459/93 MH E F 8 24-0ct-96 Legal kill second rifle season-47 172.509/93 FS H F 3 21-0ct-95 Legal kill second rifle season-47 172.542/93 FS H F 01-Feb-94 Mountain lion predation-3 6 172.549/93 PG H F 7 28-Nov-95 Legal kill late rifle season-29 172.570/93 PG H F 16 03-Nov-94 Wounding loss second rifle season-44 172.581/93 PG H F 3 17-0ct-94 Legal kill first rifle season-46 172.581194 FM B F 3 08-0ec-95 Legal kill late rifle season-29 172.590/93 PG H F 24-Apr-96 Unknown-11 5 172.610/93 PG F 04-0ct-94 Accident, birthing/calving-32 H 3 172.639193 PG F 14-Jun-96 Accident, birthing/calving-32 H 16 172.649193 PG F 30-Nov-94 Disappear first rifle season-40 H 2 172.658/93 PG F Legal kill third rifle season-48 H 5 02-Nov-97 16~Jan-95 172.670/93 PG H F Legal kill late rifle season-29 4 172.678/93 PG H F 22-0ec-94 Wounding loss late rifle season-26 9 172.690/93 FM B F 24-Jan-94 Illegal kill-7 8 172.699/93 Legal kill third rifle season-48 FM B F 7 05-Nov-95 Unknown-suspect predation-30 172.749/95 LS H F 11 07-Aug-96 172.800/93 Disappear second rifle season-41 SR B F Y 30-Nov-94 172.821/93 EG Legal kill archery season-33 B F 3 08-Sep-96 Legal kill third rifle season-48 172.890/93 F 06-Nov-96 BC C 3 Mountain lion predation-3 172.899/93 BC C F 18-Mar-94 C Legal kill first rifle season-46 172.950/93 GH 0 F 18-0ct-95 2 172.961/93 F 30-0ct-97 Disappear second rifle season-41 MG 0 4 173.000/93 25-Apr-94 Malnutrition-6 \lM G F C 07-Jan-98 Accident-fell-10 173.010/93 \lM G F 4 Legal kill first rifle season-46 173.041/93 M 19-0ct-95 GM A 2 27_'Oec-95 Legal kill late rifle season-29 173.060/93 MH E F 2 Illegal kill-7 03-Mar-98 173.070/93 MH E F 5 Legal kill second rifle season-47 173.081193 FS H F 3 22-Oct-96 Legal kill second rifle season-47 24-0ct-97 173.090/93 FS H F 4 Legal kill second rifle season-47 173.100/93 F 30-0ct-97 FS H 4 Legal kill first rifle season-46 .14-Oct-95 173.120/93 GM A M 2 Wounding loss second rifle season-44 31-0ct-96 173. 140i93 PG H F 3 Legal kill third rifle season-48 173.160/93 FS F 13-Nov-96 H 3 Illegal kill second rifle season-50 BC 29-0ct-94 173.190/93 C M Y Wounding loss thtrd rifle season-45 173.201193 GR M 30-Nov-95 B 2 Legal ki II first rifle season-ee 19-0ct-95 173.210/93 GC B M 2 Illegal kill archery/muzz. season-8 30-0ct-97 173.210/96 LM 0 M 2 Legal kill third rifle season-48 06-Nov-95 173.219/93 BC M 2 C Illegal kill second rifle season-50 15-Nov-94 M Y 173.232193 BR A 18-Mar-94 173.262/93 BC M C Unknown-" C Legal kill first rifle season-46 M 12-0ct-96 173.262/94 HM B 2 Legal kill third rifle season-48 06-Nov-95 173.269/93 MC C M 2 Legal ki LL. first rifle season-46 12-0ct-96 MC 3 173.279/93 C M Unknown-11 22-Mar-94 MC C M C 173.289/93 Legal kill muzzleloading season-34 M 18-Sep-96 173.289/94 PR A 2 Legal kill second rifle season-47 24-Oct-95 GS C M 2 173.300/93 Illegal kill first rifle season-49 30-Nov-94 HY C M Y 173.309/93 Illegal kill first rifle season-49 20-0ct-94 HY C M Y 173.320/93 Legal kill archery season-33 14-Sep-96 HM M 2 173.320/94 B !~~}~L~~ _____ ~~__E_____ ~____ ~___ ~~~~~~:?] ______ ~~2!_~L~~~!~~~~2~l~[_s:2~~~:~~ __________________ . �205 A~ndix I. (continued) Frequency 10/ Ira~ Year CaE!tured Site Zone 173.340/93 D SG 173.351/93 SG D 173.359/93 AC E 173.370/93 MH E 173.370/96 RG E 173.381/96 RG E D 173.390/93 MG 173.402193 MG D 173.410/93 WM G WM 173.420/93 G 173.429/93 MH E 173.439/93 PG H 173.450/93 PG H 173.461/93 PG H 173.461/94 CM A 173.469/93 FS H 173.469/94 PR A 173.479/93 FS H 173.492/93 FS H 173.502193 FS H 173.510/93 FM B 173.521/93 FM B 173.540/94 OG B B 173.549/94 HM 173.569/94 PR A A 173.580/94 PR 173.589/94 OG B 173.610/94 BC C 173.640/94 MC C 173.640/96 GM A 173.649/94 BC C BG 173.689/94 E 173.789/94 E MR MM F 173.819/94 MM F 173.829/94 173.859/94 SM D 173.870/94 SM D 173.919/94 BC C 173.929/94 BC C 173.939/94 BC C BC 173.959/94 C C 173.970/94 MC 173.981/94 MP D 173.990/94 BC C 174.001/94 BG E E 174.009/94 BG BG E 174.019/94 MR E 174.039/94 174.049/94 MM F E 174.059/94 MR E 174.069/94 BG 174.080/94 MR E E 174.090/94 MR ,174.101/94 F MM 174.109/94 MM F F 174.119/94 MM 174.119/95 GB B F 174.129/94 MM 174.140/94 MM F B 174~140/95 GB F 174.150/94 MM F 174.160/94 MM FM B 174.170/94 FM B 174.181/94 MS F 174.200/95 MS F 174.210/95 MS F 174.220/95 MS F 174.230/95 MD E 174.240/95 MS F 174.319/95 MD E 174.329/95 Sex M M M M M M M M M M M M M M M M M M M M M M M F F F F F F F F F F F F F F M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M F F Q~ath Date 20-0ct-97 05-Nov-95 06-Sep-95 08-Nov-95 08-Jan-97 19-Feb-97 30-Nov-95 18-0ct-95 02-Nov-95 24-0ct-95 14-Sep-95 30-Nov-95 15-0ct-95 07-Feb-94 19-5ep-95 25-Apr-94 20-0ct-96 10-Sep-95 03-Sep-95 13-Nov-96 28-Aug-95 17-0ct-95 t3-0ct-96 07-Sep-95 16-0ct-97 21-Dec-95 01-Jun-95 14-Nov-97 30-Mar-95 16-Mar-97 11-0ct-97 31-0ct-96 20-Mar-95 30-0ct-97 10-Jan-97 23-0ct-96 21-Feb-95 02-Nov-95 16-0ct-96 18-Sep-96 02-Nov-96 20-0ct-96 02-Nov-96 20-0ct-96 30-Nov-96 12-Oct-96 13-Noy-96 05-Sep-96 14-Sep-96 17-0ct-95 26-Sep-96 11-0ct-97 20-Oct-96 24-May-96 19-0ct-96 01-Mar-95 30-Noy-97 14-0ct-96 14-Mar-95 30-Noy-96 14-0ct-96 15-Sep-96 25-Apr-95 30-Nov-96 18-0ct-96 11-Oct-97 08~Apr-96 24-Apr-96 17-Apr-96 C 02-Oct-96 Y 03-Sep-96 Y Age 4 2 2 2 C C 2 2 2 2 2 2 2 C Y C 2 2 2 3 2 2 2 Y 3 Y C 3 C C 3 2 C 3 2 2 C Y 2 2 2 2 2 2 2 2 2 2 2 Y 2 3 2 Y 2 C 2 2 C Y 2 2 C 2 Y 2 C C !~~~3)2L~~ _____ ~~ ___~ ____ ~ ____ ~___ Cause of Death & Code Number Legal kill second rifle season-47 Legal kill third rifle season-48 Legal kill archery season-33 Legal kill first rifle season-46 Mountain lion predation-3 Unknown-suspect predation-30 Disappear first rifLe season-40 Legal kill first rifle season-46 Wounding loss second rifle season-44 Legal kill second rifle season-s? 'Legal kill archery season-33 Disappear first rifle season-40 Legal kill first rifle season-46 Malnutrition-6 Wounding loss archery season-50 Malnutrition-6 Legal kilL second rifLe season-47 Legal kill archery season-33 Legal kiLL archery season-33 Legal kill third rifle season-48 Legal kiLL archery season-33 Legal kill first rifle season-46 Legal' kill first rifle season-46 Legal kiLL archery season-33 Wounding loss first rifle season-43 Unknown-11 Unknown-suspect predation-30 Wounding loss third rifLe season-45 Unknown-suspect predation-30 Malnutrition-6 Legal kill first rifle season-46 Legal kiLL first rifle season-46 Unknown-suspect maLnutrition-31 Legal kill second rifle season-47 LegaL kill late rifle season-29 Legal kill second rifle season-47 Mountain lion predation-3 Illegal kill second rifle season-50 Legal kill first rifle season-46 Legal kill archery season-33 Legal ki II third riHe season-48 Legal kill second rifle season-47 Legal kill third rifle season-48 Legal kill second rifle season-47 Disappear archery/muzzLe season-21 Legal kill first rifle season-47 ILlegal kiLL rifle season-9 Legal kill archery season-33 Legal kilL muzzleloading season-34 Illegal kill first rifLe season-49 Wounding loss archery/muzzle-24 Legal kill first rifle season-46 Legal kill second rifle season-47 Unknown-suspect predation-30 Legal kilL second rifle season-47 Mountain lion predation-3 Wounding loss rifle season-25 Legal kill first rifle season-46 Mountai'n lion predation-3 Illegal kill third rifle season-51 Legal kill first rifle season-46 Legal kill muzzleloading season-34 Mountain lion predation-3 Disappear second rifle season-41 Illegal kilL first rifle season-49 Legal,kill first rifle season-46 Unknown-suspect malnutrition-31 Mountain lion predation-3 Mountain lion predation-3 Wounding Loss archery season-52 Legal kill archery season-33 lI~~~~:?~ ______ u_n!~2~JP~~~!~~:? _________________________________ : �206 Appendix I. (continued) Frequency 101 Ie!!!:! Year Captured Site Zone 174.349/95 MD E 174.360/95 MD E 174.401/95 AP D 174.420/95 AP D \JI) 174.478/95 C 174.491195 lB C 174.500/95 lB C 174.520/95 KR B 174.520/96 GM A 174.560/95 Bl A 174.609/95 GB B 174.629/95 MD E 174.639/95 MD E 174.679/95 HG E 174.689/95 AP D 174.689/96 EW G 174.700/95 AP 0 174.710/95 AP 0 VM 0 174.729/95 VM 174.740/95 0 VM 174.750/95 0 174.770/95 WD C WD C 174.780/95 174.789/95 WD C 174.800/96 BG E 174.809/95 WD C 174.820/95 WD C 174.830/95 lB C 174.851/95 lB c 174.861/95 KR B lB c 174.870/95 KR B 174.880/95 174.889/95 we G Bl A 174.899/95 lM 0 174.910/96 Be c 175.059/96 175.130/96 GM A lM 0 175.170/96 [1g!!!1:! Sex F F F F F F F F F F F M M M M M M M M M M M M M M M M M M M M M M M F F Age 2 Y F c M 2 2 2 2 C Y C y' C 2 2 Y C C 2 2 Y 2 2 2 2 C C Y 2 2 2 Y 2 2 2 2 c c c Date 13-0ct-97 22-Apr-97 22-Sep-96 11-0ct-97 13-Sep-97 10-Sep-97 16-Apr-96 21-Sep-96 25-Apr-97 14-May-97 '15-Apr-96 11-0ct-97 21-0ct-97 13-Nov-96 05-Feb-96 14-May-97 16-0ct-97 15-0ct-97 17-0ct-96 08-Nov-97 05-Nov-97 16-0ct-97 11-0ct-97 22-May-96 09-Apr-97 22-Apr-97 14-0ct-97 13-0ct-97 22-0ct-97 31-0ct-96 30-0ct-97 11-0ct-97 20-0ct-97 30-0ct-97 27-Feb-97 13-Feb-97 24-Mar-97 26-Mar-97 Cause of Death & Code Number legal kill first rifle season·46 Unknown-suspect predation-30 legal kill muzzleloading season·34 legal kill first rifle season-46 legal kill muzzleloading season-34 Wounding loss archery season-52 Unknown-suspect predation-30 legal kill muzzleloading season·34 Mountain lion predation-3 Illegal kill-7 Unknown-suspect predation-30 legal kill first rifle season-46 legal kill second rifle season-47 Illegal kill second rifle season-50 Mountain lion predation-3 Unknown-suspect malnutrition-31 Wounding loss first rifle season-43 legal kill first rifle season-46 Illegal kill first rifle season-49 legal kill third rifle season-48 legal kill third rifle season-48 Disappear first rifle season-40 legal kill first rifle season-46 Unknown-suspect predation-30 Mountain lion predation-3 Accident, fell-10 legal kill first rifle season-46 legal kill first rifle season-46 Illegal kill second rifle season-50 Illegal kill second rifle seasonc50 Wounding loss second rifle season·44 legal kill first rifle season-46 legal kill second rifle season-47 Wounding loss second rifle season·44 Unknown-suspect predation·30 Unknown-11 Mountain lion predation·3 Unknown-suspect malnutrition-31 �207 Colorado Division Wildlife Research July 1998 of Wildlife Report Job Progress state of Colorado Project No. W-153-R-ll Project No. 3002 Work Program Period Cost Center Mammals 3430 Program Elk conservation No. Covered: Report Elk Movements in Response to Early Season Hunting in the White Riyer Area July 1, 1997 - June 30, 1998 Author: Mary Conner Personnel: Dave Freddy, Gary White, Rick Kahn, Lipscomb, Bruce Gill, Jeff Madison John Ellenberger, Jim ABSTRACT The effects of early hunting seasons on the movements and distribution of elk remains controversial. Those who hunt the regular rifle seasons claim that archery and muzzle loading rifle seasons drive elk off of public lands and onto private land refuges before the regular rifle season begins, making them unavailable to most who hunt in the regular rifle season.· Landowners complain that elk responses to early hunting seasons cause elk to spend longer periods on private property at the expense of the landowner. Beginning in 1996, the Colorado Division of Wildlife sponsored a management experiment designed to test the effects of early hunting on the movements and distribution of elk residing in and around the White. River plateau. Four primary data: conclusions emerged from a preliminary analysis of 2 years of (1) In both 1996 and 1997, between 22-37% of the elk were on private land before mid-July. By mid-October, between 65-85% of the elk were on private land. The proportion of elk on private land was positively correlated with date; there was a 28-60% increase of elk on private land during the study period. (2.)The mean date of movement from public (non-refuge) to private land was not significantly different between treatments in the crossover analysis. Mean date of movement was different between treatments for 1997 but not for 1996. Mean date of movement was different between treatments for elk on the north half of the study area, but not found different for elk on the south half. The difference between late minus early date of movement was 6 days for elk receiving both treatments. �208 (3) Differences between elk on the north area and elk on the south area showed in responses of mean date of movement and proportion of elk on refuge areas. Mean date of movement between treatments was significantly different for elk on the north area but not for elk on the south area. The difference in proportion of elk on refuge areas between treatments was greater for elk on the north area than for elk on the south area. (4) Although all elk were captured on national forest land, some elk immediately moved to private land in 1996, and some elk were never located on public land during 1997 flights. "Manageable" elk include only those elk that are on public land within a month of early-season opening date. "Unmanageable" elk are those not on public land within a month of early-season opening date and are, therefore, not likely to be affected by changes in hunting patterns on public land. Between 39-64% of elk on public land one month before early-season opening date (unmanageable elk) moved to private land, and the mean date of movement including opening date in 3 of 4 cases. However, the elk that moved represent 31-41% of the entire herd (unmanageable and manageable elk). Therefore, any management decisions regarding hunter numbers is likely to affect at most 41%, and possibly less, of the White River elk. �209 ELK MOVEMENT IN RESPONSE BUNTING IN THE WHITE TO EARLY SEASON RIVER AREA Mary Conner P•N. OBJECTIVE Teat whether early-aeason hunting season causes movement of elk from areas heavy hunting pressure to areas of lighter hunting pressure. of SEGMENT OBJECTIVES 1. Estimate the mean date of movement of elk from non-refuge to refuge areas, and determine whether the timing is equivalent to the opening archery season. of 2. Evaluate alternative hypotheses livestock grazing, woodcutting, quality. about causes of elk movement such as recreationalists, weather, or forage 3. Publish analyses of elk movement in response to archery hunting in peerreviewed scientific journals. Preparation of manuscripts will begin during 1997-98. METHODS AND MATERIALS Methods and materials used in this investigation study plan reported in Conner 1996. RESULTS are described in the detailed AND DISCUSSIQN Preliminary analyses, results and conclusions of the 2-year study of the effects of early hunting seasons on the movements and distribution of elk in the White River area are summarized in Attachment 1 to this report. LITERATURE Conner, M. 1996. Elk movements White River area. Colorado Report. July 1996 Part 1. Prepared in response to early season hunting in the Division of Wildlife. Wildlife Research Pp. 43-86. by Graduate Research CITED Assistant ��211 Attachment 1 ELK MOVEMENTS IN RESPONSE TO EARLY-SEASON HUNTING IN THE WlllTE RIVER AREA: 1997 PRELIMINARY RESULTS Mary Conner EXECUTIVE March 31, 1998 SUMMARY This year was the second and fmal year of an experiment designed to determine if elk movements in the White River area were caused by early-season (archery and muzzleloading) hunting. The goal of this report is to present 1997 results in a timely manner. This interim report provides information for those involved with the White River elk to make relevant management decisions. Analyses that are more extensive will be included in my dissertation, which will be completed late spring or early summer 1998. Three main conclusions emerged from preliminary analyses of data from both years: (1) the proportion of elk located on private land (refuge) increased between July 15th - October 13th, (2) the timing of these movements weakly corresponded to the opening date of archery season, (3) elk on the north area had a stronger response to changes in archery-season opening date than elk on the south area, and (4) it is not clear how many elk in the White River area would be affected by management actions such as a reduction of archery hunting licenses. Conclusion (1): In both 1996 and 1997 between 22-37% of the elk were on private land before midJuly. By mid-October, between 65-85% of the elk were on private land. The proportion of elk on private land was positively correlated with date; there was a 28-60% increase of elk on private land during the study period. Conclusion (2): The mean date of movement from public (non-refuge) to private land was not significantly different between treatments in the crossover analysis. Mean date of movement was different between treatments for 1997 but not for 1996. Mean date of movement was different between treatments for elk on the north half of the study area but not found different for the south half elk. The difference between late minus early date of movement was 6 days for elk receiving both treatments. Conclusion (3): Differences between elk on the north area and elk on the south area showed in responses of mean date of movement and proportion of elk on refuge areas. Mean date of movement between treatments was significantly different for elk on the north area but not for elk on the south area. The difference in proportion of elk on refuge areas between treatments was greater for elk on the north area than for elk on the south area. Conclusion (4): Although all elk were captured on national forest land, some elk immediately moved to private land in 1996, and some elk were never located on public land during the 1997 flights. "Manageable" elk include only those elk that are on public land within a month of early-season opening date. "Unmanageable" elk are those elk not on public land within a month of early-season opening date and t rtherefore, not likely to be affected by changes in hunting patterns on public land. Between 39-64% of elk on public land a month before early-season opening date (manageable elk) moved to private land, and the mean date of movement included opening date in 3 of 4 cases. However, the elk that moved represent 3141 % of the entire herd (manageable and unmanageable elk). Therefore, any management decisions regarding hunter numbers are likely to affect at most 41 %, and possibly less, of the White River elk. INTRODUCTION As with the 1996 analyses, the treatment was difference in opening date of archery season and the primacy response variables were mean date of movement and proportion of elk that move from public to private land. I repeated all 1996 analyses for area x year to evaluate effects of archery hunting on elk that were on public land within a month of opening date and to compare to pilot study results (Appendix A: Methods). Because there are now 2 years of data, overall analyses including year and area effects were done to evaluate effects of changes in opening date of archery hunting. To evaluate treatment effects on mean date of movement, I did the crossover analysis described in my study plan (Appendix A: Methods). To evaluate �212 treatment effects on proportion .of elk moving to refuge areas, I included area and year effects in the logistic regression model used in 1996 to predict the proportion of elk on refuge areas (Appendix A: Methods). Data from the entire 3-month study period were used in the crossover and logistic regression analyses. PRELIMINARY RESULTS Area x Year Analyses The timing of elk movements from non-refuge to refuge areas on the early- and late-opening treatments is presented for 1996 and 1997 (Fig. lA, B). Only elk moving within a month of opening date were used for within-treatment analyses to represent effects on manageable elk.Figure 1. Histograms of number of elk moving from public (non-refuge) to private (refuge) land with respect to archery-season opening date on the early-opening (A) and late opening (B) treatments; White River elk herd, 1996 and 1997. A B 1996 north side 1996 south side Mean date of movement 8126 Historical opening date Mean Date of Movement: 915 C) c >3 o Historical opening date E ~ !2 o ZE :::l Z 1 ~ ~~ I I~ ~ ~ ~ ~ ~ ~ ~ ;~ ~ ~ ~ I~ I I~ ~ ~ ~ ~ ~ ~ ~ ;~ ~ ~ Early opening: 8124 Late opening: 1997 north side Mean Date of Movement:8118 C) c .~ Historical opening date E Mean Date of Movement: 9/6 3 E ~ ~ 2 ZE 1 :::l Z Historical opening date C) .~ ~ o 1997 south side c 3 9114 Qj 2 '0 ZE :::l Z ~ ~ ~ § ~ ~ ~ ~ ~ ~ i~ I~ I~ I~ Early opening: 8123 1 ~ ~ § ~ ~ ~ i~ ~ §~ I~ I~ I~ I Late opening 9113 For elk moving from non-refuge to refuge areas, mean date of movement was tested under the null hypothesis that the mean date of movement was not different from opening date (Table 1). Between 39-64% of elk on non-refuge areas one month before opening date moved to refuge areas within a month of opening date (Table 2). The probability that classifications are independent of opening date (Table 2) evaluated the effect archery hunting has on elk location by testing whether movements between classifications occurred independently before and after opening of archery hunting. �213 Table 1. Mean dates of movement of elk moving from non-refuge to refuge areas, opening dates of archery season, and P-values from a t-test for differences between opening date and mean date of movement for earlyand late-opening treatments; White River study area, 1996 and 1997. Early opening Late opening Mean date of movement 95% Confidence interval 26 August 17 August- 18 August 11 August- 5 September 6 September 30 August- 26 August- Opening date of early-season 24 August 23 August 14 September 13 September P = 0.578 P = 0.178 P = 0.014 P = 0.131 Ho: - d = opening date Table 2. Movements of radio-collared elk between refuge and non-refuge areas with respect to opening date, early-and late opening treatments; White River study area, 1999 and 1997. Movement Combinations: Early opening Late opening Pre-opening -> post-opening YEAR 1 - South YEAR 2 - North YEAR 1 - North YEAR 2 - South Refuge -> refuge 11 8 4 11 Refuge -> non-refuge 5 0 2 2 Non-refuge -> refuge 12 18 15 14 Non-refuge -> non-refuge 19 10 20 15 Sample size 47 36 41 42 Probability movement combinations P=0;051 P=0.047 P=0.280 P=0.027 Overall Analyses on Mean Date of Movement . The time frame for between-treatment analyses is the 3-month study period of July is- - October 13th• Results from the crossover analysis on mean date of movement indicated no area (P = 0.138) or year effect (P = 0.644); however, there was a significant random effect of elk in area (P = 0.025). There was not a significant treatment effect (P = 0.159) on the mean date of elk movement from non-refuge to refuge areas (Table 3). Sequence effect refers to the fact that each elk received either a late-early or early-late sequence of treatments. Elk on the north half received the late-early treatment sequence, while elk on the south half received the early-late treatment sequence. Area effects and sequence effects were confounded. That is, it is impossible to determine if a sequence effect is really a difference due to the fact that the north side is different from the south side because oflocation of private-land refuges, topography, etc., or due to the fact that elk getting a late-opening treatment the first year followed by an early-opening treatment the second year will behave differently than elk getting the early-late sequence. I will refer to an area effect for clarity in the remainder of the paper. There appeared to be a treatment x year and treatment x area interaction in the crossover analysis that clouded the results. In a post hoc analysis, I found that there was a difference in mean date of movement between treatments for elk moving from non-refuge to refuge areas in 1997 but not in 1996 (Fig. 2 and Table 4). �214 Table 3. Least square mean differences in days between mean dates of movement for effects specified in the crossover analysis with their 95% confidence intervals; White River elk herd 1996 and 1997. No. of days difference No. of days difference No. of days difference between areas: 8k between treatments: a, between years: /3; 7±9 2±7 6±8 Figure 2. Mean date of movement, 95% confidence interval, and archery opening date for elk on early-and late opening treatments, 1996 (A) and 1997 (B), White River elk herd. A 1996 Early open: South Mean date of movement: ~ j:::; :£ ~:£ Late open: North Aug 26 Aug 31 :£ ~ ~ ~ M ~ ~ ~ co co iii (j; Opening Date: :£ j:::;:£ :£ :£ •.... (j; ~~ (j; N C! CI> s ~ ~ ~ ~ ~ Sept 14 Aug 24 B 1997 Early open: North Mean date of movement: Opening Date: Aug 18 Late open: South Sept 3 Aug 23 Sept 13 Table 4. Differences between opening dates and mean dates of movement between early- and late-treatments, 95% confidence interval, and P-value for test of differences between mean opening dates 1996 and 1997; White River elk herd. Year Days between Days between mean P 1996 1997 opening dates 20 20 dates of movement 5 ± 11 16 ± 12 0.371 0.010 �215 I also did a post-hoc analysis to evaluate the treatment effect by area, as it appeared that elk on the north area were moving to private land more readily than elk on the south area. There was a difference in mean date of movement between treatments for elk on the north area but not for elk on the south area (Table 5). Table. 5. Differences between opening dates and mean dates of movement between early- and late-treatments for north- and south-side elk, 95% confidence interval, and P-value for test of differences between mean opening dates; White River elk herd 1996 and 1997. Area Days between Days between mean dates P Northside opening dates 20 of movement 13 ± 10 0.009 20 8 ± 14 0.267 (late-early treatment) South side (early-late treatment) I then analyzed date of movement only for individual elk that received both the early and late treatment. Only 16 elk received both treatments because some elk were not on public land for one year and some changed side of study area between years. For the elk getting both treatments, I subtracted their early date of movement from their late date of movement to account for the fact that this was a repeated measurement on the elk. I tested the hull hypothesis that: Ho: Difference in date of movement for late treatment - early treatment =0 Ha: Difference in date of movement for late treatment - early treatment> 0 . The difference in mean date of movement was 6 ± 7 days (p = 0.065). The distribution of differences in date of movement was not normal (Fig. 3); 12 of the 16 elk had a positive difference for date of movement. If movements were random, then there would be an equal number of positive and negative differences. There was a greater count of positive differences than expected if date of movement was random with respect to treatment (p = 0.038). Overall Analyses on Proportion of elk on Refuge Akaike's Information Criteria (AIC) was used to choose a model to predict the proportion of elk on refuge areas from a combination of treatment, area, year, and date effects (Fig. 4A, B). All models accounted for the repeated measurements taken on individual elk between treatments. For the lowest AIC model, the logistic regression parameters for treatment (P = 0.032), area (P < 0.001), year (P < 0.001), date (P < 0.001), and area x date (P < 0.001) were .significantly different from zero. When these factors were controlled for there was a treatment x date (P < 0.082) effect (Table 6). The borderline treatment x date effect indicated that the slope, or rate of change in proportion of elk on refuge areas, was different for the 2 treatment areas (Fig. 5). The significance of the area x date parameter indicates that the slope, or rate of change in proportion of elk on refuge areas, was greater for elk on the north side (late-early sequence) compared to elk on the south side (early-late sequence; Fig. 6). Elk on the north area showed a greater response in proportion of elk on refuge area by treatment compared to elk on the south area (Fig. 7A, B). The fitted proportion of elk on refuge for each area x year category and the corresponding predicted proportion of elk on refuge for the opposite treatment was calculated to evaluate the effect of changing archery season opening date (Fig 8A, B, C,D). �216 Figure 3. Distribution of differences in date of movement between late and early treatments for elk experiencing both treatments; White River elk herd 1996 and 1997. J!l'llBQUDlCY 7 5 3 Z 1 -35 -Z5 -15 -5 5 15 Z5 35 Figure 4. Proportion of elk on refuge areas with respect to archery-season opening date, on early- and lateopening treatments for 1996 (A) and 1997 (B); White River elk herd. 1996 A Q) 0> 0.9 ~ 0.7 - ::J 0.8 c o 0.6 .::£ Q) 0.5 o 0.4 c o 0.3 ••e a. e a.. r: Early treatment ,... .•. , , " -"'" .• , ....•.... , .• " ". " , ..• •• I I I 0.2 Late treatment North side 0.1 0.0 - - <0 en it; ,.... <0 I'- I'- en N ~ <0 s~ I'- - - -- - <0 <0 <0 <0 ~ N .•.. .•.. <0 CD en CD en en CD en ~ CD <0 en ~ en Early opening date: Aug 24 -- -- <0 en en 0) <0 en <0 ,.... 0) c <0 <0 <0 ~ ~ 0) M 0) - t: Late opening date: Sept 14 ~ 0 en 0 �217 Figure 4. (Continued) 1997 B - 0.9.--------------------------------------------. (1) ~ - , ... _- g> 0.8 , ~ ,, 0.7 c: o 0.6 .::tt:. (1) 0.5 '0 0.4 c: o Io 0:. - •. 0.3 0.2 I I ..• Early treatment "", North side 0.1 0.0 +-.--.--,--,...,r-r-,--,--.--h----.---.--,--,--.--fr-...-.--.--,--,...,r-r--l •..... m - 0 ~ •..... ~ ~ •..... m •..... ~ •..... m •..... ...,. •..... m •..... m •..... m CD CD CD C! C') CD m •..... •..... m •..... •..... m -- -.•.. -.•..- - -- se- - s- -- s- C') CD Early opening date: Aug 23 .•.. ~ ...,. .•.. ...,. m .•.. 0 m Late opening date: Sept 13 Table 6. Parameter estimates for logistic regression model used to predict the proportion of elk on refuge areas; White River elk herd 1996 and 1997. INTERCEPT SEQUENCE TREAT JULDAY YR JULDA Y*TREAT -7.2244 4.2030 l.2382 0.0305 -0.2395 -0.0041 0.5082 0.5757 0.5684 0.0021 0.0606 0.0023 0.0001 0.0001 0.0323 0.0001 0.0001 0.0821 �218 Figure 5. Treatment x date effect for elk on early- and late-opening treatments 1996 and 1997; White River elk herd. . Treatment Effect 0.9 Q) C> :::l ~ c: 0 ~ Q) 0 c: 0 1:: 0 •• • •• 0.8 Early opening 0.7 0.6 •• ••• • • • • 0.4 • 0.3 •• • • •• • • -, I • 0.5 a. 0.2 e o, ~. • • • • •• • • • •• Late opening 0.1 • Early opening • Late opening 0 7/8 7/22 8/5 8/19 9/2 9/16 9/30 10/14 Date Figure 6. Area x date effect for elk given the early-late or the late-early treatment sequence during 1996 and 1997; White River elk herd. Area Effect 0.9 Q) - 0.8 0 0.3 a. 0.2 C> :::l ~ c: 0 ~ 0.6 Q) 0.5 0 0.4 c: 1::0 e n, early-late treatment 0.7 • •• • •• South side elk • • • • • ••• • •• • • • North side elk late-early treatment 0.1 • South-side elk • North-side elk 0 7/8 7/22 8/5 8/19 9/2 Date 9/16 9/30 10/14 �219 Figure 7. Fitted curves of proportion of elk on refuge areas by treatment from logistic regression analysis for elk on the north area (A) and elk on the south area (B); White River elk herd 1996 and 1997. (A) North Area: Elk movement to refuge by treatment 0.9 0.8 Q) C) :::J ~ c 0 ~ Q) 0 c 0.7 0.6 0.5 0.4 0 1:: 0 0.3 e a, 0.2 a. -North 0.1 - early opening - fitted curve North late opening - fitted curve 0.0 7/28 7/8 9/6 8/17 9/26 10/16 Date (B) South Area: Elk movement to refuge by treatment 0.9 0.8 Q) C) :::J ~ c 0.7 ~ 0.5 ~- 0.6 0 Q) 0 c 0.4 ~ 0 1::0 0.3 e o, 0.2 a. -South 0.1 - early opening - fitted curve South late opening - fitted curve 0.0 7/8 7/28 9/6 ' 8/17 Date 9/26 10/16 �220 Figure 8. Fitted curves and predicted curves for reversed treatments, early treatment on south area 1996 (A), late treatment on north area 1996 (B), early treatment on north area 1997 (C), and late treatment on south area 1997 (D); White River elk herd. (A) (C) South Side - early treatment 1996 North Side - early treatment 1997 0.9,-----------------'---, 0.8 Q) { 0.7 c o 0.6 ~ '0 c 0.5 :E 0.3 8. e n, 0.4 0.2 I-Fitted early I - Predictedlate 0.1 0.0 +----.------r----,------r----I 7/8 7128 8117 9/6 9/26 7/8 10/16 7128 8117 Date 9/6 9/26 10/16 Date (B) (D) North Side - late treatment 1996 South Side - late treatment 1997 0.9,-------------------, 0.8 Q) Q) 0.8 0> ~ !!.! 0.7 ~ 0.7 c o 0.6 5 0.6 ~ 0.5 "'Gi " 0.5 ~ 0.4 c .2 o '0 0.4 :e 0.3 8. 0.3 e o, 0.2 £. 0.2 8. 0.1 0.0 0.1 .1---.-------,,.-----.--=::::;::====-1 7/8 7128 8/17 9/6 Date DISCUSSION 9/26 0.0 10/16 I -Fitted late . _ Predictedearly. I .I-----,r-----,-----r----==:;:::~~ 7/8 7128 8117 9/6 9/26 10/16 Date AND FUTURE WORK Although all elk were captured on national forest land, some elk moved to private land early in the summer before the ± 1 month time frame used for within-half analyses, and some elk were never located on public land during the 1997 flights. "Manageable" elk are the proportion of the herd on public land within a month of early-season opening date. "Unmanageable" elk are the proportion of the herd not on public land within a month of early-season opening date and are, therefore, not likely to be affected by changes in hunting patterns on public land. In determining effects of archery hunting activity on elk movement, two types of effects were examined; the effects on manageable elk, and the effects on total herd (manageable plus unmanageable elk). Between 39-64% of the manageable elk moved to private land within a month of archery-season opening date. For the manageable elk, confidence intervals on mean date of movement included opening date in 3 of 4 cases, and movements of elk between refuge and non-refuge areas was not independent of opening date in 3 of 4 cases. North-side elk experiencing the late-opening treatment were the exception in the tests, possibly due to several early movers that attenuated the treatment effect. In general, it appears that early season hunting activity has an effect on the manageable elk. �221 The effects of the manipulation of archery-season opening date on elk movements on the entire herd was evaluated by responses in mean date of movement and proportion of elk on refuge. The strongest test is the crossover analysis because it controls for external sources of variation when testing for treatment effects. The lack of a treatment effect in the crossover analysis is evidence that early-season hunting is not affecting the timing of elk movements from refuge to non-refuge areas. However, because there appeared to be interactions between treatment x area and treatment x year, post hoc analyses were done which reveled several pieces of evidence indicating that early-season hunting activity altered elk movements: (1) mean date of movement was different between treatments for elk on the north area, (2) mean date of movement was different between treatments for 1997, (3) for elk getting late and early treatments, there was a 6 day difference in late minus early date of movement, and (4) there was a significantly greater count of positive late minus early date of movement differences than expected if date of movement was random with respect to treatment. Also, from the logistic regression analysis of proportion of elk on refuge, the rate at which elk move from public land to private land refuge areas was borderline different for early- and late-opening treatments. Part of the ambiguity in evaluating the effects of archery hooting on elk movements may result from differences in movement patterns of elk on the north side versus elk on the south side of the study area. The steeper slope, or faster gain in proportion of elk on refuge areas over time for the north-side elk (Fig. 6) suggests that north-side elk move to private land more readily than south-side elk. This was surprising as south-side elk, which experienced the early-opening treatment the first year, were expected to move more readily the second year. Similarly, north-side elk, which experienced the late-opening treatment the first year would be taken by surprise by an early opening the second year, and were expected to move less readily. In addition to moving more readily, elk on the north area were more responsive to changes in opening of archery season. The timing of the movements of north-side elk was affected by archery season opening date, as indicated by the signiftcant difference in mean date of movement between treatments. South-side elk did not have as wide a spread in their mean date of movement between treatments and there was not a signiftcant difference in mean date of movement between treatments. Also, the proportion of elk on refuge was greater during the early treatment compared to the late treatment for elk on the north area (Fig. 7A). For elk on the south area, the proportion of elk on refuges was similar for early and late treatments (Fig. 7B). It may be that the north-side elk move more readily due to the presence of a large private-land refuge adjacent to national forest land. Between 39-64% of the manageable elk moved to private land within a month of early-season opening date. However, the elk that moved represent 31-41 % of the entire herd. Hence, any management decisions regarding hunter numbers are likely to affect at most 41%, and possibly less, of the White River elk. The question remains as to what percentage of the manageable elk would still move if hunting were reduced or eliminated. These results are not complete; I still need to analyze 1997 hunter survey data, test for elk avoidance of livestock, and calculate daily distances moved and elevational shifts with respect to early-season hunting activity. Additionally, I need to reclassify refuge areas; there needs to be a third land classiftcation for habitat refuges. Habitat refuges will be defmed as topographically steep areas where changes in elevation per linear distance are greater than some threshold (to be determined). Elk locations will be reclassifted as public, private, or habitat refuge, and reanalyzed in a multinomial model. Although it appears that north-side elk move more readily than south-side elk, results may change when habitat refuge areas are included in analyses. South-side elk may not need to move to private land because they move into habitat refuges. It may be that elk with available habitat refuges are of little management concern with respect to private-land problems. These results will be incorporated and discussed in my dissertation. ��223 APPENDIX A Methods METHODS The study area was split roughly east-west by the White River and North Fork of the White River into 2 halves for application of early- or late-opening treatments during 1996 and 1997. Game Management Unit (GMU) 12 and part of GMUs 23 and 24 comprised the north half, while GMU 33 and part of GMUs 23 and 24 comprised the south half. There was a 3-week difference in opening dates; archery hunting opened early, August 24th, on the south half, and late, September 14th,on the north half during 1996. Treatments were reversed in 1997; archery hunting opened late, September 13th on the north half, and early, August 23rd, on the south half. Although data were collected on each elk from July 15 - October 13 for both years, withintreatment analyses were restricted to locations collected within a month of opening date (i.e. July 23 September 24 for early-opening treatment, and August 13 - October 13 for the late-opening treatment). The ± l-month interval allows evaluation of treatment effect on manageable elk and allowed for comparison to pilot study results. All other analyses were done over the 3-month study period for valid comparison of between-treatments effects on the entire herd (manageable plus unmanageable elk). The primary response variables were mean date of movement and proportion of elk that change classification (non-refuge ->refuge I public ->private). All locations were labeled as either refuge or nonrefuge based on land ownership or management, and not on topographical considerations. That is, the Flat Tops Wilderness Area and all private land were labeled as refuge, while all National Forest, State Wildlife, and BLM lands were labeled as non-refuge. Area x Year Analyses This is a geographically nested design, with 2 levels of analysis. One level of analysis is within half, where the treatment is location on non-refuge or refuge areas. The time frame for within treatment analyses is ± 1 month of opening date. The ± 1 month interval allows evaluation of treatment effect on manageable elk. If hunting has no effect on elk movements, then elk movements from hunted non-refuge areas should not correlate with opening date. The primary hypothesis tested were: 1. Ho: Mean date of movement from non-refuge to refuge areas = opening date. Ha: Mean date of movement from non-refuge to refuge areas ~ opening date. 2. Ho: Location of elk on non-refuge and refuge areas was independent of the opening of archery hunting. Ha: Location of elk on non-refuge and refuge areas was not independent of the opening of archery hunting. These were the same hypotheses tested for the 1992,..1995pilot study data; they were tested separately for both north and south halves of the study area. Differences in time of movement were tested with a one sample r-test, . Hypothesis 2 evaluated the effect archery hunting had on elk movements by testing whether changes in classifications (refuge->refuge, refuge->non-refuge, non-refuge->refuge, and non-refuge->nonrefuge) were independent of the presence of archery hunting. If archery hunting had no effect, then there should be no difference in elk locations before or after the opening of archery season. A chi-square test was used to evaluate if movement of elk between non-refuge and refuge areas was independent of being before or after opening date. �224 Overall Analyses on Mean Date of Movement The experimental unit is north or south half of the study area, the sampling unit is individual elk, and the treatment is early or late opening of archery season in this 2-period crossover design. The time frame for all overall analyses is the 3-month study period of July 15th - October 15th; the 3-month interval allows valid comparison between treatments for the entire herd (manageable and unmanageable elk). The crossover analysis used data from both years of the study to test the effects of early-season hunting on the mean date of elk movement from non-refuge to refuge areas (see methods: Elk movements in response to early-season hunting in the White River area: study plan). If hunting had no effect on elk movements, then there should be no difference between the mean date of movement to refuge areas between treatments. To test for a treatment effect on mean date of movement, the area (sequence) in which the treatments were applied, .the year, and the random effect of the individual sample elk must be accounted for. The corresponding analysis of variance model is: where: = = Yjlk] J.l a; = = = f3j = t)k m(k) date of movement for the [Ih elk, in area k, treatment i, and year} overall mean date of movement (for all elk, both years) fixed effects due to area k (either north or south) random effects due individual elk I in area k fixed effects due to treatment i (early or late opening) fixed effects due to year} (year 1or year 2). Specific questions tested in the crossover ANOV A model were: I. 2. 3. Mean date of movement for north-side elk (early-late treatment sequence) = mean date of movement for south-side elk (late-early treatment sequence), t)k = O. Elk randomly chosen from the south side (early-late sequence) were the same as elk randomly chosen from the north side (Iate-early sequence); trJ(k) -= O. Mean date of elk movement for early- opening treatment = mean date of elk movement for late opening treatment; ai = Mean date of elk movement of elk for year 1of experiment = mean date of elk movement of elk for year 2; /3j = O. o. 4. Hypothesis 3 is the experimental hypothesis; hypotheses 1, 2, and 4 are needed to account for other sources of variation. Overall Analyses on Proportion of elk on Refuge The logistic regression model used in 1996 to test for treatment and date effects on the proportion of elk located on refuge areas (p): <. 10git(p)=Po + p,(date)+ P2(treatment) + P3(treatmentxdate)+&, was expanded to include area and year effects. Various combinations of treatment, area, year, and date were modeled. Akaike's Information Criterion (AIC) was used to select the best model given the precision and bias tradeoff. The DSCALE option was used in this analysis to account for the repeated measures on the individual elk and allow for inference to the entire herd. The model with the lowest Ale was: logit(p) = Po + PI (area) + P2 (year) + P3 (date) + P4 (treatmemt) + P6(areaxdate) + &. + P5 (treatment xdate) �225 From this model the following hypotheses were tested: I: 2: 3: 4: 5: 6: Proportion of elk on refuges was independent of the area, given date, treatment, and year differences were controlled for; /3, = O. Proportion of elk on refuge areas was independent of year, given date, treatment, and area differences were controlled for; /32 = O. Proportion of elk on refuge areas was independent of date, given area, year, and treatment differences were controlled for; /33 = O. Proportion of elk on refuge areas was independent of treatment, given area, year, and date . differences were controlled for; /3r O. Early and late treatment had the same slope, that is, elk moved from non-refuge to refuge areas at the same rate on early- and late-treatment areas; /3s= O. The south and north side had the same slope, that is, elk moved to refuge areas at the same rate whether they were on the south side (early-late treatment sequence) or on the north side (late-early treatment sequence); /36= O. Hypotheses 5 is the experimental hypotheses. Hypothesis 5 was a test of proportion on refuge and timing of movement to refuge; that is, if archery hunting had no effect, then elk would be expected to move to refuge areas at the same rate for early- or late-opening treatments. A likelihood ratio test of the area x date effect (/3,)was an overall test of archery effects, given that elk would move the same on the 2 areas otherwise. Hypotheses 1-4 allow for accounting of additional sources of variation. Hypothesis 6 tested whether the rate of movement to refuge areas was different by area. This test was more biologically interesting than testing for a difference in the proportion of elk on refuge between areas. For example, a difference between areas may only indicate that more elk happened to begin on refuge areas on the north or south side, which is irrelevant to archery hunting activity. However, if elk on the south side receiving the early-treatment the first .year were more wary the second year, then the south side elk should show a greater rate of movement than the north side (late-early sequence) elk. This effect was tested by the inclusion of the area x date interaction term in the logistic model. ��227 Colorado Division Wildlife Research July 1998 of Wildlife Report JOB PROGRESS state of Project No. Work Package Colorado Cost Center W-lS3-R-11 Mammals No. ~3~0~0~2~ Task No. Period REPORT _ 3430 Program Elk Conservation Monitoring and Managing-Chronic Wasting Disease in Elk Covered: July 1, 1997 - June 30, 1998 Authors: M. W. Miller Personnel: S. Berry, Wheeler, K. Larsen, K. I. O'Rourke, T. R. Spraker, S. Tracy, E. M. A. Wild, and E. S. Williams ABSTRACT Elk from throughout Colorado were examined for occurrence of chronic wasting disease using a combination of targeted surveys and harvest or road-kill surveys. We continued to develop and modify a statewide targeted surveillance program for acquiring, examining, and reporting on CWO suspects submitted from Colorado. Betweep June 1997 and May 1998, 2 chronic wasting disease (CWO) cases were diagnosed among 6 "suspect" elk submitted from endemic game management units (GMUs) in northeastern Colorado; CWO was not diagnosed in any of 6 additional "suspect" elk submitted from elsewhere in Colorado. Harvest surveys were used to estimate CWO prevalence in enzootic management units. About 0.2% of elk harvested in Larimer County data analysis units (DAUs) (E4 or E9) tested positive for CWO via immunostaining; CWO was not detected in any of the harvested elk submitted from outside Larimer County. Requiring hunters to participate in harvest surveys continued to increase the number of samples submitted for CWO examination. Data from both targeted surveillance and surveys indicate that Larimer County remains the only focus of CWO in Colorado elk, alt-hough some undetected natural spread may be occurring. Compared to deer, however, CWO is a relatively rare disease in free-ranging elk in northcentral Colorado. Targeted surveillance of clinical suspects appears to be the most sensitive approach for initially detecting CWO in elk populations throughout Colorado, and will be continued statewide. Based on low prevalence detected in 1995-1997 surveys, future harvest and road-kill surveys will be conducted at about 5-yr intervals to estimate prevalence, monitor trends, and compare rates among endemic elk DAUs. ��229 MONITORING AND MANAGING CHRONIC WASTING DISEASE IN ELK M.W. Miller P. N. OBJECTIVES (1) Design, conduct, and report results of: (a) targeted surveillance to estimate and detect changes in distribution of chronic wasting disease (CWO) in free-ranging elk populations; and (b) harvest or road-kill surveys to estimate and detect changes prevalence of CWO in enzootic elk populations. in (2) Design, conduct, and report results of experimental studies using captive elk naturally or experimentally infected with CWO. SEGMEBT OBJECTIVES (1) Conduct and report results of targeted surveillance to estimate and detect changes in distribution of CWO in free-ranging elk populations statewide. (2) Conduct and report results of harvest surveys to estimate of CWO in DAUs E4 and E9. (3) Observe elk. epizootiological features of naturally-occurring prevalence CWO in captive INTRODUCTION Chronic wasting disease (CWO) is a disease of native deer and elk characterized by behavioral changes and progressive loss of body condition that invariably lead to the death of affected animals (Williams and Young 1992). Neither the causative agent nor its mode of transmission have been identified. There are no tests currently available for diagnosing CWO in live animals, and pos~mor~em tests require microscopic examination of brain tissue. There are no known treatments for CWO. Previous attempts to eradicate CWO from research facilities failed on at least 2 occasions (Williams and Young 1992). Although similar in some respects to other transmissible spongiform encephalopathies that affect domestic sheep (scrapie) and cattle (bovine spongiform encephalopathy; BSE; "mad cow disease"), there is no evidence suggesting CWO can be naturally transmitted to domestic livestock, or that scrapie or BSE can be transmitted to native cervids. Moreover, there is no evidence suggesting that CWO presents a threat to'human health. "Chronic wasting disease" was first recognized by biologists in the 1960's as a disease syndrome of captive deer held in wildlife research facilities in Ft. Collins, CO, and was subsequently recognized in captive deer, and later in captive elk, in wildlife research facilities near Ft. Collins, Kremmling, and Meeker, CO and Wheatland, WY (Williams and Young 1980, 1982). Since 1981, 74 cases of CWO have also been diagnosed in free-ranging mule deer, white-tailed deer, and elk from northeastern Colorado; most of these diagnoses have been made since 1990 (Spraker et al. 1997; M. W. Miller, unpubl. data). At present, the known world-wide distribution of CWO in wild cervids appears to �230 be limited to northeastern Colorado and southeastern Wyoming. Although CWO was first diagnosed in captive cervids, the original source of CWO in either captive cervids or free-ranging cervids is unknown; whether cwo in captive cervids really preceded CWO in wild cervids, or vice versa, is equally uncertain (Spraker et ale 1997). In Colorado, free-ranging CWO cases in elk have all originated from along the northern Front Range. Game Management Units (GMUs) yielding infected elk include 7, 9, 19, 191, and 20. All clinical elk cases have come from two Data Analysis Units (DAUs)(E4, E9). To date, examinations of elk from other parts of Colorado have not detected CWO. Preliminary data from hunter-killed deer indicate that even in enzootic areas CWO is relatively rare: it probably affects ~1.S% of the elk in E4 and E9. No real trend in prevalence (increasing or decreasing) can be discerned from data available to date. In the absence of historical (20-30 years ago) prevalence data or reliable estimates of transmission rates, it is also unclear whether incidence of CWO in northcentral Colorado DAUs is stable or increasing, or whether short-term observations can accurately forecast long-term trends. The significance of CWO and its impacts on native elk populations are unclear. Preliminary results of simulation modeling to predict dynamics and impacts of CWO on affected deer populations suggest that, sustained at current prevalence (about 6.5% in deer), CWO could impact wild deer herds and lead to population declines (M. W. Miller and C. W. Mccarty, unpubl. data). Although CWO prevalence is lower in elk than in deer in enzootic DAUs, it is conceivable that impacts could occur if prevalence were allowed to increase. Clearly, reliable estimates of CWO prevalence in wild elk populations are needed to guide policy decisions and monitor efficacy of management efforts. In the absence of data to the contrary, and considering the difficulties inherent in eliminating CWO from captive or wild cervid populations once established, it seems most prudent to assume CWO could adversely affect native deer or elk populations and manage to reduce its occurrence and prevent its further spread. Consequently, the Colorado Division of Wildlife needs to undertake a variety of actions to further understanding about CWO and its management through surveillance and experimental research in order to reduce the occurrence of CWO and minimize the risk of its spread to other native deer or elk populations in Colorado. MATERIALS AND METHODS Surveillance We monitored elk populations throughout Colorado combination of targeted surveillance and harvest were organized and conducted as follows: for occurrence of CWO using a or road-kill surveys. These Targeted (- clinical disease) surveillance: Elk showing clinical signs consist~nt with those seen in chronic wasting disease were collected by field personnel statewide and brain tissues examined for evidence of spongiform encephalopathy. The "suspect case" profile was defined as follows: • • • Species: Age: Signs: elk ~ 18 months emaciated and abnormal behavior &/or indifference to human activity &/or �231 increased salivation &jor tremor, stumbling, incoordination &jor difficulty or inefficiency in chewing/swallowing &jor increased drinking and urination Where possible, submissions were subjected to complete necropsy; in some situations, only heads were available for examination and sampling. In all cases, histopathology of brain tissue (Williams and Young 1993) was used to diagnose CWO; in some cases, immunohistochemistry or other ancillary tests were used to confirm or support diagnoses. Harvest surveys: In order to obtain reliable estimates of CWO prevalence that will serve as a basis for monitoring responses to management interventions, we continued conducting harvest surveys on select deer populations. During the 1997-1998 hunting seasons, fresh brain and select lymphatic tissues were collected from endemic and nonendemic GMUs. Brain tissues were examined at the Colorado state University Diagnostic Laboratory for histopathological lesions (Williams and Young, 1993)or anti-PrP immunostaining reactions (O'Rourke et al., 1998) consistent with CWO infection. Because sample sizes for most individual GMUs were too small to provide reliable prevalence estimates by GMU, we pooled data by DAU for comparisons within and among species. Small sample sizes also precluded meaningful analysis of data from deer or elk harvested outside known enzootic areas. Epizootiological Studies Epidemiology of naturally-occurring CWO in captive elk (Miller and Wild): We observed captive adult elk (n = 19) held at CDOW's Foothills Wildlife Research Facility for clinical signs of CWO and submitted all mortalities for complete necropsy and histopathological examination. A manuscript describing CWO epidemiology in captive elk (Miller et al., 1998) was revised and accepted for publication. RESULTS AND DISCUSSIQN Surveillance Targeted (= clinical disease) surveillance: Between June 1997 and May 1998, 2 chronic wasting disease (CWO) cases were diagnosed among 6 "suspect" elk submitted from endemic game management units (GMUs) in northeastern Colorado; CWO was not diagnosed in any of 6 additional "suspect" elk submitted from elsewhere in Colorado. Harvest surveys: During the 1997-1998 hunting seasons, fresh brain and select lymphatic tissues were collected from 653 elk harvested in enzootic GMUs . (Table 1); 103 elk harvested or culled in other GMUs throughout Colorado were also sampled as negative controls. Estimated prevalence among harvested elk did not differ (P = 1.0) between E4 (0.2%) and E9 (0%). As in 1996, overall CWO prevalence among elk harvested in Larimer County was lower (P < 0.009) than in sympatric deer populations (see Miller, 1998). Even the foregoing prevalence estimates may be somewhat liberal because the definition of "positive" included subclinical cases where either histopathological lesions or anti-PrP immunostaining reactions in brain tissue were observed. The elk case identified in 1997, as well as all 4 elk cases in 1996, were classified as positive solely on the basis of immunostaining reactions. Although no known "false positives" were identified among the 103 elk examined from outside known enzootic DAUs in 1997, further evaluation of �232 both sensitivity and specificity appears warranted. of existing diagnostic techniques still During the 1997-1998 rifle seasons, harvest survey participation was required of successful elk hunters in both E4 and E9. In E4, 454 successful elk hunters submitted heads in 1997, compared to only 81 voluntary submissions in 1996. These observations reemphasize previous observations (Miller, 1997) that compelling participation in harvest surveys via regulation is the single most effective method for increasing sample sizes. Data from both targeted surveillance and surveys indicate that Larimer county remains the only focus of CWO in Colorado elk, although some undetected natural spread may be occurring. Compared to deer, however, CWO is a relatively rare disease in free-ranging elk in northcentral Colorado. Targeted surveillance of clinical suspects appears to be the most sensitive approach for initially detecting CWO in elk populations throughout Colorado, and will be continued statewide. Based on low prevalence detected in 1995-1997 surveys, future harvest and road-kill surveys will be conducted at about 5-yr intervals to estimate prevalence, monitor trends, and compare rates among endemic elk DAUs. Epizootiological Studies Epidemiology of naturally-occurring CWO in captive elk (Miller and Wild): None of the 19 captive adult elk held at CDOW's Foothills Wildlife Research Facility developed clinical CWO during June 1997-May 1998; one female (G86) that died showed no histological evidence of CWO. We will continue to monitor this naturally-infected captive herd and examine all mortalities for evidence of cwo. A manuscript describing CWO epidemiology in captive elk (Miller et al., 1998) was accepted for publication. Judging from requests for prepublication copies, this manuscript appears to be of considerable interest to those investigating and managing recently recognized foci of cwo in privately-owned captive elk throughout the US and Canada. ACKNOWLEDGMENTS The statewide CWO monitoring and surveillance program described here relies heavily on efforts of dedicated field personnel throughout the Colorado Division of Wildlife, and truly represents a division-wide effort to improve our understanding and management of this important disease problems. In addition to those specifically listed, we collectively thank all of those regional and area biologists, district and area wildlife managers, volunteers, deer and elk hunters, and others who assisted by submitting suspect cases, harvested animals, or road-killed animals throughout the year. LITERATURE CITED Miller, M. W. 1997. Monitoring and managing wildlife chronic wasting disease in Colorado. in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-lO, WP2, J17. Colorado Division of Wildlife, Fort Collins, Colorado, USA, pp. 37-46. �233 1998. Monitoring and managing wildlife chronic wasting disease in deer. in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, project W-153-R-11, WP3001, T3. Colorado Division of Wildlife, Fort collins, Colorado, USA, in press. , M. A. Wild, and E. S. Williams. 1998. Epizootiology of chronic wasting disease in captive Rocky Montain elk. J. Wildl. Dis. 34: 532538. O'Rourke, K. I., T. V. Baszler, J. M. Miller, T. R. Spraker, I. SadlerRiggleman, and D. p .•Knowles. 1998. Monoclonal antibody F89/160.1.5 defines a conserved epitope on the ruminant prion protein. J. Clin. Microbiol. 36: 1750-1755. Spraker, T. R., M. W. Miller, E. S. Williams, D. M. Getzy, W. J. Adrian, G. G. Schoonveld, R. A. Spowart, K. I. O'Rourke, J. M. Miller, and P. A. Merz. 1997. Spongiform encephalopathy in free-ranging mule deer (Odocoileus hemionus), white-tailed deer (Odocoileus virginianus), and Rocky Mountain elk (Cervus elaphus nelsoni) in northcentral Colorado. J. Wildl. Dis. 33:1-6. Williams, E. S., and S. Young. 1980. deer: A spongiform encephalopathy. 98. _____ , and Journal 1982. spongiform encephalopathy of Wildlife Diseases 18: 465-471. _____ , and Scientifique 567~ of Rocky Mountain 1992. Spongiform encephalopathies in Cervidae. et Technique Office International des Epizooties _____ , and 1993. Neuropathology deer (Odocoileus hemionus) and elk Pathology 30: 36-45. Prepared Chronic wasting disease of captive mule Journal of Wildlife Diseases 16: 89- Research Revue 11: 551- of chronic wasting disease in mule (Cervus elaphus nelsoni). Veterinary by c Wildlife elk. Veterinarian �234 Table 1. Results of 1997 CWO harvest surveys -- archery, muzzleloader, & rifle seasons. ELK DAU E4 E9 GMU # Examined # Positive 7 104 0 8 184 0 191 30 0 9 9 0 19 127 1 Prevalence (95% CI) Total 454 1 0.002 (0.0001-0.012) 20 199 Total 199 o o o (0-0.019) �235 Colorado Divison of Wildlife Wildlife Research Report July 1998 JOB FINAL REPORT state of project Work No. package Colorado Cost Center W-153-R-11 Mammals No. __ ~3~0~0~3 _ 3430 Program Predatory Mammals Management Use of Sport Hunting to Reduce Puma Depredation on Sheep Task No. Period Covered: Author: Thomas Personnel: July 1, 1997 - June 30, 1998 D. I. Beck T. Beck, J. Madison, J. Wallace; CDOW ABSTRACT A puma density sampling protocol was prepared based on probability sampling along line transects. A prototype sampling unit was designated. However, the appropriate snow conditions did not occur in Jan.-Mar., 1998 to allow for sampling. An evaluation of the logistical constraints on the technique, along with loss of management authority, resulted in the termination of the study. ��237 Use of Sport Bunting to Reduce Puma Depredation to Sheep P.N. Objectives 1. Evaluate the efficacy the density of puma. 2. Investigate on domestic of liberal the relationship sheep. Segment sport hunting of puma density Compile historic data on sport kill of puma on sheep in DAU L-7. 2. Prepare detailed estimation. 3. Conduct puma density on transect estimation METHODS to reduce to depredation levels Objectives 1. protocol quotas and puma depredation sampling transects and puma density on 2 study areas. AND MATERIALS Kill of puma (~ concolor) by hunters was tabulated from mandatory check files for the years 1988-1997 for DAU L-7. The frequency distribution of kill by time was calculated in one-week intervals in hopes of identifying periods where sampling would have minimal impact on hunting activities. Based on location of historic kills and terrain considerations, a pilot study site of approximately 400 km2 was delineated in the Piceance Creek drainage. A detailed sampling protocol was prepared based on the techniques described by Becker (1991) and Van Sickle and Lindsey (1991). RESULTS AND DISCUSSION Hunter kill of puma increased from 12 in 1988 to 80 in 1997 in DAU L-7. The marked increase in kill began in 1993 and was the result of greater quotas allocated in hopes of reducing puma depredation on domestic sheep. During the period 1995-1997 a total of 209 puma were taken by hunters and an additional 30 were taken by either landowners or predator control agents in DAU L-7. Game damage payments for puma depredation statewide varied from $46,000 to $60,000 per year during 1988-1991. Since 1992 the annual payments have varied from $93,000-$132,000. The relative importance of possible causative factors is unknown. Approximately 50% of the statewide puma damage payments are made in DAU L-7. A detailed sampling protocol was prepared (Appendix A) but never implemented. During the January-March 1998 sampling period adequate snow cover did not exist in the sampling block. In trying to implement the sampling scheme several serious logistical impediments became apparent. Scheduling of helicopter time was problematic because of restrictions with state contracts and the higher priority status of ungulate census and survival work. The puma census would have to compete with other on-going work for both helicopter and personnel time. �238 The logistical impediments, along with the changed management status of puma in Colorado (shared authority with Colorado Dept. of Agriculture), make the successful completion of this job improbable. Thus it was decided by participants to cease the effort at density estimation for future years. �239 Appendix PROTOCOL FOR PROBABILITY A SAMPLING OF PUMA DENSITY SAMPLING AREA: Area should be a rectangle of at least 390 km2, and must be at least 13 km wide on the shortest axis (baseline axis). The long axis (transect axis, 30 km) should be perpendicular to the majority of the major drainages in the area. You must delineate a specific rectangular area in order to make inclusion judgments. TRANSECTS: At least 3 transects should be flown in a straight line along the long axis for the entire 30 km length. Transect 1 should be randomly selected to start within the first 3 kms of the baseline axis. The remaining transects should be spaced 4 km distant (Example: Transect 1 begins at baseline point of 2 km, No.2 at 6 km, No.3 at 10 km). Flight time will be about 60 minutes to fly a 30 km transect plus about 10 minutes of search time per puma track set. MODIFICATIONS: If a 30 km transect cannqt be fitted into the areas of concern, the long axis can be shortened. However, total area should remain least 400 km2 and transects should be separated by at least 4 km. Thus a 20X20 km area would have 5 transects flown, of 20 km each. Long (30 km) transects are more efficient in terms of flight time. at SURVEY PERIOD: The helicopter surveys should be flown 2-3 days following a snowfall which is sufficiently deep as to not melt out within 48 hours and completely cover old tracks (about 8 cm). Very deep snows (>20 cm) restrict puma movement and are to be avoided. Optimum snow depth would be 8-12 cm. There must be at least one complete night for movement following the snow. A 2-night period is better unless very high winds are occurring; however, more than 2 nights creates difficulty because of other animal tracks and wind-blown snow. PROCEDURE: Surveys should be conducted by a crew consisting of a trained observer and a navigator. The navigator should have detailed knowledge of the terrain. While biologists may be excellent observers, we should try to utilize experienced puma hunters for observers as they will more easily detect the track patterns of puma. All locations should be recorded by UTM. Fly a straight line along Transect 1 until a puma track is encountered. Record location. Fly the backtrack until you find where puma started moving following snowfall; record the location of the point where the track is farthest from the transect. Fly the front track until you locate puma (often you will not see animal but will see where tracks end in heavy cover); record location of the point where the track is farthest from the transect. Because of the non-linear nature of travel, the beginning and ending points may not be the farthest distance traveled parallel to the baseline axis. Record the extreme distances from the transect. Return to original transect line at the point where track was located. Continue along the line until the next puma track is encountered. Repeat the process to estimate travel distance for each set of tracks. The objective is to obtain a straight-line distance of travel parallel to the base-transect. At the end of transect 1, move 4 kms along the baseline and return along transect 2, which parallels transect 1. Multiple puma tracks traveling together (female w/kittens or male-female pair) should only be counted as 1 track set. However, make a notation of the group size. There are separate analysis procedures which can be used if this is a common occurrence. �240 Do not count kitten unlikely occurrence quite problematic. tracks traveling separate from the mother. and distinguishing from bobcat tracks while This is an airborne is Kill sites will be characterized by lots of tracks in the near vicinity of the kill. Estimate the diameter of the circle which would include all the tracks. This diameter will be used as the travel distance. Most straight-line travel distances will be less than 3 km. However, in the event that a puma track crosses 2 transects, it should be separately recorded for each transect. Data to be recorded: location of farthest all in UTM's. Data to be calculated: each puma, in km's. location of each puma track along each transect; points of travel of puma perpendicular to the transect; straight-line distance parallel to baseline Data for any puma where >50% of the travel distance was outside rectangular survey area will not be used in the calculations. ANALYSIS: Population size is estimated from the following Becker (1991) and Van Sickle and Lindzey (1991): moved by the designated equations, from p;=x/(D/q) ~=L (lip) where Pi = probability that the ith puma i sample; Xi = distance, parallel to the basel~ofttatnedelftdtb~ 9ba aystpmaa!cD length of baseline; q = number of transects per systematic sample; and Tj = population estimate for the jth systmatic sample. Since we will not be flying replicate samples for an area, variance of the estimate will be estimated using jackknife procedures and dividing the transects into segments (McDonald and Manly 1989). This will be done by a contract statistician. While replication is the preferred methodology, helicopter costs make this prohibitive. ASSUMPTIONS 1. 2. 3. 4. 5. 6. 7. 8. FOR PROBABILITY SAMPLING PROCEDURE: All animals move during the survey period. Puma tracks are readily recognizable. All animal tracks are continuous. An~mal movements are independent of the sampling process. Post-snowstorm tracks can be distinguished. All puma tracks crossing a transect will be observed. Study area is rectangular in shape. All the transects are oriented perpendicular to the baseline LITERATURE axis. CITED Becker, E. F. 1991. A terrestrial furbearer estimator based on probability sampling. J. Wildl. Manage. 55(4):730-737. Van Sickle, W. D. and F. G. Lindzey. 1991. Evaluation of a cougar population estimator based on probability sampling. J. Wildl. Manage. 55(4):738743. �241 Colorado Division Wildlife Research July 1998 of Wildlife Report JOB PROGRESS state of Project Work Colorado No. Package Task No. Period Author: Cost Center W-15J-R-11 No. 3004 1 & 2 Covered: July REPORT Mammals 3430 Program Management Pronghorn of Other Ungulates Data Analysis and Reporting 1, 1997 - June 30, 1998 T.M. Pojar Abstract A herd structure estimate was made on the Middle Park pronghorn population during August, 1997 amd the total winter population estimate was made during January 1998. This information, along with like information since 1986 was incorporated into a spreadsheet population model. This model was provided to management personnel for. projecting effects of various management options. It was also used in public meetings for exploring public opinion of the proposed options. A manuscript was prepared summarLzLng the results of experiments in pronghorn inventory methods. It was submitted to the Journal of Wildlife Management and rejected on the basis that the topic addressed a relatively narrow issue. The manuscript will be modified and submitted either to the Wildlife Society Bulletin or the African Journal of Wildlife Management. The abstract is included in the results section. ��243 PRONGHORN DATA ANALYSIS Thomas AND REPORTING M. Pojar P.N·OBJECTIVES Task 1 Test for density dependence in Middle Park population parameters and summarize the findings in manuscript format suitable for submission to a professional wildlife management or ecological journal. Task 2 Analyze and summarize the results of experiments in pronghorn inventory methods and report the findings in manuscript format suitable for submission to a professional wildlife management journal. SEGMENT OBJECTIVES Task 1 Test for density dependence in Middle Park population parameters and summarize the findings in manuscript format suitable for submission to a professional wildlife management or ecological journal. Task 2 Analyze and summarize the results of experiments in pronghorn inventory methods and report the findings in manuscript format suitable for submission RESULTS Task 1 Key population information in the form of herd structure estimates and a count of the wintering population was collected. This was included in the population data set that began in 1986 and used to build a spreadsheet population model. This model was used to project population responses to proposed management options and was provided to managment personnel. The overall objective of Task 1 was not fulfilled during this segment. Task 2 A manuscript was prepared that analyzed and summarized the results of experiment in pronghorn invenorty methods. It was rejected by the Journal of Wildlife Management as having too narrow a focus. The manuscript will be modified and sumbitted either to the Wildlife Society Bulletin or the African Journal of Wildlife. The following is the abstract for this article. PRONGHORN DENSITY ESTIMATES: COMPARISON OF FIXED-WING LINE TRANSECT AND HELICOPTER QUADRAT SURVEYS THOMAS M. POJAR', Colorado Division of Wildlife, 317 W. Prospect Road, Fort Collins, CO 80526, USA DAVID C. BOWDEN, Department of Statistics, Colorado state University, Fort Collins, CO 80523, USA JEFF D. MADISON, Colorado Division of Wildlife, P. o. Box 1181, Meeker, CO 81641, USA Abstract: Estimates of pronghorn (Antilocapra americana) density in northwest Colorado sagebrush (Artemisia spp.) steppe habitat were compared using fixedwing line transect and helicopter quadrat surveys. Fixed-wing line transects offer wildlife managers a survey method that is both economical and statistically valid but this technique has not been evaluated with a standard or known density for pronghorn. We used the helicopter quadrat survey method as the standard and compare density estimates to fixed-wing line transect �244 surveys. Fixed-wing line transect estimates were always less than quadrat estimates (t=0.682, P = 0.050) and were 83%, 69%, and 83% of the quadrat estimates respectively, for 3 years data. In addition, we analyzed the first and second line transect intervals as narrow strips of 50 m and 100 m, respectively. The narrow strips were not different from either the quadrats or lines (P > 0.070) and may offer managers a method that combines the economy of line transects with the analysis simplicity of quadrat data. We conclude that line transect density estimates should be adjusted upward by considering them approximately 0.78 (SE = 6.32) of quadrat estimates. At the density of pronghorn encountered in this study (approximately 5/km», the cost of fixedwing line transect sampling for large scale management application is about 510% of the cost of helicopter quadrats for similar precision. Prepared Wildlife Researcher �245 Colorado Division Wildlife Research July 1998 of Wildlife Report JOB state of Project Work PROGRESS Colorado No. Plan No. Cost Center W-153-R-ll Mammals 3004 Management Task No. Period Covered: REPORT 3430 Program of other Ungulates strategies for Managing Pasteurellosis in Mountain Populations Sheep July 1, 1997 - June 30, 1998 Authors: M. W. Miller Personnel: S. Berry, and H. J. McNeil J. George, K. Larsen, J. Vayhinger, T. Verry ABSTRACT We continued investigations of multivalent Pasteurella haemolytica supernatant vaccines in captive bighorn sheep (Ovis canadensis). A vaccine lot(PhSV lot #970520) that combined bighorn and domestic strains appears to contain antigenicity comparable to that of the original multivalent vaccine, and was selected for use in future laboratory and field studies. To evaluate vaccine delivery options, 30 captive bighorns were divided into 3 groups of 10 based on vaccine history and baseline P. haemolytica leukotoxin neutralizing antibody titers; treatment groups included hand injection, biobullet implantation and oral gavage of PhSV. Serum leukotoxin-neutralizing antibody titers differed among delivery treatments (P = 0.009). Neutralizing titers were highest among hand-injected bighorns; no serum antibody response was detected among bighorns vaccinated orally. Although neutralizing titers were lower among implanted bighorns than in the hand injected controls (P < 0.021), seroconversion rates did not differ (implantation = 6/10, hand injection = 9/10; P = 0.303). Although our data demonstrated that handinjection elicits higher absolute titers than either biobullet implantation or oral vaccination, biobullet implantation may also stimulate effective antibody responses to P. haemolytica supernatant vaccines. Further evaluation of biobullet vaccination against pneumonic pasteurellosis in free-ranging populations of wild bighorn sheep appears warranted. A total of 110 free-ranging bighorn sheep from 4 different herd units in central Colorado were vaccinated with PhSV during January-March 1998. No adverse reactions or mortalities were detected among vaccinated bighorns through June 1998. Ongoing monitoring in the Tarryall-Kenosha herd complex will provide additional data on efficacy of vaccination to protect freeranging bighorns from pasteurellosis under field conditions. ��247 EXPERIMENTS TO IDENTIFY AND MANAGE STRESS IN MOUNTAIN SHEEP POPULATIONS M. W. Miller and H. J. McHeil P. H. OBJECTIVES 1. Design, conduct, and report on experiments evaluating Pasteurella haemolytica vaccines and vaccine delivery systems and identify those with potential application in managing free-ranging bighorn populations. 2. Design, conduct, and report on field experiments evaluating management strategies for preventing pasteurellosis epizootics in bighorn populations SEGMENT 1. 2. OBJECTIVES Conduct and report results of an experiment evaluating options for delivering a multivalent Pasteurella haemolytica supernatant vaccine bighorn sheep. Provide multivalent Pasteurella haemolytica supernatant vaccine in select bighorn sheep management activities statewide. to for use STRATEGIES FOR MANAGING PASTEURELLOSIS IN MOUNTAIN SHEEP POPULATIONS Inability to control infectious disease outbreaks and subsequent mortality in mountain sheep populations represents a significant obstacle to long-term success in their management. Although the "bighorn pneumonia complex" has been studied intensively for over 3 decades, little is known about many aspects of its etiology and epizootiology. Moreover, management interventions recommended for preventing or controlling this problem remain untested. Although viral, bacterial, and parasitic agents have all been incriminated in these outbreaks, Pasteurella spp. are perhaps the most common pathogens associated with bronchopneumonia in bighorns. Two species, P. haemolytica and P. multocida, and several biotypes and/or serotypes within those species, have been isolated from bighorns during epizootics. Unfortunately, despite extensive diagnostic and experimental investigation, the epizootiology of pasteurellosis in wild bighorn populations is poorly understood. In the absence of knowledge about the epizootiology of pasteurellosis, effective strategies for managing pneumonia in bighorn populations have not emerged. Here, we report on a series of ongoing research studies designed to improve knowledge about various aspects of pasteurellosis epizootiology and management in bighorn sheep. METHODS AND MATERIALS Refinement of experimental Pasteurella haemQlytica supernatant vaccine (McNeil, Miller, and Shewen): A rabbit study was previously conducted with Pasteurella haemolytica supernatant vaccine (PhSV) lot #970520 (Miller and McNeil, 1997). Rabbits were exsanguinated after receiving three 0.5 ml doses �248 of vaccine at 2-wk intervals. All rabbits responded to vaccination by producing anti-leukotoxin antibodies. Protein profiles from Western blots (blotted with vaccinated rabbit serum) transferred from 12% precast PAGE gels containing the vaccine components of both PhSV lot #970520 and multivalent vaccine lot # 940902 (Miller et al., 1997) also showed strong vaccine-induced antibody reactivity. Based on these findings, a small pilot study in bighorn sheep was conducted. Six captive Rocky Mountain bighorn sheep were used. Sheep were paired on the basis of age and vaccination history. On~ bighorn from each pair received 2 ml PhSV lot #970520 intramuscularly; the other animals received 1 ml PhSV lot #970520 and 1 ml Presponse~ mixed in the same syringe, delivered intramuscularly. (Presponse~ is a commercially available supernatant vaccine of Pasteurella haemolytica AI.) We used this approach to establish whether the presence of P. haemolytica serotype Al was necessary to drive the marked increase in leukotoxin neutralizing titer seen in previous studies (Miller et al., 1997; Kraabel et al., 1998). Animals were bled prior to vaccination and again at day 7 and day 14. Serum leukotoxin neutralizing antibody titers stimulated by the two treatments were measured and compared using methods described by Miller et al. (1997). Delivery of Pasteurella haemolytica supernatant vaccine to bighorn sheep (McNeil and Miller): Thirty captive bighorn sheep were used in this study. Animals were divided into three groups of ten based on their vaccine history and baseline neutralizing titers. The three treatment groups included hand injection, biobullet implantation and oral gavage of vaccine. Four additional animals were intramuscularly injected with saline to monitor for changes in antibody titers unrelated to vaccination during the study period. PhSV lot #970520 was used for all treatments. Vaccine was lyophilized for use in biobullets and microspheres. The standard dose for intramuscular injection, 2 ml (Miller et al., 1997; Kraabel et al., 1998), yielded about 0.025 g lyophilized vaccine. Biobullets were packed with about 0.030 g lyophilized vaccine, then filled with methylcellulose filler for remote delivery. Lyophilized vaccine was also sent to Dr. Harm HogenEsch of Purdue University for encapsulation in alginate microspheres; 5 ml of alginate microsphere suspension was recommended as equivalent to a 2 ml dose of vaccine. The hand injection group received 2 ml PhSV injected intramuscularly into the left haunch. The biobullet group were vaccinated with the use of a specialized biobullet rifle from a distance of approximately 4 meters. Animals in this group had a small area on their left hindquarters shaved to facilitate confirming implantation. Animals receiving vaccine orally were dosed with the aid of a small stainless steel gavage tube. The tube was inserted into the animal's mouth and laid along the teeth; 5 ml of vaccine was then squirted into the sheep's mouth and subsequently swallowed. Animals were bled prior to vaccination and again at 1, 2, 4, 8 and 12 weeks postvaccination. Sera were collected and used to assess serological responses. Titers were measured using a leukotoxin neutralization assay, along with three separate direct agglutination assays that incorporated formanilized Pasteurella haemolytica serotypes AI, A2 or T10 as antigen (Miller et al., 1997; Kraabel et al., 1998). Use of Pasteurella haemolytica supernatant vaccine in free-ranging bighorn ~ (Miller, George, and Vayhinger): A total of 110 free-ranging bighorn sheep from 4 different herd units in central Colorado were vaccinated with �249 PhSV during January-March 1998 (Table 1). Bighorns from Georgetown herd unit were vaccinated just prior to translocation to the Arkansas River canyon for release; nearby resident bighorn populations appear to have recurrent problems with respiratory disease in this area, and translocated bighorns were considered potentially at risk. Bighorns in the Black Canyon and Twin Eagles herd units of the Tarryall-Kenosha complex were vaccinated in an attempt to prevent spread and/or minimize impacts of a pasteurellosis epidemic that started in the Sugarloaf herd unit in December 1997 (see WP7610-T4 for details); a few sheep in the Sugarloaf herd also were vaccinated in an attempt to enhance postepidemic survival in that subpopulation. Survival of vaccinated and unvaccinated individuals was compared opportunistically in conjunction with ongoing field studies. RESULTS AHD DISCUSSION Refinement of experimental Pasteurella haemolytica supernatant vaccine (McNeil, Miller, and Shewen): We observed no adverse effects in bighorns receiving PhSV lot #970520. All vaccinated bighorns responded with an increase in leukotoxin neutralizing titer. Based on these results, PhSV lot #970520 was judged an appropriate vaccine for use in the delivery experiment and in field applications. Delivery of Pasteurella haemolytica supernatant vaccine to bighorn sheep (McNeil and Miller): other than a mild transient lameness in animals vaccinated intramuscularly or remotely, no adverse reactions were observed following vaccination. Serum neutralizing antibody titers to P. haemolytica leukotoxin differed among delivery treatments (P=0.009) as well as among baseline titer/vaccination history groups (P=0.013). Neutralizing titers were highest among hand-injected bighorns; no serum antibody response was detected among bighorns vaccinated orally. Although neutralizing titers were lower among implanted bighorns than in the hand injected controls for ~2 weeks post vaccination (P < 0.021), seroconversion rates (defined as a ~2 10g2 increase in titers) in response to implantation (6/10) and hand injection (9/10) did not differ (P=0.303). Agglutinating antibody titers to T10 were high and did not differ over time or between delivery treatments. In addition, agglutinating antibody titers to A1 did not vary over time or between groups. Agglutinating antibody titers to A2 were higher in the hand injected group than the orally vaccinated sheep during the time period between 2 and 4 weeks post vaccination (P < 0.026); A2 agglutinating titers in hand-injected and implanted sheep did not differ during the same time period (P = 0.07). Vaccination history/baseline titer categorization affected responses to serotype A2 surface antigens (P = 0.0001). Our data support previous observations (Miller et al., 1997; Kraabel et al., 1998) that hand-injected P. haemolytica supernatant vaccines stimulate marked antibody responses in bighorn sheep; it follows that delivery via projectile syringe should stimulate similar antibody responses. Although our data demonstrate that hand-injection elicits higher absolute titers than either biobullet implantation or oral vaccination, biobullet implantation may also stimulate effective antibody responses to P. haemolytica supernatant vaccines. Whether lack of serum antibody response to oral vaccination truly reflects failure to stimulate immunity remains undetermined. Further evaluation of biobullet vaccination against pneumonic pasteurellosis in free-ranging populations of wild bighorn sheep appears warranted. �250 Use of Pasteurella haemolytica supernatant vaccine in free-ranging bighorn ~ (Miller, George, and Vayhinger): As with previous captive animal studies (Miller et al., 1997; Kraabel et al., 1998; McNeil and Miller, above), no adverse effects were observed in free-ranging bighorns that received PhSV via hand-injection, projectile syringe, or biobullet implant. No mortality was been detected among vaccinated bighorns in the 4-6 mo after vaccination; however, little or no mortality occurred among unvaccinated bighorns in these populations during that time. We will continue to monitor survival rates among vaccinated and unvaccinated bighorns in the Tarryall and Kenosha subpopulations in conjunction with other ongoing studies. These data should be useful in evaluating efficacy of vaccination in protecting freeranging bighorns from pasteurellosis under field conditions. LITERATURE CITED Kraabel, B. J., and M. W. Miller, J. A. Conlon, and H. J. McNeil. 1998. Evaluation of a multivalent Pasteurella haemolytica vaccine in bighorn sheep: Protection from experimental challenge. Journal of Wildlife Diseases 34: 325-333. Miller, M. W., J. A. Conlon, H. J. McNeil, J. M. Bulgin, and A. C. S. Ward. 1997. Evaluation of a multivalent Pasteurella haemolytica vaccine in bighorn sheep: Safety and serologic responses. Journal of Wildlife Diseases 33: 738-748. Prepared by veterinarian Table 1. Free-ranging bighorn sheep treated with multivalent haemolytica supernatant vaccine during Jan-Mar 1998. Population (Herd Unit) Georgetown (Georgetown) Status Hand Deliyery Dart Pasteurella Biobullet healthy -translocated to Arkansas R. 25 o o healthy, exposure future likely 35 4 5 (Twin Eagles) healthy, exposure future likely 21 6 3 (Sugarloaf) pasteurellosis epidemic (12/97) 11 o o Tarryall-Kenosha (Black Canyon) �251 Colorado Division Wildlife Research July 1998 of Wildlife Report JOB PROGRESS State of project Colorado No. Work Program Cost Center W-153-R-11 No. __~7~6~1~0~ _ Task No. Period REPORT Mammals Program Support services Mammals Covered: Authors: Jackie and Bruce Gill July 3430 Project Support Services I, 1997 - June 3D, 1998 Boss, Nancy wild, Margaret W~ld, Michael Miller, Gary White, ABSTRACT Task 1 - During the FY 1997-1998 the following were accomplished: 53 Publications acquired by the Research Center Library of Colorado Div. of Wildlife employees, cooperators, educators, and the public. for the use wildlife 2,292 Items of information delivered to Colorado Div. of wildlife employees, cooperators, wildlife educators, and the public, resulting from requests and literature searches. 606 Items of information catalogued into the electronic and card catalogues, which expanded the Research Center Library inventory to 17,296 items. 983 Computer scanned items of information entered into the electronic catalogue for the maintenance of the circulation system of the Research Center Library. 2,261 Items checked out by Colorado Div. of Wildlife employees, cooperators, wildlife educators, and the public indicating satisfaction of library services. o Comments received from Colorado Div. of Wildlife employees, cooperators, wildlife educators, and the public, through surveys and a 'Suggestion Box' to monitor and evaluate library patron satisfaction with library services. �252 Task 2 - Progress made on each Objective objective is reported below. 1. The following publication-ready during manuscripts this Segment were developed: Draft Strategy for the Conservation and Reestablishment wolverine in the Southern Rocky Mountains Northern Wild Sheep Symposium and Goat Council: made on each Proceedings of Lynx and of the Tenth Biennial 2. Developed graphics and visual researchers and biologists. 3. Special Reports Nos. 71 and 72 and Federal Aid Abstracts for 1995, 1996, and 1997 were published and distributed to Division personnel and outside agencies and libraries. 4. No progress 5. Wildlife Research and distributed. 6. A Customer Service Survey concerning publication services was developed and distributed to 4.8customers. Twenty two were returned (46%). The results were very positive. There were 9 items for evaluation and the totals are reported as follows: Very good = 129; Good = 54; Fair = 8; Poor = 2; Very poor = 1; NA = 4. materials was made on Special Reports Report for 52 presentations by No. 73 in this Segment. for 1996 and 1997 were assembled, published, Task 3 - The Colorado Division of Wildlife's Foothills Wildlife Research Facility (FWRF) maintained captive animals (annual total: 161 wild ungulates of 5 species, 12 domestic cattle, and 34 prairie dogs) and facilities in support of research on chronic wasting disease (CWO) in deer and elk (and potential transmission to domestic cattle and pronghorn), deer contraception, pneumonia immunization of bighorn sheep, and prairie dog research for blackfooted ferret recovery. During the year, domestic cattle and additiona~ mule deer were added to the facility to support CWO research and white-tailed prairie dogs were added to support research to benefit black-footed ferret recovery. Twenty-eight animals died or were euthanatized for health problems and an additional nine animals were euthanatized as part of study protocols or facility management programs. Chronic wasting disease (CWO) was again a significant source of mortality in captive mule deer (n = 7). CWO also emerged as a significant source of mortality in white-tailed deer (n = 6). A revised FWRF CWO Management Protocol reflects a change in philosophy on CWO management--rather than attempt to control and eradicate CWO at FWRF, we will maintain the disease for research purposes under heightened biosafety guidelines. A high quality of animal care and facility maintenance was provided by temporary, work-study, YCC employees, and volunteers. Volunteers contributed 685 hr (equal to 0.34 FTE) work at FWRF. The high standard of care is in part reflected by the finding of compliance under the Animal Welfare Act during the annual USDA APHIS inspection of FWRF. In addition to routine maintenance performed at FWRF, significant facility modifications included buildings and cages to house prairie dogs, a handling/weigh area for cattle, and enclosures for mule deer feeding. �253 Task 4 - In 1990, the Colorado Division of Wildlife initiated a Federal Aid project to be a focus for coordinating a variety of actions to further understanding about diseases affecting wildlife populations throughout Colorado through surveillance, modeling, and research. The overall goal of this project is to provide data and analyses to support management programs directed toward detecting and managing important health problems affecting Colorado's wildlife resources. Surveillance We continued our program for acquiring, examLnLng, reporting on, and summarizing wildlife disease cases occurring throughout Colorado. Investigations We investigated a respiratory disease outbreak Sugarloaf subpopulation of the Tarryall-Kenosha among bighorn herd complex sheep in the (S23N, 5235). Modeling McCarty, Burnham, and Miller continued developing, refining, and evaluating models for predicting dynamics and impacts of infectious diseases in freeranging ungulate populations. Task 5 contract Colorado statement Colorado declining Accomplishments resulting from consulting services provided on from Dr. Gary C. White, Department of Fishery and Wildlife Biology, state University, Fort Collins, Colorado are summarized in bullet format. In addition, mule deer population trend data from the Division of Wildlife were analyzed for evidence of causes for deer populations. Contributions were made toward each of the objectives listed in the Segment Narrative. Assistance was provided in the experimental design of several Mammals Program studies. A draft of a scientific book on analysis of spatial distribution and statewide abundance of wildlife species was prepared and submitted to publishers. A spreadsheet model of mule deer population dynamics was developed. Consulting services were provided to Thomas D.I. Beck, Richard M. Bartmann, Thomas M. Pojar, R. Bruce Gill, David J. Freddy, and James F. Lipscomb to assist with developmental, analytical, and/or design aspects of various research projects concerning mammalian species. Analysis of mule deer population trend data revealed a significant long-term decline in fawn:doe and calf:cow ratios in several Colorado management units in both December aerial count surveys and harvest data for mule deer, elk, and pronghorn. Reasons for this decline are not entirely clear and should be investigated more critically and intensely. If the hypothesis is true that declining buck:doe ratios is a major contributor to declines in mule deer fecundity, I would have expected to see stronger relationships from both the DEAMAN and Piceance data analyses. Not much support is provided. However, the lack of support cannot be used as evidence that the hypothesis is not true. When you accept the null hypothesis of a statistical test, you don't learn much unless you have high power. None of the analyses performed to detect mule deer population trends have much power. ��2SS Mammals Jackie Project Support Services Boss, Nancy Wild, Margaret Wild, Michael Gary White, and Bruce Gill Miller, P •N. OBJECTIVE Provide effective library, publication, editorial, veterinary, pen and animal maintenance, wildlife disease diagnosis and monitoring, biometrical, supervisory, and administrative services at minimal cost by centralizing them and enhancing accountability for support services. SEGMENT Task OBJCECTIYES 1 1. Acquire reference library reference materials research staff members and cooperators. 2. Assist wildlife research staff members and in acquiring reprints of requested 3. Maintain electronic holdings. 4. Develop a process for monitoring with library services. requested in conducting materials. and card catalogues literature of all research and evaluating by wildlife customer searches library satisfaction Task 2 1. Assist 2. Assist Division of Wildlife employees materials for oral presentations. 3. Publish Special Reports No. 71 and 72 along with and distribute to Division personnel and outside 4. Create camera-ready (bighorn sheep). S. Assemble Research Task 1. Wildlife Researchers in developing copy, print, and publish Reports. the FY 9S-96 publication in developing and publish ready manuscripts. graphics and visual Federal Aid Abstracts agencies. Special Report and FY 96-97 editions No. 73 of Wildlife 3 Maintain research facilities and experimental animals in support of research on chronic wasting disease, deer contraception, and pneumonia immunization of mountain sheep. Task 4 1. Maintain systems for submitting, diagnosing, and reporting disease cases in wild animals throughout Colorado. 2. Maintain databases and update simulation models on sporadic for assimilating and �256 analying data on and/or guiding identified through surveillance 3. Task management of wildlife and surveys. Provide assistance in investigation outbreaks in Colorado. and managing disease wildlife problems disease 5 1. Provide ongoing consulting services to the Colorado Division of Wildlife (CDOW) to design and analyze harvest surveys, terrestrial wildlife inventory systems, and wildlife population models and modeling procedures. 2. Provide services and recommendations necessary to develop criteria and processes to estimate spatial distribution and statewide abundance of Colorado's wildlife species and to identify those species most in need of protection or conservation. 3. According to specifications provided by CDOW, develop and modify a mule deer population model that allows the user to sample modeled mule deer populations to mimic current CDOW management and regulatory procedures. The purpose of this model is to assist biologists in evaluating big game modeling procedures and developing and evaluating harvest and population management alternatives. 4. Update and correct problems in the DEAMAN computer data management system and conduct workshops necessary to assist Terrestrial Wildlife staff in the use of DEAMAN and population modeling procedures, and in the use of statistical techniques for monitoring spatial distribution and statewide abundance of terrestrial wildlife populations. 5. Assist in the design, analysis, and interpretation of scientific investigations to facilitate the development and implementation of conservation strategies for various terrestrial species; including Preble's meadow jumping mouse, kit fox, lynx, and black-footed ferret. 6. Assist in the design, analysis, and interpretation of scientific investigations to develop improved procedures to estimate elk abundance and survival. Task 6 1. Prepare PACE plans for all Mammals 2. Prepare performance 3. Coordinate scientific input from Mammals Program process for implementing Amendment 14 and Senate 4. Process all encumbering documents for Mammals Program staff members maintain records of those encumberances in the COFRS program. 5. Update the Administrative Standard Operating Procedures distribute copies to all mammals Program staff members. 6. Coordinate the preparation of Federal Aid Program Narratives, Segment Narratives, Job Progress Reports, and Job Final Reports and submit evaluations Program staff members. for all Mammals Program staff members. staff members Bill 52. Manual into the and and �257 required numbers of copies to the U.S. Fish and Wildlife Service in accordanced with specified formats and deadlines. 7. Develop a process for monitoring and evaluating customer satisfaction with administrative services. Results of Progress Task 1 Publications Acgyired in the Research Center Library Allen, H. E., A. W. Garrison, and G. W. Luther, III, eds. 1998. Metals in surface waters. Chelsea, MI : Ann Arbor Press. 262pp. Alverson, W. S., W. Kuhlmann, and D. M. Waller. 1994. Wild forests: conservation biology and public policy. washington, D.C. : Island Press. 300pp. Andrews, R. and R. Righter. 1992. Colorado birds: a reference to their distribution and habitat. Denver, CO : Denver Museum of Natural History. 442pp. Bailey, J. A. 1984. Principles of wildlife management. New York: John wiley & Sons. 373pp. Beebee, T. J. C. 1996. Ecology and conservation of amphibians. New York : Chapman & Hall. 214pp. Bennett, L. E. 1994. Colorado gray wolf recovery : a biological feasibility study: final report. [Washington, D.C. : U.S. Fish & Wildlife Service]. 3l8pp. Bergman, H. L. and E. J. Dorward-King, eds. 1993. Reassessment of metals criteria for aquatic life protection. Pensacola, FL SETAC Press. SETAC technical publications series. 114pp. Carbyn, L. N., S. H. Fritts, and D. R. Seip, eds. 1995. Ecology and conservation of wolves in a changing world. Canadian Circumpolar Institute, Occasional Publication No. 35. Edmonton, Alberta, Canada : University of Alberta. 620pp. Claar, J. J. and P. Schullery, eds. 1994. Bears - their biology and management : a selection of papers from the 9th International Conference on Bear Research and Management held at Missoula, MT February 1992. Yellowstone National Park, WY : Yellowstone Center for Resources. 586pp. Clark, T. W. 1997. Averting extinction: reconstructing endangered species recovery. New Haven: Yale University Press. 270pp. Cvancara, V. 1997. Current references in fish research; vol. 22. Eau Claire, WI : Univ. of Wisc.-EC. 163pp. Daily, G. C., ed. 1997. Nature's services: societal dependence on natural ecosystems. washington, D.C. : Island Press. 392pp. Darling, L. M. and W. R. Archibald, eds. 1990. Bears - their biology and management : a selection of papers from the conference held at Victoria, British Columbia, Canada: February 1989. Victoria, BC : T. D. Mock & Associates, Inc. 448pp. Fitzgerald, J. P., C. A. Meaney, and D. M. Armstrong. 1994. Mammals of Colorado. Niwot, CO : Univ. Pres~ of Colo. 467pp. Franzmann, A. W. and C. C. Schwartz, eds. 1997. Ecology and management of the North American moose. Washington, D.C. : Smithsonian Institution Press. 733pp. Geist, V. 1993. Wild sheep country. Minocqua, WI : NorthWord Press. 175pp. �258 Geist, V. and I. McTaggart-Cowan, eds. 1995. Wildlife conservation policy : a reader. Calgary, Alberta: Detselig Enterprises Ltd. 308pp. Goodwin, B. 1994. How the leopard changed its spots the evolution of complexity. New York: Simon & Schuster. 252pp. Greene, H. W. 1997. Snakes: the evolution of mystery in nature. Berkeley, CA : Univ. of Calif Press. 351pp. Hadidian, J., G. R. Hodge, and J. W. Grandy, eds. 1997. Wild neighbors : the humane approach to living with wildlife. Washington, D.C. : Humane Society of the u.S. 253pp. Haigh, J. C. and R. J.•Hudson. 1993. Farming wapiti and red deer. st. Louis : Mosby. 369pp. Hartshorne, C. 1973. Born to sing: an interpretation and world survey of bird song. Bloomington, IN : Indiana univ. Press. 304pp. Hummel, M. and S. Pettigrew. 1992. Wild hunters : predators in peril. Niwot, CO : Roberts Rinehart. 25lpp. Kellert, S. R. 1996. The value of life : biological diversity and human society. Washington, D.C. : Island Press. 263pp. Knopf, F. L. and F. B. Samson, eds. Ecology and conservation of Great Plains vertebrates. New York : Springer-Verlag. 320pp. Litvaitis, J. A., ed. [1997]. Northeast wildlife: transactions of the Northeast Section: The Wildlife Society: Vol. 52 - 1995. Amherst, MA : U.W. Forest Service. 116pp. Manly, B. F. J., L. L. McDonald, and D. L. Thomas. Resource selection by animals : statistical design and analysis for field studies. New York: Chapman & Hall. 177pp. Martinka, C. J. and K. L. McArthur, eds. 1980. Bears - their biology and management : a selection of papers from the 4th International Conference on Bear Research and Management held at Kalispell, MT, USA: Feb. 1977. [Washington, D.C.] : U.S. Gov. Printing Office. 375pp. McCullough, D. R., ed. 1996. Metapopulations and wildlife conservation. Washington, D.C. : Island Press. 429pp. McGowan, C. 1997. The raptor and the lamb : predators and prey in the living world. New York : Henry Holt & Co. 272pp. McShea, W. J., H. B. Underwood, and J. H. Rappole, eds. 1997. The science of overabundance : deer ecology and population management. Washington, D.C. : Smithsonian Institution Press. 402pp. Morrison, M. L. and L. S. Hall, eds. [1997]. Transactions of the Western Section of The Wildlife Society: 1996 - Vol. 32. Oakland, CA : The Wildlife Society, Western Section. 88pp. National Research Council. 1994. Rangeland health : new methods to classify, inventory, and monitor rangelands. Washington, D.C. National Academy Press. 180pp. Rappole, J. H. 1995. The ecology of migrant birds: a neotropical perspective. Washington, D.C. : Smithsonian Institution. 269pp. Samson, F. B. and F. L. Knopf, eds. 1996. Prairie conservation: preserving North America's most endangered ecosystem. Washington, D.C. : Island Press. 339pp. Searfoss, G. 1995. Skulls and bones : a guide to the skeletal structures and behavior of North American mammals. Mechanicsburg, PA : Stackpole Books. 277pp. Stebbins, R. C. and N. W. Cohen. 1995. A natural history of amphibians. Princeton, NJ : Princeton Univ. Press. 316pp. Tiedeman, J. A. and C. Terwilliger, Jr. 1978. Phyto-edaphic classification of the Piceance Basin. Fort Collins, CO : Colorado State University. Range Science Department series; no. 31. 265pp. �259 University of Michigan. School of Natural Resources. 1997. The endangered species UPDATE : special issue : habitat conservation planning: Vol.14(7 & 8). 70pp. Nowak, R. M. 1991. Walker's mammals of the world. 5th edition. Baltimore : John Hopkins University Press. 2 vols. Waters, T. F. 1995. Sediment in streams : sources, biological effects and control. Bethesda, MD : Am. Fisheries Society. 251pp. Webb, J. W., ed. 1965. Proceedings of the sixteenth annual conference: Southeastern Association of Game and Fish Commissioners : October 14-17, 1962 : Charleston, North Carolina. Columbia, SC : Southeast. Assoc. of Game and Fish Commissioners. 521pp. Webb, J. W., ed. 1965. Proceedings of the seventeenth annual conference : Southeastern Association of Game and Fish Commissioners : September 29, 30, October 1, 2, 1963 : Hot Springs, Arkansas. Columbia, SC : Southeast. Assoc. of Game and Fish Commissioners. 452pp. Webb, J. W., ed. 1967. Proceedings of the eighteenth annual conference : Southeastern Association of Game and Fish Commissioners : October 18-21, 1964 : Clearwater, Florida. Columbia, SC : Southeast. Assoc. of Game and Fish Commissioners. 584pp. Webb, J. W., ed. 1966. Proceedings of the nineteenth annual conference : Southeastern Association of Game and Fish Commissioners : October 10-13, 1965 : Tulsa, Oklahoma. Columbia, SC : Southeast. Assoc. of Game and Fish Commissioners. 471pp. Webb, J. W., ed. 1967. Proceedings of the twentieth annual conference: Southeastern Association of Game and Fish Commissioners : October 24-26, 1966 : Asheville, North Carolina. Columbia, SC : Southeast. Assoc. of Game and Fish Commissioners. 493pp. Western Association of Fish and Wildlife Agencies. [1995]. Western proceedings : 74th annual conference : Western Association of Fish and Wildlife Agencies: Anchorage, Alaska: May 14-19, 1994. 98pp. Western Association of Fish and Wildlife Agencies. [1996]. Western proceedings : Western Association of Fish and Wildlife Agencies 75th annual conference: Big Sky, Montana: July 15-20, 1995. 256pp. Western Association of Fish and Wildlife Agencies. [1997]. Western proceedings : 76th annual conference : Western Association of Fish and Wildlife Agencies : Honolulu, Hawaii : July 22-26, 1996. 385pp. Wilson, D. E. and D. M. Reeder, eds. 1992. Mammal species of the world : a taxonomic and geographic reference. 2nd edition. Washington, D.C. : Smithsonian Institution Press. 1,207pp. Wood, F., Jr. 1997. The delights and dilemmas of hunting the hunting versus anti-hunting debate. New York: Univ. Press of America, Inc. 237pp. Zager, P., ed. 1986. Bears - their biology and management: a selection of papers from the conference held at Grand Canyon, Arizona, USA: Feb. 1983. Washington, D.C. : Port City Press, Inc. 226pp. Zager, P., ed. 1987. Bears - their biology and management: a selection of papers from the conference held at Williamsburg, virginia, USA and Plitvice Lakes, Yugoslavia : February and March 1986. 394pp. �260 Theses and Books Obtained on Interlibrary Loan Literature Searches and Information Deliyered American society of Ichthyologists and Herpetologists (ASIH), The Herpetologists' League (HL), and the Society for the Study of Amphibians and Reptiles (SSAR). 1987. Guidelines for use of live amphibians and reptiles in field research. [Lawrence, KS : ASIH, HL, & SSAR.] J. of Herpet., Suppl., Vol. 4:1-14. Beebee, T. J. C. 1996. Ecology and conservation of amphibians. New York : Chapman & Hall. 214pp. Behle, W. H. 1990. Utah birds : historical perspectives and bibliography. Salt Lake City, UT : Utah Museum of Natural History. Occasional publication; no. 9. 355pp. Bromley, M., L. H. Graf, P. L. Clarkson, and J. A. Nagy. 1992. Safety in bear country: a reference manual. Rev. ed. [Yellowknife, N.W.T.] : Northwest Territories, Dept. of Renewable Resources. 134pp. Clark, T. W. 1997. Averting extinction: reconstructing endangered species recovery. New Haven : Yale University Press. 270pp. Cogger, H., E. Cameron, R. Sadlier, and P. Eggler. 1993. The action plan for Australian reptiles. Canberra, Australia : Australian Nature Conservation Agency. Australian Nature Conservation Agency Endangered Species Program : Project No. 124. 254pp. Cook, C. W. 1974. Surface rehabilitation of land disturbances resulting from oil shale development. Fort Collins, CO : Colorado State University. Technical report series; no. 1. 255pp. Corbet, G. B. 1978. The mammals of the Palaearctic Region : a taxonomic review. Ithaca, NY : Cornell University Press. 314pp. DeGraaf, R. M., V. E. Scott, R. H. Hamre, L. Ernst, and S. H. Anderson. 1991. Forest and rangeland birds of the United States : natural history and habitat use. [Washington, D.C. : U.S. Dept. of Ag.] Agriculture handbook; no. 688. 625pp. Duffy, E. and A. S. Watt, eds. 1971. The scientific management of animal and plant communities for conservation : the 11th symposium of the British Ecological Society : University of Eas Anglia, Norwich : 7-9 July 1970. Oxford: Blackwell Scientific Publications. 652pp. Essex Institute. 1874. Bulletin of the Essex Institute. Salem, MA Essex Institute. var. pagination. Fitzgerald, J. P. 1970. The ecology of plague in prairie dogs and associated small mammals in South Park, Colorado. Ph.D. Dissertation, Fort Collins, CO, Colo. State Univ. 90pp. Gipps, J. H. W. 1991. Beyond captive breeding: re-introducing endangered mammals to the wild : the proceedings of a symposium held at the Zoological Society of London on 24th and 25th November 1989. Oxford, England: Clarendon Press. Symposia of the Zoological Society of London; 62. 284pp. Heinemeyer, K. S. 1988. Temporal dynamics in the movements, habitat use, activity, and spacing of reintroduced fishers in northwestern Montana. M.S. Thesis, Missoula, MT, Univ. of Montana. 158pp. Holechek, J. L., R. D. Pieper, and C. H. Herbel. 1995. Range management : principles and practices. Englewood Cliffs, NJ : Prentice Hall. 2nd edition. 526pp. Hoover, R. L. 1955. Beaver ecology in the Longs Peak area of Colorado. M.S. Thesis, Fort Collins, CO, Colorado Agricultural & Mechanical College. 262pp. �261 James, V. H. T. 1992. The adrenal gland. 2nd edition. New York: Raven Press. Comprehensive endocrinology; revised series. 513pp. Kiesecker, J. M. 1991. Acidification and its effects on amphibians breeding in temporary ponds in montane Colorado. M.S. Thesis, Greeley, CO, Univ. of Northern Colorado. 79pp. Koopman, M. E. 1995. Food habits, space use and movements of the San Joaquin kit fox on the Elk Hills naval Petroleum Reserves, California. M.S. Thesis, Berkeley, Univ. of California. 41pp. Kuhr, R. J. and H. W. Dorough. 1976. Carbamate insecticides: chemistry, biochemistry and .toxicology. Cleveland, OH : CRC Press. 301pp. Lyons, L. J. 1984. Field tests of elk/timber coordination guidelines. Ogden, UT : u.S. Forest Service. Intermountain Forest & Range Experiment Station. Research paper; INT-325. 10pp. Manly, B. F. J., L. L. McDonald, and D. L. Thomas. 1993. Resource selection by animals : statistical design and analysis for field studies. New York: Chapman & Hall. 177pp. Martinka, C. J. and K. L. McArthur, eds. 1980. Bears - their biology and management : a selection of papers from the 4th International Conference on Bear Research and Management held at Kalispell, MT, USA: Feb. 1977. [Washington, D.C.] : U.S. Gov. Printing Office. 375pp. McCullough, D. R., ed. 1996. Metapopulations and wildlife conservation. Washington, D.C. : Island Press. 429pp. Nielsen, L. A. 1992. Methods of marking fish and shellfish. Bethesda, MD : American Fisheries Society. American Fisheries Society Special Publication; 23. 208pp. Orlans, F. B., ed. 1988. Field research guidelines: impact on animal care and use committees. Bethesda, MD : Scientists Center for Animal Welfare. 23pp. Palmer, W. C. 1965. Meteorological drought. Washington, D.C. : U.S. Dept. of Commerce. Weather Bureau. Research paper; no. 45. 58pp. Petersen, D. 1995. Ghost grizzlies. New York: Henry Holt & Co. 296pp. Robinson, J. G. and K. H. Redford, eds. 1991. Neotropical wildlife use and conservation. Chicago, IL : Univ. of Chicago Press. 520pp. Rutherford, W. H. 1954. Abstract of thesis : interrelationships of beavers and other wildlife on a high-altitude stream in Colorado. Fort Collins, CO : Colorado Agricultural & Mechanical COllege. 12pp. Rutherford, W. H. 1954. Interrelationships of beavers and other wildlife on a high-altitude stream in Colorado. M.S. Thesis, Fort Collins, CO, Colorado Agricultural & Mechanical College. 134pp. Spiegel, L. K. 1996. Studies of the San Joaquin kit fox in undeveloped and oil-developed areas. [California]: California Energy Commission. 131pp. Seigel, R. A., L. E. Hunt, J. L. Knight, L. Malaret, and N. L. Zuschlag. 1984. Vertebrate ecology and systematics : a tribute to Henry S. Fitch. Lawrence, KS : University of Kansas. University of Kansas Museum of Natural History. Special publication; no. 10. 278pp. South African Veterinary Association. Wildlife Group and World Association of Wildlife Veterinarians. 1994. Proceedings of an International Symposium on Capture, Care and Management of Threatened Mammals, Skukuza, Kruger National Park, South Africa, 14-18 Sept. 1993. [Onderstepoort, South Africa: SAVA Wildlife Group]. 84pp. Stanley Price, M. R. 1989. Animal re-introductions : the Arabian oryx in Oman. New York: Cambridge University Press. 291pp. �262 Taylor, T. G. and B. C. Pittman. 1933. Game management development and needs. Logan, UT : Utah Agricultural Experiment Sta~ion and Extension Service. Miscellaneous publication; 10. 51pp. von Ahlefeldt, J. P. 1992. The landscape ecology of the Palmer Divide, central Colorado. Ph.D. Dissertation, Fort Collins, CO, Colo. state Univ. 372pp. Wardell-Johnson, G., J. D. Roberts, D. Driscoll, and K. Williams. 1995. Orange-bellied and white-bellied frogs recovery plan. 2nd edition. Como, Western Australia : Western Australia Dept. of Conser. and Land Manage. Western Australian wildlife management program; no. 19. 28pp. Warren, E. R. 1942. The mammals of Colorado their habits and distribution. 2nd edition. Norman, OK University of Oklahoma Press. 330pp. Wedge, D. C. and A. J. Long. 1995. Key areas for threatened birds in the neotropics. Washington, D.C. : Smithsonian Institution. 311pp. CDQH Employees and Cooperators The Research Center Library staff also located and delivered approximately 2,292 individual items or free documents on request for Colorado Div. of Wildlife employees and cooperators during this segment. Maintain Electronic and Card Catalogues of all Research Library Holdings 606 is the total number of items cataloged during this period of time. This includes not only new acquisitions, but also older materials from the library collection being entered into the electronic catalog for the first time. Among the new acquisitions are Federal Aid : Job Progress Reports and manuscripts written by Colorado Div. of Wildlife Researchers and other employees. 983 is the total number of computer scanned items added to the electronic circulation system during this period. This includes not only the above mentioned newly cataloged items, but also newly acquired serials, volumes, and items being assigned scanning numbers for the electronic circulation system for the first time. . CDQW Manuscripts Published Job Progress Reports; July. 1997 - June. Federal Aid. 1998 All studies. Andelt, W. F. and T. D. I. Beck. 1998. Effect of black-footed on behavior and reproduction of prairie dogs. Southwest. 43:. Baker, ferret Nat. (in press) D. L., G. W. stout, and M. W. Miller. 1998. A diet supplement for captive wild ruminants. J. Zoo Wildl. Med. 29(3):. (in press) Beck, T. D. I. 1997. Citizen ballot initiatives: a failure of the wildlife management profession. In: The Wildlife Society : Fourth Annual Conference: Sept. 21-27, 1997 : Snowmass, Colorado. Bethesda, MD : The Wildlife Society. p.73. (abstract) Beck, T. D. I. 1998. Citizen ballot initiatives : a failure of the wildlife management profession. Human Dim. Wildl. 3(2):21-28. �263 Beck, T. D. I., R. B. Gill, W. F. Andelt, R. H. Kahn, and J. A. Fitzgerald. 1997. Putting the public back into public agencies lessons from the Colorado fur trapping controversy. In: The Wildlife Society: Fourth Annual Conference: sept. 21-27, 1997 : Snowmass, Colorado. Bethesda, MD : The Wildlife Society. p.72. (abstract) Bright, A. D., J. F. Lipscomb, and L. Sikorowski. 1997. A cognitive analysis of intergroup attitudes of state wildlife agency personnel and wildlife rehabilitators. Human Dim. Wildl. 2(1):4767. Eussen, J. T., M. A. Link, J. A. Fitzgerald, and T. D. I. Beck. 1997. Kit fox distribution and status in Colorado. In: The Wildlife Society: Fourth Annual Conference: Sept. 21-27, 1997 : Snowmass, Colorado. Bethesda, MD : The Wildlife Society. p.99. (abstract) Freddy, D. J. and G. C. White. 1997. Implications of elk survival rates in Colorado. In: The Wildlife Society : Fourth Annual Conference: sept. 21-27, 1997 : Snowmass, Colorado. Bethesda, MD : The Wildlife Society. 103-104pp. (abstract) Giesen, K. M. and G. D. Kobriger. 1997. Status and management of sharp-tailed grouse Tympanuchus phasianellus in North America. Wildl. Biol. 3(4):286. Gill, R. B. 1997. Professionalism, advocacy, and credibility: a futile cycle? In: The Wildlife Society : Fourth Annual Conference : Sept. 21-27, 1997 : Snowmass, Colorado. Bethesda, MD : The Wildlife Society. p.108. (abstract) Green, A. L., N. M. DuTeau, M. W. Miller, J. Triantis, and M. D. Salman. 1998. Differentiating cytotoxic and noncytotoxic Pasteurella trehalosi from Rocky Mountain bighorn sheep : polymerase chain reaction primers to the leukotoxin A gene. Am. J. Vet Res (in press) Hoffman, R. W., M. P. Luttrell, W. R. Davidson, and D. H. Ley. 1997. Mycoplasmas in wild turkeys living in association with domestic fowl. J. Wildl. Dis. 33(3):526-535. Kraabel, B. J. and M. W. Miller. 1997. Effect of simulated stress on susceptibility of bighorn sheep neutrophils to Pasteurella haemolytica leukotoxin. J. Wildl. Dis. 33(3):558-566. Kraabel, B. J., M. W. Miller, J. A. Conlon, and H.J. McNeil. 1998. Evaluation of a multivalent Pasteurella haemolytica vaccine in bighorn sheep: protection from experimental challenge. J. Wildl. Dis. 34(2):325-333. Larsen, R. S., and M. A. Wild. 1997. Surgical correction of urethral obstruction in an elk (Cervus elaphus) by perineal urethrostomy. In: Proceedings, American Association of Zoo Veterinarians annual conference: October 26-30, 1997. [S.l.): The Assoc. of Am. Zoo Veterinarians. p.51 (abstract) McCarty, C. W., and M. W. Miller. 1998. Modeling population dynamics of bighorn sheep the facts of life for bighorn in North America. Fort Collins, CO : Colo. Div. of Wildl. Special Report; no. 73. (in press) McCarty, C. W., and M. W. Miller. 1998. A versatile model of disease transmission applied to forecasting bovine tuberculosis dynamics in white-tailed deer. J. Wildl. Dis. 34: (in press) Meadows, L. E., W. F. Andelt, and T. D. I. Beck. 1998. Managing black bear damage to beehives in Colorado. Fort Collins, CO : Colo. State Univ. Coop. Ext. Bull. (in press) �264 Miller, M. W. 1999. Pasteurellosis. In: Infectious diseases of wild mammals, 3rd edition, E. S. Williams, et ale (eds.). Ames, IA : Iowa state Univ. Press. (in press) Miller, M. W., J. A. Conlon, H. J. McNeil, J. M. Bulgin, and A. C. S. Ward. 1997. Evaluation of a multivalent Pasteurella haemolytica vaccine in bighorn sheep: safety and serologic responses. J. Wildl. Dis. 33(4):738-748. Miller, M. W. and S. M. Schmitt. 1997. Bovine tuberculosis in captive and free-ranging cervids : implications for wildlife management. In: The Wildlife Society : Fourth Annual Conference: Sept. 21-27, 1997 : Snowmass, Colorado. Bethesda, MD : The Wildlife Society. p.145. (abstract) Miller, M. W., M. A. Wild, and E. S. Williams. 1998. Epidemiology of chronic wasting disease in captive Rocky Mountain elk. J. Wildl. Dis. 34(4):532-538. Peles, J. D., F. W. Weathersbee Jr., P. E. Johns, J. Griess, D. L. Baker, and M. H. Smith. 1998. Genetic variation in a recently isolated population of mule deer (Odocoileus hemionus). Southwestern naturalist 43:. (in press) Pojar, T. M. 1997. American pronghorn: uniquely ours. Colorado country life 28(9):16-18. Pojar, T. M. 1998. The American pronghorn - a unique species. Colorado outdoors 47(1):16-20. pojar, T. M. 1998. Interstate pronghorn survey: comparison of fixedwing line transect and helicopter quadrat surveys. S.l.: North Am. Pronghorn Foundation. Pronghorn Antelope Workshop Proceedings 18:. (in press) Snyder, W. D. 1997. Sandsage-bluestem prairie renovation to benefit prairie grouse. Fort Collins, CO : Colo. Div. of Wildl. Special report; no. 71. 34pp. Spraker, T. R., M. W. Miller, S. E. Williams, D. M. Getzy, W. J. Adrian, G. G. Schoonveld, R. A. Spowart, K. I. O'Rourke, J. R. Miller, and P. A. Merz. 1997. Spongiform encephalopathy in re-ranging mule deer (Odocoileus hemionus), white-tailed deer (Odocoileus virginianus) and Rocky Mountain elk (Cervus elaphus nelsoni) in northcentral Colorado. J. Wildl. Dis. 33(1):1-6. Stubblefield, W. A., S. F. Brinkman, P. H. Davies, T. D. Garrison, J. R. Hockett, and M. W. McIntyre. 1997. Effects of water hardness on the toxicity of manganese to developing brown trout (Salmo trutta). Environ. Tox. & Chern. 16(10):2082-2089 Thompson, K. G., E. P. Bergersen, and R. B. Nehring. 1997. Injuries to brown trout and rainbow trout induced by capture with pulsed direct current. N. Am. J. Fish. Manage. 17(1):141-153. Thompson, K. G., E. P. Bergersen, R. B. Nehring, and D. C. Bowden. 1997. Long-term effects of electrofishing on growth and body condition of brown trout and rainbow trout. N. Am. J. Fish. Manage. 17(1):154-159. Walker, P. G. 1997. Whirling disease problem reveals need to redirect agency fish health programs. Fisheries 22(8):4 White, G. C. and R. M. Bartmann. 1997. Density dependence in deer populations. 120-135 In: The science of overabundance: deer ecology and population management. eds. McShea W. J., H. B. Underwood, and J. H. Rappole. Washington, D.C. : Smithsonian Institution Press. pp.120-135. �265 Task 1. 2 The following publication-ready manuscripts were developed: Draft Strategy for the Conservation and Reestablishment Wolverine in the Southern Rocky Mountains Northern Wild Sheep Symposium and Goat Council: Proceedings of the Tenth Biennial Developed graphics and visual researchers and biologists. 3. Special Reports Nos. 71 and 72 and Federal Aid Abstracts for 1995, and 1997 were published and distributed to Division personnel and outside agencies and libraries. 4. No progress 5. Wildlife Research and distributed. 6. A Customer Service Survey concerning publication services was developed and distributed to 48 customers. Twenty two were returned (46%). The results were very positive. There were 9 items for evaluation and the totals are reported as follows: Very good = 129; Good = 54; Fair = 8; Poor = 2; Very poor = 1; NA = 4. was made on Special Reports Report for 52 presentations and 2. Task materials of Lynx No. 73 in this for 1996 and 1997 were by 1996, Segment. assembled, published, 3 Operation of Foothills Wildlife Research Facility (FWRF) during FY1998 was in support of research on chronic wasting disease in deer and elk (and potential for natural transmission of CWO to domestic cattle and pronghorn), deer contraception, pneumonia immunization of bighorn sheep, and prairie dog research for black-footed ferret recovery. Animal care and facility maintenance standard operating procedures (SOP's) were followed for routine animal husbandry and facility procedures. An SOP was developed for care of prairie dogs at FWRF. Given this general guidance, and the direction required to meet terrestrial research needs, we performed the following tasks: Animal Maintenance General: Again this year, routine feeding and caretaking of research animals, including health observations, training, weighing, and clean-up, was performed primarily by well trained work-study and temporary employees, as well as volunteers. FWRF was inspected by USDA APHIS for compliance with federal animal welfare regulations on 27 May 1998. To maintain optimal population size of captive wildlife species for use in research, we euthanized some pronghorn, but added to the herd size of mule deer, elk, and added cattle and white-tailed prairie dogs to the facility. Nutritional Maintenance Feeding protocols: Feeding protocols were as previously described (reviewed by Wild 1997). Prairie dogs were maintained on a base diet of rat chow �266 (Harlan Teklad #8640, Madison, WI) provided ad libitum and supplemented with fresh vegetables (e.g., carrots, lettuce, dandelions) and a horse sweet grain mixture. Health Maintenance General: We continued to monitor animal health using FWRF SOP's. Animal health care was provided as required and as mandated by the preventive medicine program and chronic wasting disease protocols. Four mule deer does were ovariectomized in preparation for evaluation of a contraceptive. Chronic wasting disease: We followed protocols for the preventive medicine program (Wild 1995) and prepared a revised chronic wasting disease (CWO) management protocol (Attachment 1) to increase biosecurity at FWRF. The protocol was prepared in response to a change in objectives for CWO management at FWRF--rather than attempting to control and eliminate the disease in captive cervids, we will maintain CWO for research purposes. Additionally, domestic cattle were added to the captive herd to investigate the potential for natural transmission of CWO from cervids to cattle (See Work Package 30013). These changes mandated the increase in biosecurity to minimize risk to personnel and the potential spread of CWO outside the facility. The guidelines established in the protocol were reviewed by scientists with the USDA Agricultural Research Services and Veterinary Services branches, the Colorado State Veterinarian's Office, and the Colorado State University Biosafety Committee. Since 1994, we have attempted to minimize the potential spread of CWO between captive mule deer by euthanatizing CWO suspects early in the course of the disease; however, to assure that research cattle at FWRF are allowed ample opportunity to contact CWO affected deer, we have increased our criteria for euthanasia of CWO suspect deer. Deer are now allowed to progress after initial clinical signs of CWO have been noted (moderate behavioral changes and weight loss) until they die or are euthanatized for humane reasons after weight loss >20%, concurrent disease, or severe behavioral changes. In conjunction with research investigating the pathogenesis of chronic wasting disease in mule deer (See Work Package 3001-3), pronghorn at FWRF are exposed to mule deer with CWO through fenceline contact and a shared water source. No evidence of CWO has previously been found in pronghorn. We will continue to examine brains and other tissues of pronghorn that die at FWRF to monitor for evidence of CWO after this planned longterm close exposure to affected deer. Facility/Maintenance/Repairs/Improvements A variety of scheduled and unscheduled maintenance and repair activities were necessary to support facility operation and ongoing research programs. We worked toward a conservation-oriented approach for facility care by undertaking preventive maintenance projects, and performing high-quality new construction and repairs to existing facilities. Facility repair and construction projects were prioritized based on animal welfare concerns and anticipated research needs. Research Projects Facility projects operations offered support for pilot studies and for research conducted by CDOW personnel and other collaborators that were �267 initiated, throughout Educational conducted, the year. or continued using FWRF animals and facilities Contributions Facility tours and educational lectures were provided to school, university, and professional groups visiting FWRF. We emphasized the importance of maintaining captive wildlife for performing controlled experiments and the contributions made by research projects performed at FWRF. FWRF animals and facilities were also used occasionally for hands-on training for professional groups. RESULTS Animal AND PISCUSSION Maintenance General: In FY1998, temporary, work-study, YCC employees and volunteers performed the majority of tasks involving animal care and facility maintenance at FWRF. Nine volunteers contributed 685 hr work to FWRF. These volunteers performed primarily caretaker tasks and also assisted in weighing and collecting samples from animals. Contributions by volunteers represented a savings to FWRF of about 0.34 TFTE and $6749 (vs. cost of temporary employees). The animal welfare inspection by USDA APHIS found FWRF in compliance with AWA standards. This finding highlights the high standard of animal care provided by FWRF employees and volunteers. all At the close of FY1998, FWRF maintained 25 elk, 34 bighorn sheep, 5 whitetailed deer, 51 mule deer, 11 pronghorn, 33 white-tailed prairie dogs, and 11 domestic cattle. During the year a colony of white-tailed prairie dogs was added to the facility in support of black-footed ferret recovery research (Work Package 0880-1). Prairie dogs were captured at the Arapaho National Wildlife Refuge, Colorado. Domestic cattle from the Padlock Ranch, Wyoming and an additional 32 mule deer from Rocky Mountain Arsenal, Colorado were also added to the herd in support of CWO research (Work Package 3001-3). Five cow elk (2 from Sybille Wildlife Research Unit and 3 from Wyoming and Idaho Fish and Game) and 2 viable elk calves born at FWRF. were added to the herd in support of brucellosis RB51 vaccine research. Twenty-eight natural mortalities occurred in addition to planned euthanasia of 5 pronghorn for population reduction and 4 mule deer as per study protocol (Table 1). Nutritional Maintenance Feeding protocols: Individuals of all species maintained reasonable body condition on available diets with the exception of some mule deer fawns. Death of four fawns <2 mo of age (Table 1) may have been associated with general poor condition of the does. Addition of cattle to the mule deer pens in July 1997 disrupted access of lactating mule deer does and their fawns to feed and water areas. Despite mitigation measures employed to improve access, disturbance from cattle may have contributed to poor body condition of mule deer. Ali adult prairie dogs accepted the laboratory diets (and water bottles) within 2-3 days of capture and increased body mass from that point on. The adult males gained an average of 500 g between capture and 30 June, which represents a body mass increase of approximately 36%. Juvenile prairie dogs �268 gained an average of 63 g between time of capture and 30 June, which represents an increase of approximately 12%. Wild white-tailed prairie dogs become hyperphagic during the summer months in order to accumulate sufficient fat stores for hibernation, therefore the weight gains we observed were not unusual. Health Maintenance General: Overall, captive wildlife maintained at FWRF remained healthy throughout the year. Chronic wasting disease (CWO) continues to be a significant source of mortality in captive mule deer 'and has emerged as a significant source of mortality in white-tailed deer (see below). Chronic wasting disease: All animals at FWRF were monitored closely for clinical signs of cwo. Tissues from all mortalities occurring at FWRF were examined for evidence of infection with CWO. Of the eight adult mule deer that died or were euthanized in FY1998, seven were due to CWO (Table 1). In July, chronic wasting disease was diagnosed for the first time in FWRF whitetailed deer. Four of a cohort of 11 white-tailed deer (2 bucks, 1 castrate, 1 doe) died or were euthanized with CWO in a 1 mo period (July 1997). Two more white-tailed deer (one castrate, one doe) were euthanized with cwo in the following 11 mo. Based on information in other cervid species (Williams et al. 1982), exposure of white-tailed deer likely occurred ~18 mo prior to the July deaths. Interestingly, the current outbreak of cwo in captive mule deer at FWRF began 18 mo prior to the July deaths. Due to the occurrence of cwo in white-tailed deer, the USDA Animal Damage Control group (for whom the deer were maintained for immunocontraceptive research) terminated involvement and financial support for maintenance of the deer. The white-tailed deer will now be used primarily in cwo research. The change in management at FWRF to maintain CWO and maximize potential exposure by cattle may also effect the epizootiology of CWO observed in FWRF mule deer, i.e., affected deer may have an increased opportunity to expose other deer before removal from the herd. Clinical observations suggest that CWO-affected captive deer, if not euthanatized early in the course of disease, often develop aspiration pneumonia or die of hypothermia. Facility Maintenance/Repairs/Improvements As older portions of the facility are repaired and replaced, the need for unscheduled daily repairs appears to be decreasing. Maintenance projects continue to be important for animal safety and facility function. Significant maintenance/repair/ improvement projects completed at FWRF this year included: - Renovation of "duck lab" to small mammal facility - Construction of new small mammal building - Design and construction of 9 individual prairie dog cages Construction of handling/weigh area for cattle - Modifications to mule deer feeding enclosures �269 Research Projects The following pilot studies and research experiments were conducted, or continued using FWRF animals and facilities initiated, this year: Evaluation of antemortem tests to diagnose chronic wasting disease cervids--M.A. Wild, M.W. Miller, T.R. Spraker, and K. O'Rourke. - Susceptibility of cattle to cervid Williams, M.W. Miller, et ale - Survey of rodents for evidence of chronic -A. McNeil, M.W. Miller, D. Gould. pathogenesis M.W. Miller Efficacy Kreeger, of chronic wasting spongiform disease of Brucella abortus strain M.W. Miller, et ale in encephalopathy--E.S. wasting in mule RB51 vaccine disease deer--E.S. in adult prion protein- Williams and cow elk--T.J. Evaluation of a multivalent Pasteurella haemolytica supernatant vaccine in bighorn sheep: efficacy of alternative delivery approaches--H. McNeil, M.W. Miller, et ale - Regulation of mule deer population growth by fertility control: laboratory, field, and simulation experiments--D.L. Baker and T. Nett. - Use of insect growth regulators to control dogs--K.T. Castle and M.A. Wild. Development of fecal indicators Robinson, J. Byers, M.A. Wild. Educational of pronghorn fleas on white-tailed diet quality prairie and stress--M. Contributions Numerous informal tours of FWRF were provided individually to visiting professionals. FWRF provided formal educational instruction for special interest grade school through university groups, in addition to professional groups. - CDOW Youth in Natural Resources - Denver Youth Naturally The Wildlife Society Annual Meeting-Animal Immobilization Training CSU Zoological Medicine Society-Animal Immobilization Training CSU Wildlife Techniques class Rocky Mountain High School advanced biology class - Loveland High School advanced biology class - Front Range Community college forestry and wildlife classes Several special interest grade school groups LITERATURE Wild, M. A. Colorado Collins. CITED 1993. Animal and pen support facilities for mammals research. Div. Wildl. Res. Rep., WP1a, Jl, Jul 1992 - Jun 1993, Fort �270 Wild, M. A. 1995. Animal and pen support facilities for mammals research. Colorado Div. Wildl. Res. Rep., WP1a, J1, Jul 1994 - Jun 1995, Fort Collins. Wild, M. A. 1997. Animal and pen support facilities for mammals research. Colorado Div. Wildl. Res. Rep., WP1a, J1, Jul 1996 - Jun 1997, Fort Collins. Williams, E. 5., S. Young, and R. F. Marsh. 1982. Preliminary evidence of transmissibility of chronic wasting disease of mule deer. Wildlife Disease Association Conference, Madison Wisconsin, Abst. #22. Task 4 In 1990, the Colorado Division of Wildlife initiated a Federal Aid project to be a focus for coordinating a variety of actions to further understanding about diseases affecting wildlife populations throughout Colorado through surveillance, modeling, and research. The overall goal of this project is to provide data and analyses to support management programs directed toward detecting and managing important health problems affecting Colorado's wildlife resources. Surveillance We continued our program for acquiring, examining, reporting on, and summarizing wildlife disease cases occurring throughout Colorado. Between July 1997 and June 1998, we received 14 noncervid submissions for diagnostic evaluation (all deer and elk submissions are now reported under targeted CWO surveillance; see WP3001-T3 and WP3002-T3). Aside from canine distemper in a coyote pup and pasteurellosis in bighorn sheep (see below), all cases appeared to represent isolated incidents of trauma, intoxication, or disease. Investigations We investigated a respiratory disease outbreak among bighorn sheep in the sugarloaf subpopulation of the Tarryall-Kenosha herd complex (S23N, 5235). Carcasses or partial carcasses of 16 bighorns were recovered. Of those suitable for postmortem examination, all showed evidence of acute to peracute fibrinous pneumonia. Pasteuralla haemolytica was recovered from several of the bighorns sampled; further characterizations of isolates by Dr. Al Ward, University of Idaho, are in progress. A male yearling domestic sheep was observed running with wild sheep from the affected subpopulation about 2 wk after the first mortality was detected; this animal was strongly implicated as the source of the pathogen(s) responsible for this epizootic. No estimate of overall mortality was made, but >75% of the lambs present in this herd in late October had disappeared by late January. Additional follow-up information will be reported as it becomes available. Research Experimental FY97/98. studies: No experimental studies were planned or funded for Modeling (McCarty, Burnham, and Miller): We continued developing, refining, and evaluating models for predicting dynamics and impacts of infectious diseases in free-ranging ungulate populations. A manuscript by McCarty and Miller (1998) on forecasting tuberculosis dynamics in free-ranging whitetailed deer was accepted for publication in the Journal of Wildlife Diseases (see FY97/98 publications list for complete reference). �Table __ 1. Wildlife 7-17-97 golden health surveillance ;Jl~_ submissions, eagle w 7-24-97 coyote 3 mos 9-12-97 desert 9-16-97 raccoon bhs mt. lion canine 978-02833 distemper unknown 978w1058 trauma 978-08896 mt. 12-11-97 gr. horned owl lion 978-12976 978-14542 fungal or higher bacterial infection shot 978-15232 sw attacked shot 978-16553 ne disseminated bacterial infection 978-18222 subacute suppurative bronchopneumonia 978-20381 se gunshot 978-24070 ne electrocution ne emaciation and aspergillosis 978-27248 ne no encephalitis unknown 978-30255 f bhs se 9 (? ) focal enteric lesion with secondary septicemia and complications. coyote 11-25-97 2-5-98 978-01985 se m 1-5-98 pleural urate deposition poisoned m f 11-12-97 1998. ne bhs yrlg 11-5-97 1997-June w a 10-22-97 July 271 f goat a person a 2-13-98 3-4-98 3-30-98 bald eagle a f a f snowy owl coot (? ) 978-25095 �272 Task 5 RESULTS Bulleted Accomplishment ~ regarding Highlights the objectives of Accomplishments of Task 5 include: Consulting assistance to CDOW on harvest surveys, terrestrial inventory systems, and population modeling procedures was provided. Estimates of fall turkey, grouse, and squirrels were computed from survey data. Input on the design and analysis of the Harvest Information Program was provided on several occasions. A book on methods to estimate the spatial distribution and statewide abundance of Colorados's wildlife species is being published by Academic Press. This book should be available by September, 1998. ~ A spreadsheet model of mule deer and elk populations was developed CDOW, and distributed to terrestrial biologist via email. ~ No updates or problems were identified. ~ were corrected in DEAMAN software because for none Two 1-day workshops were conducted with region personnel in the use of DEAMAN and population modeling procedures, mainly to instruct region personnel on the use of spreadsheet models for ungulate population dynamics. In addition, numerous questions were answered via meetings with biologists, and via email. Assistance was provided data regarding estimates Uncomphagre Plateau and data will be summarized publication, a draft of to Thomas D.I. Beck with the analysis of of black bear population size on both the the Middle Park North study areas. These and reported in a technical, scientific which will be prepared during Segment 12. ~ Building on experience gained with the use of TrailMaster camera surveillance systems triggered by infrared beams to estimate abundance and population size of swift fox, the system was modified for use with kit fox initially to detect kit fox presence at suspected dens sites. Eventually the system will be used to monitor activities, litter sizes, and family group dynamics. ~ Assistance provided to David J. Freddy regarding the analysis of elk sightability models resulted in a more economical sightability model (the model required fewer variables to be measured) and a more efficacious model (the population was estimate with greater precision and less bias). ~ A research proposal to examine the impact of elk competition winter survival of mule deer fawns was developed. ~ Additional analyses to understand the decline of mule deer fecundity were conducted. Proposals for research on impact of predation and habitat changes were developed. ~ The results on over- from the study Compensatory Effects of Harvest in a Mule (Federal Aid Project W-153-R Work Plan 2 Job 15) were Deer population �273 published 225. ~ of Wildlife Management, Volume 62, pages 214- Locations of elk radio-collared in the White River Data Analysis Unit were monitored during the August-September period to determine their response to the opening of the archery hunting season. ~ ~ in the Journal A preliminary report providing an evaluation of the impact of archery hunting season and the use of All Terrain Vehicles (ATVs) on the movements of radio-collared elk in the White River Qata Analysis Unit was prepared. In addition to these routine consulting outcomes, Division of Wildlife supervisors Bruce Gill and Rick Kahn requested an analysis of existing mule deer data files to examine for clues to the cause(s) of the recent mule deer population decline in Colorado. Following is a detailed report of those analyses. RESULTS OF DATA ANALYSES TO EXAMINE FOR CAUSES OF MULE DEER POPULATION DECLINES G. C. White OBJECTIVES 1. Evaluate changes 2. Summarize 3. Develop 4. Develop research projects and reverse the trend. evidence hypotheses in mule deer fecundity of the decline during in fecundity about why this decline to further RESULTS the last 20 years. in mule deer. may have occurred. understand the cause of the decline, AND DISCUSSION I examined state-wide trends in deer age ratios. Fawn:doe ratios in the December age ratio surveys, and the antlerless harvest were regressed against year, 1972-95. I've included the details of the analysis below, but the conclusion is undeniable. Fawn:doe ratios have declined about 1.5 fawns per 100 does per year since 1972. This result requires further attention. Deer populations are generally perceived to have declined. Is this population decline real or imagined, and if real, does reduced recruitment explain the decline in population size? The following is a tabulation of ideas, many without merit or support. However, I want to define a broad view of the issue, with the purpose to focus thinking after evaluating multiple hypotheses, and develop research scenarios to understand the cause of the decline, and hence management tactics to reverse the trend. The goal is to suggest research needed to understand why the decline has taken place so that corrections can be developed. Hypotheses 1. a. about why the decline has occurred. Predation Increased coyote predation due to declining coyote control especially with the use of toxicants such as 1080. �274 b. 2. a. b. 3. a. b. 4. a. b. 5. a. b. 6. 7. Increased black bear populations due to less effective harvests and control measures as a result of the passage of Amendments 10 and 14. Elk-deer competition Summer-range competition -- social interactions? Winter-range competition, lower nutrition for does, resulting in smaller fawns born, with lower neonatal survival Habitat degradation Winter-range browse in poor condition (hedged, low vigor) from 50 years of high deer populations -- does in poor condition after winter, small fawns born, low neonatal survival. Urbanization Cattle/sheep competition Summer range Winter range Density-dependence operating on deer population High deer populations since 1950's, habitat changes, lowered recruitment High over-winter fawn mortality observed at Little Hills is symptom of the problem of fawns with low vigor. Buck-doe ratios have declined, as a result,fecundity has dropped. Survey methodology has changed, making the decline an artifact. Data to evaluate 1. 2. 3. 4. 5. Evidence each hypothesis: December age/sex surveys January Quadrat counts Pregnancy rates Harvest estimates Other pertinent data and Analysis December Age Ratios I took the available data from DEAMAN, and only used DAU's with the first age ratio survey prior to 1985 or before. This criterion ensures a reasonably long time span, and enough years to perform a valid test of trend across time. I regressed the number of fawns counted against the number of does counted, plus year. Thus, the slope of year is zero under the null hypothesis, but negative under the alternative hypothesis of declining age ratios. Following is a summary table of the slopes for year for each of the DAU's used in the analysis. The variable FEMALES is the slope for fawns per doe, thus gives an average age ratio for the DAU. DAU D-3 D-4 D-5 D-6 D-7 D-8 D-9 D-10 D-11 D-12 D-13 D-14 FEMALES 0.48569 0.57404 0.55244 0.62674 0.70465 0.85948 0.92723 0.72607 0.62462 0.6845 0.68873 0.64283 YR -5.0426 -1. 7103 4.0531 -7.1713 -54.4899 -22.1644 -21. 2133 2.2341 -4.9147 -42.4126 -7.0638 -7.4644 SE(YR) 9.26908472 5.26367626 7.36844542 2.88838843 10.4325489 7.00784334 6.44471111 6.47837271 3.78410461 13.091934 3.7686302 1.50876603 T -0.544 -0.325 0.550 -2.483 -5.223 -3.163 -3.292 0.345 -1.299 -3.24 -1.874 -4.947 PR«T) 0.5983 0.7519 0.6375 0.0324 0.0002 0.0082 0.0054 0.7403 0.2302 0.0048 0.0936 0.0006 �275 D-18 D-19 D-24 D-41 D-42 D-43 D-44 0.93523 0.38938 0.52392 0.64904 0.66242 0.68422 0.63453 -9.1734 -35.7564 9.874 -9.1554 -8.6565 -13.4339 -10.9093 4.19828097 28.8925014 15.5965841 4.94391518 2.04052268 2.16800259 6.95426285 -2.185 -1.238 0.633 -1.852 -4.242 -6.196 -1. 569 0.0604 0.2510 0.5443 0.0938 0.0017 0.0002 0.1607 In the above table, only 3 of the 19 DAU's have positive slopes. Under the null hypothesis of no decline, ~ of the DAU's should have had positive slopes. None of the positive slopes are significantly different than zero. Eight of the 16 negative slopes are significantly less than zero at Q=0.05, and 3 more at Q=0.10. An overall test of the year effect is significant (P = 0.009), with an overall slope of -21.21333. Thus, the evidence is overwhelming that age ratios have declined in the last 2 decades. Doing a less rigorous, but more easily interpreted analysis, is to regress fawns:100 does ratio against time. The resulting equation is fawns:100 suggesting a decline is shown in Fig. 1. does = 2952.8 - 1.4543 the year of 1.45 fawns per 100 does per year. This relationship Trends in age ratios for Deer DAUs RATIO = 2952.8 -1.4543 YR 140 .... ""0 0 0 0 .... s::: ~ 120 • 100 239 •• • • • --.- ..•... -•.-1-1_, _-_I • 60 •• • .. • .• • 0.3177 I· I ., • • T:-•• --rf- r I 40 I I •• • I • •• • I I I I •••• I. • 20 •• .:: I • I : •I I I. . -I -1--•.. •• CI •.... Rsq • -- _. 00 N I I I AdjRsq · .. • • 'I -1-1_ t· ••• -r- -1-..•.. • I • I I • I 0.3149 Rt I'ISE 12.364 ---- • I I 1972 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 YR Figure 1. Regression offawns per 100 does against year for deer DAU's in Colorado with the first year s 1985. The only question that I would pose about the above analysis is how these DAU's came to have enough data in DEAMAN. The most likely explanation is that these DAU's are the major deer DAU's, and biologists have entered the data because they were doing DAU plans. Also, I selected several of these for past modeling efforts, so entered the data myself. However, I don't see that the DAU's in the analysis are particularly non-representative of the state, so don't really believe that a serious bias exists. �276 Anterless Haryest Age Ratios A similar analysis as above was performed for the estimated harvest of fawns and does. Harvest data in OEAMAN were used, generally starting in 1971 or 1972 and ending in 1994. The following is a summary table of the slopes for year for each of the OAU's used in the analysis. OAU 0-1 0-2 0-3 0-4 0-5 0-6 0-7 0-8 0-9 0-10 0-11 0-12 0-13 0-14 0-15 0-16 0-17 0-18 0-19 0-20 0-21 0-22 0-23 0-24 0-25 0-26 0-27 0-28 0-29 0-30 0-31 0-32 0-33 0-34 0-35 0-36 0-37 0-38 0-39 0-40 0-41 0-42 0-43 0-44 0-45 0-46 0-47 0-48 0-49 FEMALES 0.0672 0.0903 0.0821 0.0843 0.1203 0.0319 0.1068 0.1203 0.1258 0.0803 0.0442 0.1221 0.1139 0.1211 0.1131 0.0505 0.0773 0.1537 0.0963 0.114 0.1533 0.0784 0.1283 0.1086 0.0577 0.0595 0.136 0.1207 0.096 0.0371 0.0604 0.0901 0.055 0.1316 0.0739 0.0668 0.0859 0.2178 0.0949 0.0887 0.1135 0.0882 0.0631 0.1447 0.1334 0.0445 0.0369 0.0319 0.1068 YR -0.3393 -5.4372 0.2216 -2.5455 0.3853 -0.1742 -5.3818 -5.8868 -3.9294 -0.2350 -0.5698 -6.9460 -1.3295 -2.8266 -0.9491 -1.0610 -0.4544 -1.1029 -3.8147 -2.0384 -1.6571 -1.2947 -0.3343 -4.2936 -0.2437 0.1596 -0.2813 -0.3183 -3.2409 -0.4482 -0.1235 -0.0675 -0.0846 -0.7979 -0.1965 0.0045 -0.4092 0.0079 -0.3891 -3.2332 -2.0231 -0.8045 -0.4502 -0.0107 -0.2075 0.0349 -0.4437 -0.1929 1.0048 SE~YRl 0.2564 2.4892 0.1813 0.8227 0.9096 0.0709 2.9437 1.4775 0.9649 0.3294 0.5960 1.3582 0.4862 0.5277 0.4492 0.3552 0.1424 0.9602 1.1224 0.9405 0.3566 0.5077 0.1328 0.8760 0.2154 0.1613 0.3076 0.5125 1.0891 0.5560 0.1421 0.0829 0.0449 0.5468 0.1878 0.1326 0.1629 0.1463 0.2919 1.2844 0.5319 1.0680 0.5698 0.7746 0.0900 0.2952 0.2076 0.2612 0.8273 T -1.323 -2.184 1.222 -3.094 0.424 -2.458 -1.828 -3.984 -4.072 -0.713 -0.956 -5.114 -2.735 -5.357 -2.113 -2.987 -3.19 -1.149 -3.399 -2.167 -4.647 -2.55 -2.517 -4.901 -1.132 0.989 -0.914 -0.621 -2.976 -0.806 -0.869 -0.814 -1. 884 -1.459 -1.046 0.034 -2.512 0.054 -1.333 -2.517 -3.804 -0.753 -0.79 -0.014 -2.305 0.118 -2.137 -0.738 1.214 p 0.2032 0.0424 0.2394 0.0057 0.6764 0.0266 0.0851 0.0007 0.0006 0.4859 0.3533 <0.0001 0.0128 <0.0001 0.0497 0.0076 0.0054 0.2650 0.0032 0.0425 <0.0001 0.0231 0.0247 <0.0001 0.2756 0.3383 0.3749 0.5415 0.0075 0.4328 0.4035 0.4269 0.0768 0.1627 0.3109 0.9735 0.0224 0.9576 0.2000 0.0215 0.0013 0.4616 0.4388 0.9892 0.0320 0.9069 0.0451 0.4688 0.2387 �277 The evidence is again overwhelming that a decline in the fawn:doe ratio in the harvest has occurred since 1972. Of the 49 DAU's, 42 have negative slopes. The overall test of the year effect is highly significant (P < 0.001), with a slope of -3.929412. Again doing fawns:100 a state-wide does = analysis, 1099.5 the regression - 0.5486 equation is year meaning that 0.5486 fewer fawns are shot for every each year. This relationship is shown in Fig. 2. 100 does in the harvest Trends in harvest age ratios for Deer DAUs RATIO 100 rn ., 80 = 1099.5 -0.5486 •.• •• YR • N I o c o 60 o rn .,. Rt MSE 40 153.59 r::: c •.... 1009 Rsq 0.1700 AdjRsq 0.1692 20 o 1970 1972 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 YR Figure 2. Regression offawns per 100 does from harvest estimates against time for deer DAU's in Colorado. Elk and Pronghorn In thinking about what is causing the decline in deer age ratios, I checked to see if a similar trend was present in pronghorn and elk. The idea was to eliminate a methodological explanation, i.e., that the same methods are used for all 3 species, and only deer show the downward trend. Surprise! Pronghorn and elk show very similar trends in both the age ratio surveys and the harvest surveys. Pronghorn and elk age survey age ratios were explained by year at ~ = 0.0328 and ~ < 0.0001, respectively, and in the harvest at E < 0.0001 for both species. Thus, the results are just as conclusive as for deer that age ratios have been declining state-wide in both of these populations also. Below, I've included the graphs for the 2 species and 2 approaches. One interpretation of these results is that subtle changes in methods have caused the decline. However, I'm at a loss as to what these changes might be. Another hypothesis is that predation is operating on all 3 species, but deer show the biggest impact because of poor overwinter survival of fawns. �278 Trends in age ratios for DAUs SPEC IES"A RAT 10 •• 2752.8 -1.354 VR 100 • In U 90 --, 80 --- .•.. --...,. ..•.. "" E IIJ ... _- -- ..... ..._ 70 , 0 60 0 • • 50 40 &:: • co 30 >20 - • .- ---- -- _....._. - N • • • = I I I I • • I --- --! ....• ...,. - • • • 0"1 I _.. • .---- _-- --- - - ....• -'- •.. - 29 Rsq 0.1580 AdjRsq 0.1268 Rt I'1SE 17.018 • I I I I I I I 1972 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 VR Trends in age ratios for DAUs SPEC IES=E RATIO = 1331.3 -0.6442 VR 100 In 90 IIJ • 80 "" E • • IIJ 70 u, 0 60 0 -.--~-I'-:,-,--- .1 50 IC7' • • C 40- - .. = •. , -_. •• I • I I I I I I , 1_ I • I • !. II 11 1- i -- I • I • I • •• • • I . • I • 30 20 N ri '1-1 · -. I· I,!,-:-.J __1,i-~1 .. co >- • I I I __ I • I I TI -----, I I 1972 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 VR 282 Rsq 0.2059 AdjRsq 0.2031 Rt I'1SE 9.384 �279 Trends In harvest age ratios for DAUs RATIO 100 = 1568 . 8 -0.7841 ,... rn u c 0 0 ::::I 60 : I •• 40 • •• • • • I·· .. .. • 587 Rsq 0.1540 AdjRsq 0.1526 Rt I'ISE 151.28 • • • • I • • • •• •••• 20 0 N •• •• 0 >- YR • • C" c . 80 E u u.. SPECIES=A • ~• · • • · -1~-r-!-!---:-i-i-!-1!1I1-!t!tltit-t-i-f-i-l-i-I-I-~--I I I I I I I I I I I I I I 1970 1972 1974 1976 1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 YR Trends In harvest age ratios for DAUs SPEC IES=E RATIO = 518.59 100U1 -0.2539 • • 80 .... 0 0 ,. • • u u, • N 1061 Rsq 0.1019 AdjRsq 0.1010 Rt I'ISE 105.21 • U c E YR 60 • • • • • • I 40 •. I ,. • ,. · · C" c ::::I 20 0 >- 0 ~-tit-t-!-!-'-I-i-i-I-i-l-j-i-f-i-~ir~~~~-l-i-i--I I I I I I I I 1970197219741976197819801982198419861988 I I I I I I 1990 199219941996 YR Possible Decline in Buck:Doe Ratios One possibility for the decline in deer age ratios is that declining buck:doe ratios allow fewer mature males to do the breeding. I've been able to address some aspects of this hypotheses with data from DEAMAN. For the analyses that follow, I have not removed DAU's with no data entered in DEAMAN prior to 1985, as I did for the previous analyses. Part of the reason the old NW region was overly represented in that analysis is that they had entered old data for developing DAU plans. However, I know that there is still a bias towards that region, even without censoring. possibly, unlimited hunting of bucks in most areas of the state over the past 20 years has resulted in significant declines in the buck/doe ratios of many hunt units. The buck:doe ratio plotted against year looks like this: �280 SPEC IES=Deer M 70 " \. a 1 60 e s 50 o () \ \. o <> ~ '\ •• ~ <> •• <> 1 o <> o (> ,, S· ••.•. ",fi o 40 o ..... o 30 F e 20 <> ¢ c ¢ <> o o o m a 1 10 e s 0 ..... <> o 40 ~'-~-r-r~~~~r-r-r-~~'-~-r-r-r~~~~r-r-r-~~'-~-r-r-. 1970 1980 1990 2000 fR Although the decline is evident prior to 1980, we've basically had 15 years of nearly constant buck:doe ratios. The above plot is a cubic relationship. If you do a linear regression, the slope is -0.620818 (P<O.OOl) with R2 = 13%. Heavy hunting pressure would reduce the proportion of mature in the population. The proportion of yearling bucks to total against year looks like this: bucks/young bucks bucks plotted SPEC IES=Deer P r 0.9 () .(> ¢ 0.0 o 0 p 0.7 0.6 Y 0.5 r 1 0.4 g 0.3 M a 0.2 1 e 0.1 s 0.0 1970 1900 1990 2000 YR Again, conditions have been relatively constant for the last 15 years. The slope for a linear regressions is 0.002 (P = 0.115) with R2 = 1%.· Thus, there is really no evidence that the proportion of yearling bucks has changed consistently across the last 20 years. �281 I also looked at some other comparisons. If buck:doe ratios affect fawn:doe ratios, the previous year's buck:doe ratio should be a predictor the current year's fawn:doe ratio. The following graph depicts this relationship for the data from DEAMAN. of SPEC IES=Deer AGERATIO 120 = 52.551 +0.5566PREY_SA • • N .. ._ -.,..-. . .. .... - ':" -::· ·.....,"'.11 .• I. • • ,. -. --• •• • . ...•...,:-. ... ··~:fl~t' • • • •._. •• • • I • .: \. .,_ • ••• ",,;,'! •• "SI& • •• •• • .'" • •• III. =-:.•••. "Iiii " • • 210 Rsq 0.1726 AdjRsq 0.1686 Rt MSE 513.02 •• • I I I 10 20 30 I 40 I I I 50 60 70 Previous Sex Ratio (P < 0.001), and in the right went the opposite direction! The relationship is significant Interestingly enough, pronghorn The age ratio should decline in the previous year: as a function of the proportion direction. of yearling bucks SPEC IES=Deer AGERATIO 110 = 64.771 -4.9566PREY_YLG N • • •• • • • • •• -.1-• --..• ....#...1' .. • •• • ••• •• .. •• •• •• ,- :.. •• ~. • •• •• -e .• •••• I • .. ~....r.".. .. _..!.--• •• •••~.I'" • .., • • •• :. • A • ." • • 1.1•• • •• w,L: _. ••, • I ••• ••• I , • .• •I',,-:' '. • I I I I I I I I I 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 Previous Yrlg Ratio 184 Rsq 0.0040 AdjRsq -.0015 Rt MSE 531.6 �282 This relationship was not significant (P = 0.393). Results from the above analysis that support the breeding hypothesis are the relationship between age ratios and sex ratios. However, I think this result can be explained just as easily by confounded variables. Both the sex ratio and the age ratio have declined through time. The age ratio shows a fairly constant decline through time, whereas the sex ratio shows a strong decline prior to 1980, and then a fairly constant level for the last 15 years. Because both decline through time, or at least some portion of time, they correlate. The change in sex ratio over time is logically explained by hunting pressure. I question whether sex ratio is the cause of the decline in age ratios, because age ratios have continued to decline even when the sex ratio has been constant. Sex ratios may be part of the answer, but I don't think they are all of it. Just for comparison, the following is the graph of age ratios versus year for all the data from DEAMAN, not just DAU's with observations prior to 1985. The decline is not just in the period prior to 1980 as is the case for sex ratios. SPEC IES=Deer V 130 0 u n g 1 0 0 F e m a 1 e s 120 110 100 90 80 70 60 50 40 30 20 ¢ ¢ ¢ ¢ 0 ¢ ~ 0 ~ 1980 1970 ~ ~ ~ ¢ 1990 2000 VR Piceance Study Results I've also examined some of these relationships with data from the Piceance mule deer study, where we have weights and survival for fawns each year since 1980. I've taken the age and sex data from GMU 22 out of DEAMAN. A significant relationship (P = 0.034) between previous year's survival and age ratio is found. I think this negative decline is because of the addition of yearling females to the population, causing the observed age ratio to decline. �283 Relation of GMU 22 age ratios and CB and LH Fawn survival estimates 0.80 y o 0.75 u n 0.70 g : 0.65 1 o o F ...... - ...• ....• _ ... .- _- .•....•.. <> 0.60 0.55 e s o. 1 0.0 0.2 0.3 0.4 0.5 0.6 0.7 0.8 Previous Survival Rate The following are the relationships between survival (SHAT) and fawn weights as a function of the previous year's buck:doe ratio or proportion of yearling males of all males. Relation of GMU 22 age ratios and CB and LH Fawn survival estimates SHAT 0.8 0.7 0.6 ----.-- ._-_.__ 0.5 ._-. --.~----------.-.------.-~.-- ---------- 0.4j_--~~~-- "'_ • w .. --- _ . o 4- _ 0.3 0.2 o. 1 0.0 Q _----- - - _--~------ ~.-----------M-.-v_. A 0 ------ _ <> - - ~- - - - <> ~~~~~~~~TT~~~~~~~~TT,,~~~~~rr~ O.06 0.07 0 .08 0.09 "--"'- .... ,. ---~--. •• ~~~ o. 10 o. 11 o. 12 0 .13 Previous Sex Ratio 0.14 o. 15 o. 16 o. 17 �284 Relation of GMU 22 age rates and CB and LH Fawn survival estimates SHAT 0.8 0.7 0.6 0.5 0.4 - - _ _ _ __ --- -- .•...•. ----- V A 0.3 - ••- ••• - .•••••••. o 0.2 <> .•.••.• ••••••• ¢ 0.1 •••• .•••.•••• <> 0.0 ~-. •••• -rro••-.",.-r",.-r" ••-." ••-r""rrTT"-'''''-'ro~ 0.35 0.40 0.45 0.50 0.55 0.60 0.65 0.70 0.15 0.80 0.85 Previous Yrlg Prop. Relation of GMU 22 age rates and CB and LH Fawn survival estimates 36.0 W e 35.0 34.0 g h 33.0 t ( k g ) ._ ••••.• -~ <> ....-.~ -_ .... -._ ... ~--.. c ------~~---_ -~--_--_-----------.-------.---.--¢ 32.0 31.0 30.0 0.06 0.07 0.08 0.09 0.10 0.11 0.12 0.13 0.14 0.15 0.16 0.17 Previous Sex Ratio �285 Relation of GMU 22 age ratios and CB and LH Fawn survival estimates 36.0 W e i 35.0 ---. "'-_ 34.0 --_ - •. -_ g h - •. ----. -._- ---~--- ----~---.---------------------.--¢ 33.0 t ( k g ) 32.0i-----~o~------..••• ---------:¢:¢r-------------------~---..-----------..-----------------.. 31.0 30.0 ~----- ---_.-- . __ 0.40 •.- ----------.----~.--. ----<:>--.. _----- o .45 0 .50 0 .55 O. 60 0.65 0.70 -- ..---~-A ~-. 0.75 v 0.80 0.85 Previous Yrlg Prop. Only survival predicted by the previous proportion of yearlings is significant (P = 0.020). The rest are P > 0.15. We have observed large variations from year to year during this study. Whether the significant relationship is real, I don't know. If the breeding hypothesis is true and operating, I would have expected all 4 of the graphs to have been significant. However, because 3 are not does not imply that the breeding hypothesis is false. Low power from the limited sample size probably explains these results more than anything else. Also, the range in buck:doe ratios over the period of this study is somewhat limited, so reduces the power of the study to detect trends. Conclusion Fawn:doe and calf:cow ratios have declined in both the December surveys and in the estimated antlerless harvest for deer, pronghorn, and elk. Reasons for this decline need to be enumerated, and evidence gathered to support or dismiss them. If the hypothesis that declining buck: doe ,ratios is causing problems with fecundity is true, I would have expected to see stronger relationships from both the DEAMAN and Piceance analyses. Not much support is provided. However, the lack of support cannot be used as evidence that the hypothesis is not true. When you accept the null hypothesis of a statistical test, you don't learn much unless you have high power. None of the analyses performed above have much power. Task 6 PACE Plans were prepared and reviewed for and with all of the Mammals Program staff in August 1997. PACE evaluations were prepared and reviewed with Mammals Program staff members in July 1997. Signed review forms were forwarded to Jim Lipscomb, Terrestrial Wildlife Section Leader on 8-10-98. Several discussions and working group meetings were held to discuss the ramifications of Senate Bill 96-52 and Amendment 14 to Division operations and �286 relationships with the Colorado Department of Agriculture. The result of these discussions was an letter of instruction to all Division field staff signed by the Director. The letter of instruction detailed the Division's role in: a) dealing with claims for damage from bears and pumas and, b) the Division's role in certifying that landowners had unsuccessfully employed unrestricted damage control measures so they could qualify for restricted methods. All COFRS documents relating to Mammals Program expenditures were processed within 48-56 hours of the time receipts of expenditures were received by the Mammals Program Administrative Assistant. One potential problem arose because of discrepancies between CDOW's accounting records and CSU's accounting records in which CSU claimed up to $20,000 in unpaid balances. However, the CSU accounting section could provide no documentation of the unpaid invoices. Many of CSU's original records were destroyed in the flood of July 1997 so, to date, the issue remains unresolved. A complete procedures manual was completion update of the Mammals Program manual of standard operating was begun in January 1998 and completed in March 1998. The revised distributed to all Mammals Program staffers immediately upon of the revisions. April 1998 was the renewal period for the Application for Federal Assistance (AFA) for Federal Aid Project W-138-R. All Federal Aid project documentation was reviewed, revised, and resubmitted within the required deadlines. The format of W-153-R documents was modified to accomodate project identification by the u.s. Fish and Wildlife Service and the Division's new PBE planning and budgeting system. The new format should make it easier to account and audit federal aid expenditures under Project W-153-R for both agencies. A questionnaire was sent to each 'of the Mammals Program staff members asking for their evaluations of customer service. Only about ~ of the questionnaires were returned, but all of those returned rated customer service very good to excellent. Two staffers requested the administrative staff to develop and maintain a database of projects reviewed and approved by the Research Section's Animal Care an Use Gommittee. The database is being developed. Prepared by chael Mille~----- M~~ _.~~ Gary C. White td-J<<. �287 ATTACHMENT REVISED 1 PROTOCOL FOR MANAGING CHRONIC AT FOOTHILLS WILDLIFE RESEARCH WASTING DISEASE FACILITY HISTORY Chronic wasting disease (CWO) is a transmissible spongiform encephalopathy of cervids (deer and elk). CWO is in the same family of diseases as scrapie of sheep, bovine spongiform encephalopathy (BSE; mad cow disease), and Cruetzfeld-Jacob disease of humans. The disease causes behavioral changes and loss of body condition and is invariably fatal in affected deer and elk. Despite a comprehensive program initiated in 1985 to eradicate CWO from cervids at Foothills Wildlife Research Facility (FWRF), CWO remains endemic in the facility. After the 1985 clean-up, CWO was first diagnosed in elk in 1989 and in mule deer in 1994. Sporadic cases continue to occur, with cases in mule deer being markedly more common than those in elk. Based on these observations, guidelines established in 1985 (and revised in 1993) for maintaining a CWO-free facility are largely obsolete. Here, we provide revisions to those guidelines that are directed at maintaining the disease for research purposes in captive deer and elk while minimizing the risk to personnel and the potential spread of CWO outside the facility. OBJECTIVES 1. Prevent transmission of CWO to animals outside FWRF. 2. Minimize potential for exposing personnel to CWO or other potential pathogens. 3. Maintain endemic CWO in deer and elk at FWRF; however, animals showing classic clinical signs of CWO will be euthanized to avoid undue suffering. 4. Minimize potential spread of CWO between species of captive wildlife (deer, elk, and noncervid research animals). ASSUMPTIONS 1. CWO is an infectious disease of deer and elk likely caused by a prion; CWO is not widespread in free-ranging cervids. 2. Mode of transmission for CWO is not known, and may be direct, via animal/animal contact, or indirect, through contact with excreta (saliva, urine, feces); animate and inanimate objects may serve as fomites (vehicles) in transmitting CWO. 3. Noncervid wildlife and domestic species are potentially inapparent carriers of CWO; however, no data have been found to support this assumption. 4. If CWO is transmitted to a new host species, then the likelihood of further transmission to others within that species is increased. 5. There is no evidence that CWO is transmissible to humans; however, it is prudent to minimize exposure to this, and all, animal pathogens. APPROACH Overview: 1. Follow general guidelines that prevent contact of captive research animals with animals outside FWRF (wild and domestic). 2. Minimize potential transmission of CWO between species of captive animals (especially transmission from deer/cattle pens) via contaminated materials, equipment, or clothing. �288 3. 4. Maintain each species of animal in isolation from others, unless directed by research protocol (e.g., deer with cattle). Educate animal caretakers about CWO (hazards, protocols, and clinical signs exhibited by affected animals). Perform daily animal observations and maintain detailed records of animal health as a portion of the CWO surveillance program. Animals: 1. Exclude wild or captive cervids from endemic CWO areas from entering the captive herd, unless directed by a research, protocol. Endemic areas will now include: northeastern and northcentral Colorado, Park and Albany Counties, Wyoming, and Denver Zoo. 2. orphans, and neonates raised outside FWRF, will occasionally be accepted from areas that are not CWO endemic. 3. Raise and maintain each animal species in isolation from others, unless directed by a research protocol. 4. To prevent transmission of CWO from FWRF to facilities where CWO is not endemic, animals from FWRF will be transferred or donated to other facilities only if the following criteria are met: 1) animals are scheduled for a specific research project, 2) the destination is a closed facility (no egress of live animals) or FWRF-source animals are kept in isolation and sanitation measures followed. Recipients will be notified of CWO risks associated with accepting animals from FWRF. Animal Maintenance: 1. House and maintain each species in isolation from other species, unless directed by a research protocol. 2. Maintain accurate records for all animals. This information includes (but is not limited to): birth date, origin, body weights (monthly), vaccinations, health problems, research projects, travel. Additionally, i. Tag all animals for easy individual identification. ii. Train FWRF personnel to recognize clinical signs of CWO. FWRF personnel will maintain daily animal observation records describing animal status and will report abnormal observations to the facility manager. 4. Weigh and/or briefly examine every animal at least once monthly. 5. Follow a preventative medicine program that includes routine vaccination, anthelmintic treatment, hoof trimming, nutritional evaluation, and other measures to optimize overall health of research animals. Use of Research Animals Outside FWRF: 1. The loaning of animals to facilities outside FWRF is discouraged. 2. Procedures for isolating cervids at other CDOW facilities will be the same as those at FWRF. 3. RFAC will approve movement of FWRF research animals out of, and back in to, FWRF. 4. Animals maintained at FWRF will not be released into the wild. S. During field studies, precautions should be taken to minimize chances of exposure to free-ranging wildlife and livestock. 6. Researchers are responsible for maintaining accurate records of animals taken outside FWRF. General Facilities and Equipment: 1. Exclude free-ranging wildlife or livestock from the facility or from contact with captive animals using interior and perimeter fencing. �289 2. 3. 4. 5. 6~ 7. 8. Maintain each species of animal separately and allow no direct or fenceline contact (unless directed by a research protocol). Minimize runoff between pens housing different species through appropriate pen assignment and drainage control. Minimize common use of equipment between pens housing different species. Feed and handle animals or clean pens using the following traffic pattern: noncervids, elk, white-tailed deer, mule deer alone, mule deer with cattle. Clean animal pens (especiaLly feed areas and waterers) weekly. Dispose of waste from all pens, except those where deer and cattle are housed together, in the main dumpster. Isolation pens, and other areas where animals are held for extended periods, will be cleaned of organic matter and disinfected with a chlorine solution after use. The researcher last using the area will be responsible for cleanup. Cooperative compliance will be made a condition of all study plans which use FWRF ungulates. Different species may be held concurrently in isolation pens if a buffer zone (empty pen) is used. Feed: 1. Hay will not be accepted cultivated pastures. from areas where domestic sheep have grazed on Personnel: 1. Wash hands before and after handling each species of animal. 2. No eating or drinking allowed in animal areas. 3. coveralls, boots, gloves, and masks are available for use if desired and are recommended when handling animals showing clinical signs of C~. 4. Unsupervised access to FWRF will be limited to authorized personnel. Unauthorized persons will not enter animals pens or be permitted direct contact with research animals. The facility will be locked except when attended during normal business hours. 5. All researchers and collaborators and their subordinates will comply with this protocol. All personnel working at FWRF will be required to read this protocol and other appropriate literature and to sign the attached sheet of informed consent. Additional Requirements for Deer/Cattle Pens: 1. Protective clothing must be worn when entering deer/cattle pens, either: i. Designated coveralls and boots/shoe covers. ii. Personal clothing changed and washed after working in deer/cattle pens. 2. Place waste feed and manure from deer/cattle pens in compost pile at FWRF (NOT in dumpster). Dispose of compost by burial or incineration as needed. 3. Dedicated (separate) equipment should be used for cleaning deer/cattle pens (wheelbarrow, rakes, shovels, etc.). Alternately, equipment could be cleaned of organic material and disinfected with 10% bleach solution prior to use in other areas of FWRF. 4. Vehicles should be cleaned after use in deer/cattle pens. Wash organic material from tires. Remove all organic material from bed of truck (sweep, wash down if needed). 5 Keep gates to pens, hub/working area, and west side of the facility closed at all times except when passing through. �290 6. 7. 8. Animal carcasses should be enclosed with a protective cover to contain potentially infectious materials during transportation to the necropsy lab. Cattle will not leave the facility alive unless transfer to a biosecurity level 2 or greater facility is required as part of the research protocol. Report any abnormalities or accidents immediately to facility supervisor. CWO, SURVEILLANCE PROGRAM 1. Euthanitize any animal showing clinical signs of CWO and examine tissues grossly and histologically. 2. Perform complete postmortem examination and histologically examine brain tissue of any animal that dies at FWRF. 3. Carcass disposition will be by incineration (required for cattle) or appropriate burial. 4. If CWO is diagnosed in any noncervid species at FWRF, this protocol will be immediately revised and biosecurity at FWRF further increased. 5. RFAC will evaluate and amend this program as necessary. �291 INFORMED CONSENT I, , have read the Foothills Wildlife Research Facility (FWRF) protocol concerning chronic wasting disease (CWO) and agree to follow the protocol. Although there is no evidence that CWO is transmissible to humans, I realize that I will be working with research animals potentially infected with cwo. I understand that this protocol reflects current knowledge on measures for minimizing exposure to and spread of CWO and other potential pathogens at FWRF. Signature Date � https://cpw.cvlcollections.org/files/original/bc03b166e23990fdc566a2faf9b59c14.pdf 5cc208b4d29a8b7001d4a45186d39e86 PDF Text Text 1 Colorado Division of Wildlife Wildlife Research Report April 1999 JOB PROGRESS REPORT State of: ____,!C~o~l~or~a~d~o Peregrine Falcon Investigations (Avian Research) W..:....:.,_-..:,_17"""3'--'-R=-:-2=---_ Project: Work Plan: _ ___L Job _1_ Job Title: ---'P:...,:e=r""'egn=·n=e:...::F:....:al=co=n:...;:R=e=s=to=r=atI=·o=n=-- _ Period Covered: 01 January through 31 December, 1998 Author: Gerald R Craig Personnel: James H. Enderson, of the Colorado College and Rob Clemens, Gerald RCraig, Michelle Lockwood, Terry Meyers, Emile McCain, Ryan Pleune, Sara Wight, and Catherine Wightman of the Colorado Division of Wildlife. ABSTRACT In the 1998 breeding season, 88 territories were occupied by peregrines (Falco peregrinus anatum) and 76 pairs fledged 155 young for an overall productivity of l.91 young per occupied site. The rate of occupancy remained above 80% and a limited sample of eggshells averaged -8.4% of pre-DDT era eggs. The expanding population and limited resources necessitated development of a monitoring protocol that uses a sample of 41 preselected sites that serve as surrogates to represent the total population. Although site occupancy and eggshell condition are parameters that also require monitoring, annual productivity may be the most sensitive measure of Colorado peregrine population viability. Should chemical contamination reoccur, reduced fledging success will be the first manifestation of population decline. The protocol developed and tested this year appears to be a viable method for monitoring Colorado's peregrine falcons. �2 �3 PEREGRINE FALCON RESTORATION PROGRAM Gerald R Craig P. N. OBJECTIVES 1. Annually survey all documented peregrine breeding sites throughout Colorado to establish the presence of nesting peregrines. 2. Annually monitor a statistically significant sample of breeding pairs to document their reproductive success. 3. Annually monitor organochlorine pesticide levels. 4. Investigate population dynamics. ·5. Evaluate, characterize, and protect breeding habitat. 6. Document and protect important migration and wintering areas. SEGMENT OBJECTIVES 1. Whole, nonviable eggs that are encountered during eyrie visits will be collected, preserved and submitted to the appropriate U.S. Fish and Wildlife Service approved laboratory for pesticide analysis. 2. Compile data from previous years and prepare final report of eggshell thinning and pesticide residues in Colorado peregrine falcons from 1974 through 1997. 3. Compile and analyze data on peregrine falcon recovery and population dynamics and prepare manuscripts and annual report. 4. Moni tor selected peregrine falcon eyries for occupancy and productivity when funded. RESULTS Survey Effort Three teams comprised of 2 observers each were fielded for the 1998 season. Each team was assigned 20 priority territories to monitor. Among the 60 sites were 45 surrogate sites considered to be representative of the Colorado peregrine population. Selection was based upon occupancy longevity (prior to 1990), accessibility, and distribution. In addition to monitoring the priority sites for productivity throughout the season, the teams also attempted to visit all registered sites as well as survey potential cliffs throughout the state as time permitted. The teams were able to check 99 of 109 sites that were on record at the beginning of the field season. Time constraints and accessibility restricted them from visiting the �7 Colorado Division of Wildlife Wildlife Research Report April 1999 JOB PROGRESS REPORT State of: ----=C::.;°=l=°r""'a=d=° Project Migratory Bird Investigations W:..:,_-1~6:<..:::6~-Ro..::...... _ Work Plan: _1_: Job Title: Job_n_ ---'M=o=n:.:..:it=or:....:B=an=d=in:..:.lg=>-=of"-'M=al=l=ar=d=s..::in~C=ol=o;:_:ra=d=o'-_ Period Covered: Author: _ 01 January through 31 December 1998 Michael R. Szymczak Personnel: 1. Broderick, P. Creeden, V. Graham, 1. Gumber, T. Mathieson, J. Miller, 1. Olterman, R. Olterman, M. Szymczak, and S. Yamashita, Colorado Division of Wildlife. ABSTRACT Ducks were trapped in modified Salt Plains bait traps and banded at 1 wetland location near Grand Junction in western Colorado in August and September 1998 and at 4 locations during the same period near Cortez. Seven hundred and sixty-six mallards (Anas platyrhnchos) were banded; 343 near Grand Junction and 425 near Cortez. r[~[11 BDOW013538 ��9 PRESEASON MONITOR BANDING OF MALLARDS IN COLORADO Michael R Szymczak INTRODUCTION In 1990, the Pacific Flyway Study Committee formulated a 5-year cooperative mallard and northern pintail (Anas acuta) preseason banding program that was endorsed by the Pacific Flyway Council. This program was designed to address banding needs throughout the western U. S., including Alaska, and in the provinces of British Columbia and Alberta. Through the first 5 years, about 9,000 ducks were banded in Colorado under this program. Following the 5th year of banding, an analysis of recoveries of all mallards banded during the 5-year period in the western U. S. and Canada showed that cohorts of mallards banded in southeastern Idaho, western Wyoming, northern Utah, and western Colorado had similar recovery distribution properties. Since trapping and banding efforts in the 4-state region had been most successful in southeast Idaho and western Colorado, those 2 areas were selected for continued banding in relation to the Pacific Flyway Council efforts to establish a western mallard management unit. Banding activities in 1998 marked the third year of monitor banding activities. P. N. Objective Band mallards in Colorado's portion of Banding Reference Areas in the Pacific Flyway that will contribute information on harvest rates, survival rates, and distribution of harvest for use in Adaptive Harvest Management of western mallard populations. SEGMENT OBJECTIVES 1. Trap and band mallards in the Grand JunctionlDeltalOlathe and Cortez area of western Colorado in late August-early September using salt plains bait traps (Szymczak and Corey 1976). Banded ducks will be classified according to age and sex using accepted techniques (Carney 1964, Weller 1976: 35). Banding schedules and recapture reports will be submitted to the U. S. Fish and Wildlife Services' Bird Banding Laboratory. Band return reports will be summarized and remain on file with the Colorado Division of Wildlife. METHODS Trap Area Selection The selection of wetland locations for continued banding were based on number of birds banded per unit of effort during 1991 through 1995, and on the availability of personnel to operate the banding stations. The Walker State Wildlife Area (SWA) near Grand Junction, Markley's Pond near Olathe, and wetlands near Cortez were selected sites. �10 Trapping Ducks were trapped and banded near Grand Junction from 26 August through 3 September 1998 and in the Cortez area from 29 August through 8 September 1998. All birds were trapped in modified Salt Plains bait traps (Szymczak and Corey 1976) using whole shelled com for bait. Traps were visited daily. Mallards were the target species. Banded birds were recorded by wetland site. Band numbers of all birds captured that were banded in previous years or outside the specific area of trapping were recorded. RESULTS Trapping, Banding and Record Keeping Trapping occurred only on the Walker SWA near Grand Junction and 4 wetlands near Cortez (Table 1). A trapping crew was not assembled to trap in the Uncomphragre River Valley. Trap sites At Walker were in backwater areas in side channels of the Colorado River. Three small perenial ponds were selected for trapping in the Cortez area along with a bay on Totten Reservoir. All Cortez sites were trapped during the early 1990's. A total of 766 mallards was banded during trapping in western Colorado in 1998 (Table 1). The addition of the Cortez banding sites doubled the previous years total when birds were banded only on the Walker SWA. Immatures comprised 65 % of the sample. Females comprised only 25% of the adult sample. The number of birds trapped annually for the past 3 years have been consistent (Table 2) Band Reporting and Record Keeping All band numbers of newly banded birds and recaptures were submitted to the U. S. Fish and Wildlife Service's Bird Banding Laboratory on standard forms. Computer files containing the number of birds banded by area, site, day, age and sex were constructed at the Colorado Division of Wildlife's Research Center. LITERATURE CITED Carney, S. M. 1964. Preliminary keys to waterfowl age and sex identification by means of wing plumage. U. S. Dep. Inter., Fish and Wildl. ServoSpec. Sci. Rep.- Wildl. 82. 47pp. Szymczak, M.R., and 1. F. Corey. 1976. Construction and use of the Salt Plains duck trap in . Colorado. Colorado. Div. Wildl., Div. Rep. 6. 13pp. Weller, M. W. 1976. Molts and plumages of waterfowl. Pages 34-38 in F. C. Bellrose. Ducks, geese and swans of North America Stackpole Books, Harrisburg, PA. Prepared by: ~ '£ ~ MiChaclRSZYlnCZ3k General Professional V �11 Table 1. Number of mallards banded by age and sex during pre-season trapping in western Colorado, 1998. IF Area Site Colorado River WaikerSWA 92 127 33 91 343 Cortez Totten Res. 46 43 8 37 134 Nolan's Pd. 21 59 11 29 120 Merritt's Pd. 21 53 3 18 95 Williamson's Pd. 22 20 13 21 76 110 175 35 105 425 202 302 68 196 766 Subtotal Grand Total AM Age/sex 1M AF Totals Table 2. Number of mallards banded by age and sex during pre-season trapping at the Walker SWA in western Colorado 1996-98 Site Year AM Age/sex 1M AF IF Walker SWA 1996 68 143 18 133 362 1997 93 120 40 116 369 1998 92 127 33 91 343 Totals �14 �15 Colorado Division of Wildlife Wildlife Research Report April 1999 JOB PROGRESS REPORT State of: ____::C~o~lo~r.::;ad!:!:o~---,- _ Project: .....:W_,_....• 1;..:::6..:::,6-...••R.::...._ _ Migratory Bird Investigations Work Plan: _lQ_: Job _1_ Job Title: ~C~oo~p~er~a~ti~ve~~~an~a~ge~m~e=n~t~P~ro~w~ams~ Period Covered: Authors: _ 01 January through 31 December 1998 . James H. Gammonley and Michael R Szymczak Personnel: Matthew Reddy, Robert Sanders and Mike Shannon, Ducks Unlimited, Inc.; William Noonan, U. S. Fish and Wildlife Service; and Clait E. Braun, Alex Chappell, James H. Gammonley, arid Michael R Szymczak, and Jeff Yost, Colorado Division of Wildlife. ABSTRACT Grant applications were prepared, submitted and grants received from Great Outdoors Colorado, the North American Wetland Conservation Council, and the Playa Lakes Joint Venture Management Board. Wetland projects were reviewed and selected for funding for the Wetland Initiative. Ten wetland projects were selected for funding with Colorado Duck Stamp and Ducks Unlimited, Inc. MARSH money. Work with the Wetland Focus Area committees continued in relation to wetland project development proposals. Responsibilities as Colorado's representative on Pacific Flyway Study Committee and Council and Central Flyway Technical Committee, were fulfilled. Flyway responsibilities included providing support for cooperatively funded flyway studies. Recommendations for wetland habitat improvements andlor management were provided for public and private land managers throughout Colorado. Presentations on wetland ecology and waterfowl identification were given at workshops. No post-breeding Canada goose (Branta canadensis) banding was conducted during this segment �22 expertise in wetland project planning. The resources provided by project personnel will insure that money raised through the Colorado Duck Stamp program or any other funding initiative will be spent in accordance with the objectives of the program. Continued participation on Pacific and Central Flyway committees ensures that Colorado will remain informed on migratory bird matters, have input into migratory.bird hunting regulations, and influence habitat programs affecting migratory game birds. Increasingly, studies on migratory birds that have flyway-wide implications are being funded cooperatively through the Flyway councils. Conducting and/or formulating surveys and banding efforts and informing management agency personnel about aspects of waterfowl and wetland ecology provides a valuable service to management agencies, the waterfowl resource and, in some cases, the hunting public. Prepared by: 7Jl;.J<,..f p. ~ Michael R Szymczak James H. Gammonley LSSRN LSSRID �25 COlorado Division of Wildlife Wildlife Research Report April 1999 JOB PROGRESS REPORT Smteof =C=o=lo=ra=d=o~ Project: W.....,__-.:...;16:::..:6 •... -R ••.••.... Work Plan: _]Q_: Job Title: _ _ Migratory Game Bird Investigations Job _1_ Spring Stopover Food Resources and Land Use Patterns of Rocky Mountain Population Sandhill Cranes in the San Luis Valley, Colorado Period Covered: 01 January through 31 December 1998 Author: James H. Gammonley Personnel: James H. Gammonley and Michael R Szymczak, Colorado Division of Wildlife, and Murray K. Laubhan, USGS Biological Resources Division ABSTRACT In response to recent concerns over whether current or projected supplies of waste grain in the San Luis Valley (SL V) are adequate to meet the spring nutritional demands of Rocky Mountain Population (RMP) sandhill cranes (Grus canadensis tabida), we initiated a study to determine food habits, body condition, and distributional patterns ofRMP sandhill cr;:mes during their spring stopover in the SLY, and the amount and distribution of waste grain availble during the spring stopover period. In 1998, we produced a base GIS map of the SLY and over-layed locations of grain fields (n = 944) and major crane roosting and loafing areas. We conducted a mail survey of grain producers to determine postharvest practices used that may influence waste grain availability. We sampled 20 grain fields representing the range of post-harvest practices and estimated the amount of waste grain in each field (range = 0-33 kg/ha). We also established a road survey route to monitor the numbers and distribution of cranes weekly in the SL V, and collected diurnal time-activity budget data on cranes in fields, pastures, and wetlands. This study will continue through 2001. �26 �27 SPRING STOPOVER FOOD RESOURCES AND LAND USE PATIERNS ROCKY MOUNTAIN POPULATION SANDHILL CRANES IN THE SAN LUIS VALLEY, COLORADO OF James H. Gammonley INTRODUCTION Virtually the entire population of Rocky Mountain Population (RMP) of greater sandhill cranes (Grus canadensis tabida) uses the San Luis Valley (SLY) of Colorado as a spring stopover area RMP cranes in the SLV forage on unharvested grain provided on Monte Vista National Wildlife Refuge, and on waste grain in privately-owned fields. In recent years, however, fall tillage and irrigation of grain fields has become increasingly widespread in the SLV. These changes in farming practices have resulted in an unmeasured reduction in waste grain availability for RMP cranes during spring and have prompted concem over whether current or projected food supplies are adequate to meet spring nutritional demands of a target population size of 18,000-20,000. Changes have also occurred in the availability, distribution, and quality of wetland habitats in the SLV. PROGRAM NARRATIVE OBJECTIVES 1. Determine food habits ofRMP cranes on grain fields, pasturelands, and wetlands in the SLY during the spring stopover period. 2. Determine the body condition of RMP cranes in the SLV during the spring stopover period. 3. Determine the amount and distribution of waste grain available to RMP sandhill cranes during the spring stopover period. 4. Estimate the amounts of each major food item (particularly small grains) consumed by RMP cranes required to meet the cumulative energy requirements of a population of 18,000-20,000 during the spring stopover period in the SLV, and compare these estimates to the estimates of abundance of each food item 1) during the study and 2) under projected trends in farming practices and available public land management practices. SEGMENT OBJECTIVES 1. Determine food habits of Rocky Mountain Population (RMP) sandhill cranes on grain fields, pasturelands, and wetlands in the San Luis Valley (SLV) during the spring stopover period. 2. Analyze the body condition ofRMP cranes in the SLV at arrival and immediately prior to departure in spring. 3. Determine the amount and distribution of grains (barley, wheat) used by RMP cranes upon arrival and after departure of cranes in spring. �28 4. Estimate the amounts of each major food item (particularly cultivated grains) consumed by RMP cranes required to meet the cumulative energy requirements of a population of 18,000-20,000 during the spring stopover period in the SLY. 5. Prepare annual reports. RESULTS In 1998, we produced a GIS· map of agricultural fields, crane roosting and loafing areas, and other habitats in the SLY. Of the 2,068 center-pivot irrigated fields in the SLY, 944 (46%) were planted to grain in 1997. We conducted a mail survey of grain producers (response rate = 47%) to determine post-harvest practices and soil characteristics of grain fields. Preliminary analysis indicates untilled fields were rare, and approximately 50% of respondents indicated fields were irrigated after harvest in 1997. We sampled 20 grain fields that represented the range of post-harvest practices used on private lands. Estimates of waste grain available on the soil surface of individual fields ranged from 0 to 33 kg/ha, Based on survey responses and estimates of waste grain in individual fields, post-harvest treatment/soil type combinations will be categorized into high, medium, and low categories of waste grain reduction, with corresponding estimates waste grain abundance (kg/ha) assigned to each category. This will enable mapping the abundance and distribution of waste grain across the entire SLY. We also established a road survey route to monitor crane numbers and distribution. Based on weekly count totals, the survey route accounted for >90% of the RMP cranes present in the SL V. We collected time-activity budget data on cranes in fields, unflooded pastures, and wetlands throughout the stopover period. Preliminary analysis indicates that cranes spent 70->90% of diunal hours foraging in fields and pastures. PLANS FOR 1999 We will continue field sampling, weekly surveys, and time budgets in 1999. We also secured state and federal collection permits and will begin collections of cranes for food habits and body condition information in 1999. . . Prepared by: ~ James H. Gammonley, GP4 �29 Colorado Division of Wildlife Wildlife Research Report April 1999 INTERIM FINAL REPORT State of: -""C""'o..."lo<!,;ra""'d::;,:o<-_ Project .....:W~-1~6:..::::6....!-R;.!:..._ _ Migratory Game Bird Investigations (Avian Research) Work Plan: __]J_: Job _1_ Job Title: __ ~Inc:.!t~egr=at::::e~d.....:W~at~e~rb~ir~d~M~an~a~g~e~m~e~n:.!:..t .!:!S~tu~d~ie~s~ __ Period Covered: 0 1 January through 31 December 1998 Author: James H. Gammonley Personnel: James H. Gammonley and Michael R Szymczak, Colorado Division of Wildlife; Murray K. Laubhan, USGS Biological Resources Division ABSTRACf Laboratory analyses of waterbird body condition, and data analyses, were continued. We completed . analyses of selection among cover types by eight species of foraging waterbirds. We used line-transect distance sampling to determine the abundance of each species in each cover type. We used water depth measurements in random plots Within each cover type in combination with a GIS-based contour map of the study area to estimate the area of each cover types that was flooded to available foraging depths for each species. Each species selected from among available cover types, and all species differed in habitat selection patterns. Manuscript preparation continued, and a final report for the project will be completed in 1999. �30 �31 INTEGRATED WATERBIRD MANAGEMENT STUDIES James H. Gammonley PROGRAM NARRATIVE OBJECTIVES 1. Map the location of wetlands and wetland communities on Russell Lakes State Wildlife Area 2. Document the hydrologic regime and water, soil and vegetation characteristics of each wetland type. 3. Identify the aquatic invertebrates associated with each wetland community, and document seasonal trends in invertebrate diversity, abundance and biomass. 4. Quantify the abundance, spatial and temporal use patterns, behaviors, and food habits of waterbirds in different wetland types. Relate the dynamics of endogenous lipid and protein reserves to food habits and migration and breeding ecology. 5. Determine the seasonal wetland habitat requirements for waterbirds, and consolidate these needs into a conceptual design for an optimum wetland community. 6. Determine the water management protocol and wetland development guidelines needed to produce the optimum wetland community. Prepare a wetland development and water management plan for Russell Lakes State Wildlife Area SEGMENT OBJECTIVES 1. Use data collected at Russell Lakes State Wildlife Area (RLSW A) during 1994-96 to develop predictive models for estimating the quality of foraging and nesting habitats used by 6 species of breeding waterbirds, and test the application of these models to habitat management at RLSW A and to habitat features used by waterbirds at other wetland sites in Colorado, as follows: a Determine food selection of mallards, redheads, cinnamon teal, American avocets, killdeer, and Wilson's phalaropes, i.e., compare food use (gut ·Contents)to availability (invetebrate mid 'seed biomass at collection sites) for each species, with age, sex, reproductive status, niolt intensity, size of nutrient reserves, and collection date as co-variables. b. Analyze time-activity budget and abundance data collected during 1994-96 for waterbird species in l a, Determine foraging habitat selection, and the influence of habitat features (cover type, water depth, vegetation structure) on food availability and use offoraging habitat. Model habitat management options that maximize foraging habitat for each species and optimize foraging habitat for all species. c. Analyze nest habitat and nest success data collected during 1995-96 for each waterbird species listed in Ia Determine nest habitat selection, and the influence of habitat features (cover type, water depth, vegetation structure) and spatial attributes (distance to water; distance to roads, �32 ditches, etc.; cover type patch size) on nest success. Model habitat management options that maximize nesting habitat for each species and optimize nesting habitat for all species. 2. d. Manipulate water and vegetation on selected sites at RLSWA in 1998 and 1999. Following habitat manipulations, measure a) nest locations and nest success, b) locations used by foraging waterbirds, and c) invertebrate and seed biomass to see if patterns are as predicted in 1b and Id. e. Develop a draft long-term management plan for RLSW A that includes monitoring protocols, and use results from la-ld to prepare manuscripts for submission to peer-reviewed publications. Prepare final report. RESULTS Waterbird Carcass Analyses We completed dissections and continued proximate analyses of waterbird carcasses collected in 1994-96. Dissections and nutrient analyses have been completed on 245 of the 272 waterbirds collected. Summary results of proximate analyses are presented in Table 1. Food Selection Preliminary results of food habits and food availability have been presented in previous reports (Gammonley 1995, 1996). Because age, sex, reproductive status, molt intensity, and size of nutrient reserves will be used as co-variables in the analyses of food selection by each species, final analyses will not be completed until all dissections and nutrient analyses are completed. Foraging Habitat Selection The spatial distribution of habitats, including levees and water transfer ditches, was mapped by interpreting 1:4000 aerial color photographs and incorporated irito geographic information system (GIS). We used the GIS to select a stratified-random sample of 303. 0.25-ha2 permanent plots in seasonal wetlands with no emergent vegetation (SW), short emergent (8E), semipermanent and permanent wetland with no emergent vegetation (SPOW), tall emergent (TE), inland saltgrass (SG), and upland shrub (US) habitats. We measured water depth (± 1 em) in each plot four times annually during 1995 and 1996 (Table 2).' Sample periods (18 April-02 May, 09-23 May, 30 May-13 June, and 20 June-04 July) were selected to overlap the prenesting and nesting periods of most breeding waterbirds at RLSW A. We estimated the area of each habitat type flooded to different depths during each sample period by intersecting the habitat coverage with a hydrologic grid (5-m cell size) generated using TOPOGRID (ESRI 1996). Elevated roads and levees that restricted movement of surface water were input as contour data to generate a generalized surface morphology ofRLSW A. The mean water depth in each 0.25-ha plot, calculated for each sample period, was assigned to the center of the plot and used as point data Water depths in areas between plots were interpolated using TOPOGRID, which is based on the ANUDEM program (Hutchinson 1988). a �33 For species included in the analysis, we used the water depth maps generated byTOPOGRID to determine the area of each habitat available for foraging during each sample period. We defined available water depths as follows based on observed use of foraging depths: killdeer, >0-5 em; American avocet (hereafter avocet) and white-faced ibis (hereafter ibis), >0-30 em; cinnamon teal (hereafter teal), gadwall, mallard, and Wilson's phalarope (hereafter phalarope), >0 em; and redhead, >5 em. We assumed all habitat types in appropriate water depths were accessible to ducks and ibis, but we did not consider TE habitats available to shorebirds because we did not observe shorebirds using TE during the study. We used distance sampling (Buckland et al. 1993) and program DISTANCE (Laake et al. 1994) to estimate the abundance of waterbird species within each habitat type. We randomly placed a set of twelve parallel line-transects (range = 0.66-2.4 Ian, total length = 28.5 km) spaced at 400-m intervals across the study area Surveys were conducted 8 times/year, on the beginning and ending dates of each sampling period. Survey times alternated between morning (0700-1000 hrs) and afternoon (15001800 hrs), when foraging was the dominant activity of species included in our analysis. During each survey, observers walked and recorded the perpendicular distance, habitat type, and number in each group (cluster size) of each waterbird species visible from the transect line. We analyzed each species and habitat type separately to account for differences in sightability, and pooled data for each species-habitat combination across counts to facilitate model development and selection. To facilitate modeling the detection function, we truncated 5% of the objects detected at the longest distances in each habitat prior to analysis (Buckland et al. 1993). For each species-habitat .. combination, we examined histograms of the data and used Akaike's Information Criterion (AlC) to select the most parsimonious model (Buckland et al. 1993). We estimated the abundance of each species for each count based on the amount (ha) of each habitat sampled (determined by GIS) and the appropriate model. For each species, we used univariate Analysis of Variance (ANOVA) tests to determine if densities during the pre-nesting and nesting periods differed (pROC GLM, SAS Inst. Inc. 1988). We assigned counts conducted before and after the date at which 25% of nests were initiated each year to the pre-nesting and nesting period, respectively, for all species except phalaropes and ibis. We searched for nests on 219 ha (12.6%) of the study area, including permanent plots and four leveed areas (range = 66-147 ha) known to support high nest densities. Nest searches were conducted three times (09-23 May, 30 May-13 June, and 20 June-04 July) annually and required 7-10 days to complete. Areas were searched on foot by two or more observers spaced 10-m apart. For duck nests, we used number of eggs laid and stage of incubation (Weller 1956) to establish nest initiation date by back-dating. We monitored avocet and killdeer nests daily and determined initiation dates of successful nests (>1 egg hatched) .by subtracting the number of eggs laid and average incubation time (28 days) from the hatch date. Because only 4. successful phalarope nests were located, we based nest initiation dates on the reproductive status of 25 females collected for nutritional ecology information. For ibis, we used nesting colony observations and long-term monitoring data (Schreur 1987, Ron Ryder, unpubl. data) to determine nest initiation dates. We used compositional analysis to determine habitat selection by waterbirds because habitat proportions were not independent (Aebischer et al. 1993). We constructed log-ratios for each species from each count by dividing the proportional use and availability of each habitat by the proportional use and availability of SW and transforming the resulting ratios to logarithms. We assigned habitat availability calculated during each sample period to the counts conducted on the first and last day of the sample period. We replaced zero values for use with 0.001 (a value less than the least nonzero proportions) for compositional analysis (Aebischer et al. 1993). We used multivariate analysis of variance (MANOV A) on the log-ratios to determine ifhabitat use varied from availability (pROC ANOVA, SAS Inst. Inc, 1988). We conducted an initial test to determine if habitat selection differed �34 between pre-nesting and nesting periods for each species. If a difference was detected, we analyzed counts for each period separately; otherwise, we pooled all counts for analysis. All tests were considered significant atP < 0.05 using the Wilk's lambda criterion (Johnson and Wichern 1988). Following a significant MANOVA (i.e., nonrandom use), we assigned ranks to each habitat type and used t tests to determine significant differences among ranks for each species (Aebischer et al. 1993). Of 27 species of nonpasserine, wetland-dependent birds breeding at RLSW A during the study, we obtained adequate data from line-transect sampling to estimate abundance and determine habitat selection for 8 species, including 3 shorebirds (avocet, killdeer, and phalarope), 4 ducks (teal, gadwall, mallard, and redhead), and 1 wading bird (ibis). Mean densities of teal, mallard and ibis were greater during the pre-nesting period than during the nesting period, whereas killdeer densities were greater during the nesting period than during the pre-nesting period (fable 3). Mean densities of other species did not differ between periods. The amount (ha) of available foraging habitat remained relatively stable within and among years. During both years of the study, total available foraging habitat ranged from 228-320 ha for killdeer, 362-440 ha for phalaropes, 461-541 ha for redheads, 490-563 ha for avocets, 610-692 ha for ibis, and 775-867 ha for teal, gadwall, and mallards. Due to differences among species in water depths used for foraging and differences among habitats in flooding patterns, proportionate habitat availabilty differed among species. Availability of SE exceeded that of any other habitat for all species except redheads (Table 4). SW and SG each comprised <10% of available foraging habitat for all species (fable 4). All species used habitats in a nonrandom manner (all Ps < 0.05). Foraging habitat use differed between the prenesting and nesting periods only for teal (F= 3.37; 5, 10 df; P = 0.048) and redheads (F = 7.76; 3, 12 elf; P = 0.004). Prenesting teal used SPOW more and SE less than nesting teal. An insufficient number of surveys (n = 4) during the prenesting period precluded determining differences in use of specific habitats between periods for redheads. Overall use of available habitats differed among all species (F= 7.65; 45,514 df; P = 0.001), but there interspecific similarities in patterns of habitat selection (fable 3). SW was among the most preferred habitat, and US among the least preferred habitat of all species. SW, SE, and TE were equally preferred by mallards; SW and TE were equally preferred by prenesting teal; and SW and SE were equally preferred by ibis. SPOW was selected over all other habitats except SW by redheads, and SPOW and SE were selected equally by gadwall and prenesting teal. Shorebirds and nesting teal selected SE over all other habitats except SW. SG was among the least preferred habitats of all species except killdeer, which used SG in similar proportions to SE. We continued entering and checking time-activity budget data During 1994-96, over 1,550 individual, 1O-min· time-activity bouts were collected on the 7 waterbird species we studied at RLSW A. We recorded habitat variables (water depth, conductivity, vegetation structure, cover type) at 338 sites where Collected time-activity data on birds that were foraging. This habitat data has been entered in a database and will be merged with the time-activity data when that database is completed. Habitat availability data from 330 random plots on RLSWA has been entered on computer files and checked, and a habitat map of the area has been completed (Gammonley 1995, 1996, 1997). Analysis of habitat selection by foraging waterbirds we Nest Habitat Selection and Nest Success Data on nest site checked for accuracy. software (ARC-INFO) cover types, distances data habitat attributes and success of nests have been entered into computer files and Preliminary attempts have been made to use geographical information system to calculate spatial attributes of individual nest locations (e.g., distance to other to other nests). No additional work was done in 1998 on analyses of nesting �35 Habitat Manipulations at RLSW A The Colorado Division of Wildlife began work on a management plan for RLSW A. We provided the GIS map showing the distribution of habitats, ditches, roads and levees. We also provided descriptions of habitats and recommendations for additional habitat developments. These recommendations have been incorporated into a draft management plan outline (Appendix). PLANS FOR 1999 Laboratory analyses of waterbird carcasses will be completed. Statistical analyses of all data sets will continue. The final report for this study will be completed. Manuscripts will be prepared for publication in peer-reviewed technical journals. Results will be incorporated into a revised management plan for RLSW A, as well as management plans for other SWAs in the San Luis Valley. LITERATURE CITED Aebischer, N. J., P. A. Robertson, and R E. Kenward. 1993. Compositional analysis of habitat use from animal radio-tracking data. Ecology 74:1313-1325. Buckland, S. T., D.R Anderson, K P. Burnham, and J. L. Laake. 1993. Distance sampling: estimating abundance of biological populations. Chapman and Hall, London, England. ESRI. 1996. Arc!Info and Arc/Grid, version 7.0.1. Redlands, California Gammonley, J. H. 1995. Integrated waterbird management studies. Colorado Division of Wildlife Federal Aid Progress Report. Gammonley, J. H 1996. Integrated waterbird management studies. Colorado Division of Wildlife Federal Aid Progress Report. Gammonley, J. H. 1997. Integrated waterbird management studies. Colorado Division of Wildlife Federal Aid Progress Report. Hutchinson, M F. 1988. Calculation of hydrologically sound digital elevation models. Third International Symposium on Spatial Data Handling. International Geographical Union, Columbus, Ohio. Laake, J. L., S. T. Buckland, D. R Anderson, and K. P. Burnham. 1994. DISTANCE user's guide, version 2.1. Colorado Cooperative Fish and Wildlife Research Unit, Colorado State University, Fort Collins, Colorado. SAS Institute, Inc. 1988. SAS/STAT User's guide, release 6.03 edition. SAS Institute Inc., Cary, North Carolina Schreur, J. L. 1987. Ciconiiform reproductive success and public viewing in the San Luis Valley, Colorado. Thesis, Colorado State University, Fort Collins, Colorado. Weller, M. W. 1956. A simple field candler for waterfowl eggs. Journal of Wildlife Management 20:111-113. PreparedbY:~ James H. Gammonley, GP4 �36 Table 1. Mean (SE) proportions of water, lipid, protein, and mineral in carcasses of waterbirds collected at RLSW A, 1994-96. Component Species Water Lipid Protein Mineral Avocet 0.664 (0.005) 0.064 (0.009) 0.220 (0.005) 0.049 (0.004) Teal 0.684 (0.010) 0.066 (0.010) 0.211 (0.008) 0.037 (0.003) Killdeer 0.643 (0.008) 0.072 (0.009) 0.231 (0.007) 0.055 (0.003) Mallard 0.710 (0.011) 0.036 (0.012) 0.211 (0.010) 0.033 (0.005) Phalarope 0.624 (0.005) 0.092 (0.007) 0.228 (0.005) 0.056 (0.004) Table 2. Sampling effort within each habitat on RLSWA used to determine water depths (0.25-ha permanent plots) and animal abundance (line transects) during each sample period. Area Plots b Transects Habitat 8 C 72 (4.1) 13 (25) 9 (1.5) Short emergent (SE) 628 (36.1) 49 (55) 12 (7.9) Semipermanent open water (SPOW) 231 (13.3) 30 (25) 10 (2.9) Tall emergent (TE) 223 (12.8) 21 (20) 11 (3.9) 74 (4.3) 62 (65) 9 (1.8) 511 (29.4) 128 (50) 10 (10.5) 303 (14,620) 12 (28.5) Seasonal open water (SW) Saltgrass (SG) Upland shrub (US) Total 1,739 • Hectares (percent of total area). b Number of plots (water depth measurements/plot). c Number of transects (combined length [km] of transect segments) that intersected habitat type. Table 3. Nest initiation dates and comparison of mean (± SE) densities between pre-nesting and nesting Eeriods for 8 common waterbird sEecies at RLSWA, 1995 and 1996. Nest initiation Density 1996 Pre-nesting nb 1 Jun 16 May 0.17(0.02) 8· 0.14(0.02) . Teal 16 May 10 May L39(0.21) 6 0.51(0.06) Gadwall 30 May 24 May 0.37(0.05) 9 0.28(0.03) Killdeer 31 May 7 May 0.07(0.01) 7 Mallard 2 May 25 Apr 0.87(0.18) Redhead 5 May 2 May 21 May 1 Jun Species, Avocet Ibis Phalarope 1995 Nesting nb F P 1.65 0.22 10 . . 24.43 0.0002 7 1.84 0.20 0.19(0.03) 9 9.63 0.008 3 0.38(0.10) 13 4.72 0.04 0.16(0.03) 4 0.13(0.01) 12 1.39 0.26 21 May 0.65(0.17) 6 0.25(0.06) 10 6.70 0.02 1 Jun 0.16(0.04) 10 0.14(0.02) 6 0.11 0.74 "See methods for procedures used to determine nest initiation dates. b Number of counts. 8 �37 Table 4. Mean (± SE) proportions of habitat use and availability, and habitat preference ranks for 8 common waterbird species at RLSWA, 1995 and 1996. Habitats ranks are in ascending order (e.g., 1 is most preferred). For each species, preference ranks with the same capital letter are not significantly (P> 0.05) different. NA indicates that the habitat was considered not available «25 ha of the habitat at appropriate water depths was Eresent or that no use occurred during the study). Seasonal open water Species Use Available Short emergent Rank Use Semipermanent open water Available Rank Use Available Rank Avocet 53.5 ± 5.3 6.1±0.1 lA 35.1 ± 5.7 41.3 ± 0.3 2B 7.3 ± 3.5 10.9 ± 0.1 3C Killdeer 45.2 ± 4.5 6.1 ± 0.1 lA 34.2 ± 5.9 40.2 ± 0.3 3B 0.0 ± 0.0 7.5 ± 0.1 4C Phalarope 53.0 ± 7.1 7.1 ± 0.0 lA 31.6±6.9 41.3 ± 0.2 2B 14.0 ± 7.1 10.9 ± 0.1 4CO Teal (pre-nesting) 28.5 ± 4.1 4.8 ± 0.1 lA 38.4 ± 6.0 28.2± o.s 2B 23.9 ± 6.3 21.5 ± 0.2 3B Teal (nesting) 22J ± 4.3 4.9 ± 0.1 lA 56.1 ± 3.6 27.5 ± 0.3 2B 10.1 ± 1.4 21.8 ± 0.2 3C Gadwall 15.1 ± 1.1 4.9 ±O.O lA 41.2 ± 3.5 27.7 ± 0.2 2B 28.9 ± 3.7 21.7±0.1 3B Mallard 14.1 ± 2.1 4.9 ±O.O 2A 57.5 ± 5.1 27.7± 0.2 lA 8.1 ± 1.5 21.7 ± 0.1 4B Redhead (nesting) 20.5 ± 3.3 4.8 ± 0.1 lA 12.6 ± 5.2 23.2 ± 0.7 3C 56.5 ± 6.2 31.8±0.3 2B Ibis 11.0 ± 2.2 5.7·± 0.1 2AB 70.6± 6.2 33.0 ± 0.2 lA 3.6 ± 1.7 Saltgrass Tall emergent Species Use Available Rank Use Available 8.7 ± 0.1 4C Upland shrub Rank Use Available Rank Avocet NA 4.1 ± 1.8 6.1 ± 0.1 4B 0.0 ± 0.0 34.6 ± 0.3 5C Killdeer NA 18.7 ± 2.7 5.8 ± 0.3 2B 1.8 ± 1.8 40.4 ± 0.3 4C Phalarope NA 1.5±0.1 6.1 ± 0.1 4C 0.0 ± 0.0 34.6 ± 0.3 50 Teal (pre-nesting) 9.2 ± 2.5 18.9 ± 0.0 4AB O.O±O.O 4.2± 0.2 5B 0.0 ± 0.0 22.4 ± 0.3 6C Teal (nesting) 9.7 ± 2.4 19.1 ± 0.2 4C 0.1 ± 0.1 4.0 ± 0.1 5C 1.7 ± 0.0 22.6 ± 0.3 60 Gadwall 6.3 ± 1.4 19.0 ± 0.1 4C 0.7 ± OJ 4.1 ± 0.1 .6C 7.8 ± 2.1 26.8 ± 0.2 5C Mallard 17.6 ± 2.5 19.0 ± 0.1 4AB 0.5 ± 0.4 4.1 ± 0.1 6C 2.2 ± 0.8 22.5 ± 0.2 5C Redhead (nesting) 10.3 ± 4.1 21.6 ± 0.2 4C O.O±O.O 3.5 ± 0.1 5C 0.0 ± 0.0 15.0 ± 0.4 6D 14.5 ± 4.2 20.0 ± 0.1 3B 0.1 ± 0.0 4.9 ± 0.1 0.3 ± OJ 27.7 ± 0.2 60 Ibis 5C �38 Appendix. Draft management plan outline for RLSW A. Introduction: Historically, the primary purpose of this State Wildlife Area has been to support breeding waterbird populations. This includes a variety of species, each with unique habitat requirements. Most common breeding waterbirds include pied-billed grebe, Western grebe, Clark's grebe, American bittern, snowy egret, black-crowned night heron, white-faced ibis, Canada goose, mallard, gadwall, cinnamon teal, redhead, ruddy duck, American avocet, killdeer, spotted sandpiper, common snipe, Wilson's phalarope, Virginia rail, sora, American coot, northern harrier, marsh wren, yellow-headed blackbird, and red-winged blackbird. The importance of the area as a migration stopover, particularly during spring, is also an important function of the area; at least 25 other waterbird species use RLSW A primarily during periods other than breeding. This will be another main component of the management of the property. Our goal is to maintain the benefits of wetland habitats on RLSW A for breeding and migrating waterbirds over the long term. In order to accomplish this goal, habitats will require periodic disturbance, primarily in the form of water management and mechanical manipulation of vegetation. Our objective is to provide a diversity of habitat types across the area, each at the appropriate time for the waterbirds that benefit from the conditions. Management of the property can be viewed in terms of three main categories. These categories reflect the level of management approach, and are as follows: 1. Un-managed wetlands complexes 2. Lake complex 3. Management impoundments The first segment includes wetlands that are not actively managed. This segment includes natural "playa" basins in the area between Trites, Davey, and Harrence lakes. These basins are primarily influenced by groundwater fluctuations; they are influenced by water management elsewhere on RLSW A and surrounding lands, but these relationships are complex and would be difficult to manage. These areas will be monitored to ensure that other management activities does not significantly alter these important wetland types. This segment also includes the historic Russell Creek drainage; once water enters this drainage, there little or no ability (legally) to manage thewater. is The second segment includes the six large lakes at RLSW A. These are connected to the Russell Creek drainage, but have some management capabilities. Many of the management concerns are similar among these lakes. The third segment includes the impounded areas at RLSW A. Most of these leveed areas have some degree of water control. Also included are several sites that are not leveed, but where water can be actively managed (e.g., west of Davey Lake, saltgrass flats south of the shop). All habitat types at RLSW A are present in these impoundments. These are the portions ofRLSWA that can and should be most actively managed. Within the framework of these three broad categories, for management purposes, the property will be divided into 5 units. These units reflect the water sources and management opportunities for �39 each unit. The water sources on the property are primarily derived from wells, and as outlined in the report from Leonard Rice, 1998, each well has constraints in its usage. The three largest wells provide approximately 75% of the total yield of well water for the property. Management objectives will focus on these wells and corresponding units. These units are identified as follows: 1. ArgyUnit 2. Wetherill Unit 3. Davey Unit 4. Southwest Unit 5. DFK Unit IMPOUNDMENTS Existing impoundments are designated as 1-1 through 1-16. These impoundments are found within 4 of the 5 units. These impoundments will become the focus of active management efforts on the property. We will develop a schedule of management actions that provide a rotational approach to habitat conditions; The impoundments will be allowed to succeed into dense vegetation stands, then periodically returned to early stages of wetland conditions by burning, mowing, or other methods. Timing will be such that a diversity of wetland conditions will be available at any given point in time, and monitoring stations will provide the framework for managers to evaluate the time to act. Water level management will be the key tool in these wetland impoundments. New development of impoundments will be incorporated into the rotational scheme, so that no additional acres of wetlands or impoundments are created at any given point in time. WELLS All wells should be identified and tagged, and documented in the Russell Lakes SW A database for future reference. Most project focus should be on the large capacity wells (for re-drilling or relocation), as well as for the use of water for wetlands. Direction for management activity should be to providing seasonally flooded wetlands, in order to provide water use that best follows decreed well water rights (i.e., away from year round diversions and permanent flooding). HABITAT MANAGEMENT MANAGEMENT OUTLINE OBJECTIVES: l. ARGY UNIT: This unit involves impoundments 1-5,1-6,1-12,1-13,1-14. These areas are fed by wells W2465, W2475 1&2. Water from the Wetherill Unit also can service 1-13 and 1-14. This unit currently contains a good mixture of short and tall emergents, semipermanent open water, salt grass and uplands. A. Upgrade, repair, replace all water control structures in the Unit. All replaced structures will be with designs intended to reduce management manpower. Well W2465 (Argy) should have the "plumbing" upgraded. �40 B. Develop impoundment 1-12 to expand the wetland acres and provide additional short emergent wetland types by small dikes and improved water delivery. Currently, only the southern end of the unit is wetland habitat. Develop a new structure on the east side ofI-13. C. Develop impoundment 1-5 to expand the wetland acres and provide additional short emergent wetland types by small dikes and improved water delivery. Currently, this SE end of the unit is upland habitat. D. Evaluate the ability to by-pass impoundments in this unit, in order to dry the impoundments. E. Implement rotational schedule for the impoundments in this unit. 2. WETIIERILL UNIT: This unit involves impoundments 1-1, 1-2, 1-3, 1-4, 1-8, 1-9, 1-10, 1-11, 1-13, and 1-14. These areas are fed by wells W1632, # 1-9. Water from the Argy Unit also can service 1-13 and 1-14. This unit currently contains a good mixture of short and tall emergents, semipermanent open water, shallow water wetlands, salt grass and uplands. Surface water rights from Russell Creek may also be available for some use on parts of these units within sections 20 and 28 .. A. Upgrade, repair, replace all water control structures in the Unit. Especially structures at the junction ofI-3, 1-4, 1-8, and 1-9. All replaced structures will be with designs intended to reduce management manpower. B. Evaluate the ability to by-pass impoundments in this unit, in order to dry the impoundments. C. Evaluate the potential to develop ephemeral shallow water wetlands in sections 20 and 28. D. Evaluate the potential to utilize surface water rights from Russell Creek into existing impoundments, and/or use in new developments in section 20/28. E. Implement rotational schedule for the impoundments . '. . in this unit. . 3. DAVEY UNIT: This unit includes the 6 large lakes within the property. This unit involves impoundments 1-7 and 1-15. These areas are primarily fed by wells W2023, # 1-8. It includes mainly permanent and semipermanent open water, short emergent wetlands, and uplands. Some shallow water wetlands, tall emergents, and salt grass habitats are also within the unit. Based on recommendations from Gammonley and Cooper, we will explore opportunities to periodically drawdown the water levels in the lakes. This is intended to promote the productivity of the lakes, and stimulate more aquatic vegetation, invertebrate production, and invigorate bulrush for the benefit of waterbirds. A. Upgrade, repair, replace all water control structures in the Unit. Repair the levee and discharge structures on the pool near well W2023#1. All replaced structures will be with designs intended to reduce management manpower. �41 B. Evaluate and initiate a project proposal to either renovate, replace, or pump water from the large well W2023#1 (Davey well), to bring it up to its decreed flowrate. Currently, it is flowing at about 800 g.p.m. less than its decreed flowrate. C. Maintain the wet meadow complex (short emergent) on the west side of Davey Lake. Determine if periodic management action will be required to maintain the viability of the wetlands, and implement a schedule of activity if necessary. D. Explore the feasibility of controlling water levels at all 6 lakes, and providing the ability to lower the lake levels. The lakes will need to be independently managed for drawdown, and plans developed for what managers would do with the water from the lakes, as well as re-fill issues. E. Evaluate the potential of developing additional playa wetland habitat in the areas around Harrence Lake. F. Evaluate the potential of developing additional wet meadow (short emergent) wetland habitat in the areas around northwest Harrence Lake, north Trites~ and northwest Davey Lake. G. Evaluate the potential of releasing water from Island Lake into private lands south of the unit. Inorder to periodically dry Island Lake, it may be necessary to move water from the lake into the lands south of the lake. Frequency, timing of releases, and amounts of water, will all be factors. SOlITHWEST UNIT: This unit involves impoundment 1-16; a series of small dikes and impoundments. This area is fed by wells W2576, 1-3. This unit currently contains mainly short emergent wetlands, with some small areas of semipermanent open water, and tall emergents. A. Upgrade, repair, replace all water control structures in the Unit. All replaced structures will be with designs intended to reduce management manpower. B. Provide dike modifications to impoundment 1-16, to provide better sheet flows, protect dikes from ice and overflows, and enhance water movement throughout the impoundment. C. Evaluate the ability to by-pass individual impoundments productivity. D. Implement rotational schedule for the impoundments in this unit, in order to dry the wetlands for in this unit, if feasible. DFK UNIT: This unit involves no existing impoundments, but plans for development will be initiated. This area is fed by wells W1919, 1-13. Most of these wells are small producers, the largest being W1919 #1. This well is a large capacity well, with potential to provide flows to develop the unit. It is strategically located to provide water to this unit, has no specified area of use but was associated with the sections in this unit, and has a relatively senior priority compared to other wells on the SW A. Currently, it is �42 producing at about 1000 g.p.m. less than its decreed fIowrate. No vegetation mapping is currently available for this unit. A. Evaluate and continue the project proposal to either renovate, replace, or pump water from the large well W1919#1, to bring it up to its decreed flowrate. Currently, it is flowing at about 1000 g.p.m. less than its decreed flowrate. B. Plan development of ephemeral shallow water wetlands in sections 33 and 34. Contour mapping may be needed to plan for dike locations. Wetlands will be developed that are temporary, and have the ability to be dried on a rotational basis. C. Develop water control structures, as planned in B, in the Unit. All structures will be with designs intended to reduce management manpower. D. Implement rotational schedule for the impoundments in this unit. FISHERIES: A. Evaluate the potential for utilizing RLSWA for conservation needs of TIE fishes. We need to discuss what, if any , potential exists, and how it fits with wetland management objectives. B. Evaluate the potential for carp control, and implement a control plan on the property. GENERAL: A. All wells should be identified and tagged, and documented in the Russell Lakes SW A database for future reference, as suggested in the Leonard Rice report. B. All water control structures should be identified and tagged, and documented in the Russell Lakes SW A database for future reference and maintenance scheduling. C. Photographic benchmarks for management decision making will be established, and incorporated into the working MMP to aid RLSW A personnel in management activities. This will be done in the summer of 1999. �43 Colorado Division of Wildlife Wildlife Research Report April 1999 JOB PROGRESS REPORT Smteof ~C~o~lo~r~a~do~ Project: ---'Wc.:........:-1:...,::6=6--'-R=---_ Work Plan: _ Migratory Game Bird Investigations .u.. Job _L_ Job Title: __ ....::M=o!.!;ni~to~n~· n!.l:g...:an=d:..:E:..:v~al~u~a~ti~on~of~W..!.!...!:e~tl~an~d~D:.!:e:..!.v:::::el~op~m=en~t:..:P~r~o~je~cts~in~C~ Period Covered: Author: 01 January through 31 December 1998 James H. Gammonley Personnel: James H. Gammonley and Michael R Szymczak, Colorado Division of Wildlife, and Matthew A. Reddy, Duck Unlimited Inc. ABSTRACT We developed a database of wetland development proj ects in Colorado that were funded by Colorado Division of Wildlife (CDOW), Ducks Unlimited Inc. (DU), and/or Partners for Wildlife (PFW). We selected a stratified random sample of60 of these projects for monitoring of structural, biological, landscape, and management attributes during 1999-2001. In 1998, initial site visits were made to 39 of these projects. Protocols for monitoring were developed, and monitoring of all selected projects will begin in 1999. �44 �45 MONITORING AND EVALUATION OF WETLAND DEVELOPMENT PROJECTS IN COLORADO James H. Gammonley INTRODUCTION It is estimated that half of Colorado's wetlands have been destroyed in the last 100 years (Dahl and Johnson 1991). Many other wetlands in the state have been degraded or their functions have been greatly modified, particularly palustrine habitats with intermittently flooded, temporarily flooded, and seasonally flooded hydroperiods (Cowardin et al. 1979). These highly productive wetlands provide important habitats for waterfowl and other wetland-dependent wildlife (Fredrickson and Taylor 1982, Eldridge 1992, Mitsch and Gosselink 1993). . In recent years, wetland conservation efforts have increased in Colorado, led by the Colorado Division of Wildlife (CDOW), Ducks Unlimited Inc. (DU), and the Partners for Wildlife (PFW) program of the U.S. Fish and Wildlfe Service (Chappell 1997). To date, more than 200 wetland conservation projects have been completed by these organizations in Colorado. The focus of most of these projects has been to protect, restore, enhance, or create habitat for waterfowl and other wetlanddependent birds (e.g., Colorado Division of Wildlife 1989, Colorado Division of Wildlife et al. 1996). Portions of Colorado were included in the Intermountain West Joint Venture (IWJV) and Playa Lakes Joint Venture (pLJV) of the North American Waterfowl Management Plan (NAWMP). Within the IWJV, seven focus areas were established west of the continental divide where most IWJV conservation efforts in Colorado would take place (YampalWhite River [YWR], Lower Colorado River [LCR], Gunnison River [GR], North Park [NPl, Middle Park[MP], South Park [SP], and San Luis Valley [SLV]). In 1995, CDOW created three additional focus areas in eastern Colorado (playa Lakes!Arkansas River [PLAR], South Platte River [SPR], and Front Range [FR]) to facilitate wetland conservation efforts on a statewide basis. These ten focus areas collectively form the infrastructure for .... ;.. CDOW's Wetlands Initiative, a partnership between CDOW, DU, PFW, The Nature Conservancy, and the Colorado Department of Parks plan to spend over S10 million over the next three years on wetland conservation projects in Colorado (Chappell 1997). DU also has an ambitious Colorado Conservation Plan (Ducks Unlimited 1997) that focuses on conserving waterfowl habitat in the SL V, NP, and SPR focus areas. A goal of all wetland conservation projects is to provide protection from further threats to wetland ecological functions. Naturally functioning wetland systems, requiring protection only, are relatively rare in Colorado, except at the highest elevations. Rather, natural or historic functions at most existing wetland sites have been lost or degraded. Consequently, 'many conservation projects '. have included structural developments, such as levees and water control structures, that provide greater hydrological control, allowing managers to more reliably restoreand maintain historic wetland conditions (i.e., restoration projects). Usually, the broad goal of a wetland development project is to improve capabilities to emulate historic, natural wetland functions at the site, including hydrologic regimes (Fredrickson and Taylor 1982, Fredrickson and Reid 1990, Galatowitsch and Van der Valk 1996). Alternatively, developments can be used to produce wetland habitats at sites where they did not historically occur (i.e., creation projects), or to alter existing wetlands to produce greater benefits for specific wetland functions, such as providing habitat for wetland-dependent wildlife (i.e., enhancement projects) (Fredrickson and Reid 1986, Weller 1990). Many procedures have been developed to assess wetland functions (Harris 1988, Adamus and Brandt 1990, D' Avanzo 1990, Erwin 1990, Marble 1990, Kentula et al. 1992, World Wildlife Fund 1992, Brinson 1993). Most of these approaches have focused on mitigation projects, but the �52 CO-SE-6-004 Wilkinson" PLAR 4/11/96 100 CO-SE-7-002 CO-SE- 7-011 Davis * PLAR 12/18/96 10 CO-SE-6-007 Jackson" PLAR 5/30/96 51 CO-SE-5-001 Foreman" PLAR 4/11/96 8 CO-SE-6-0 12 Hinds PLAR 7/10/96 18 Wooten PLAR 1998 ? Wetland Intitiative X-YRanch* PLAR 1998 100 SE-8990-003 SE-9192-00 1 Lathrop CO-SV-7-2 CO-SV-7-4 Wesley* SLY 5/14/97 51 CO-SV -7-001 Shriver" SLY 9/19/96 35 CO-SV -6-102 CO-SV -7-003 Walters* SLY 10/13/95 105 CO-SV -4-009 Woodman* SLY 8/1/94 CO-SV-6-014 Anderson" SLY 9/17/96 250 CO-SV -6-009 Cheslock" SLY 8/1196 5 Wetland Initiative Higel SWA* SLY 1998 80 CO-SV -4-005 Chiles SWA* SLY 7/1/94 175 CO-SV -5-007 Swift SLY 1112/94 Wetland Initiative Monte Vista NWR* SLY 1998 800 CO-GO-6-0 17 Gordon* SP 1995 ? SP 1997 ? se- Wakem* 20 PLAR - 230 4 Project code Project name CO-GO-7-012 Sublette SPR 5/5/97 25 CO-GO-6-012 Wind* SPR 8/2/96 7 - DT-Ranch* SPR 7112/96 6 .CO-GO-7-007 Treadway* SPR 11112/96 31 NE-9495-003 Jackson SWA * SPR 5/1194 55 Wetland Initiative Elliott SWA* SPR 1998 200 RedLionSWA SPR 1998 ? CO-GO-6-0 15 Focus Area Initiation date .- Wetland acres CO-GO-6-0 19 Walsley YWR ? ? CO-NP-6-040 Hankin YWR 4/1/95 2 CO-GO-5-007 Price YWR 10/30/95 20 CO-GO- 7-003 Harris YWR 10/4/96 3 Wetland Initiative YampaSWA YWR 1998 ? �53 Colorado Division of Wildlife Wildlife Research Report April 1999 JOB PROGRESS REPORT State of: Colorado ------======~------------- Project W..:...:,_-.,:_16::..;7;_-=R'-- Work Plan _1_: Job Avian Research _ 24 Job Title: __ ~E:..!,;val~u~atI:!.:o· 0~n~02f~H~a~b~it~a~t D~ev.!..!e~lo~p~m!.!e~n.!.!:t..!;.fo~rwRi=·n~g_..!-n.!!:e~c~ke~d~P~h~e~as~an=ts~in~E~as=te::!. Period Covered: Author: 0 1 January through 31 December 1998 Thomas E. Remington ABSTRACT There were no research results reported during this Segment. later. A Final Report will be written �54 �55 Colorado Division ofWildIife Wildlife Research Report April 1998 .. FINAL REPORT State of: Colorado -----======-----Avian Research ----'-W'--...,:.1.=..67.;_-..:..R=-_ Project: Work Plan: __ --=3::..__ Job Title: Job 19 Implications of Habitat Loss and Fragmentation for Gunnison Sage Grouse Period Covered: Author: _ 0 1 January through 31 December on Conservation Strategies 1997 Sara J. Oyler-McCance Personnel: Sara J. Oyler-McCance, Colorado State University; Clait E. Braun, Colorado Division of Wildlife; Thomas Quinn, University of Denver ABSTRACf The newly recognized Gunnison sage grouse (Centrocercus minim us) has declined markedly with extirpations in 12 of the 17 counties in southwestern Colorado which supported them in the early 1900's. Populations that remain are small and isolated, and exist in degraded and fragmented habitats. As a result, conservation of this species has become a significant concern. Particular issues of concern involve habitat quality and quantity, and genetic isolation from other populations. I developed a habitat-based model to: (1) identify the relative importance of landscape and micro-level variables, (2) examine the suitability of any sagebrush (Artemisia spp.) Patch in southwestern Colorado, and (3) identify which patches have the highest probability of occupancy by sage grouse. The best model to make inferences from the data included patch area and distance to the nearest paved road. I quantified loss and fragmentation of sagebrush-dominated habitat using aerial photographic analysis. Between the mid-50's and the mid-90's, 20% of habitat was lost and sagebrush in 37% of the plots was fragmented. The Gunnison Basin had the lowest rate of habitat loss. I examined whether genetic data supported the new species designation of Gunnison sage grouse, and documented relative amounts of gene flow and genetic diversity between Gunnison sage grouse populations and northern sage grouse (c. uophasianus) populations from northern Colorado. My genetic data supported the species distinction, and I found that Gunnison sage grouse populations have less genetic diversity and gene flow than northern sage grouse. Incorporating data from the habitat and genetic studies, I developed a Geographic Information System (GIS) based model which consolidated current knowledge about Gunnison sage grouse so that managers could prioritize conservation strategies. ��57 DISSERTATION GENETIC AND HABITAT FACTORS UNDERLYING CONSERVATION FOR GUNNISON SAGE GROUSE Submitted by Sara 1. Oyler-McCance Department of Fishery and WIldlife Biology In partial fulfillment of the requirements for the Degree of Doctor of Philosophy Colorado State University Fort Collins, Colorado Summer 1999 STRATEGIES �58 COLORADO STATE UNIVERSTIY 25 March 1999 WE HEREBY RECOMMEND mAT THE DISSERTATION PREPARED UNDER OUR SUPERVISION BY SARA 1. OYLER-MCCANCE ENTIILED GENETIC AND HABITAT FACTORS UNDERLYING CONSERVATION STRATEGIES FOR GUNNISON SAGE GROUSE BE ACCEPTED AS FULFILLING IN PART REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PlllLOSOPHY. Committee on Graduate Work Cfi I~,I! Advisor" ~ t (1, ~---&~J~ tLUi ' ' '1 �59 ABSTRACT OF DISSERTATION GENETIC AND HABITAT FACTORS UNDERLYING CONSERVATION STRATEGIES FOR GUNNISON SAGE GROUSE The newly recognized Gunnison sage grouse (Centrocercus minim us) has declined markedly with extirpations in 12 of the 17 counties in southwestern Colorado which supported them in the early 1900's. Populations that remain are small and isolated, and exist in degraded and fragmented habitats. As a result, conservation of this species has become a significant concern. Particular issues of concern involve habitat quality and quantity, and genetic isolation from other populations. I developed a habitat-based model to: (1) identify the relative importance of landscape and micro-level variables, (2) examine the suitability of any sagebrush (Artemisia spp.) patch in southwestern Colorado, and (3) identify which patches have the highest probability of occupancy by sage grouse. The best model to make inferences from the data included patch area and distance to the nearest paved road. I quantified loss and fragmentation of sagebrush-dominated habitat using aerial photographic analysis. Between the mid-50's and the mid-90's, 20% of habitat was lost and sagebrush in 370/0 of the plots was fragmented The Gunnison Basin had the lowest rate of habitat loss. I examined whether genetic data supported the new species designation of Gunnison sage grouse, and documented relative amounts of gene flow and genetic diversity between Gunnison sage grouse populations and northern sage grouse (c. urophasianus) populations from northern Colorado. My genetic data supported the species distinction, and I found that Gunnison sage grouse populations have less genetic diversity and gene flow than northern sage grouse. Incorporating data from the habitat and genetic studies, I developed a Geographic Information System (GIS) based model which consolidated current knowledge about Gunnison sage grouse so that managers could prioritize conservation strategies. Sara 1. Oyler-McCance Department of Fishery and Wildlife Biology Colorado State University Fort Collins, CO 80523 Summer 1999 ACKNOWLEDGMENTS Funding for research in this dissertation was from Colorado Federal Aid in Wildlife Restoration Project W-167-R through the Colorado Cooperative Fish and Wildlife Research Unit. Much of this research could not have been completed without the support of landowners in near Dove Creek, Dry Creek Basin, Hamilton Mesa, Miramonte Reservoir, Crawford, and in the Gunnison Basin. I am particularly grateful to my advisor, Kenneth P. Burnham and the members of my graduate committee, Michael F. Antolin, Clait E. Braun, Dale A. Hein, and Kenneth R Wilson who all played integral roles in my research and development as a scientist. I could not have asked for an advisor better than Ken Burnham. He constantly made the time to help, gave me encouragement and support throughout the entire project, and helped me grow as a scientist. Clait Braun gave me endless support and advice, and let me find my own niche. Mike Antolin taught me much about molecular genetics and was extremely helpful in the analysis of my genetics data. Dale Hein was concerned with making sure I had a good experience as a graduate student and helping me see the big picture. I am also grateful to him for welcoming me into his classroom and helping me learn more about teaching. Ken Wilson also �60 helped me with my teaching skills, and was always willing to share his knowledge, advice, and good humor. I also am thankful to David R Anderson who offered inspiration and advice. My genetics research was conducted in the lab of Thomas W. Quinn at the University of Denver. I thank Tom for graciously welcoming me into his lab and providing me with endless training and support. He has not only been a great mentor, but also a great friend. I am also grateful to Nate W. Kahn and Nick Benedict for their help and encouragement. Judy St. John was also a great source of motivation and a great friend. The fieldwork associated with my research was greatly enhanced by the advice and camaraderie of Michelle L. Commons, Christy A. Brigham, and Christopher P. Woods. Many thanks also to Jessica R Young for her inspiration and advice, Kim M. Potter whose work made mine a lot easier, and Christian A. Hagan who allowed me to get out of the lab and into the field every once in awhile. lowe special thanks to Allison B. Shoemaker who worked endless, tedious hours with me on the aerial photography portion of this research. I also thank Barb White from the Midcontinent Ecological Sciences Center for allowing me to use their photographic interpretation equipment and Dawn Brownne for training me on that equipment. I especially thank the staff of the Colorado Cooperative Research Unit who provided me with resources, help, and good humor. Beverly A. Klein, Bridgit Williams, and Karen 1. Adleman were particularly helpful and fun to work with. I also thank the staff of the Colorado Division of Wildlife for their support. E. Lee Olton, in particular, helped me through numerous problems and logistical nightmares. My fellow graduate students at Colorado State University were a source of advice, humor, and stress relief. I particularly thank my office mates, Gail S. Olson and Craig W. McCarty for their support and friendship. I also thank Alan B. Franklin for his willingness to help and his advice. I especially thank Laura E. Ellison for her sincere friendship. She and Bill W. Iko kept reminding me that there was a life outside graduate school. My love for science and nature began at an early age though numerous camping trips, nature hikes, and science fairs. These experiences have been extremely influential in my life and career development. For that, I lovingly thank my parents, John F. and Nancy L. Oyler, who showed me the amazement of science and how awe-inspiring the natural world can be. I particularly thank my father for helping me develop an inquisitive mind and my mother teaching me that I could do whatever I aspired to. I thank my sister, Elizabeth A. Oyler, for her unconditional love and inspiration, and for her camaraderie as we both worked on our dissertations. I also thank my brother John V. Oyler who, in his own humorous way, was always supportive of me and my research on 'the sacred grouch'. lowe special thanks to my dear friend, Diane M. Owens, who has always been there for me through the most important times in my life. Her friendship is unfaltering and is truly cherished. Finally, I am eternally grateful to my superhero husband, .James L. McCance, whose enduring love and support have sustaID.ed me throughout this entire project, His wonderful attitude, unique outlook life, and great sense of humor, have inspired me and made my life much brighter .. '. ' on �61 TABLE OF CONTENTS ABSlRACT . ACKNOWLEDGMENTS . IN1R.ODUCTION Literature Cited . . CHAPTER ONE: A HABITAT-BASED MODEL TO PREDICT GUNNISON SAGE GROUSE OCCURRENCE IN SOUTHWESTERN COLORADO Introduction Study Area Methods Selection of Sites Variables Measured Data Collection Data Analysis Results , Discussion Literature Cited Tables Figures '" , . . . . . . . . . . . . . CHAPTER TWO: A POPULATION GENETIC COMPARISON OF LARGE AND SMALLBODIED SAGE GROUSE IN COLORADO Introduction Methods Tissue Collection DNA Extraction and Microsatellite Genotype Scoring mtDNA Sequencing Data Analysis Results Microsatellite Data mtDNAData Discussion Literature Cited Tables Figures Appendix . . . . . . . . . . . . . . . CHAPTER THREE: POPULATION GENETICS OF GUNNISON SAGE GROUSE: IMPLICATIONS FOR MANAGEMENT Introduction Study Area Methods Data Analysis Results . . . . . . �62 Discussion Management Implications Literature Cited Tables Figures CHAPTER FOUR: QUANTIFYING CHANGES IN SAGEBRUSH HABITAT IN SOUTIIWESTERN COLORADO FROM THE MID-50'S TO THE MID-90'S Introduction Methods Plot Selection Aerial Photography Acquisition and Interpretation Data Analysis Results Habitat Loss Habitat Fragmentation Discussion Literature Cited Tables " Figures - . . . . . ' -, . . . . . . . . . . . . . CHAPTER FNE: DEVELOPMENT OF A MODEL TO ASSESS MANAGEMENT AND CONSERVATION STRATEGIES FOR GUNNISON SAGE GROUSE IN COLORADO Introduction Methods Results Discussion Literature Cited Tables Figures . . . . . . . . DISCUSSION . �63 INTRODUCTION The Gunnison sage grouse (Centrocercus minim us) is a newly recognized species (Braun and Young 1995) whose range is restricted to southwestern Colorado and southeastern Utah. The distribution and abundance of Gunnison sage grouse in Colorado has declined markedly, with extirpations in 12 of the 17 counties in southwestern Colorado which once supported them (Rogers 1964, Braun 1995). Declines are thought to be the result of habitat loss (conversion of big sagebrush [Artemisia tridentata] into farmland or housing developments), habitat degradation (heavy grazing, sagebrush removal, road and powerline development through sagebrush areas), and habitat fragmentation (Braun 1995). The majority of populations that remain are small, and exist in isolated, degraded patches of sagebrush habitat. One large population does remain, however, in the Gunnison Basin. Because of its restricted range and small population size, the conservation of this species has become a significant concern. The conservation of Gunnison sage grouse requires knowledge of certain issues which have not yet been addressed. First, little is known about landscape level habitat requirements of sage grouse living in fragmented habitats. It is not known how large a sagebrush patch must be to support sage grouse, or if patch edge, or distance to the nearest road affect the probability of sage grouse persistence. Second, it is not known how much sage grouse habitat has already been lost and how much might be lost in the future, given human population growth and land development This is essential information if a balance between human population growth and sage grouse conservation is to be achieved. Third, little is known of the dispersal movements of sage grouse, as only one study has addressed this issue. Dunn and Braun (1985) measured natal dispersal of sage grouse in contiguous but altered habitats of northwestern Colorado and found average dispersal distances of8.8 km for juvenile females and 7.4 km for juvenile males. It is not known, however, whether Gunnison sage grouse move among fragmented habitats (across distances up to 300 km) or whether some populations in southwestern Colorado are truly isolated. Knowledge of movement among patches and the levels of genetic diversity would provide essential information and aid in any conservation plan which addresses translocations and reintroductions. .. In this dissertation I address three issues for which information is lacking. In Chapter One, I develop a habitat-based model which can be used to identify the relative importance of landscape and micro-level variables (or combinations of them) in sustaining Gunnison sage grouse. This model can be used to examine the suitability of any sagebrush patch in southwestern Colorado using the variables deemed important by the model. This gives biologists information on which occupied patches are most at risk of extinction and also allows unoccupied sagebrush patches to be ranked in order to.identify which patches have the highest probability of occupancy by sage grouse. This is important because sage grouse population expansion could involve translocation into unoccupied sagebrush patches. Chapters Two and Three address genetic issues concerning Gunnison sage grouse. Chapter Two is a population genetic analysis of nine sage grouse populations in Colorado using two different molecular genetic markers. In this chapter I address the question of whether genetic data support the new species designation of Gunnison sage grouse, and I compare relative amounts of gene flow and genetic diversity between Gunnison sage grouse populations from southwestern Colorado and sage grouse (c. urophasianus) populations from northern Colorado. Management implications of the genetic data are addressed in:Chapter Three. In Chapter Four I document the loss and fragmentation of sagebrush-dominated habitat in southwestern Colorado using aerial photographic analysis. This is important because if this species is listed as threatened, quantitative documentation of habitat loss and fragmentation (thought to be a contributing factor in the species' decline) is essential. Also, rates of habitat loss and fragmentation can be used to make predictions about future habitat loss given current rates of human population �68 e) distance to nearest oakbrush, pinon/juniper. wet meadow. and fence post (within 100 m); and 1) number of sage grouse pellets in the plot and the belt outside of the plot. Observations of sage grouse or grouse sign along the transect between stops were recorded. Occupancy or vacancy of a patch was based on whether or not sage grouse pellets were seen or whether sage grouse were flushed. Sage grouse pellets last for up to a year (C.E. Braun. Colorado Division of Wildlife. personal communication). The area of each patch. the area/perimeter ratio, distance to nearest occupied patch. and the distance to the nearest road (paved and unpaved) were determined from satellite data in a GIS operated by the Western Region of the Colorado Division of Wildlife. Data Analysis A logistic regression framework (proc GENMOD; SAS® Institute Inc. 1993) was used for analysis since the dependent variable (occupancy) was binary. The general form of logistic regression IS 1 0=--- 1+ e= where A are variables in the model, Po... PIc-l are coefficients fit by the model, and 0 is the predicted probability given the model. Because the number of actual data points (patches) was small, only a limited number of candidate models should be considered for model selection (Burnham and Anderson 1998). Thus. I developed a number of composite variables from the raw data The habitat requirements of sage grouse are well known and are generally categorized into winter. breeding and nesting. and summer habitat. I created composite variables representing the percent of a patch in winter habitat, breeding and nesting habitat. and summer habitat. Further, I created a variable representing the area of habitat in a patch that was preferable (meaning that it represented either winter, breeding and nesting, or summer habitat). Winter habitat was defined by greater than 20% cover oflive sagebrush taller than 20 em (Eng and Schladweiler 1972. Beck 1977). Breeding and nesting habitat was defined by 20 - 40 % cover of live sagebrush between 17 and 119 em in height. 7 - 10 % cover of grass, and > 4% cover offorbs. (patterson 1952:114, Kebenow 1969, Wallestad and Pyrah 1974. Connelly et al. 1991, Gregg et al. 1994; Musil etal, 1994, Young "1'994). Summer habitat was characterized by J 4 - 30 % cover of live sagebrush. 1 - 17% forb cover, and 1 - 22% grass cover (Martin 1~}70"Wallest3.d 1971, Klebenow 1969, Klott and Lindzey 1990, Young 1994). I proposed seven a priori candidate models (Table 1.1) for the model selection process. These seven models included three explanatory variables: patch area (as a measure of habitat quantity), distance to the nearest paved road from the centroid of the patch (as a measure of human disturbance and fragmentation). and the area of the patch considered to be suitable winter. breeding and nesting or summer habitat (as a measure of habitat quality). To choose the "best" model that is supported by the science of the situation. by the data, with enough parameters to avoid bias but not so many as to lose precision. I used Aikaike's Information Criterion for small sample sizes (AlCc) (Buckland et al. 1997, Burnham and Anderson 1998): X I' .. X "_I 2K(K+ 1) Alec = -2(ln-E) + 2K + n- K-l �69 where .In(.13) is the natural logarithm of the likelihood function evaluated at the maximum likelihood estimates for a given model, K is the number of estimable parameters from that model, and n is sample size. To address uncertainty in the model selection process, models were compared and ranked using AAIC (Lebreton et al. 1992;Bumham and Anderson 1998) and Akaike weights (Buckland et al. 1997, Burnham and Anderson 1998). AAlC was calculated as: AAIC. I = AlC. I - minAIC where AlCj was the Alec value for the ith model in a suite of models being compared and minAIC was the minimum AlCc value among those models. Akaike weights were computed as 1 exp(- 2"A;) 7 1 L exp(- -Ar) r=1 2 where Ai is the AAIC value for the ith model and A r is the AAIC value for the rth model as AAIC values are summed from one to seven. A model is considered to be competitive by Burnham and Anderson (1998) if the AAIC is less then two or if the ratio of the Akaike weight of the best model to the weight of a candidate model is less than eight. RESULTS The results of the model selection procedure for the seven candidate models vaned (Table 1.2). Four models had AAlC values less than two and ratio of Akaike weights less than eight. They were considered to be competing models. The remaining three models had large AAIC values and negligible weights and were dropped from consideration (Table 1.2). The top two models had almost identical AlCc values (23.530 and 23.614) and, hence, similar weights (0.335 and 0.321). The third and fourth ranked models also had similar AlCc values and weights. Inspection of the parameter estimates for the four top models (Table 1.3), however, revealed that the parameter estimate for area of suitable habitat was negative in one case (technically meaning that the less area of suitable habitat, the more likely it would be Occupied). This obviously makes no biological sense. There are several reasons why this may have occurred. First, I estimated area of suitable habitat by calculating the percentage of plots with either winter, or breeding and nesting, or summer habitat and multiplying it by the patch area, which may not estimate this parameter well. Second, the definition of winter, breeding and nesting, and summer habitats, came from other studies of sage grouse (in most cases large-bodied C. urophasianus, not with small-bodied C. minim us) in other areas. The habitat requirements for the small-bodied Gunnison sage grouse may be somewhat different than for the large-bodied sage grouse. Finally, area of suitable habitat was highly correlated with patch area Adding area of suitable habitat to the model already containing patch area and distance to the nearest road did not improve the model and, because of its high correlation with patch area, may cause a spurious parameter estimate. �70 As a result, I eliminated any models containing the variable area of suitable habitat and recalculated Akaike weights for the remaining three models (Table 1.4). The top model with patch area and distance to the nearest paved road received more weight than models with either variable alone. The model with distance to the nearest paved road, however, was a close second. Because the Akaike weights of the top two models were similar (0.507,0.486), suggesting that both models were competitive, I concluded that none of the three remaining models alone was sufficient to make predictions about occupancy given patch area and distance to the nearest paved road. Instead, I used model averaging (which accounts for uncertainty in model selection), to estimate the probability of occupancy given a patch size and distance to road (Burnham and Anderson 1998). " I calculated the model averaged prediction Oa as: A R A (}a = L w.O. j=1 I I " where OJ is the predicted value for occupancy from model i, and Wj is the Akaike weight for model i. Variance was calculated as: and confidence intervals were calculated using: and where Thus, for a range of patch areas and distances to the nearest paved road I can predict the probability of occupancy with appropriate confidence intervals (Table 1.5). I also examined the relationship between the probability of occupancy and distance to the nearest paved road for a series of different patch areas (Fig. 1.4) and developed a surface of probability of occupancy for a range of patch areas and distances to paved roads (Fig. 1.5). I determined the importance of each predictor variable by summing the Akaike weights for all models containing each predictor variable (Burnham and Anderson 1998) and found that the distance to the nearest paved road was almost twice as important as the patch area (0.994 for road distance vs. 0.513 for patch area on a scale from zero to one). �71 DISCUSSION Despite all the micro-scale data collection which was used to quantify the amounts of winter, breeding and nesting, and summer habitat, the two variables included in the two best models were patch size and distance to the nearest paved road. There are several reasons why this may have occurred. First, the number of available patches meeting my criterion was small (25) which considerably limited not only the number of candidate models and variables that could be considered, but also the ability to detect real effects. Second, the composite variable area of suitable habitat (which measured the area of each patch which was comprised of either winter, breeding and nesting, or summer habitat) might not have been estimated well given my sampling scheme of measuring habitat characteristics in a sample of 1 m2 plots in each patch, and deciding whether or not that plot could be characterized as winter, breeding and nesting, or summer habitat. Finally, the winter, breeding and nesting, and summer habitat classifications were taken from previous studies of sage grouse in Colorado and other states. Most studies which were used to define habitats were of "large-bodied" sage grouse (c. urophasianus), and only one (Young 1994) included data from "small-bodied" sage grouse (c. minimus). Perhaps the definitions of winter, breeding and nesting, and summer habitat for large-bodied sage grouse are somewhat different than for small-bodied sage grouse. However, it is most likely that small sample sizes impeded identification of effects from microscale variables. A measure of habitat quality (as a function of the probability of occupancy) can be determined for any patch in southwestern Colorado using patch size, the distance to the nearest paved road from the centroid of the patch, and model averaging over my three models. This produces a model averaged estimate of probability of occupancy with appropriate confidence intervals. Using my models and model averaging, I can also rank patches as to their suitability which may prove helpful if reintroductions or population augmentation become management options (Chapter Five). Further, the effects of habitat reduction and road construction can be assessed. While inferences of the importance of certain habitat variables may be weak due to small sample size, the utility of the models and model averaging is still tenable. Because the variables included in the models are large scale variables, they can be measured with a minimum of effort and cost. The time and money involved in measuring habitat variables to the extent to which they were measured in this study may not be a reasonable option. Thus, these models and model averaging can be used as a coarse grained, quick method to predict the probability of occupancy and can be incorporated into a broad scale management scheme for Gunnison sage grouse. LITERATURE CITED Aldrich, J. W. 1963. Geographic orientation of American Tetraonidae. Management 27:529-545. Beck, T. D. 1977. Sage grouse flock characteristics Management 41: 18-26. Journal of Wildlife and habitat selection in winter. Journal of Wildlife Boecklen, W. J., and C. W. Bell. 1987. Consequences offaunal collapse and genetic drift for the design of nature reserves. Pages 141-149 in D. A. Saunders, G. W. Arnold, A. A. Burbridge, and A. 1. M. Hopkins, editors. Nature conservation: the role of remnants of native vegetation. Surrey and Beatty and Sons, Chipping Norton, Australia Braun, C. E. 1995. Status and distribution of sage grouse in Colorado. Prairie Naturalist 27: 1-9. �74 Table 1.1. Candidate models used for model selection, within a logistic regression framework to predict the probability of occupancy of Gunnison sage grouse in southwestern Colorado. Model Model structure Area Po + PI (area) Distance to road Po + PI (distance) Area of suitable habitat Po + PI (area suitable) Area, distance to road Po + PI (area) + P2 (distance) Area, area of suitable habitat Po + PI (area) + P2 (area suitable) Distance to road, area of suitable habitat Po + PI (distance) + P2 (area suitable) Table 1.2. Selection of models using AlCc as a model selection criterion. Models in this logistic regression framework predict the probability of occupancy of Gunnison sage grouse in southwestern Colorado. K represents the number of parameters. Model K AlCc aAIC Akaike weight Area, distance to road 3 23.530 0.000 0.335 Distance to road 2 23.614 0.085 0.321 Distance to road, area of suitable habitat 3 24.578 1.048 0.19& Area, distance to road, area of suitable habitat 4 25.340 . 1.811 0.135 Area, area of suitable habitat 3 32.050 8.520 0.005 Area 2 32.321 8.791 0.004 Area of suitable habitat 2 36.974 13.448 0.000 �75 Table 1.3. Parameter estimates and model conditional standard errors for the four best logistic regression models to predict the probability of occupancy of Gunnison sage grouse in southwestern Colorado. Estimate SE Intercept -5.8699 2.8930 Area 0.0845 0.0609 Distance to road 0.0016 0.0008 Intercept -4.5271 1.8817 Distance to road 0.0015 0.0006 Intercept -5.3861 2.5611 Distance to road 0.0016 0.0007 Parameter Model with area, distance to road Model with distance to road Model with distance to road, area of suitable habitat Table 1.4. Three logistic regression models remaining after model selection using AlCc with updated Akaike weights. These models can be used to predict the probability of occupancy of sagebrush patches by Gunnison sage grouse in southwestern Colorado. '. Model K Alec AAIC Area, distance to road 3 23.530 0.00 0.507 Distance to road 2 23.614 0.085 0.486 Area 2 32.321 8.791 0.006 Akaike weight . �76 Table 1.5. Model averaged predictions of patch occupancy by Gunnison sage grouse for a series of different theoretical patch areas and distances to the nearest paved road. Area (km') 1 1 1 1 1 1 1 10 10 10 10 10 10 10 50 50 50 50 50 50 50 90 90 90 90 90 90 90 500 500 500 500 500 500 500 Distance to nearest paved road (m) Predicted probability of occupancy 95%CI 10 100 1000 2000 3000 4000 5000 10 100 1000 2000 3000 4000 5000 10 100 1000 2000 3000 4000 5000 10 100 1000 2000 3000 4000 5000 10 100 1000 2000 3000 4000 5000 0.0086 0.0096 0.0310 0.1194 0.3694 0.7208 0.9196 0.0113 0.0126 0.0405 0.1561 0.4588 0.7953 0.9442 0.0942 0.1055 0.2771 0.5077 0.7181 0.8916 0.9693 0.4434 0.4529 0.5175 0.5906 0.7375 0.8958 0.9704 0.5183 0.5190 0.5346 0.5942 0.7382 0.8960 09704 0.0002 - 0.3118 0.0002 - 0.3235 0.0014 - 0.4227 0.0157 - 0.5362 0.1152 - 0.7251 0.2989 - 0.9399 0.4070 - 0.9948 0.0002 - 0.3610 0.0003 - 0.3721 0.0021 - 0.4562 0.0280 - 0.5433 0.2082 - 0.7321 0.3956 - 0.9584 0.4913 - 0.9966 0.0005 - 0.9519 0.0007 - 0.9539 0.0058 - 0.9619 0.0405 - 0.9618 0.1500 - 0.9735 0.3273 - 0.9929 0.4169 - 0.9993 0.0094 - 0.9853 0.0112 - 0.9838 0.0255 - 0.9778 0.0454 - 0.9777 0.1364 - 0.9804 0.3331 - 0.9933 0.4922 - 0.9991 0.0218 - 0.9811 0.0220 - 0.9811 0.0258 - 0.9803 0.0447 - 0.9786 0.1357 - 0.9806 0.3320 - 0.9934 0.4901 - 0.9991 �77 • ._.. "10>", _._ _----. •. Figure 1.1. Historic (top) and current (bottom) distribution of sage grouse and Gunnison sage grouse (lower left cut out) in Colorado. �78 Figure 1.2. Sampling scheme for micro-scale variables in a small patch. Starting points were chosen randomly (represented here by X), transects were run in north/south directions, measurements were taken from I-m2 sampling frames (represented by the white box) every 200 m. Figure 1.3. Sampling scheme for measurements in a large patch. Starting points were randomly chosen, transects were established in north/south directions, and measurements sampling plot (represented by the white boxes) every 200 m. were taken in a I-m2 �79 AREA=1 KM' '.0 D.' >- 0.1 ~ 0.7 .. 0 :> 0 U 8 ~ ~ /-- 0.1 0.4 :I if 0.3 / 0.2 D.' 0.0 ../ -====---_'-".~ .. ......,..... - 3DDO .000 DlS1»fCE AREA=30 FROM ROAD ~ ... M IODO KM' IJ) ..• ~ ..• ~ 0.7 o 0.1 ~ 0.1 5 OA D ~ 0.3 f 0.2 D.' OJ) .0•• 20 •• :sooo 4••0 5000 "STANCE FROM ROAD.,) AREA = 500 KM 2 '.1) ..• ~ ..• ~ 0.7 g 0.5 •.. ._---------_ . 0'" 5u .. :i o f 0.3 0.2 0.' 1000 2000 DISTANCE 40 •• 3000 FROM RQ\D $000 (II) Figure 1.4. Theoretical relationship between the probability of occupancy and distance to the nearest paved road from the centroid of the patch for three different patch areas. Dotted lines represent upper and lower 95% confidence intervals for the predicted value (non-dotted line). �80 Figure 1.5. Relationship between patch area (in krrr'), distance to the nearest paved road (in m), and the model averaged prediction of occupancy. �81 CHAPTER1WO A POPULATION GENETIC COMPARISON OF LARGE AND SMALL-BODIED GROUSE IN COLORADO SAGE INTRODUCTION Sage grouse (Centrocercus urophasianus) have experienced marked declines in their distribution and abundance throughout their entire range (Braun et al. 1994). Their historic distribution included at least 16 states and three provinces in North America (Aldrich 1963, Johnsgard 1973, Braun 1998) and has since been extirpated from five states and one province (Braun 1998). In Colorado, the distribution and abundance of sage grouse have also been greatly reduced (Braun 1995) as they have been extirpated from 12 of the 27 counties in Colorado in which they occurred in the 1900's (Braun 1995) and populations in nine of the remaining 15 counties are thought to number less than 500 breeding birds. Because of this marked decline, sage grouse have become the focus of management and conservation concerns. Sage grouse have historically been classified into two subspecies: C. u. urophasianus (Eastern sage grouse) and C. u. phaios (Western sage grouse) .. This subspecies distinction was based on plumage and coloration differences (Aldrich and Duvall 1955), yet its validity has been questioned (Johnsgard 1983). Studies in southwestern Colorado (Hupp and Braun 1991) and southeastern Utah (Barber 1991) found sage grouse to be approximately 33% smaller than sage grouse from northern Colorado and throughout the rest of the species' range. Further, these "small-bodied" sage grouse have longer filoplumes and different tail banding patterns. Young (1994) and Young et al. (1994) compared strutting displays from the small-bodied sage grouse in southwestern Colorado to "large-bodied" sage grouse populations in northern Colorado and in California and found that many of the ritualized components of the strutting display differed. Further, Young (1994) found that small-bodied females avoided tape-recorded vocalizations oflarge-bodied males. Based on morphological and behavioral differences between large and small-bodied sage grouse, Braun and Young (1995) proposed that small-bodied sage grouse from southwestern Colorado and southeastern Utah be recognized as a new species, based on the biological species concept. To determine whether genetic evidence is consistent with this new species designation, Kahn et al. (1999) compared the genetic variation among five populations of large-bodied sage grouse from northern Colorado, one population of large-bodied sage grouse from Utah, and one population of smallbodied sage grouse from southwestern Colorado. To document this variation, they sequenced 141 base pairs ofa rapidly evolving portion (region I) of mitochondrial DNA (mtDNA) and showed that sequences from the seven populations included 21 .haplotypes that formed two monophyletic clades. Several different haplotypes from both clades were found in all six large-bodied populations, while within the small-bodied population, all but one of the 31 individuals analyzed were genetically identical, and both observed haplotypes were members of the same clade. They concluded that the unusually low level of genetic variation and absence of several haplotypes that were common in the large-bodied populations in Colorado provided evidence of a lack of gene flow between the two proposed species. While their study provides evidence that can be construed to support the new species designation, I expanded it to include individuals from three additional small-bodied populations not included in Kahn et aI.'s (1999) study, and supplemented their mtDNA data with data from the nuclear DNA. This was done to more completely characterize the mtDNA data and to eliminate any concern that male biased gene flow would not be elucidated using the maternally inherited mtDNA. The nuclear �82 molecular markers that I chose were microsatellite markers which are areas in the nuclear genome characterized by short, tandem repeats with a high rate of variation in copy number among individuals. Microsatellites are highly variable and are generally considered to be among the most powerful molecular genetic markers for population genetic studies (Goldstein and Pollock 1997). METHODS Tissue CoUection Extracted DNA from 20 birds from the Gunnison Basin and from the five large-bodied populations that were used in Kahn et al. 's (1999) study, were also used in this study. These five northern Colorado populations include Cold Springs, Blue Mountain, Eagle, Middle Park, and North Park (Fig. 2.1). Blood samples and feathers were obtained from small-bodied sage grouse which were captured using a spotlight trapping method (Giesen et al. 1982) in the following populations in Colorado: Dove Creek (N = 15), Dry Creek Basin (N = 22), Crawford (N = 20), and Guimison Basin (N = 9) (Fig. 2.1). Blood samples were obtained by clipping a toe nail of each sage grouse and placing 2-3 drops of blood into a microfuge tube previously coated with EDTA. These blood samples, as well as feather samples from each sage grouse were frozen at _20DC. The nine Gunnison Basin samples were from the same area sampled by Kahn et al. (1999) and were used to augment the 20 Gunnison Basin samples collected by Kahn et al. (1999). DNA Extraction and MicrosateUite Genotype Scoring DNA extractions from blood or the bottom 2 em of the feather shaft, followed the procedure of Quinn and White (1987). Over 30 microsatellite primers from the chicken genome project were used to screen for polymorphism of microsatellites as well as 12 primers developed for red grouse (Lagopus Zagopus scoticus). I found four microsatellites with clean, scorable products that were polymorphic in both the large and small-bodied sage grouse. These four loci proved to be informative and allowed me to sufficiently address the objectives of this study. Primers for those four microsatellites (LLSTIF, LLSTIR, LLSD3F, LLSD3R, LLSD4F, LLSD4R, LLSD8F and LLSD8R) were developed by Piertney and Dallas (1997). One primer (either the forward or reverse primer) was chosen and radioactively labeled for later visualization on autoradiography film using the T 4 Polynucleotide Kinase (PNK) labeling procedure. In a 0.5 ~l microfuge tube, i ~110 J.!Mprimer, 1 III lOX Buffer, 0.25 III T4 PNK (10 U/f.ll), 0.25 III A-33p-ATP(10~Q.mCilml), and 7.5 III H20 were mixed and incubated at 37DC for 15 minutes. The reaction was stopped by heatingto 70DC for 10 minutes. ... . Polymerase chain reactions (PCR) were performed in a Perkin-Elmer DNA thermal cycler. Approximately 30 ng of genomic DNA (in a I IIIvolume) was used as template in each 25 IIIPCR (as described in Quinn 1992), using one forward and one backward primer, with the following thermal profile: 2 min denaturation at 94 DC followed by 35 cycles of "touchdown" ramping: 30 seconds denaturation at 94 DC and 30 seconds annealing while stepping from 60DC to 50DC. A 20 minute extension at 74 DC was performed at the end of the 35th cycle. PCR products and a size standard were electrophoresed at 55 watts for two hours through 6% denaturing poly-acrylamide gels as described in Sambrook et al. (1989). Autoradiographs were made of each dried acrylamide gel by exposure to X-ray film (Fuji RX). Individuals were assigned genotypes (corresponding to microsatellite fragment length) based on banding patterns on the autoradiographs. In some cases samples containing alleles of similar sizes were rerun in adjacent lanes. The distribution of allele frequencies for each population was recorded. �83 mtDNA Sequencing The procedures were described in detail previously (Kahn et al. 1999). I identified new haplotypes by comparison to those designated previously by Kahn et al. (1999). Data Analysis Microsatellite genotypes were tested for departures from Hardy-Weinberg equilibrium within each population at each locus using the computer program Arlequin (Schneider et al. 1997). Arlequin uses a Markov-chain random walk algorithm (Guo and Thompson 1992) which is analogous to Fisher's exact test but extends it to an arbitrarily sized contingency table. Population genetic structure was investigated using pairwise population F ST significance tests. F tests (Tjur 1998) for each locus were conducted to determine whether the distributions of alleles were significantly different between the large and small-bodied birds. An Ftest is a ratio of mean squares (analogous to ANOYA) which is used here because it is robust to overdispersed data Genetic distance for all pairs of populations was estimated using two different distance metrics. Both metrics assume an infinite alleles model of mutation. Although Goldstein and Pollock (1997) advocate using stepwise mutation models to estimate genetic distances for phylogenetic reconstruction using microsatellite data, D. B. Goldstein (personal communication) suggests that population genetic studies using microsatellites should use genetic distances based on the infinite alleles model (specifically the proportion of shared alleles [Bowcock et al. 1994]) because they are linear over short periods of time and have a low variance. I calculated the proportion of shared alleles (Bowcock et al. 1994) and also Cavalli-Sforza and Edwards' (1967) chord distance because Takezaki and Nei (1996) showed it to have a higher probability of obtaining correct tree topologies than other distance measures with microsatellite markers. Chord distance, De, was calculated as r Dc = (2/ trr)'L j where Xij andYij are the frequencies of the ith allele at thejth locus in populations X and Y respectively, mj is the number of alleles at the jth locus, and r is the number of loci examined. The proportion of shared alleles, Ps, was calculated as Ps = s /(2/) where s is the number of shared alleles summed over loci, and 1 is the number of loci compared. I calculated genetic distance between all pairs of populations and constructed neighbor joining trees describing the relationship among populations using the microsatellite data and both distance measures. For the mtDNA analysis, I documented population subdivision in Arlequin (Schneider et al. 1997) using significance tests of pairwise population FST values. An F test was calculated to determine whether the distribution ofhaplotypes among the large and small-bodied birds differed. I conducted an analysis of molecular variance (AMOYA) as described by Excoffier et al. (1992) which produces estimates of variance components to reflect haplotype diversity at different levels. of a hierarchy. I documented the variation due to large vs. small bodied birds as one level of hierarchy, the variation among populations within the two body sizes as a second level, and the variation among individuals in a population as the third level. The molecular distances between haplotypes were modeled following Tamura (1992) because my haplotypes had unequal frequencies of A, C, T, and G and because my transition/transversion ratio was much higher than the expected (mathematically) ratio �84 of 1:2. I calculated pairwise population genetic distances which incorporate both the Tamura (1992) corrected molecular distance between haplotypes and the haplotype frequencies in each population. Neighbor joining trees were constructed showing the relationship of the nine populations. RESULTS MicrosateHite Data There were 19 different haplotypes across all individuals. Kahn et al. (1999) found that the five large-bodied populations all had at least five different haplotypes in each population. They found four dominant haplotypes (A. B, C, and D) with haplotypes B, C, and D common in all large-bodied populations and haplotype A found in all but one. In the small-bodied populations, I found only two or three haplotypes per population (Fig. 2.3). Only one of the haplotypes dominant in the large-bodied birds, A. was found and haplotype G was found to be unique among the small-bodied birds (Appendix 2.A). I found significant population subdivision using population pairwise Fsr significance tests (Table 2.3). As with the microsatellite data, all possible pairwise comparisons between small and largebodied sage grouse populations showed significant differences. Further, I found that within the largebodied sage grouse, no two populations were significantly different and among the small-bodied sage �85 grouse, only one population pair was not significantly different (Dry Creek and Dove Creek, P = 0.072) (Appendix 2.A). To test whether the distribution ofhaplotypes from the large-bodied populations differed from the distribution ofhaplotypes from the small-bodied populations, I used an F test. There was a statistically significant difference between the distribution ofhaplotypes in the large and small-bodied populations (FI8•70 = 3.82, P < 0.001). Further, I used AMOVA to examine components of variance between the large and small-bodied groups, among populations within groups, and among individuals within populations. I found that 65% of the variance could be explained by the large vs. small-bodied group distinction, only 2% of the variance was explained by between population variation within body size, and the remaining 33% of the variance was explained by within population variation (Table 2.4). The pattern noted in the trees from the microsatellite data is similar to the mtDNA tree (Fig. 2.4) suggesting a separation between the large and small-bodied sage grouse. DISCUSSION In all four microsatellites and in the 141 bp control region of the mtDNA high variability was found even at my smallest sampling level (within populations) which provided me with a powerful tool to detect population subdivision. The only significant departure from Hardy-Weinberg equilibrium (Eagle locus LLSD3) was a case of heterozygote deficiency which could be the result of many factors including null-alleles, Wahlund effect, and inbreeding. Null alleles occur when a mutation causes one oligonucleotide primer not to amplify one allele which is manifested by a deficiency ofheterozygotes (pemberton et al. 1995). Null alleles are also sometimes detected when PCR products cannot be amplified for certain individuals (Lehman et al. 1997). I doubt null alleles were the cause for the heterozygote deficiency in Eagle because I had no problem getting PCR products from Eagle individuals for any loci. Also, I had two family groups of known mother and offspring which I tested over all loci and found no evidence of null alleles. Further, a heterozygote deficiency was found only in one population and I might expect to find deficiencies in other populations if null alleles were the cause. The heterozygote deficiency in Eagle might be the result of the Wahlund effect of pooling separate populations into one population or of inbreeding. However, if either was the case I would expect to find this effect among the three other loci which I did not. Pairwise population Fsr significance tests showed similar patterns in the microsatellite and mtDNA analyses (Tables 2.2, 2.3). Both markers revealed significant differences between all large vs. small-bodied population comparisons supporting a distinction between these two groups of birds. Both markers also revealed there were no significant differences among any of the large-bodied bird populations suggesting substantial gene flow among them. Within the small-bodied bird populations, most pairwise population comparisons showed significant differences among populations with a few exceptions (Gunnison and Dry Creek P = 0.007, Dry Creek and Dove Creek P = 0.025 for microsatellites; Dry Creek and Dove Creek P = 0.054 for mtDNA). Also, the F ST value calculated among the large-bodied populations (FST = 0.0266,95% CI -0.0016 - 0.0528) is significantly smaller than the value calculated among the small-bodied populations (FST = 0.2153, 95% CI 0.1230 - 0.3339). This suggests there is some amount of subdivision among the small-bodied birds likely due to their small population sizes (~2600 birds in Gunnison Basin, ~ 175 birds in Crawford, ~ 75 in Dove Creek, and ~ 300 in Dry Creek, (C. E. Braun, Colorado Division of Wildlife, unpublished data» and isolation (Fig. 2.1). This is consistent with Braun's (1995) assertion that clearing of sagebrush for cultivated crops, highway construction, ranch development, powerline placement, reservoir construction, and other facets of human settlement have resulted in the fragmentation and loss of sagebrush habitats such that sage grouse populations in southwestern Colorado are small and isolated (also see Chapter Four). This reduction of habitat is evident when comparing the historic range of sage grouse in Colorado with �86 its current distribution (Fig. 2.1). A comparison of these two distributions reveals that the majority of fragmentation and loss of habitat has occurred in southwestern Colorado resulting in small, isolated populations, and that populations in northern Colorado remain relatively large and contiguous, all of which is supported by my genetic data The three of four significant F tests for the microsatellite loci and the significant F test for the mtDNA data reveal that the distribution of allele and haplotype frequencies are different for the large and small-bodied sage grouse populations. Further, in both the microsatellite and mtDNA data there are alleles and a haplotype unique to the small-bodied sage grouse thereby supporting the idea that gene flow between the two groups is likely absent and some amount of divergence has occurred. This supports Braun and Young's (1995) recognition of small-bodied sage grouse as a new species based on the biological species concept. In addition, the mtDNA AMOV A (Table 2.4) indicates that 65% of the total variation in the mtDNA data can be explained by the large vs. small-bodied sage grouse distinction and that only 2 % of the variation can be attributed to differences among populations within the large or small-bodied group. Kahn et al. (1999) discuss the ancestry of the mtDNA haplotypes and profess two different explanations for the establishment of the small-bodied sage grouse. They believe that either a founder population of large-bodied birds diverged rapidly from other large-bodied populations likely due to sexual selection or that the small-bodied sage grouse evolved across a widespread portion of the southwestern range (remaining unnoticed as a separate taxon) and underwent a severe bottleneck recently due to habitat fragmentation and habitat loss. My data are consistent with the founder hypothesis because in the microsatellite analysis the majority of the alleles present in the small-bodied populations are also present in the large-bodied populations, yet the diversity in the small-bodied populations (17 alleles) is much less than in the large-bodied populations (44 alleles). The mtDNA analysis also supports this hypothesis in that the dominant haplotype in the small-bodied populations (A) is well represented in the large bodied birds. The haplotype unique to the small-bodied birds is close to the A haplotype (one transition) representing a recent mutation. As in the microsatellite analysis, the genetic diversity in the large-bodied populations is much higher (17 haplotypes) than in the small-bodied populations (three haplotypes). All genetic distances from both markers show a similar broad pattern of a distinction between the large and small-bodied populations. From the mtDNA tree I can conclude that within the largebodied group populations are more closely related (shorter branch lengths) than within the smallbodied group (longer branch lengths). This was also apparent from the pairwise population F ST significance tests (Tables 2.2, 2.3) in which populations within the large-bodied group were not significantly different whereas, within the small-bodied group they were different. This study has provided valuable additions to the study conducted by Kahn' et al. (1999) in that there is now nuclear data to corroborate the mtDNA data Further, I expanded the survey of smallbodied sage grouse to include informatiori from three additional populations which is essential to the conservation of the small-bodied sage grouse. I not only extended Kahn et al.' s (1999) picture of the distinction between large and small-bodied sage grouse, but I documented the isolation and low genetic diversity of the small-bodied sage grouse populations. This is important information for the management of the small-bodied sage grouse as a species. Future research on sage grouse should include more microsatellite loci and population surveys throughout the entire range of sage grouse. This would provide a much deeper knowledge base for the understanding and management of sage grouse. �87 LITERATURE CITED Aldrich, J. W. 1963. Geographic orientation of American Tetraonidae. Journal of Wildlife Management 27:529-545. Aldrich, J. W., and J. W. Duvall. 1955. Distribution of American gallinaceous game birds. United States Department of Interior, Fish and Wildlife Service, Washington, D. C., USA. Circular 34. Barber, H. A. 1991. Strutting behavior, distribution and habitat selection of sage grouse in Utah. Thesis, Brigham Young University, Provo, UT, USA. Bowcock, A. M, A. Ruiz-Linares, J. Tomfohrde, E. Minch, J. R Kidd, and L. L. Cavalli-Sforza. 1994. High resolution of human evolutionary trees with polymorphic microsatellites. Nature 368:455457. Braun, C. E. 1995. Status and distribution of sage grouse in Colorado. Prairie Naturalist 27:1-9. Braun, C. E. 1998. Sage grouse declines in western North America: what are the problems? Proceedings of the Western Association ofFish and Wildlife Agencies 78:000-000. Braun, C. E., and J. R Young. 1995. A new species of sage grouse in Colorado. Joint Meeting Wilson Ornithological Society and the Virginia Society of Ornithology, Abstract #23. May 4-7, 1995, Williamsburg, VA, USA. Braun, C. E., K Martin, T. E. Remington, and J. R Young. 1994. North American grouse: issues and strategies for the 21st Century. Transactions of the North American Wildlife and Natural Resources Conference 59:428-438. Cavalli-Sforza, L. L., and A. W. F. Edwards. 1967. Phylogenetic analysis: models and estimation procedures. American Journal of Human Genetics 19:233-257. Excoffier, L.~ P. E. Smouse, and J. M. Quattro. 1992. Analysis of molecular variance inferred from metric distances among DNA haplotypes: application to human mitochondrial DNA restriction data Genetics 131:479-491. Giesen, K M., T. J. Schoenberg, and C. E. Braun. 1982. Methods for trapping sage grouse in Colorado. Wildlife Society Bulletin 10:224-231. Goldstein, D. B., and D. D. Pollock. 1997. Launching microsatellites. Journal of Heredity 88:335-342. Guo, S., and E. Thompson. 1992. Performing the exact test of Hardy-Weinberg multiple alleles. Biometrics 48:361-372. proportions for Hupp, J. W., and C. E. Braun. 1991. Geographic variation among sage grouse in Colorado. Wilson Bulletin 103:255-261. �88 Johnsgard, P. A. 1973. Grouse and quails of North America University of Nebraska Press, Lincoln, NE, USA. Johnsgard, P. A. 1983. Grouse of the world. University of Nebraska Press, Lincoln, NE, USA Kahn, N. W., C. E. Braun, J. R Young, S. Wood, D. R Mata, and T. W. Quinn. 1999. Molecular analysis of genetic variation among large and small-bodied sage grouse using mitochondrial control region sequences. Auk (in press). Lehman, T, N. J. Besansky, W. A. Hawley, T G. Fahley, L. Kamau, and F. H. Collins. 1997. Microgeographic structure of Anopheles gambiae in western Kenya based on mtDNA and microsatellite loci. Molecular Ecology 6:243-253. Pemberton, J. M., J. Slate, R Bancroft, and J. A. Barrett. 1995. Nonamplifying alleles at microsatellite loci: a caution for parentage and population studies. Molecular Ecology 4:249-252. Piertney, S. B., and J. F. Dallas. 1997. Isolation and characterization ofhypervariable the red grouse Lagopus lagopus scoticus. Molecular Ecology 6:93-95. microsatellites in Quinn, T. W. 1992. The genetic legacy of Mother Goose - phylogeographic patterns of lesser snow goose Chen carulescens carulescens maternal lineages. Molecular Ecology 1:105-117. Quinn, T. W., and B. N. White. 1987. Identification of restriction- fragment-length polymorphisrns in genomic DNA of the lesser snow goose (Anser carulescens carulescens). Molecular Biology and Evolution 4:126-143. . Sambrook, J., E. F. Fritsch, and T. Maniatis. 1989. Molecular Cloning: a Laboratory Manual, 2nd edition. Cold Spring Harbor Laboratory Press, New York, NY, USA. Schneider, S., J. Kueffer, D. Roessli, and L. Excoffier. 1997. Arlequin ver. l.1: software for population genetic data analysis. Genetics and Biometry Laboratory, University of Geneva, Switzerland Takezaki, N., and M. Nei. 1996. Genetic distances and reconstruction of phylogenetic trees from microsatellite DNA. Genetics 144:389..:399. . Tamura, K. 1992. Estimation of the number of nucleotide substitutions when there are strong transition-transversion and G+C content biases. Molecular Biology and Evolution 9:678-687. Tjur, T 1998. Nonlinear regression, quasi likelihood, and overdispersion The American Statistician 52:222-227. in generalized linear models. Young, J. R 1994. Sexual selection of sage grouse. Dissertation, Purdue University, West Lafayette, lN, USA. Young, J. R, J. W. Hupp, J. W. Bradbury, and C. E. Braun. 1994. Phenotypic divergence of secondary sexual traits among sage grouse, Centrocercus urophasianus, populations. Animal Behavior 47:1353-1362. �89 Table 2.1. Polymorphism of microsatellite loci among nine populations of sage grouse in Colorado. Mean Heterozygosity Population Mean sample size per locus (SO) Mean#of alleles per locus (SO) Polymorphic loci (%) Observed (SO) Expected from HdyWbg(SD) Gunnison Basin 28.5 (0.5) 3.8 (1.4) 75 0.386 (0.123) 0.374 (0.120) Crawford 17.3 (0.6) 2.3 (0.6) 75 0.299 (0.138) 0.297 (0.151) Dry Creek 17.5 (1.6) 2.5 (0.6) 50 0.179 (0.135) 0.283 (0.177) Dove Creek 14.5 (0.3) 1.8 (0.5) 50 0.193 (0.135) 0.221 (0.142) Cold Springs 20.5 (0.6) 5.5 (2.5) 100 0.631 (0.118) 0.611 (0.114) Blue Mountain 21.5 (1.2) 6.5 (3.2) 100 0.596 (0.120) 0.600 (0.144) North Park 22.8 (1.0) 5.5 (2.2) 100 0.643 (0.080) 0.619 (0.098) Middle Park 19.3 (0.8) 5.5 (1.6) 100 0.701 (0.089) 0.639 (0.078) Eagle 20.3 (0.8) 5.5 (2.5) 100 0.748 (0.145) 0.636 (0.103) Small-bodied Large-bodied Table 2.2. Significance (P < 0.005) of pairwise population Fsr tests for microsatellite data from sage grouse in Colorado. Pairs of populations significantly different are shown by significantly different are shown by-. Small-bodied Gunnison Basin Crawford Crawford those not Large-bodied Dry Creek Dove Creek + + Dry Creek + and Dove Creek + + Cold Springs + + + + Blue Mountain + + + + North Park + + + + , Cold Springs Blue Mountain North Park Eagle �90 Table 2.3. Significance (P < 0.005) of pairwise population F sr tests for mtDNA sequencing data from sage grouse in Colorado. Pairs of populations significantly different are shown by + and those not significantly different are shown by-. Large-bodied Small-bodied Gunnison Basin Crawford Dry Creek Dove Creek Crawford + Dry Creek + + Dove Creek + + Cold Springs + + + + Blue Mountain + + + + North Park + + + + Eagle + + + + Middle Park + + + + Cold Springs Blue Mountain North Park Eagle Table 2.4. AMOV A design and results for mtDNA analysis of nine populations of sage grouse in Colorado. Source of variation Sum of Squares Variance components 584.06 5.88 64.84 7 43.57 0.15 1.63 Within populations 192 584.14 3.04 33.53 Totals 200 1211.77 9.07 elf Among groups Among groups, within populations P~~eofVariation �91 Park Blue Mountain -bodied sage grouse Crawford Gunnison Basin Figure 2.1. Historic (left) and current (right) distribution oflarge and small-bodied sage grouse and sample locations in Colorado. The boundary between the ranges of large and small-bodied birds is shown on the right. �92 PROPORTION OF SHARED ALLELES Bluoliounllin CHORD DISTANCE Figure 2.2. Neighbor joining trees of microsatellite data using two different genetic distance measures. Small-bodied populations are identified by a box around the name. �93 Eagle (26) Cold Springs (25) B L Z "C H " ... : ...... : ::~. ::..:w...$'1;'''.':' ,'.. "','.. ' . North Park (23) " H X " o .::::.: o B Blue Mountain (21) "F" Middle Park (21) C 0 L AL s ~ • .....:.,':: : -, ~.::,' C . . ", B . D o Dove Creek (13) A Gunnison Basin (40) G Dry Creek (17) Crawford (15) Figure 2.3. Distribution of 19 mtDNA haplotypes among nine populations of sage grouse in Colorado. Number in parentheses represents sample size for each population. TAMURA Figure 2.4. Neighbor joining tree of mtDNA genetic distances calculated using allele frequencies and haplotype distances (Tamura 1992). Small-bodied populations are identified by a box around the name. �Appendix 2A. Distribution of alleles (reported as fragment length in base pairs) for four microsatellite loci and mtDNA haplotypes among nine populations of sage grouse in Colorado. The first four populations are small-bodied sage grouse and the last five populations are large-bodied sage grouse. Table 2A.I. Allele distributions for microsatellite LLSTI among nine populations of sage grouse in Colorado. Crawford Gunnison Drv Creek Cold Springs Dove Creek Blue Mountain 154 MP1 154 154 EG1 154 154 NP2 154 163 MP2 154 157 EG2 154 154 NP3 154 163 MP3 154 154 EG3 154 157 154 154 NP4 154 163 MP4 154 157 EG4 154 157 154 157 . NPS MP5 154 154 EG5 157 157 BM7 154 154 NP6 154 154 MP6 154 154 EG6 154 154 154 BM8 154 154 NP7 154 163 MP7 154 154 EG7 154 154 154 157 BM9 154 154 NP8 154 154 MP8 154 154 EG8 154 154 CS9 154 154 BM10 154 154 Npg 154 154 Mpg 154 163 EG9 154 157 154 CS10 154 154 BM11 154 157 NP10 154 154 MP10 154 CS11 154 154 BM12 154 154 NP11 154 157 MP11 154 154 EG11 154 154 154 CS12 154 157 BM13 154 157 NP12 154 154 MP12 151 157 EG12 154 157 154 154 CS13 154 154 BM14 154 157 NP13 154 157 MP13 154 157 EG13 154 154 DVCF2 154 154 CS14 154 157 BM15 154 157 NP14 154 157 MP14 154 154 EG14 154 157 DVCF3 154 154 CS15 154 157 BM16 154 157 NP15 154 154 MP15 154 154 EG16 154 154 CS16 154 154 BM17 154 154 NP16 154 154 MP16 151 157 EG18 154 154 BM18 154 154 NP17 154 157 MPH 154 157 EG19 154 157 154 CR1 154 157 DYC1 154 154 DVC1 154 154 CS1 GB2 154 154 CR2 154 154 DYC2 154 154 DVC2 154 154 CS2 154 154 BM3 GB3 154 154 CR3 154 154 DYC3 154 154 DVC3 154 154 CS3 154 157 BM4 154 154 GB4 154 157 CR4 154 154 DYC4 154 154 DVC4 154 154 CS4 154 154 BM5 GB5 154 157 CR5 154 154 DYC5 154 154. 154 154 CS5 154 154 BM6 GB6 154 154 CR6 154 157 DYC6 154 154 ovcs ovce 154 154 CS6 154 154 GB7 154 157 CR7 154 154 DYC7 154 154 DVC7 154 154 CS7 154 GB8 154 157 CR8 154 157 DYC8 154 154 DVC8 154 154 CS8 GB9 154 154 CR9 154 154 DYC9 DVC9 154 154 GG1 154 154 CR10 154 154 DYC10 154 154 DVC10 154 GG2 154 154 CR11 154 157 DYC11 154 154 DVC11 154 GG3 154 157 CR12 154 157 DYC12 154 154 DVC12 154 GG4 157 157 CR13 154 154 DYCF1 154 154 DVCF1 CR14 154 154 DYCF2 154 154 154 157 DYCF3 GG6 154 154 CR15 GG7 154 157 CR16 GG8 154 157 CEF1 GG9 154 154 FM2 GG10 154 157 FM3 GG11 154 154 GG12 154 154 GG13 154 154 GG14 DYCF4 154 154 154 157 154 154 CS17 DYCF5 154 154 CS18 154 157 BM19 154 154 NP18 154 154 MP18 154 154 EG20 157 157 DYCF7 154 154 CS19 154 154 BM20 154 154 NP19 154 154 MP19 151 154 EG21 154 157 FM4 DYCF8 154 154 CS20 154 157 BM21 154 154 NP20 154 154 MP20 154 154 EG22 154 154 FM5 DYCF9 154 154 CS21 154 154 BM22 154 154 NP21 154 154 MP21 154 157 EG23 154 154 DYCF10 154 154 CS22 BM23 154 154 NP22 154 157 EG24 154 154 CS23 BM24 154 154 NP23 BM25 154 154 NP24 154 157 NP25 154 157 154 154 CS24 154 157 GG16 154 157 CS25 157 157 GG17 154 157 CS26 154 157 GG18 154 154 154 157 GG20 154 154 EG10 DYCF6 GG15 GG19 Eagle Middle Park 154 154 154 154 GG5 North Park NP1 BM2 GB1 \0 ~ �Table 2A.2. Allele distributions for microsatellite LLSD8 among nine populations of sage grouse in Colorado. Crawford Gunnison Drv Creek Dove Creek Cold Springs Blue Mountain GB1 143 143 CR1 143 143 DYC1 143 143 DVC1 143 143 CS1 137 137 BM2 GB2 137 143 c~ 143 143 DYC2 143 143 DVC2 143 143 CS2 137 143 BM3 GB3 143 143 CR3 143 143 DYC3 143 .143 DVC3 143 143 CS3 143 157 BM4 GB4 143 143 CR4 143 143 DYC4 143 143 DVC4 143 143 CS4 137 143 137 157 GB5 143 143 CRS 143 143 DYC5 143 143 DVes 143 143 CS5 137 157 BMS BMO 137 GB6 143 143 CR8 143 143 DYCB 143 143 DVCB 143 143 CS6 143 157 BM7 GB7 143 143 CR7 143 143 DYC7 143 143 DVC7 143 143 CS7 137 157 GB6 GOO 143 143 CR8 143 143 DYCB 143 143 DVCB 143 143 CS6 143 143 143 CR9 143 143 DYC9 143 143 DVC9 143 143 CS9 GG1 143 143 CR10 143 143 DYC10 143.143 DVC10 143 143 CS10 GG2 143 143 CR11 143 143 DYC11 143 143 DVC11 143 143 GG3 143 143 CR12 143 143 DYC12 143 143 DVC12 143 143 GG4 143 143 CR13 143 143. DYCF1 143 ·143 DVCF1 143 143 GGS GGO 143 143 CR14 143 143 DYCF2 143 143 DVCF2 143 143 143 CR15 143 143 DYCF3 DVCF3 143 GG7 143 143 CR18 143 143 DYCF4 143' 143 GG8 143 143 FM1 DYCF5 CS17 GG9 143 143 FM2 DYCF6 CS18 137 GG10 143 143 FM3 DYCF7 143 ,143 CS19 GG11 143 143 FM4 DYCF8 143 143 GG12 143 143 FM5 DYCF9 143 '143 GG13 143 143 CRF1 GG14 143 143 CS23 GG15 143 143 CS24 GG16 143 143 CS25 GG17 143 143 CS26 GG18 143 143 GG19 143 143 GG20 143 143 143 143 DYCF10 Eagle Middle Park 137 143 MP1 137 143 EG1 137 143 NP2 137 157 MP2 137 157 EG2 137 143 NP3 137 137 MP3 137 143 EG3 137 143 NP4 137 157 MP4 137 157 EG4 137 157 137 NPS 137 157 143 157 EG5 137 157 137 137 NP6 137 157 MPS MP6 137 157 EG6 137 157 BM8 137 183 .NP7 137 143 MP7 137 143 EG7 137 137 137 BM9 137 157 NP8 137 137 MP8 137 157 EG8 137 137 157 157 BM10 143 143 NP9 143 1557 MP9 137 137 EG9 137 157 137 143 BM11 137 137 NP10 137 137 MP10 137 143 EG10 CS11 137 137 BM12 137 157 NP11 137 157 MP11 137 143 EG11 137 137 CS12 137 157 BM13 137 157 NP12 137 157 MP12 137 157 EG12 137 137 CS13 137 143 BM14 137 137 NP13 137 137 MP13 137 157 EG13 137 157 143 CS14 137 157 BM15 137 137 NP14 137 143 MP14 137 143 EG14 137 157 143 CS15 137 157 BM16 137 157 NP15 137 143 MP15 137 137 EG16 137 157 CS16 157 157 BM17 143 157 NP16 137 137 MP16 137 143 EG18 137 143 BM18 137 157 NP17 143 143 MP17 137 157 EG19 137 143 143 BM19 137 143 NP18 143 157 MP18 137 137 EG20 137 157 137 143 BM20 137 157 NP19 137 157 MP19 137 143 EG21 137 137 CS20 157 157 BM21 137 157 NP20 137 143 MP20 137 143 EG22 143 157 CS21 137 137 BM22 157 157 NP21 137 137 MP21 EG23 143 157 BM23 137 157 NP22 137 157 EG24 137 143 BM24 137 143 NP23 137 143 BM25 137 137 NP24 137 143 NP25 143 143 143 143 143 North Park NP1 CS22 137 \0 Vl �\0 0\ Table 2A.3. Allele distributions for microsatellite LLSD3 among nine populations of sage grouse in Colorado. Crawford Gunnison Dove Creek DrvCreek Cold Springs Blue Mountain North Park Middle Park Eagle GB1 135 137 CR1 133 133 DYC1 135· 135 DVC1 133 135 CS1 133 137 BM2 GB2 133 135 CR2 133 133 DYC2 133 135 DVC2 135 135 CS2 133 141 BM3 GB3 133 133 CR3 133 133 DYC3 135 135 DVC3 135 135 CS3 133 133 BM4 141 153 NP3 133 GB4 133 135 CR4 133 133 DYC4 133 135 DVC4 135 135 CS4 133 133 BM5 133 133 NP4 133 GBS 133 133 CR5 133 135 DYC5 135 135 ovcs 133 135 CS5 133 133 BM6 133 137 NPS 133 GB6 133 133 CR6 133 135 DYC6 133 133 DVC6 135 135 CS6 133 133 BM7 141 141 NP6 133 GB7 133 135 CR7 133 133 DYC7 135 135 DVC7 135 135 CS7 133 137 BM8 141 141 NP7 133 141 MP7 141 153 EG7 133 141 GB6 135 135 CR8 133 133 DYC6 135 135 DVC6 135 135 CS6 133 141 BM9 133 133 NP8 133 133 MP8 133 141 EG8 133 141 GB9 133 135 CR9 133 133 DYC9 135 135 DVC9 135 135 CS9 133 135 BM10 133 141 NP9 133 133 MP9 135 141 EG9 133 137 GG1 133 135 CR10 133 133 DYC10 133 135 DVC10 133 133 CS10 133 137 BM11 133 137 NP10 141 141 MP10 133 141 EG10 133 141 G02 133 135 CR11 133 133 DYC11 135 135 DVC11 133 135 CS11 133 133 BM12 133 141 NP11 133 133 MP11 133 133 EG11 133 137 003 133 133 CR12 133 133 DYC12 135 135 DVC12 135 135 CS12 133 133 BM13 133 141 NP12 141 153 MP12 133 141 EG12 133 133 G04 133 135 CR13 133 135 DYCF1 135 135 DVCF1 135 135 CS13 133 133 BM14 133 141 NP13 141 153 MP13 133 137 EG13 133 137 GG5 133 135 CR14 133 133 DYCF2 135 135 DVCF2 135 135 CS14 133 133 BM15 133 133 NP14 133 141 MP14 133 153 EG14 133 137 GG6 135 135 CR15 133 133 DYCF3 133 133 DVCF3 135 135 CS15 133 133 BM16 133 133 NP15 133 141 MP15 133 133 EG16 133 137 GG7 135 135 CR16 133 133 DYCF4 135 135 CS16 135 141 BM11 133 141 NP16 133 153 MP16 133 141 EG18 133 153 133 141 NP1 133 133 MP1 133 153 EG1 133 141 MP2 141 153 EG2 133 141 143 MP3 133 141 EG3 133 137 133 MP4 133 153 EG4 133 137 133 MP5 133 133 EG5 141 153 153 MP6 133 137 EG6 133 137 NP2 GG8 133 133 FM1 DYCF5 135 135 CS17 BM18 133 153 NP17 153 153 MP17 133 153 EG19 133 137 GG9 133 135 FM2 133 133 DYCF6 135 135 CS18 135 141 BM19 133 141 NP18 133 141 MP18 133 135 EG20 133 137 GG10 133 135 FM3 133 133 DYCF7 133 133 CS19 133 141 BM20 133 141 NP19 133 133 MP19 133 137 EG21 133 141 GG11 133 135 FM4 133 133 DYCF8 133 133 CS20 133 133 BM21 133 131 NP20 133 141 MP20 133 133 EG22 133 137 GG12 133 133 FM5 DYCF9 135 135 CS21 133 141 MP21 GG13 133 133 CRF1 DYCF10 GG14 133 G015 GG16 BM22 153 153 NP21 133 133 CS22 BM23 133 153 NP22 133 153 133 CS23 BM24 133 141 NP23 133 141 135 137 CS24 BM25 133 133 NP24 133 141 133 135 CS25 CS26 G017 133 135 GG18 133 137 G019 133 135 GG20 135 135 NP25 EG23 EG24 133 141 �Table 2A.4. Allele distributions for microsatellite LLSD4 among nine populations of sage grouse in Colorado. Crawford Gunnison Dove Creek DrvCreek GBl 191 191 CRl 191 203 DYCl GB2 191 201 CR2 203 203 DYC2 GB3 191 191 CR3 191 191 DYC3 191 205 DVC3 GBC 191 225 CR4 193 193 DYC4 203 207 DVC4 GBS 191 191 CR5 191 225 DYC5 215 215 DVC5 Gee 191 191 CR6 193 203 DYC6 215 215 GB7 203 225 CR7 191 203 DYC7 191 GBS 191 203 CR8 191 203 DYCB 191 GOO 191 215 CR9 191 203 DYC9 191 215 DVC9 GGl 191 191 CR10 191 191 DYC10 191 217 DVC10 191 203 Cold Springs Blue Mountain DVCl 199 201 CSl 195 197 BM2 DVC2 191 201 CS2 195 243 BM3 191 201 CS3 189 189 BM4 187 201 201 CS4 193 225 BM5 191 199 CS5 195 243 BMB 185 DVC6 201 201 CS6 189 195 BM7 191 DVC7 201 201 CS7 193 203 191 DVC6 201 201 CS6 195 215 191 201 CS9 189 191 201 CSl0 205 193 193 North Park Eagle Middle Park NPl 187 207 MPl 207 207 EGl 185 329 NP2 195 215 MP2 193 193 EG2 185 329 197 NP3 189 215 MP3 189 189 EG3 189 215 1111 289 NP4 191 205 MP4 193 215 EG4 189 245 329 NPS 203 215 MPS 183 209 EG5 215 ')137 281 329 NP6 205 245 MP6 EG6 187 ')137 BM8 189 357 NP7 203 245 MP7 193 205 EG7 219 297 BM9 ')137 321 NP8 193 205 MP8 183 189 EG8 197 BM10 189 195 NP9 203 207 MP9 187 187 391 BMll 189 191 NP10 207 207 MP10 187 309' EG10 215 ')137 187 225 EG9 GG2 191 191 CRll 191 225 DYC11 DVC11 191 201 CSll 195 243 BM12 195 195 NPll 205 215 MP11 189 189 EG11 GG3 191 203 CR12 193 203 DYC12 DVC12 191 191 CS12 197 391 BM13 189 195 NP12 205 205 MP12 183 207 EG12 GG4 1111 191 CR13 191 191 DYCFl 191 203 DVCFl 191 191 CS13 197 215 BM14 309 329 NP13 205 245 MP13 189 189 EG13 GG5 1111 215 CR14 193 203 DYCF2 191 203 DVCF2 199 201 CS14 197 215 BM15 189 193 NP14 205 205 MP14 189 193 EG14 GG6 215 225 CR15 193 203 DYCF3 CS15 197 215 BM16 223 ')137 NP15 191 245 MP15 189 193 EG16 195 ')137 GG7 191 191 CR16 203 203 DYCF4 191 217 CS16 183 197 BM17 185 323 NP16 195 205 MP16 183 195 EG18 195 215 GG8 191 205 FMl DYCF5 CS17 BM18 187 289 NP17 205 239 MP17 189 205 EG19 187 ')137 GG9 191 191 FM2 DYCFB CS18 187 193 BM19 NP18 191 205 MP18 189 193 EG20 187 193 GG10 191 205 FM3 DYCF7 217 217 CS19 197 255 BM20 187 187 NP19 187 189 MP19 187 189 EG21 189 ')137 GGll 193 219 FM4 DYCF8 191 191 CS20 189 255 BM21 189 329 NP20 209 245 MP20 EG22 195 ')137 GG12 191 191 FM5 DYCF9 CS21 BM22 NP21 MP21 EG23 187 205 GG13 191 203 CRFl DYCF10 CS22 BM23 NP22 EG24 187 215 GG14 191 203 CS23 BM24 NP23 GG15 191 191 CS24 BM25 NP24 GG16 191 191 CS25 GG17 193 215 CS':lJ3 GG18 191 191 GG19 191 205 GG20 191 191 DVCF3 NP25 \0 -..I �Table 2A.S. Distribution of mitochondrial DNA haplotypes among nine populations of sage grouse in Colorado. P012ulation A Gunnison Basin Crawford Dry Creek Dove Creek Cold Springs Blue Mountain Middle Park North Park Eagle B C D E G 38 2 2 15 4 6 H L Ha12lo~ S X Z M AC AD \0 00 AE AF 1 AL AM 8 11 3 AI 2 7 10. 1 8 1 . 1 7 9 2 1 2 4 5 6 3 2 2 15 4 2 1 1 3 1 1 1 2 1 3 Table 2A.6. P values of pairwise population F ST tests for microsatellites among all pairs of populations of sage grouse in Colorado. The first four populations are small-bodied and the last five populations are large-bodied. The average P value of comparisons among smaIl-bodied bird is 0.0055, of comparisons between large vs. small-bodied birds is 0.0000, and of comparisons among large-bodied birds is 0.1171. Crawford Dry Creek Dove Creek Cold Springs Blue Mountain North Park Population Gunnison Crawford 0..0.0.0.9 Dry Creek 0..0.0.73 0..0.00.0. Dove Creek 0..0.0.0.0. 0..0.0.0.0. 0..0.250. Cold Springs 0..0.0.0.0. 0..0.0.0.6 0..0.0.0.0. 0..0.0.0.0. Blue Mountain 0..0.0.0.0. 0..0.0.0.0. 0..0.0.0.0. 0..0.0.0.0. 0..1746 North Park 0..0.0.0.0. 0..0.0.0.0. 0..0.0.0.0. 0..0.0.0.0. 0..0.538 0..0.90.3 Middle Park 0..0.0.0.0. 0..0.0.0.0. 0..0.0.0.0. 0..0.0.0.0. 0..1666 0..2320. 0..10.58 Eagle 0..0.0.0.0. 0..0.0.0.0. 0..0.0.0.0. 0..0.0.0.0. 0..0.929 0..1672 0..0.239 Middle Park 0..0.641 �Table 2A.7. P values of pairwise population FSf tests for mtDNA among all pairs of populations of sage grouse in Colorado. The first four populations are small-bodied and the last five populations are large-bodied. The average P value of comparisons among small-bodied bird is 0.0124, of comparisons between large vs. small-bodied birds is 0.0000, and of comparisons among large-bodied birds is 0.3739. Crawford Dry Creek Population Gunnison Dove Creek Cold Springs Blue Mountain Crawford 0.0000 Dry Creek 0.0000 0.0005 Dove Creek 0.0020 0.0000 0.0717 Cold Springs 0.0000 0.0000 0.0000 0.0000 Blue Mountain 0.0000 0.0000 0.0000 0.0002 0.5890 North Park 0.0000 0.0000 0.0000 0.0000 0.3888 0.1832 Middle Park 0.0000 0.0000 0.0000 0.0000 0.4118 0.4737 0.0787 Eagle 0.0000 0.0000 0.0000 0.0000 0.5838 0.3796 0.1481 North Park Middle Park 0.5019 \0 \0 �100 �101 CHAPTER mREE POPULATION GENETICS OF GUNNISON SAGE GROUSE: IMPLICATIONS MANAGEMENT FOR INTRODUCTION The distribution and abundance of sage grouse in Colorado have been greatly reduced primarily due to habitat loss and fragmentation (Braun 1995). Sage grouse have been extirpated from 12 of the 27 counties in Colorado in which they occurred in the 1900's and populations in nine of the remaining 15 counties are thought to number less than 500 breeding birds (Braun 1995). Sage grouse in southwestern .Colorado have been the most severely impacted by destruction and fragmentation of habitat and, as a result, populations are small and isolated (Fig. 3.1). Recently, sage grouse in southwestern Colorado and southeastern Utah have been described as a new species of sage grouse, i.e., the Gunnison sage grouse (Centrocercus minimus) (Braun and Young 1995). This new species distinction was based on morphological and behavioral data (Hupp and Braun 1991, Young 1994, Young et al. 1994), and later supported by genetic data (Kahn et al. 1999, Chapter Two). Because Gunnison sage grouse are 33% smaller than all other sage grouse, I refer to them as either Gunnison sage grouse or more generally as small-bodied sage grouse. In Chapter Two I compared five large-bodied populations from northern Colorado with four small-bodied populations from southwestern Colorado using mitochondrial and nuclear markers. I found that smallbodied sage grouse have much less genetic diversity than large-bodied sage grouse and that there was markedly less gene flow among the four Gunnison sage grouse populations in southwestern Colorado than among the five populations of large-bodied sage grouse in northern Colorado. As a result, I argue that genetic data should be considered in management decisions for Gunnison sage grouse. This chapter uses the results from Chapter Two to address specific management implications for Gunnison sage grouse. Microsatellites are thought to be among the most powerful markers in population genetic studies today because of their high rate of mutation (Goldstein and Pollock 1997), which makes them extremely useful in distinguishing differences among populations thought to have low genetic diversity. Mitochondrial DNA, while rapidly evolving, is maternally inherited and thus, masks any effect of male dispersal. Because I was interested in the implications of relative amounts of gene flow and isolation, I refer only to the microsatellite data presented in Chapter Two. The specific objectives of this chapter were to examine the genetic diversity of each Gunnison sage grouse population and gene flow among these populations and to make management recommendations based on this information. STUDY AREA Gunnison sage grouse have an extremely limited range as they are restricted to southwestern Colorado and southeastern Utah. In southwestern Colorado, five populations have been studied (Gunnison Basin, Dove Creek, Dry Creek, Crawford, and Glade Park). Only one other population near Poncha Pass has recently been documented consistently by lek surveys. Samples were obtained from all five studied populations, but the Glade Park population was omitted due to insufficient sample size. The largest area of contiguous habitat and consequently the largest population occurs in the Gunnison Basin (Fig. 3.2) which supports approximately 2,600 birds in the breeding season and is thought to be a stable population (C. E. Braun, Colorado Division of Wildlife, unpublished data). The �102 Crawford population (Fig. 3.2) underwent a severe decline until 1994 but as a result of a habitat manipulation, has rebounded (Commons 1997, Commons et al. 1999) to approximately 175 birds (C. E. Braun, Colorado Division of Wildlife, unpublished data). The Dry Creek population (Fig. 3.2) is stable or declining with approximately 300 birds and the Dove Creek population (Fig. 3.2) is declining with approximately 75 birds (C. E. Braun, Colorado Division of Wildlife, unpublished data). The Gunnison Basin is an intermontane basin ranging in elevation from 2,300 to 2,900 m with several flat-topped mesas. The lower lands consist of broad, alluvial flood plains which abut major streams, and the uplands have moderate to steep slopes dissected by intermittent streams. Dominant vegetation includes mountain big sagebrush (Artemisia tridentata vaseyana), and black sagebrush (A. nova), intermixed with antelope bitterbrush (Purshia tridentata), and mountain snowberry (Symphoricarpos oreophilus) (Hupp and Braun 1989). The Crawford population in Montrose County is northwest of the Gunnison Basin separated by the Black Canyon of the Gunnison River and Black Mesa with elevations ranging from 1,968 to 2,952 m. Large mesas dominate the landscape around the town of Crawford and the area is bisected north to south by deep canyons. The dominant vegetation consists of a mix of mountain big sagebrush, black sagebrush, pinon pine (Pinus edulis), andjuniper (Juniperus spp.). At higher elevations, gambel oak (Quercus gambelii) and serviceberry (Amelanchier spp.) intermix with mountain big sagebrush (Commons 1997). The Dry Creek population in San Miguel County is southwest of Naturita and Norwood. This area is semi-arid high desert and ranges in elevation from 1,936 m in Dry Creek Basin to 2,385 mat Miramonte Resevoir. Dry Creek Basin is an old glacial bed which consists of flats, gently rolling hills, and deep drainages surrounded by mesas to the north, south, and east. This area is dominated by basin big sagebrush, low sagebrush (A. arbuscula), and winterfat (Eurotia lanata). The sagebrush dominated area north of Miramonte Reservoir is characterized by gently rolling hills and shallow drainages. The dominant vegetation surrounding the reservoir is black sagebrush and mountain big sagebrush with pinon pine and juniper invading (Commons 1997). The Dove Creek population is approximately 58 km southwest of Dry Creek isolated by the Dolores Canyon and Disappointment Valley. Dove Creek, in Dolores County, is semi-arid desert ranging in elevation from 2,020 to 2,303 m. The town of Dove Creek bisects the population with the northern area dominated by pinto bean, alfalfa, and wheat production, and the southern half used mainly for wheat production. Portions of the agricultural areas were enrolled in the Conservation Reserve Program (CRP) in the late 1980's. The area consists mostly of rolling hills and deep drainages bounded to the south by Squaw Canyon and to the northeast by Dolores Canyon. The northern part of the Dove Creek area consists of farmland and areas dominated by mountain big sagebrush, black sagebrush, gambel oak, pinon pine, juniper, ponderosa pine (P. ponderosa), mountain snowberry, serviceberry, antelope bitterbrush, and chokecherry (Prunus spp.). The southern Dove Creek area is dominated by farmland with small areas of basin and mountain big sagebrush, rabbitbrush (Chrysothamnus spp.), and broom snakeweed (Gutierrezia sarothrae). METHODS Complete methods for blood and feather sample collection, DNA extraction, PCR, and visualization are presented in Chapter Two. �103 Data Analysis Two measures of genetic distance were calculated for all pairs of small-bodied populations, the proportion of shared alleles (Bowcock et al. 1994), and chord distance (Cavalli-Sforza and Edwards 1967). Each genetic distance metric was calculated by determining the genetic distance for each locus and averaging across loci. I used a Mantel test to determine whether there was a relationship between genetic difference (FsrJ and geographic distance. To compare the amount of population subdivision within the small-bodied birds to the amount of population subdivision within the large-bodied birds, I calculated Wright's (1951) FST statistic for both groups of populations. Pairwise population FST significance tests were also conducted among all populations to test whether pairs of populations were statistically different. I documented the amount of genetic diversity per population by calculating mean heterozygosity and mean number of alleles per locus for each population, and by counting the number of unique alleles for each population. RESULTS The two measures of genetic distance show somewhat similar patterns (Table 3.1). Using the proportion of shared alleles distance, the smallest genetic distance was between Dove Creek and Dry . Creek and the largest was between Crawford and Dove Creek. The chord distance metric showed a similar ranking, yet found the pairs of Gunnison Basin/Crawford and Gunnison BasinlDry Creek to be closer than the Dove CreeklDry Creek pair that the other metric ranked as closest. Both metrics agreed that Gunnison BasinlDove Creek, CrawfordlDry Creek, and CrawfordIDove Creek should be ranked at four, five, and six. The relationship among these four populations was represented using a neighbor joining tree (Fig. 3.3). Both genetic distance measures produced similar trees with Gunnison Basin and Crawford clustering together and Dove Creek and Dry Creek clustering together. Separate Mantel tests for the large and small-bodied birds both revealed no significant isolation by distance for the small (P = 0.3127) or large-bodied bird (P = 0.4356) populations. A comparison ofF sr values between large and small-bodied birds revealed that the smallbodied birds were significantly more subdivided (Fsr = 0.2178, 95% CI 0.1230 - 0.3339) than the large-bodied birds (Fsr = 0.0266,95% CI -0.0016 - 0.0528). Pairwise population Fj, significance tests also indicated significant population subdivision (Table 3.2). A P value of 0.005 was used to indicate statistical significance because of the multiple comparisons nature of the analysis. As would be expected for comparisons of populations from different species, all small vs. large-bodied populations were significantly different. There were no differences between pairs of large-bodied populations suggesting substantial gene flow among the five large-bodied populations. Only two pairs of populations within the small-bodied birds were not significantly different at P = 0.005 (Dry Creek and Gunnison Basin, P = 0.0073; Dry Creek and Dove Creek, P = 0.025) suggesting isolation and reduced gene flow among the small-bodied birds. In comparing the genetic diversity oflarge and small-bodied sage grouse, the small-bodied sage grouse populations had less genetic diversity, reduced heterozygosity and fewer polymorphic loci (Table 3.3). The amount of diversity of Gunnison sage grouse populations varies substantially with the Gunnison Basin having the highest number of unique alleles, the highest mean number of alleles per locus and the highest mean heterozygosity. The Dove Creek population had the fewest number of unique alleles, the lowest mean number of alleles per locus, and the second lowest heterozygosity. �104 DISCUSSION My comparison of large and small-bodied sage grouse in Colorado has shown that smallbodied Gunnison sage grouse populations are isolated with relatively little gene flow among populations and much less genetic diversity than the large-bodied sage grouse. Furthermore, three of the four Gunnison sage grouse populations are small, and at risk of extinction. Together these factors provide evidence suggesting that the viability of at least three of the Gunnison sage grouse populations should be addressed. There has been much concern about the viability of small populations and how it might be affected by demographic, environmental, and genetic stochasticity, as well as catastrophes (Shaffer 1981, Soule 1987). Although minimum viable population sizes vary enormously among different species, it is generally thought that populations smaller than a few hundred individuals warrant at least investigation into possible negative effects that accompany small populations (Shaffer 1987). The persistence of wild populations is usually influenced more by ecological effects (such as the direct effects of catastrophes and environmental and demographic stochasticity) than by genetic effects. Yet when wild populations are reduced to small populations by artificial means such as habitat destruction, genetic factors and their interaction with ecological factors become increasingly important(Lande 1995). Historically, Dove Creek, Dry Creek, and Crawford all had much larger populations which were somewhat connected through more contiguous areas of sagebrush habitat (Fig. 3.1). It is Braun's (1995) assertion that clearing of sagebrush for cultivated crops, highway construction, ranch development, powerline placement, reservoir construction, and other facets of human settlement have resulted in fragmentation and loss of sagebrush habitats in southwestern Colorado leading to the current isolation of these populations which is consistent with the relatively low amounts of gene flow documented in this dissertation. This human-induced reduction in population sizes of Gunnison sage grouse leads me to believe that the Dove Creek, Dry Creek, and Crawford populations are at risk from the direct effects of catastrophes, environmental and demographic stochasticity, and, potentially, to the effects of inbreeding. Being a lek breeding species, sage grouse have less genetic diversity than other non-lekking grouse (Leberg 1991, Young 1994). Because only approximately 10 - 20 % of males on leks actually breed and up to 80% of all matings on a lek each year are by one or two males (Wiley 1973, Vehrencamp et al. 1989,1. R Young, Western State College, unpublished data) the effective size of sage grouse populations is likely much lower than the actual population size. Many equations exist to quantify effective population size which take into account different scenarios. A simple equation for effective population size which takes into account unequal sex ratios is described by Hartl and Clark (1989) as Ne= 4( Nmales)( /&ma1es) NmalU+ Ntemales . This is a simplified model for calculating effective population size which assumes discrete, non overlapping generations, and accounts only for differences in sex ratio. However, it can be used to provide an idea of the reduction in effective population size of sage grouse due to the lek breeding system. If a breeding population has 300 birds, 100 males and 200 females (the sex ratio for sage grouse is generally two females to one male), the effective population size can be calculated to be between 38 and 73 birds depending on whether it is assumed that 10 or 20% of the males breed. Either way, this is a substantial reduction from the natural population size of 300. �105 It is highly debated whether reduced genetic variation reduces the viability of a population. Avise (1994) warns that caution should be used in interpreting low variation in populations for a variety of reasons including knowledge that at least a few successful, widespread species have low genetic variation; in some endangered species like the elephant seal, (Mirounga angustirostris) (Bonnel and Selander 1974), lack of genetic variation has not seemed to seriously inhibit population recovery, and the effect of inbreeding on fitness differs widely among species with some being highly affected and some seemingly unaffected (Price and Waser 1979, Ralls and Ballou 1983, Ralls et a1. 1988, Laikre and Ryman 1991). Lande (1988) argues that low reproductive output in small populations may be due to non-genetic factors such as the Allee effect (Andrewartha and Birch 1954). Furthermore, small populations, (regardless of the amount of genetic variation) are at risk of extinction because of demographic fluctuations (Gilpin and Soule 1986). Because of such factors, Lande (1988) argued that, for conservation plans, demographic and behavioral concerns should be a higher priority than genetic concerns. Other authors, however, are adamant that genetic variation is extremely relevant to the health and viability of populations and that it must be addressed and monitored in management plans (O'Brien and Evermann 1988, Quattro and Vrijenhoek 1989). Examples of how inbreeding have affected some characters offitness include the survival, growth, early fecundity, and developmental stability of the Sonoran topminnow (Poeciliopsis occidentalis sonorensis) (Quattro and Vrijenhoek 1989), fertility and hatching success of greater prairie-chickens (I'ympanuchus cupido) (Westemeier, et a1. 1998), pair bonding behavior in wolves (Canis lupus) (Wayne et a1. 1991), and high instances of abnormal sperm in cheetahs (Acinonyxjubatus) (O'Brien and Evermann 1988). Further, O'Brien and Evermann (1988) found low variation in the major histocompatibility complex (MIIC) in cheetahs and documented a 50 - 60% mortality in cheetahs over a three year period due to a corona virus. They advocate that genetically depauperate populations face enhanced susceptibility to infectious disease or parasitic agents. MANAGEMENT IMPLICATIONS Acknowledging Lande's (1988) assertion not to emphasize genetic concerns over other concerns and O'Brien and Evermann's (1988) belief that genetic considerations are vital to conservation, I advocate addressing both genetic concerns and other ecological concerns (habitat loss, fragmentation) in management of Gunnison sage grouse. This echos Soule and Mills' (1998) idea of not isolating genetics from other factors affecting small populations, but rather addressing all factors (demographic, genetic, habitat-based) together in the preservation of small populations. I believe every attempt should be made to prevent further loss of habitat in any area inhabited by Gunnison sage grouse. Further, I believe the habitat of Gunnison sage grouse should be managed according to established general procedures (Braun et al, 1977), with case by case specifications to address specific problems in different areas (Commons 1997). Conservation plans have been developed for the Gunnison Basin, Dove Creek, Dry Creek, and Crawford populations, and working groups are developing plans for Glade Park and Poncha Springs. The purpose of these community-based conservation working groups is to provide coordinated management across jurisdictional/ownership boundaries and to develop community-wide support necessary to assure the survival of Gunnison sage grouse. With the support of the conservation working groups, population sizes in Dove Creek, Dry Creek, and Crawford should stabilize andlor increase, lessening the extinction risk from demographic and environmental stochasticity. Because Gunnison sage grouse overall have much lower genetic diversity than other sage grouse, I feel that a management plan for Gunnison sage grouse should address genetic concerns. �106 Within the Gunnison sage grouse, the Gunnison Basin population has the most diversity followed by Dry Creek, Crawford, and Dove Creek. Because these populations are isolated and because there is little gene flow (relative to the amount of gene flow among large-bodied birds), I suggest translocating birds from the Gunnison Basin into at least the Dove Creek population, and potentially into the Crawford and Dry Creek populations. The Dove Creek population is of most concern with the lowest genetic diversity (only seven unique alleles and an average of 1.8 alleles per locus) and the smallest actual population size (approximately 75 birds) and effective population size (11 birds, as calculated from equation one, with 50 females, 25 males, and 10% of the males breeding). I advocate translocating four to six females from the Gunnison Basin population into the Dove Creek population every few years to increase the genetic variability in Dove Creek. My recommendation to translocate females rather than males is because only a few males actually breed in each population whereas most females breed. The number of birds to translocate is based on the idea that loss of genetic variability can be countered by immigration from an outside population, assuming it is large enough to maintain genetic variability. Immigration from other populations also has the effect of slowing genetic drift and fixation of alleles (Lande 1995). Wright (1931, 1951, 1969) has shown that analysis of the 'island model' indicated that immigration of a few individuals per generation will prevent loss of genetic variability. I believe that if four to six birds are translocated, then two or three will likely survive the translocation and successfully reproduce. Similar translocations from the Gunnison Basin to Crawford or Dry Creek are advised, yet are not as high of a priority. Natural gene flow is more likely to occur between Gunnison Basin and Crawford since they are geographically closer and there is evidence that gene flow between the Gunnison Basin and Crawford is higher than between the Gunnison Basin and either Dry Creek or Dove Creek (fable 3.1, Fig. 3.2). Some may argue that the translocation of birds from the Gunnison Basin to Dove Creek may have a negative impact on the Dove Creek population because of potential outbreeding effects (Dove Creek birds may be highly adapted to the Dove Creek area and birds from the Gunnison Basin would lower the overall fitness of the population). Such effects have been well documented in fisheries management. For example, Chum salmon (Onochynchus ketal eggs introduced from a foreign stock resulted in a total decline in stock size to 5% of the original number (Altukhovand Salmenlova 1987). I do not believe this will be the case for sage grouse for several reasons. First, habitat destruction has caused Gunnison sage grouse populations to be isolated. Historically, populations were much larger and well connected (Fig. 3.1). It is conceivable even today, however, that interchange among these four populations could occur naturally as sage grouse have been documented to move up to 114 km between autumn and spring (Berry and Eng 1985). My data show some population differentiation (Table 3.2) and less gene flow relative to large-bodied sage grouse, yet movements between Dry Creek and Dove Creek and between Gunnison Basin and Crawford are not impossible. I do not believe these populations have been truly isolated long enough to develop real genetic differences that might be associated with outbreeding depression. Second, the birds that I advocate translocating are from wild populations, not birds in captivity that have been manipulated by humans. Third, ifbirds in Dove Creek were extremely well adapted to their habitat and birds from the Gunnison Basin were not adapted to that environment, the birds from the Gunnison Basin would be at a selective disadvantage and would be outcompeted by the Dove Creek birds. Because I only advocate moving a few birds into an area (contrasted with the huge numbers of eggs moved in fishery studies), I do not believe this would have a negative impact on the population. The decision whether to translocate birds to increase the genetic diversity in a population like Dove Creek is difficult. There are several scenarios to consider in this situation. First, genetic diversity may not be associated with fitness at all, in which case translocating birds has no effect on the fitness of the population. Second, birds in an area like Dove Creek might be highly adapted to the environment and better suited to succeed than the few birds moved from an area like the Gunnison �107 Basin. Although I believe this scenario is least likely, even if it were true, the few birds translocated would quickly be selected against and their genes eliminated from the population. This should have little or no negative impact on the population. Finally, if genetic diversity is associated with fitness or with the ability for birds to adapt to future environmental changes, then translocating birds into the population will have a positive impact on the population. The low genetic diversity among all Gunnison sage grouse and the extremely low effective population sizes in Dove Creek, Dry Creek, and Crawford are causes for concern. Maintaining/improviog habitats and translocating birds may be insufficient to assure viability of this species. Characteristics of fitness as they relate to genetic diversity must be more closely examined. Research on reproductive features (sperm function, egg normality), parasite load, and disease resistance (e.g., MHC) should be conducted, comparing both within the small-bodied Gunnison sage grouse and between the large and small-bodied sage grouse. While genetic concerns may not be the highest priority for Gunnison sage grouse conservation and management, I believe that along with other issues (habitat loss and quality) they should at least be considered. An overall management plan including monitoring and maintaining genetic diversity, preventing future habitat loss and fragmentation, and sound management of current populations and habitat must be implemented to assure the viability of Gunnison sage grouse. LITERATURE CITED Altukhov, y. P., and E. A. Salmenkova 1987. Stock transfer relative to natural organization, management, and conservation offish populations. Pages 333-343 in N. Ryman and F. Utter, editors. Population Genetics and Fishery Management. University of Washington Press, Seattle, W A, USA. . Andrewartha, H. G., and L. C. Birch. 1954. The distribution and abundance of animals. University of Chicago Press, Chicago, IL, USA. Avise,1. C. 1994. Molecular markers, natural history and evolution. Chapman and Hall, New York, NY, USA. Beny,1. D., and R L. Eng. 1985. Interseasonal movements and fidelity to seasonal use areas by female sage grouse. Journal of Wildlife Management 49:237-240. Bonnel, M. L., and R K. Selander. 1974. Elephant seals: genetic variation and near extinction. Science 184:908-909. Bowcock, A. M., A. Ruiz-Linares, 1. Tomfohrde, E. Minch, 1. R Kidd, L. L. Cavalli-Sforza 1994. High resolution of human evolutionary trees with polymorphic microsatellites. Nature 368:455-457. Braun, C. E.1995. Status and distribution of sage grouse in Colorado. Prairie Naturalist 27: 1-9. Braun, C. E., and 1. R Young. 1995. A new species of sage grouse in Colorado. Joint Meeting Wilson Ornithological Society and the Virginia Society of Ornithology, Abstract #23. May 4-7, 1995, Williamsburg, VA, USA. Braun, C. E., T. Britt, and R O. Wallestad. 1977. Guidelines for maintenance of sage grouse habitats. Wildlife Society Bulletin 5:99-106. Cavalli-Sforza, L. L, and A. W. F. Edwards. 1967. Phylogenetic analysis: models and estimation procedures. American Journal of Human Genetics 19:233-257. Commons, M. L. 1997. Movement and habitat use by Gunnison sage grouse (Centrocercus minimus) in southwestern Colorado. Thesis, University of Manitoba, Winnipeg, Canada �108 Commons, M. L., R K Baydack, and C. E. Braun. 1999. Sage grouse response to pinon-juniper management. Pages 000-000 in R E. Stevens and S. B. Monsen, editors. Proceedings of ecology and management of pinon-juniper communities within the interior western United States. Department of Agriculture, Forest Service, RM 000. Gilpin, M. E., and M. E. Soule. 1986. Minimum viable populations: processes of species extinction. Pages 19-34 in M. E. Soule editor. Conservation biology: the science of scarcity and diversity. Sinauer Associates, Sunderland, MA, USA. Goldstein D. B., and D. D. Pollock. 1997. Launching microsatellites. Journal of Heredity, 88:335-342. Hartl, D. L., and A. G. Clark. 1989. Principles of population genetics. Sinauer Associates, Sunderland, MA, USA. Hupp.T, W., andC. E. Braun. 1989. Topographic distribution of sage grouse foraging in winter. Journal of Wildlife Management 53:823-829. Hupp.T, W., and C. E. Braun. 1991. Geographic variation among sage grouse in Colorado. Wilson Bulletin 103:255-261. Kahn N. W., C. E. Braun, J. R Young, S. Wood, D. R Mata, and T. W. Quinn. 1999 Molecular analysis of genetic variation among large and small-bodied sage grouse using mitochondrial control region sequences. Auk, (in press). Laikre, L. and N. Ryman. 1991. Inbreeding depression in a captive wolf (Canis lupus) population. Conservation Biology 5:33-40. . Lande, R 1988. Genetics and demography in biological conservation. Science 241:1455-1460. Lande, R 1995. Breeding plans for small populations based on the dynamics of quantitative genetic variance. Pages 318 - 341 in 1. D. Ballou, M. Gilpin, and T. Foose, editors. Population management for survival and recovery: analytical methods and strategies in small population conservation. Columbia University Press, New York, NY, USA. Leberg, P. L. 1991. Influence of fragmentation and bottlenecks on genetic divergence of wild turkey populations. Conservation Biology 5:522-530. O'Brien, S.· 1. and 1. F. Evermann. 1988. Interactive influence of infectious disease on genetic diversity of natural populations. Trends in Ecology and Evolution 3:254-259. Price, M. V, and N. M. Waser. 1979. Pollen dispersal and optimal outcrossing in Delphinium nelsoni. Nature 277:294-297. Quattro, J. M, and R C. Vrijenhoek. 1989. Fitness differences among remnant populations of the endangered Sonoran topminnow. Science 245:976-978. Ralls, K, and 1.Ballou. 1983. Extinction: lessons learned from zoos. Pages 35-56 in C. M. Schonewald-Cox, S. M. Chambers, B. MacBryde, and L. Thomas editors. Genetics and conservation: a reference for managing wild animal and plant populations. Benjamin Cummings, Menlo Park, CA, USA. Ralls, K, 1.D. Ballou, and A. Templeton. 1988. Estimates of lethal equivalents and the cost of inbreeding in mammals. Conservation Biology 2:185-193. Shaffer, M. L. 1981. Minimum population sizes for species conservation. Bioscience 31: 131-134. Shaffer, M. L. 1987. Minimum viable populations: coping with uncertainty. Pages 69 - 87 in M. E. Soule, editor. Viable populations for conservation. Cambridge University Press, New York, NY, USA. Soule, M. E. 1987. Viable populations for conservation. Cambridge University Press, New York, NY, USA. Soule, M. E. and L. S. Mills. 1998. No need to isolate genetics. Science 282:1658-1659. Vehrencamp, S. L., 1.W. Bradbury, and R M. Gibson. 1989. The energetic cost of display in male sage grouse. Animal Behavior 38:885-896. �109 Wayne, R K., N. Lehman, D. Girman, P. J. P. Gogan, D. A. Gilbert, K. Hansen, R O. Peterson, U. S. Seal, A. Eisenhawer, L. D. Mech, and R J. Krumenaker. 1991.Conservation genetics of the endangered Isle Royale gray wolf. Conservation Biology 5:41-51. Westemeier, R L, J. D. Brawn, S. A. Simpson, T. L. Esker, R W. Jansen, J. W. Walk, E. L. Kershner, 1. L. Bouzat, and K. N. Paige. 1998. Tracking the long-term decline and recovery of an isolated population. Science 282:1695-1698. Wiley, R H. 1973. Territoriality and non-random mating in sage grouse, Centrocercus urophasianus. Animal Behavior Monographs. 6:85-169. Wright, S. 1931. Evolution in Mendelian populations. Genetics 16:97-159. Wright, S. 1951. Evolution and genetics of populations. Annals of Eugenics 15:323-354. Wright, S. 1969. Evolution and genetics of populations, the theory of gene frequencies. Volume 2. University of Chicago Press, Chicago, IL, USA. Young, J.R 1994. Sexual selection of sage grouse. Dissertation, Purdue University, West Lafayette, IN, USA. Young,1. R, J. W. Hupp, 1. W. Bradbury, and C. E. Braun. 1994. Phenotypic divergence of secondary sexual traits among sage grouse, Centrocercus urophasinus, populations. Animal Behavior 47:1353-1362. �110 Table 3.1. Two different genetic distance measures calculated from four microsatellite loci for all pairs of small-bodied sage grouse populations in Colorado. The number in parentheses represents the rank among all populations, one being the pair of populations with the smallest genetic distance and six being the pair of populations with the largest genetic distance. Population Pairs Proportion of Shared Alleles Chord Distance Gunnison Basin and Crawford 0.252 (2) 0.250 (1) Gunnison Basin and Dry Creek 0.264 (3) 0.280 (2) Gunnison Basin and Dove Creek 0.344 (4) 0.367 (4) Crawford and Dry Creek 0.411 (5) 0.384 (5) Crawford and Dove Creek 0.456 (6) 0.471 (6) Dry Creek and Dove Creek 0.188 (1) 0.349 (3) Table 3.2. Significance (P < 0.005) of pairwise population FST tests for the microsatellite data for sage grouse populations in Colorado. Pairs of populations significantly different are shown by + and those not significantly different are shown by -. Large-bodied Small-bodied Gunnison Basin Crawford Crawford Dry Creek Dove Creek + + Dry Creek Dove Creek + + Cold Springs + + + + Blue Mountain + + + + North Park + + + + Eagle + + + + Middle Park + + + + Cold Springs Blue Mountain North Park Eagle �III Table 3.3. Genetic diversity measures for each sampled population of sage grouse in Colorado. Population % Loci Polymorphic Mean # of alleles per locus Heterozygosity X SD X SD Unique Alleles (N) Large-bodied Cold Springs 100 5.5 2.5 0.631 0.118 22 Blue Mountain 100 6.5 3.2 0.596 0.120 26 Middle Park 100 5.5 2.2 0.701 0.089 22 North Park 100 5.5 1.6 0.643 0.080 22 Eagle 100 5.5 2.5 0.748 0.145 22 Gunnison Basin 75 3.8 1.4 0.386 0.123 15 Crawford 75 2.3 0.6 0.299 0.138 9 Dry Creek 50 2.5 0.6 0.179 0.135 10 .:Dove Creek 50 1.8 0.5 0.193 0.135 7 Small-bodied �112 Figure 3.1. Historic (top) and current (bottom) distribution of sage grouse and Gunnison sage grouse (lower left cut out) in Colorado. �113 LARG&BODffiDSAGEGROUSE OOVECREEK DRY CREEK GUNNISON BASIN Figure 3.2. Populations of sage grouse sampled in Colorado. �114 PROPORTION OF SHARED ALLELES Gunnison Basin CHORD DISTANCE Dove Creek Gunnisoo Basin ~M~ ~ )------DryCreek Figure 3.3. Neighbor joining trees of micro satellite data of Gunnison sage grouse populations using two different distance measures. �115 CHAPTER FOUR QUANTIFYING CHANGES IN SAGEBRUSH HABITAT IN SOUTHWESTERN COLORADO FROM THE MID-50'S TO THE MID-90'S INTRODUCTION Sage grouse, Centrocercus urophasianus, historically occurred in at least 15 states and three provinces (Aldrich 1963, Johnsgard 1973), they currently occupy only 11 states and two provinces (Braun 1998). In Colorado, the distribution and abundance of sage grouse has been dramatically reduced (Braun 1995). Sage grouse have been extirpated from 12 of the 27 counties in Colorado in which they occurred in the 1900's and populations in nine of the remaining 15 counties are thought to number less than 500 breeding birds (Braun 1995). Population declines appear to be related to habitat loss (conversion of big sagebrush, Artemisia trldentata, into farmland or housing developments), habitat degradation (heavy grazing, sagebrush removal, road and powerline development through sagebrush, and human disturbance), and habitat fragmentation (Braun 1995). Sage grouse habitat in southwestern Colorado, the range of the newly described Gunnison sage grouse, Centrocercus minimus (Braun and Young 1995), has been most severely impacted by these processes (Fig. 4.1). In winter, sage grouse are dependent solely on sagebrush leaves (primarily big sagebrush) for food (patterson 1952, Wallestad et al. 1975). Due to lack of a grinding gizzard, sage grouse cannot digest plant fiber well (Remington 1989) and, as a result, are dependent upon sagebrush because it retains nutritious leaves all winter. Thus, the loss of sagebrush habitat is likely linked to the decline of Gunnison sage grouse in southwestern Colorado. Historical records document the occurrence of six species of sagebrush in Colorado (James 1823). Cary (1911:246) described sagebrush to be "omnipresent on the higher plains of western Colorado and also in most of the higher mountain parks up to 10,000 feet". In southwestern Colorado, sagebrush areas included by Cary (1911) were: Debeque to Glenwood and Dotsero, Wolcott, Roaring Fork Valley to Aspen, Uncompahgre Plateau, Lone Cone, Lone Mesa, Naturita, Cerro Summit, Somerset, Sapinero, Gunnison, Creede, Poncha Pass, Buena Vista, Leadville, Hotchkiss, Saguache, Bayfield, Arboles, and McElmo Canyon. Rogers (1964) reported that all sagebrush areas listed by Cary (1911) still contained some amount of sagebrush in the early 1960's, yet due to human activities, many no longer were dominated by sagebrush. Human activities mentioned by Rogers (1964) included overgrazing, irrigation projects, and dry-farming. The distribution of sagebrush and sage grouse in the. early 1960's (Fig. 4.2) was documented by Rogers (1964). Braun (1995) compared the distribution of sage grouse in 1993-94 to the range of sage grouse described by Rogers in 1964. Braun (1995) reported extirpation of sage grouse from 12 of 17 counties in southwestern Colorado which once supported them. This led me to believe that sagebrush habitats in southwestern Colorado had been lost to other land uses. Changes in vegetation types and land uses have often been documented successfully using aerial photography. For example, the National Wetlands Inventory mapped large scale changes in wetland distributions (Tiner 1990, Dahl and Johnson 1991) using this technology. Other examples include documenting tree invasion into grasslands (Mast et al. 1997), monitoring land cover change of a heathland region (Csaplovics 1992), quantifying temporal changes in seagrass areal coverage. (Robbins 1997), and inventorying and monitoring arid rangeland vegetation (Knapp et al. 1990). I used aerial photographic analysis, to document and quantify changes in sagebrush-dominated habitats in southwestern Colorado which may be affecting the persistence of Gunnison sage grouse. �116 MEmODS Plot Selection I identified 10 areas in southwestern Colorado which in the early 1960's (Rogers 1964) contained sagebrush-dominated habitat. Rough polygons (Fig. 4.3) were digitized around the 10 sagebrush areas in a geographic information system (GIS). I constructed a grid of sampling plots (sampling frame) covering each of the 10 polygons, with each sampling plot being a square, 4 km on a side (16 krnvplot). Although I did not have prior data suggesting what the change in habitat might be, I did expect habitat loss. Thus, I chose to compute a total sample size based on a worst case scenario that the proportion of sagebrush habitat in the first time period was 0.5 (worst case because if it was higher or lower the precision would be greater). I decided that a coefficient of variation (CV) ofless than 10% on my estimate of habitat change would be acceptable for this study. I then calculated the appropriate sample sizes to achieve a CV of 4 or 10% of the estimate of the proportion of sagebrush habitat. The sample sizes calculated were 625 for a CV of 4% and 100 for a CV of 10%. I chose a target sample size of 200 since it was between 100 and 625 and the CV would be around 7%. Thus, my sampling fraction was approximately 9% (200 plots sampled from 2274 total plots). The number of plots to be sampled per stratum were calculated (rounding this number to the nearest integer) such that the sampling fraction was approximately 9% in each stratum. I then randomly chose the appropriate number of plots within each stratum to achieve a stratified random sampling design (Table 4.1). Because I rounded the number of plots to the nearest integer, the total number of plots sampled increased to 202. Aerial Photography Acquisition and Interpretation I attempted to obtain low level (between 1:20,000 and 1:30,000) black and white aerial photographs of each plot in the 1950's, 1970's, and 1990's. I chose my sampled plot size such that an entire plot could fit on one low level photo (occasionally a plot was covered by a group of photos from the same flight). Aerial photographs (either black and white film positive or color infrared) for all plots in the 1990's time period were obtained. Color infrared photos were used only when black and white photos were not available. Aerial photos from the early time periods were more difficult to obtain. For each plot, I developed a list of available photos (from different time periods) covering that plot and chose the earliest available photos for each plot. If there were photos approximately mid way between the earliest date and the 1990's date, I chose those photos as well. In most cases, photos for only two time periods could be obtained I did obtain photos from three time periods for 37 plots which allowed me to examine rates of habitat change over time. I omitted eight plots because there was insufficient photography covering those plots. Aerial photos were obtained from the U.S. Geological Survey (USGS) Eros Data Center. Each plot boundary was identified and traced onto a 1:24,000 7.5-minute USGS topographic quad map (or groups of maps if needed). From features on the quad map, the plot was identified on the corresponding photo (or group of photos). A photo adjacent (along the same flight line) to the one containing the plot was identified for use on a stereoscope to visualize the plot in three dimensions. Acetate was then overlaid and taped to the appropriate photo (or groups of photos). The plot was then photo-interpreted to identify sagebrush-dominated (big sagebrush> than 50%) areas. These areas were traced onto the acetate using a Koh-i-noor Rapidograph pen with a tip to draw lines no thicker than 0.25 m, using Rapidograph Rapidraw 3084-F ink in the drawing pen. Forty-three of the 194 plots �117 were ground truthed by first interpreting the photo, then going to the area on the ground and confirming its classification as sagebrush-dominated habitat. A zoom transfer scope was used to standardize the scale and georeference the data because the photos from different time periods were taken at different scales (photos taken at different elevations). A mylar sheet was taped to each quad map and the appropriate plot was traced onto the mylar correctly overlaying the plot traced onto the quad map. Each photo with interpretation was placed on the zoom transfer scope and focused to the appropriate scale so that features in the photo were lined up with features on the quad map. The interpreted sagebrush areas were traced onto the mylar sheet, removed from the quad map, scanned into a computer, and converted into a bitmap image using Adobe Photoshop. Each bitmap image was edited to correct anomalies such as closing polygons, deleting stray marks picked up by the scanner, and thinning polygon edges. The bitmap images were then imported into the GIS software Arc View (ESRI 1996) where total area of sagebrush, number of sagebrush polygons, and area and perimeter of each sagebrush polygon were calculated. Data Analysis I calculated the total amount of sagebrush-dominated habitat in each stratum and overall using standard methods for a stratified, simple random sampling design (Thompson 1992, Thompson 1997). " The total estimated amount of sagebrush, T was calculated as L T= I «s, j=l x~ whereL is the number of strata (10), Nj is the total number of plots in stratumj, and is the mean amount of sagebrush habitat in sampled plots from stratumj. The sampling variance of 1 was calculated as where nj is the number of plots sampled in stratumj, and s~is the sampling variance estimate for stratumj. From the estimated total area of sagebrush I estimated the proportion of area that represented sagebrush-dominated habitat in each stratum and also overall. I considered the most recent time period for each plot to be the late photo and the earliest time period for each plot to be the early photo. To determine the average time span between early and late photos, I calculated the average difference between the years of the early and late photos. I calculated the annual change in proportion of habitat between the early and middle photos, and between middle and late photos for the 37 plots with three time periods. Annual change r annual was calculated as 1 rannual = 1- (1- RperiOd)A where A is the time period in years and R riod is the estimated rate of change over a given time period. I then subtracted the two annual rat:S of change and tested whether this difference was different from zero. Because I did not find a significant difference between the annual rates of change from the two time periods (early to middle and middle to late), I assumed that annual rates of change were reasonably constant over the 35 year time period. �118 Photos representing each time period (early and late) were taken in different years across plots (e.g., the early time period could be represented by a photo from mid-40's to late-60's). This made it difficult to compare changes in habitat across plots. Thus, I chose a model-based approach to standardize the data to a common early and late year for comparisons across plots. I used the average early year (1958) as my standard early year and the average late year (1993) as my standard late year. Using the assumption of montonicity of change between time periods, I calculated the proportion of habitat available per plot in the standard early and standard late years using a logistic function log( 1 Pearly - log( )= aear/y Pearly Plale I-p ) = a late late where Pearly was the proportion of area on a plot which was sagebrush-dominated habitat in the early time period and Plate was the proportion of area per plot which was sagebrush-dominated habitat in the late time period. Computing a early and a late ,I then used the following equations aearly alale to solve for a and " b. = a + btearly = a + btlate I then set a specific year (e.g., t = 58), computed ass using ass = a + b(58) and then computed an estimated proportion of sagebrush in the given plot in year 58, following equation "PS8 = PS8 , using the 1 1+e (-ass)· Thus, for every plot I used the logistic function and estimated the proportion of sagebrush-dominated habitat in 1958 and in 1993. Similar standardization and projection to a given year are explained in Terrazas-Gonzalez (1997). To obtain better estimates of within stratum variance and confidence intervals on the amount of sagebrush for strata with small sample sizes (strata 3,4,5,6,8,9, and 10) I calculated the CV of the amount of sagebrush habitat in 1958 and 1993 for each stratum. Because CVs tend to be stable (Eberhart 1978) I calculated the average CV and used it as an estimate ofCV for strata with small sample sizes. This methodology, while not commonly used, is valid and documented in Carroll and Ruppert (1988) and Buckland et al. (1993). This allowed me to calculate the z 12 multiplier for a confidence interval using 175 degrees of freedom, instead of much smaller deg~ees of freedom if this procedure is not used (Tukey 1977). The actual confidence intervals around" the estimates of the amount of sagebrush in 1958 and 1993 for each stratum were calculated using a log transform approach (Burnham et al. 1987). This is primarily to assure that the lower bound of the confidence interval cannot be less than the actual amount of sagebrush seen per stratum, aj , calculated as �119 nJ aj = L e.,*1600 i=1 where nj is the number of plots in strata}, and Pij is the proportion of sagebrush habitat in plot i, stratum} (the quantity is multiplied by 1600 to give the area in ha). This quantijy, aj , was then subtracted fr.Qmthe estimated tot~ amount of sagebrush for a given stratum, to give a normalized lower limit ~ for that stratum (T. T. - a .)..... For ea~h stratum, this lower limit was then used to calculate upper and lower confiderice intervalS: Tu and ~ using the following equations 1j = i; = (fIC)+a and i; = (fC)+a where and " cv(T)= SErf) " . T-a Because it is logical that confidence intervals around the estimate of the amount of habitat lost could be negative, confidence intervals for this parameter were calculated in a traditional way, i.e., ± 1.96 (SE [loss]). RESULTS Habitat Loss The years of the early photos ranged from 1944 to 1976 and the late photos from 1988 to 1995. The average date for the early photos was 1958 (SD = 6.7) and 1993 (SD = 1.3) for the late photos. The average number of years between the early and late photos was 35.2 (SD = 6.7). A difference of approximately 35 years should reflect changes in sagebrush habitat. The difference in annual rate of change in habitat between the early to mid time periods for the 37 plots from three time periods and the mid to late time periods was not significantly different from zero (T= 0.83, P = 0.4124). Thirty-one of the 194 plots had no sagebrush-dominated habitat in either early or late photos. Of those plots with some amount of sagebrush in the early date, 10 plots had an increase in the amount of sagebrush and 153 had a decrease. Without standardizing to a given early and late year, the mean proportion of sagebrush habitat in the early years was 0.212 (SE = 0.016) and the mean proportion in the late years was 0.173 (SE = 0.015). This corresponds to 772,358 ha (SE = 59,307) in the early years and 630,274 (SE = 55,944) in the late years. This represents an 18% loss in sagebrush habitat between early and late years. After adjusting the data based on the logistic method, the mean proportion of sagebrush habitat available in 1958 was 0.2161 (SE = 0.0166) and in 1993 was 0.1734 (SE = 0.0154) (Table 4.2) which converts to 786,411 ha in 1958 and 630,725 ha in 1993 with a loss of 155,673 ha (95% CI 124,819 - �120 186,527) (Table 4.3). Overall, this represents a 20% loss of sagebrush-dominated habitat in the 35 years measured or a 0.64% annual loss rate (95% CI 0.49% - 0.77%). Habitat loss per stratum varied (Tables 4.2, 4.3), yet only some strata gave reliable estimates because of small sample size. Of those strata with greater than 10 plots sampled, the rate of habitat loss was variable with rates as high as 50% in stratum two and as low as 11% in strata seven and eight (Table 4.2). A comparison of the historic and current distributions of sage grouse reveals that only one area in southwestern Colorado seems not to have changed much (Fig. 4.1). This area is the Gunnison Basin which, in this study, is represented by stratum seven. Because of this a priori knowledge, I combined data from all strata except stratum seven and compared the rates of habitat loss from the Gunnison Basin to all other areas. The proportion of sagebrush habitat available was much higher in the Gunnison Basin than in the rest of the areas (Table 4.4). In 1958, the estimated proportion of sagebrush habitat in the Gunnison Basin was over twice the proportion in all other areas combined (0.3673 in Gunnison vs. 0.1552 in all other areas), whereas, in 1993 the proportion of habitat in the Gunnison Basin was almost three times higher than in all other areas (0.3267 in Gunnison vs. 0.1116 in all other areas). The Gunnison Basin experienced a loss rate of only 11% compared to the combined loss rate of 28% elsewhere. . Habitat Fragmentation My results clearly document habitat loss, yet habitat fragmentation was more difficult to document. In each plot I recorded the number of sagebrush polygons, the total area of sagebrush and the total amount of edge (total perimeter). Holding the area of sagebrush constant, it is intuitive that as fragmentation occurs, the number of polygons should increase and the ratio of the square root of total area to total perimeter (AlP ratio) should decrease. When area is not held constant, however, it is not clear what these variables will do (Groom and Schumaker 1993). With decreasing area, for example, the number of polygons could either increase or decrease (Fig. 4.4) as could the AlP ratio. In general terms, example A (Fig. 4.4) seems to be affected mostly by fragmentation. This is the case when the number of polygons increases (large areas broken into smaller areas) and AlP ratio decreases (more perimeter per unit area). Example C, however, is affected by habitat loss rather than fragmentation (Fig 4.4). Here, the number of polygons decreases and the AlP ratio increases. Thus, with habitat loss and fragmentation both occurring, merely reporting trends in the AlP ratio and the number of polygons would be misleading. I looked at the relationship between the change in number of polygons for each plot and the change in AlP ratio (Fig. 4.5) and found that most of my data fell into one of two categories. Sixty-six plots (37%) had an increase in the number of polygons and a decrease in AlP ratio. This represents cases where fragmentation tends to be the dominant process. Eighty-one plots (50%) had increases in the AlP ratio and decreases in the number of polygons. In these plots, habitat loss was presumably the dominating process. DISCUSSION I found little difference in estimates of the proportion of habitat lost between the analysis with the raw and the standardized data (0.039, SE = 0.0034 for raw data and 0.0428, SE = 0.0043 for standardized data). This gave me confidence that the model-based standardization using a logistic function represented the data in a reasonable way. Although I did not find a difference in the annual rate of change between the early and middle time period and between the middle and late period for 37 plots in this analysis, it seems reasonable �121 that rates might differ and be higher in more recent years due to tremendous increases in human population growth in Colorado. It has been estimated that human population growth in western Colorado was 3.1% per year between 1990 and 1996, much higher than the national average of 0.9% (Theobald in press). I documented substantial amounts of sagebrush-dominated habitat loss throughout southwestern Colorado. It is important to note that much of what was once sagebrush habitat was already lost to other land uses before the oldest photos in this study were taken. While not quantified, the loss of sagebrush habitat before the 1950's appeared substantial. This is in agreement with Rogers (1964) statement that much of the area once abundant with sagebrush, had been converted to other land uses by 1964. I could only document habitat loss since 1958. Overall, the change in the proportion of sagebrush-dominated habitat between 1958 and 1993 was 0.0428 (SE = 0.0043). This translates to a loss rate of20% with 155,673 ha lost over 35 years. Average loss of habitat per year was 0.64% or 5,033 ha (95% CI 3,853 - 6,055). Certain areas had much higher loss rates, especially stratum two (an area south of Durango and Pagosa Springs) which had an estimated loss rate of almost 50% (SE = 13.54). Sage grouse have been extirpated from this area The Gunnison Basin had the highest proportion of existing sagebrush habitat and had one of the lowest rates of habitat loss. My comparison of Gunnison Basin with all other data combined showed that the loss rate of 11% (SE = 1.14) in the Gunnison Basin was much lower than the loss rate of28% (SE = 3.64) elsewhere. This is not surprising in light of Braun's (1995) comparison of historic and current sage grouse distributions (Fig. 4.1) in which the Gunnison Basin seems to be the only population which has not been severely reduced. Habitat fragmentation also was considerable in this study. I found 66 plots in which habitat fragmentation was a dominating process in that there were more polygons in the late time period . (evidence of fragmentation into smaller polygons) than in the early time period and lower NP ratios (evidence of more perimeter per unit area). Fragmentation typically results in a few remnant sagebrush patches surrounded by a matrix of land that is unsuitable for sage grouse use due to development and land-use changes. This makes movement among patches potentially dangerous as sage grouse are more vulnerable to predators in these instances. In this study, fragmentation was often the result of road development which is known to have a negative impact on Gunnison sage grouse (Braun 1995, Oyler et al. 1997). Powerlines often line roads and provide perches for avian predators. Sage grouse have also been known to fly into and be killed by powerlines. While this study documented the amount of habitat loss and the prevalence of habitat fragmentation, it did not measure habitat quality (with respect to sage grouse). Certainly fragmenting once continuous sagebrush habitat can influence the quality of that habitat for sage grouse by allowing the invasion of non native plants, and creating perches and travel corridors for predators. Road development also affects the quality of sage grouse habitat because it is associated with increased human activity within or near sagebrush patches. Paved roads specifically, and all human activities associated with them, have been negatively associated with Gunnison sage grouse (Oyler et al. 1997, Chapter One). Habitat requirements for sage grouse are well documented (Klebenow 1969, Wallestad 1971, Eng and Schladweiler 1972, Wallestad and Pyrah 1974, Beck 1977, and others). Such habitat characteristics (e.g., % cover of sagebrush, height of sagebrush, % cover offorbs) were not measured in this study. Thus, a portion of the remaining sagebrush habitat is likely not suitable for Gunnison sage grouse, making estimates of the total amount of 'suitable' sagebrush less than the amount of sagebrush documented here. The decline in the distribution and abundance of Gunnison sage grouse is alarming (Braun 1995). I believe this decline to be the direct result of habitat loss and fragmentation, and to a decline in the quality of the remaining habitat. While this study could not address habitat quality, I have been able to document a steady loss of sagebrush habitat since 1958 and habitat fragmentation in a substantial �122 number of areas. If current trends of habitat loss and fragmentation continue, Gunnison sage grouse will undoubtedly become extinct. Protecting the remaining habitat from further loss and fragmentation is paramount to the survival of this species (Chapter Five). LITERATURE CITED Aldrich, J. W. 1963. Geographic orientation of American Tetraonidae. Journal of Wildlife Management 27:529-545. Beck, T. D. 1977. Sage grouse flock characteristics and habitat selection in winter. Journal of Wildlife Management 41:18-26. Braun, C. E. 1995. Status and distribution of sage grouse in Colorado. Prairie Naturalist 27: 1-9. Braun, C. E. 1998. Sage grouse declines in western North America: what are the problems? Proceedings of the Western Association ofFish and Wildlife Agencies 78:000-000. Braun, C. E., and 1. R Young. 1995. A new species of sage grouse in Colorado. Joint Meeting Wilson Ornithological Society and the Virginia Society of Ornithology, Abstract #23. May 4-7, 1995, Williamsburg, VA, USA. Buckland, S. T., D. R Anderson, K. P. Burnham, and 1.L. Laake. 1993. Distance sampling: estimating abundance of biological populations. Chapman Hall, London, UK. Burnham, K. P., D. R Anderson, G. C. White, C. Brownie, and K. Pollock. 1987. Design and analysis of methods for fish survival experiments based on release-recapture. American Fisheries Society. Monograph 5. Carroll, R 1., and D. Ruppert. 1988. Transformation and weighting in regression. Chapman and Hall, New York, NY, USA. Cary, M 1911. A biological survey of Colorado. North American Fauna 33. Csaplovics, E. 1992. Analysis of colour infrared aerial photography and SPOT satellite data for monitoring land cover change of a heathland region of the Causse du Larzac (Massif Central, France). International Journal of Remote Sensing 13:441-460. Dahl, T. E., and C. E. Johnson. 1991. Status and trends of wetlands in the coterminous United States, mid-1970's to mid-1980's. U. S. Department of the Interior, Fish and Wildlife Service, Washington, D.C., USA. Eberhart, L. L. 1978. Appraising variability in population studies. Journal of Wildlife Management. 42:207-238. Eng, R L., and P. Schladweiler. 1972. Sage grouse winter movements and habitat use in central Montana Journal of Wildlife Management 36:141-146. ESRI. 1996. ArcView GIS users manual. Environmental Systems Research Institute, Redlands, CA, USA. Groom, M., and N. Schumaker. 1993. Evaluating landscape change: patterns of worldwide deforestation and local fragmentation. Pages 24-44 in P. M. Kareiva, 1. G. Kingsilver, and R B. Huey, editors. Biotic interactions and global change. Sinauer Associates Inc, Sunderland, MA, USA. James, E. 1823. An account of an expedition from Pittsburgh to the Rocky Mountains. Volume 1. H. C. Cary and 1. Les, Philadelphia, P A, USA. Johnsgard, P. A. 1973. Grouse and quails of North America University of Nebraska Press, Lincoln, NE, USA. Klebenow, D. A. 1969. Sage grouse nesting and brood habitat in Idaho. Journal of Wildlife Management 33:649-662. �123 Knapp, P. A, P. L. Warren, and C. F. Hutchinson. 1990. The use of large-scale aerial photography to inventory and monitor arid rangeland vegetation. Journal of Environmental Management 31 :29- 38. Mast, 1. N., T. T. Veblen, M. E. Hodgson. 1997. Tree invasion within a pine/grassland ecotone: an approach with historic aerial photography and GIS modeling. Forest Ecology and Management 93:181-194. Oyler, S. J., C.E. Braun, and K. P. Burnham. 1997. Use of a habitat-based model to predict sage grouse (Centrocercus urophasianus) occupancy of patches in southwestern Colorado. Wildlife Biology 3:282. (Abstract). Patterson, R L. 1952. The sage grouse of Wyoming. Sage Books, Inc., Denver, CO, USA Remington, T. E. 1989. Why do grouse have ceca? A test of the fiber digestion theory. Journal of Experimental Zoology. Supplement 3:87-94. Robbins, B. D. 1997. Quantifying temporal change in seagrass areal coverage: the use of GIS and low resolution aerial photography. Aquatic Botany 58:259-267. Rogers, G. E. 1964. Sage grouse investigations in Colorado. Colorado Game, Fish, and Parks Department. Technical Publication 16. Denver, CO, USA Terrazas-Gonzalez, G. 1997. Evaluation ofprojection methods to predict wetland area sizes: the wetlands inventory of USA Dissertation, Colorado State University, Fort Collins, CO, USA Theobald, D. M. In press. Fragmentation by inholdings and exurban development. Pages:OOO-OOO,in R L, Knight, F. W. Smith, S. W. Buskirk, W. H. Roming, and W. L. Baker, editors. Forest fragmentation in the southern Rocky Mountains. University Press of Boulder, Boulder, CO, USA Thompson, M. E. 1997. Theory of sample surveys. Monographs on statistics and applied probability 74. Chapman and Hall, London, UK. Thompson, S. K. 1992. Sampling. Wiley, New York, NY, USA. Tiner, R W. 1990. Use of high-altitude aerial photography for inventorying forested wetlands in the United States. Forest Ecology and Management 33/34:593-604. Tukey, 1. W. 1977. Exploratory data analysis. Addison-Wesley Publishers, Reading, MA, USA. Wallestad, R 1971. Summer movements and habitat use by sage grouse broods in central Montana Journal of Wildlife Management 35:129-136. Wallestad, R, and D. Pyrah. 1974. Movement and nesting of sage grouse hens in central Montana Journal of Wildlife Management 38:630-633. Wallestad, R, 1. G. Peterson, and R L. Eng. 1975. Foods of adult sage grouse in central Montana Journal of Wildlife Management 34:628-630. �124 Table 4.1. Characteristics of strata sampled for sagebrush in southwestern Colorado. Area (ha) Plots per stratum Plots sampled per stratum 1,364,800 853 74 2 476,800 298 25 3 44,800 28 3 4 238,400 149 12 5 102,400 64 5 6 49,600 31 3 7 1,044,800 653 54 8 222,400 139 13 9 52,800 33 3 10 41,600 26 2 Stratum �Table 4.2. Differences in the proportion of sagebrush habitat in southwestern Colorado between 1958 and 1993. Mean difference in available habitat is the proportion of habitat available in 1958 minus the mean proportion of habitat available in 1993. Rate of habitat loss (%) Sampling fraction (sampled/total) Mean proportion of habitat available (1958) SE Mean proportion of habitat available (1993) SE Mean difference in available habitat (1958-1993) SE 74/853 0.1640 0.0237 0.1289 0.0221 0.0351 0.0052 21.40 3.19 2 25/298 0.1777 0.0385 0.0895 0.0265 0.0882 0.0241 49.63 13.54 3 3/28 0.0485 0.0458 0.0207 0.0196 0.0278 0.0263 57.32 54.11 4 12/149 0.2366 0.0737 0.1650 0.0578 0.0716 0.0221 30.26 9.33 5 5/64 0.0000 0.0000 0.0000 0.0000 0.0000 0.0000 0 0 6 3/31 0.0041 0.0039 0.0013 0.0013 0,0028 0.0027 68.29 64.70 7 54/653 0.3673 0.0414 0.3267 0.0407 0.0406 0.0053 11.05 1.44 8 13/139 0.0867 0.0401 0.0773 0.0353 0.0095 0.0052 10.84 5.99 9 3/33 0.1099 0.0447 0.0968 0.0528 0.0131 0.0109 11.92 9.95 10 2/26 0.2448 0.0065 0.1957 0.0286 0.0490 0.0220 20.06 8.99 19412274 0.2161 0.0166 0.1734 0.0154 0.0428 0.0043 19.80 1.99 Stratum Overall SE •.... N VI �Table 4.3. Differences in the amount of sagebrush habitat in southwestern Colorado between 1958 and 1993. Habitat lost is the amount of habitat available (ha) in 1958 minus the amount of habitat available (ha) in 1993. Area (ha) Habitat available in 1958 (ha) 95% Confidence interval 1,364,800 223,827 168,988 - 296,588 175,923 126,051 - 245,632 47,896 33,658 - 62,134 2 476,800 84,732 55,735 - 128,974 42,669 24,193 -75,352 42,065 18,441 - 65,688 3 44,800 2,174 701 - 6,920 4 238,400 56,415 30,581 - 104,342 5 102,400 0 6 49,600 205 7 1,044,800 383,786 8 222,400 19,289 10,703 - 34,872 17,180 8,699 - 34,043 2,109 676 - 3,542 9 52,800 5,801 1,861 - 18,436 5,111 1,409 - 18,952 690 -286 -1,667 10 41,600 10,182 2,664 - 39,868 8,142 1,806 - 37,762 2,039 3,638,400 786,411 Stratum Overall Habitat available in 1993 (ha) 929 39,338 0-0 0 66 - 651 65 308,079 - 478,346 667,337 - 905,484 341,368 630,725 95% Confidence interval 258 - 3,451 19,397 -79,999 0-0 18 - 241 267,748 - 435,457 520,220 -741,230 Habitat lost (ha) 1,245 17,076 0 139 42,414 155,673 95% Confidence interval -516 - 3,006 4,997 - 29,156 0-0 -58 - 337 31,335 - 53,492 -1,494 - 5,573 124,819 - 186,527 - N 0\ �Table 4.4. Differences in the proportion of sagebrush habitat in southwestern Colorado between 1958 and 1993, when data from the Gunnison Basin (stratum seven) was compared to all other strata combined. Data were standardized to 1958 and 1993. Mean difference in available habitat is the proportion of habitat available in 1958 minus the mean proportion of habitat available in 1993. Stratum Sampling fraction (sampled/total) Mean proportion of habitat available (1958) SE Mean proportion of habitat available (1993) SE Mean difference in available habitat (1958-1993) SE Rate of habitat loss (%) SE Gunnison Basin (7) 54/653 0.3673 0.0414 0.3267 0.0407 0.0406 0.0053 11.05 1.44 All others 140/1621 0.1552 0.0163 0.1116 0.0141 0.0437 0.0056 28.09 3.64 Overall 19412274 0.2161 0.0166 0.1734 0.0154 0.0428 0.0043 19.80 1.99 Table 4.5. Differences in the amount of sagebrush habitat in southwestern Colorado between 1958 and 1993, when data from the Gunnison Basin (stratum seven) was compared to all other strata combined. Data were standardized to 1958 and 1993. Habitat lost is the amount of habitat available (ha) in 1958 minus the amount of habitat available (ha) in 1993. Area (ha) Habitat available in 1958 (ha) 95 % Confidence interval Habitat available in 1993 (ha) 95 % Confidence interval Habitat lost Cha) 95 % Confidence interval Gunnison Basin (7) 1,044,800 383,786 297,137 - 470,436 341,368 256,111 - 426,624 42,414 31,337 - 53,491 All others 2,593,600 402,624 318,434 - 486,814 289,358 216,497 - 362,218 113,260 84,032 - 142,488 Overall 3,638,400 786,411 667,337 - 905,484 630,725 520,220 -741,230 155,673 124,819 - 186,527 Stratum •.... N -..l �128 Figure 4.1. Historic (top) and current (bottom) distribution of sage grouse and Gunnison sage grouse (lower left cut out) in Colorado. �129 Figure 4.2. Distribution of sagebrush-dominated habitat in Colorado in the early 1960's (From Rogers 1964). stratum 1 stratum 2 stratum 3 stratum 4 stratumS stratum 6 stratum 7 stratum 8 stratum 9 stratum 10 Counties Figure 4.3. Strata used as a sampling frame for the distribution of sagebrush habitat in the early 1960's (From Rogers 1964). �130 Area = 8366 Sqrt(Area)lPerimeter Ratio = 0.2476 Number of Polygons =4 DDD DD D B. c. Area = 7333 Area = 600 Sqrt(A)1P Ratio = 0.221 Sqrt(A)1P Ratio = 0.225 Sqrt(A)1P Ratio = 0.250 # Polygons # Polygons = 4 # Polygons = 1 ~ D DD A. Area = 7571 = 6 Figure 4.4. Misleading relationship between habitat loss and fragmentation. In this example the top scenario is one large patch and three small patches. As habitat is lost and fragmented there are many different scenarios. In example A, there is some amount of fragmentation and loss. The ratio of square root of area to perimeter decreases and the number of polygons increases. In example B, the ratio decreases and the number of polygons stays the same. In example C the ratio increases and the number of polygons decreases. �131 0.10 ....... • • • .s ~ I >. 1: 0.05 •• C'G .e 0 • +> !! ~.OO •... • .sGl ••• • •••••••• Ie·. ~ . !rl:t L.• : ... ••".Ir. •••• • , I 1.1 E •• •• ·c Gl Q. a ~.05 • C'G I!! C'G c ~.10 • Br::: I • • • • I!! ~ (5 ~.15 • • -40 -30 -20 -10 o 10 20 30 Difference in number of polygons (early -late) Figure 4.5. Relationship between the difference in number of polygons and the area/ratio perimeter. Data in upper left portion of the graph represents plots affected primarily by fragmentation and data in the lower right portion of the graph represent plots affected primarily by habitat loss. �132 �l33 CHAPTER FIVE DEVELOPMENT OF A MODEL TO ASSESS MANAGEMENT AND CONSERVATION STRATEGIES FOR GUNNISON SAGE GROUSE IN COLORADO INTRODUCTION The distribution and abundance of sage grouse (Centrocercus urophasianusj have markedly declined throughout its entire range. The historic distribution. which included at least 15 states and three provinces (Aldrich 1963, Johnsgard 1973) has been reduced, with extirpation from four states and one province (Braun 1998). In those areas where sage grouse still exist, their range has declined markedly (Braun et al. 1994, Braun 1998). Populations in Colorado have also been greatly reduced as sage grouse have been extirpated from 12 of the 27 counties in which they occurred in the 1900's and breeding populations in nine of the remaining 15 counties are thought to number less than 500 breeding birds (Braun 1995). Declines appear to be related to habitat loss (conversion of big sagebrush [Artemisia tridentata] into farmland or housing developments), habitat degradation (heavy grazing, sagebrush removal, road and powerline development through sagebrush, and human disturbance), and habitat fragmentation (Braun 1995, Braun 1998). Populations in southwestern Colorado, the range of almost all the newly described Gunnison sage grouse (c. minim us) (Braun and Young 1995), have been most severely impacted by these processes (Fig. 5.1). Populations that remain are small, widely scattered, and exist in degraded, fragmented habitats isolated from a larger population in the Gunnison Basin. For this reason. Gunnison sage grouse have become a focus of management and conservation concerns. Management options for Gunnison sage grouse include actions such as habitat protection. land mitigation. translocation among existing populations, and reintroduction into unoccupied areas. Habitat improvement is another management option. but could not be used in my model due to the coarse-scale nature of the data available. Often managers have to make decisions about specific issues using only a limited amount of information (usually that which is specific to the local population). My goal was to consolidate what is known about Gunnison sage grouse, represent it spatially, and make this information accessible to managers so they can assess how their decisions might affect not only a specific population, but the entire group of populations. Thus, I developed a Geographic Information System (GIS)-based computer model using the GIS software ArcView (ESRI 1996) which utilizes a variety of different layers of information to assess potential management strategies. The use of GIS technology is rapidly becoming an integral part of conservation and wildlife studies. GIS models have been used to identify areas of highest biodiversity (Scott et al. 1993), assess existing conservation units in nature reserves (peres and Terbough 1995), and address effects of forest management on species (Liu et al. 1995, Rempel et al. 1997). Homer et al. (1995) used GIS to model structural and compositional attributes of sage grouse winter habitat in Utah. Their GIS model was on a much smaller scale and described the structural components of an area 2,548 km2 in size. My model operates on a much larger scale (covering an area roughly 100,000 krrr') and addresses broad management questions of Gunnison sage grouse across all of southwestern Colorado. MEmODS The model included information on all Gunnison sage grouse populations, information on all sagebrush patches in southwestern Colorado, and information on current and future human housing densities. The data on specific Gunnison sage grouse populations included population size (C.E. �134 Braun, Colorado Division of Wildlife, unpublished data), distance to the centroid of the nearest population, and genetic diversity (Chapters Two and Three). Information on sagebrush patches included patch area, perimeter, distance to the nearest paved road from the centroid of each patch, distance to the nearest occupied patch, rate of habitat loss (Chapter Four), and the probability of occupancy (caiculated using methods from Chapter One). Data on current and future human housing densities were classified into one of six groups based on the number of housing units per ac (Theobald in press). I represented all available information spatially by obtaining or developing GIS coverages. I obtained a GIS coverage of the present distribution of Gunnison sage grouse from the Colorado Division of Wildlife. This coverage was edited to reflect recent extirpation of populations so that it represented the most current distribution information available. Data on population size (C.B. Braun, Colorado Division of Wildlife, unpublished data), and genetic diversity (Chapters Two and Three) were entered so the information could be accessed by selecting a given population. The distance to the centroid of the nearest population from each population's centroid was calculated in ArcView (ESRI 1996) .. A coverage of paved roads in southwestern Colorado was developed using 1:100,000-scale digital line graphs obtained from the U. S. Geological Survey (USGS). These files were derived from cartographic source materials using manual and automated digitizing methods from USGS topographic maps published as 30 x 60-minute quadrangles. I downloaded transportation data corresponding to the following USGS 1:100,000 scale maps: Grand Junction, Delta, Nucla, Dove Creek, Cortez, Durango, Silverton, Montrose, Paonia, Carbondale, Glenwood Springs, Vail, Leadville, Gunnison, Saguache, Del Norte, Antonito, Alamosa, Blanca Peak, Canon City, and Pikes Peak. I then merged all of the data in ArclInfo (ESRI 1987) into one coverage and selected only paved roads (attribute codes 170201 170208). To represent sagebrush patches in southwestern Colorado, I obtained a GIS coverage of vegetation types in southwestern Colorado from the Colorado Gap Analysis project. This coverage was developed for relatively coarse scale (1:100,000) projects with each mapping unit in non-riparian areas equal to 100 ha, This coverage (currently being tested and validated) maps only generalized distributions of vegetation types based on the USGS 1:100,000 mapping scale. These data seemed appropriate because my goal was to assess management strategies of Gunnison sage grouse across southwestern Colorado. From this coverage, I selected only those polygons described as sagebrush habitat. Two different classifications were found in this coverage, Wyoming or Mountain big sagebrush (A. t. wyomingensis or A. t. vaseyana) and Basin Big sagebrush (A. t. tridentata). The habitat was classified as Wyoming or Mountain big sagebrush if it comprised more than 25% of the total vegetative cover. This classification was variable and included dense, homogenous sagebrush stands as well as more sparsely vegetated areas. Often, patches of sagebrush were found with patches of mixed grasses. In these cases, the area was classified as sagebrush if sagebrush patches occupied more than 50% of the total ground cover. The area and perimeter of each sagebrush patch were given in the coverage. Using the sagebrush and sage grouse coverages, I calculated the distance from each sagebrush polygon centroid to the centroid of the nearest population. To report the average annual rate of habitat loss for each sagebrush polygon, I determined the stratum (Chapter Four) to which each polygon belonged. Because the sample size in some strata were small, (strata 3,4,5,6,9, and 10) estimates of habitat loss from those strata were poor, yet when I pooled data across strata with small sample sizes, the estimates of habitat loss were much better. Thus, I calculated a weighted average (0.98%) across those six strata and assigned each polygon that value. Polygons in strata with adequate estimates of loss rates were assigned the appropriate annual loss rate (stratum 1 = 0.68%, 2 = 1.9%, 7 = 0.33%, and 8 = 0.33%). I also calculated the probability of occupancy for each patch using the model averaging approach and the three models discussed in Chapter One. �135 Two coverages of human housing density (one from 1990 and one projected to 2020) were obtained from the Natural Resource Ecology Laboratory (NREL) at Colorado State University. These raster maps were based on 1990 U.S. Census Bureau block-group level data (a block-group is a subdivision of census tract containing between 250 and 500 housing units). Theobald (in press) mapped housing density at quarter-section (65 ha) resolution into one of six categories: no data, rural « 1 unit per 80 ac), ranchette « 1 unit per 40 ac), exurb an « 1 unit per 10 ac), suburban « 1 unit per 2 ac), and urban (> 1 unit per 2 ac) for the 1990 map. The 2020 map was classified the same way and is a projection from the 1990 data (Theobald and Hobbs 1998). To address the issue ofland protection and mitigation I decided to prioritize patches of sagebrush for protection which were currently occupied by Gunnison sage grouse. My criterion for ranking patches was a combination of the size of the sage grouse population in that patch, the distance to the nearest population, and the probability of occupancy of that patch. There are many other ways in which these areas could be prioritized; this is just one example of an application of this model. To prioritize patches for this example, I converted the vector sage grouse distribution coverage into two raster coverages/one with population size and one with the distance to the nearest population. I then standardized the distances so that they ranged from zero to one (with zero being the farthest and one being the closest) by dividing each value by the largest distance and subtracting this value from one. Population sizes were standardized by dividing each value by the largest value such that large populations had values close to one. I also converted my vector sagebrush coverage into a raster coverage using the probability of occupancy attribute as a cell value. Using Arc View's Spatial Analyst (ESRI 1996), I averaged the values of probability of occupancy, standardized population size, and standardized distance to the nearest population to create a coverage which prioritized areas for land mitigation and protection. This, in effect, gives equal weight to the three factors considered. Unequal weighting could be used if there was a reasonable basis for its use. I was also interested in determining which unoccupied sagebrush patches might be good reintroduction sites. Sites could be prioritized in different ways, but for this example I defined good sites as patches with a high probability of occupancy which were close to existing populations so that a network of somewhat connected populations could be established. I calculated the distance between the centroid of each sagebrush patch and the centroid of the nearest sage grouse population to determine which patches would make the best reintroduction sites. I then converted the vector sagebrush coverage to two raster coverages, one with each cell representing the probability of occupancy and one with each cell representing the distance to the nearest population. I then standardized the distances so that they ranged from zero to one (with zero being the farthest and one being the closest) by dividing each value by the largest distance and subtracting this value from one. In ArcView's Spatial Analyst (ESRI 1996), the two raster coverages were then combined by averaging the probability of occupancy and the standardized distance to the nearest population. This produced a coverage which represented the suitability of each patch with values near one being the most suitable and those near zero being the least suitable. An issue which might also be important for land mitigation and reintroduction is to detemiine which patches of sagebrush and which populations of sage grouse would potentially be impacted most by hwnan population densities. Thus, I created four new coverages; two represented overlays of the sagebrush coverage and two (one each) of the human housing densities (1990 and 2020) coverages. Two additional coverages represented overlays of the sage grouse distribution coverage and each of the hwnan housing densities. These coverages were developed by converting the sagebrush and sage grouse vector coverages into raster coverages and combining them with hwnan housing densities in 1990 and 2020. To investigate the possibility of trans locating birds among populations to increase genetic diversity, I decided that areas with the lowest genetic diversity, furthest from the area with the highest �136 genetic diversity would be considered a priority for translocation. Many other criterion could also be used to prioritize populations for translocation. From the sage grouse vector coverage, I made two raster grid coverages, one with the attribute called number of unique alleles as the grid value (populations with no genetic data received a zero), and one with the distance from the centroid of the Gunnison Basin attribute as a grid value. I then standardized both values so they ranged from zero to one (zero being the lowest diversity for the genetic coverage and the furthest distance from Gunnison Basin for the distance coverage). These coverages were then combined in ArcView's Spatial Analyst (ESRI 1996) by averaging the two values. RESULTS I classified the distribution of Gunnison sage grouse into eight populations based on knowledge of movements (Commons 1997, C. Woods and C. E. Braun. 1995. Sage grouse investigations, Colorado Division of Wildlife, Fort Collins, CO, USA) and expert judgement (Fig. 5.2). Polygons shown in the same color belong to the same population. For ease of discussion I developed a map showing key townsin southwestern Colorado (Fig. 5.3). I found similar patterns in size and genetic diversity of populations (i.e., large populations had high genetic diversity and small populations had low diversity). The largest population in the Gunnison Basin (Fig. 5.4) had the most genetic diversity (Fig. 5.5) with approximately 2000 breeding birds and an average of3.75 alleles per locus. The Dry Creek Basin population and the Crawford population are intermediate in size and in genetic diversity. The Dove Creek population, with approximately 100 birds had the lowest genetic diversity with an average of 1.75 alleles per locus. Sagebrush patches with a high probability. of occupancy (Fig. 5.6) generally occurred in areas . where sage grouse currently exist (Gunnison Basin, Dry Creek Basin, Glade ParklPinon Mesa and Crawford). An exception is the sagebrush patch near Poncha Pass which has a relatively low probability of occupancy. This population is small and was a transplant from the Gunnison Basin. Unoccupied areas with a high probability of occupancy include areas east of Dry Creek Basin near the town of Norwood, areas west of Montrose, areas southeast of Pinon Mesa, areas north of Crawford, an area south of interstate 70 near Glenwood Springs and Gypsum, a few patches near the New Mexico border west of Durango, and areas near Del Norte. Areas of low probability of occupancy generally occurred in areas near towns and cities such as Grand Junction, Paonia, Aspen, Alamosa, and Cortez. Areas with the lowest annual rate of habitat loss occur in the Gunnison Basin and north of it (Fig. 5.7). Highest loss rates occur southeast of Cortez along the border of New Mexico. Areas east of the Gunnison Basin also have high annual rates of habitat loss. I identified those sagebrush areas within the existing sage grouse distribution which had a high priority for protection and land mitigation (Fig. 5.8). Areas with a high priority for mitigation and protection included most of the Gunnison Basin, most of Dry Creek Basin, Glade ParklPinon Mesa, Crawford, and Sim's Mesa The area with the lowest priority was the Poncha Pass population. The Dove Creek population was classified as "no data" because the sagebrush coverage did not identify the area as sagebrush because the sagebrush in Dove Creek occurs in a patchy distribution intermixed with agricultural fields. Potential reintroduction sites with a high probability of occupancy and close to an existing population included areas mostly west of the Gunnison Basin (Fig. 5.9). Patches west of Montrose and southeast of Pinon Mesa had a high suitability as did sites east of Dry Creek Basin near Norwood. Other potential reintroduction sites include patches east of Dove Creek and patches north of Crawford. Areas within the current distribution of Gunnison sage grouse with the highest human housing density (urban with > 2 unit per 2 ac, suburban with < 2 units per ac) in 1990 occurred exclusively in �137 the Gunnison Basin in the town of Gunnison (Fig. 5.10). The Gunnison Basin was mostly categorized as rural « 1 unit per 80 ac) and ranchette « 1 unit per 40 ac) and rural. All other areas inhabited by Gunnison sage grouse were classified as rural. The 2020 projection of human housing densities (Fig. 5.11) reveals that the Gunnison Basin will likely experience great increases in human development and disturbance particularly along highway 50 which bisects the Gunnison Basin. North of the town of Gunnison toward Crested Butte is also projected to have a substantial increases in human density. The only other area within the current distribution of Gunnison sage grouse which will likely experience such increases is on Sim's Mesa south of Montrose which is projected to receive a ranchette classification by 2020. All other areas are projected to remain in the rural classification. Overall, changes in human density within areas currently occupied by sage grouse will be greatest in the the categories of rural and exurban (Table 5.1) with the amount of area classified as urban decreasing by 2059 km2 and the amount of area classified as exurban increasing by 1694 knr', Of all the sagebrush areas in southwestern Colorado, areas with the highest human housing density (urban and suburban) in 1990 occurred near Aspen and Glenwood Springs (Fig. 5.12). The Gunnison Basin had a few areas classified as ranchette and small area around the town of Gunnison classified as exurban, suburban, and urban. A few other areas were classified higher than rural including areas around Cortez and Durango. For the most part all other areas were classified as rural. The 2020 projection is for substantial increases in human density in the Gunnison Basin (particularly between Gunnison and Crested Butte), Aspen and Glenwood Springs, Cortez, Durango, areas east of Dry Creek Basin near Norwood, Paonia, and areas east of Grand Junction (Fig. 5.13). Changes between 1990 and 2020 (Table 5.2) will be greatest in the categories of rural (decrease of6528 km"), ranchette (increase of2635 knr'), and exurban (increase of3791 km'). The Dove Creek population was determined to have the most potential for translocating birds from the Gunnison Basin to increase genetic diversity (Fig. 5.14). Four populations, however, were omitted from consideration due to a lack of genetic data DISCUSSION I found the highest number of birds with the highest genetic diversity in the largest sagebrush areas (Figs. 5A, 5.5). Of the populations for which I had genetic data, the lowest diversity was found in the smallest population, Dove Creek. This is not surprising in light of the theories of inbreeding and genetic drift (Hartl and Clark 1989). I advocate translocating four to six females from the Gunnison Basin to the Dove Creek population every few years to increase its genetic diversity (Chapter Three). The Crawford and Dry Creek Basin populations are also candidates for translocation (Fig. 5.14). Further, genetic data (Chapters Two, Three) suggest that Gunnison sage grouse populations are isolated with relatively low gene flow among populations. Thus, protecting existing habitat and preventing further loss and fragmentation is paramount. Potential reintroduction sites should be large enough to sustain a considerable number of birds and be close enough to an existing population so that natural dispersal is possible. Large sites that are far from an existing population could still be considered, yet dispersal would need to be facilitated by humans which is a less favorable option. I found that the areas with the highest probability of occupancy occurred mostly in areas already occupied by Gunnison sage grouse. The Poncha Pass population is an exception, yet the population size is less than 50 birds (C. E. Braun, Colorado Division of Wildlife, unpublished data) and that population was a transplant from the Gunnison Basin. Unoccupied areas with a high probability of occupancy include areas between Dry Creek Basin, the Gunnison Basin, and Glade Park/Pinon Mesa, and areas north of Crawford, areas south of Glenwood Springs, areas between Cortez and Durango, and areas near Del Norte. These probabilities are based on the area of the patch and the distance to the �138 nearest road from the centroid of the patch (Chapter One). This represents a coarse scale approach to finding potentially suitable sites which should be supplemented by ground measurements to assure that the habitat characteristics of the habitat meet known requirements of sage grouse (Klebenow 1969, Wallestad 1971, Eng and Schladweiler 1972, Wallestad and Pyrah 1974, Beck 1977, and others). Historically, the highest rates of habitat loss occurred in areas removed from the Gunnison Basin including areas near Durango, and areas east of Alamosa and north to Poncha Pass. Interestingly, these areas of high annual rates of habitat loss are not areas which are predicted to have. high human densities by 2020 and those areas with the highest predicted human densities occur in areas where the rate of habitat loss is low (Figs. 5.7, 5.13). This is likely because historic habitat loss occurred from the conversion of sagebrush habitat into farmland (Rogers 1964), not housing developments; Future human impacts, however, appear to be concentrated in the Gunnison Basin (Fig. 5.13) which is the one area with a currently stable population. This is cause for concern and efforts should be made to minimize housing developments in areas crucial to Gunnison sage grouse survival. Land protection and mitigation is an important management option. The areas within the current distribution of Gunnison sage grouse which I found to have a high priority for protection and mitigation were Glade ParklPinon Mesa, Dry Creek Basin, Crawford, and Sim's Mesa, arid much of the Gunnison Basin (Fig. 5.8). Of those areas, the only area that is projected to have a substantial increase in human population density is in the Gunnison Basin (Fig. 5.11). The other areas, however, may be subject to habitat loss from changing land use patterns other than human development such as conversion into farmland or destruction of sagebrush to promote growth of grass for grazing. I identified potential reintroduction sites by determining sites with a high probability of occupancy that were close to existing populations (Fig. 5.9). On the ground measurements need to be collected in these sites, however, to assure that the characteristics of the habitat meet the known requirements of sage grouse (Klebenow 1969, Wallestad 1971, Eng and Schladweiler 1972, Wallestad and Pyrah 1974, Beck 1977, and others). Further, there may be other factors not measured here which could make these sites less suitable such as powerlines or oil and gas development. The ownership of these sites might also make reintroduction unlikely. My recommendations of potential sites here should be considered as a first step for considering reintroduction. Many other issues need to be considered before a reintroduction should be implemented. The best sites I found included areas between Dry Creek Basin, the Gunnison Basin, and Glade ParklPinon Mesa, areas east of Dove Creek, and areas north of Crawford. The areas east of Dry Creek Basin near Norwood seem to be a logical place to reintroduce birds because a population in this area would bridge the gap between the Gunnison Basin and Dry Creek populations and hopefully facilitate gene flow among the populations. The human population projection (Fig. 5.13), however, shows considerable increases in human densities in these areas. The areas north of Crawford and east of Dove Creek, however, are not predicted to have substantial increases in human densities. This model contains current information about Gunnison sage grouse populations and sagebrush patches in southwestern Colorado. It was designed so that information could be updated or added as it becomes available. Various scenarios can be addressed with this model and I have shown examples of some questions that can be addressed including prioritizing areas for land protection and mitigation, prioritizing areas for reintroduction, and prioritizing areas for translocation to increase genetic diversity. These examples are not exhaustive and are used to illustrate different aspects of the model. Other applications of the model include determining how the probability of occupancy changes with changes in patch size or distance to the nearest paved road, and investigating specific scenarios such as identifying all sagebrush patches within a given distance from a population that satisfy certain size and probability of occupancy criterion. Additional data (e.g., data on land ownership, fine scale habitat quality measured on the ground, and population data) can be easily added to strengthen the utility of the model. This model, when used by managers with specific questions, will help prioritize strategies for the conservation and management of Gunnison sage grouse. �139 LITERATURE CITED Aldrich, J. W. 1963. Geographic orientation of American Tetraonidae. Journal of Wildlife Management 27:529-545. Beck, T. D. 1977. Sage grouse flock characteristics and habitat selection in winter. Journal of Wildlife Management 41:18-26. Braun, C. E. 1995. Status and distribution of sage grouse in Colorado. Prairie Naturalist 27:1-9. Braun, C. E. 1998. Sage grouse declines in western North America: What are the problems? Proceedings from the Western Association of State Fish and Wildlife Agencies 78:000-000. Braun, C. E., K. Martin, T. E. Remington, and J. R YOWlg.1994. North American grouse: issues and strategies for the 21st Century. Transactions from North American Wildlife and Natural Resources Conference 59:428-438. Braun, C. E. and J. R YOWlg.1995. A new species of sage grouse in Colorado. JoiIit Meeting Wilson Ornithological Society and the Virginia Society of Ornithology, Abstract #23. May 4-7, 1995, Williamsburg, VA, USA. Commons, M. L. 1997. Movement and habitat use by Gunnison sage grouse (Centrocercus minimus) in southwestern Colorado. Thesis, University of Manitoba, Winnipeg, Canada . Eng, R L. and P. Schladweiler. 1972. Sage grouse winter movements and habitat use in central Montana Journal of Wildlife Management 36:141-146. ESRI. 1987 ..ARCIINFO users manual. Environmental Systems Research Institute, Redlands, CA, USA. ESRI. 1996. ArcView users manual. Environmental Systems Research Institute, Redlands, CA, USA. ESRI. 1996. Arc View Spatial Analyst users manual. Environmental Systems Research Institute, Redlands, CA, USA. . Hartl, D. L., and A. G. Clark. 1989. Principles of population genetics. Sinauer Associates, Sunderland, MA, USA. Homer, C. G., T. C. Edwards, RD. Ramsey, and K. P. Price. 1995. Use of remote sensing methods in modeling sage grouse winter habitat. Journal of Wildlife Management 57:78-84. Johnsgard.P, A. 1973. Grouse and quails of North America University of Nebraska Press, Lincoln, NE, USA. Klebenow, D. A. 1969. Sage grouse nesting and brood habitat in Idaho. Journal of Wildlife Management 33 :649-662. Liu, J., J. B. Dunning, and H. R Pullium. 1995. Potential effects ofa forest management plan on Bachman's Sparrows (Aimophila aestivalis): Linking a spatially explicit model with GIS . . Conservation Biology 9:62-75. Peres, C. A., and J. W. Terbough. 1995. Amazonian nature reserves: an analysis of the defensibility status of existing conservation units and design criterion for the future. Conservation Biology 9:34-46. Rempel, R S., P. C. Elkie, A. R Rogers, and M. J. Gluck. 1997. Timber management and natural disturbance effects on moose habitat: landscape evaluation. Journal of Wildlife Management 61:517-524. Rogers, G. E. 1964. Sage grouse investigations in Colorado. Colorado Game, Fish and Parks Department. Technical Publication 16. Denver, CO, USA. Scott, J. M., F. Davis, B. Csuti, R Noss, B. Butterfield, C. Groves, H. Anderson, S. Caicco, F. D'Erchia, T. C. Edwards, Jr., J. Ulliman, and G. Wright. 1993. Gap analysis: a geographic approach to protection of biological diversity. Wildlife Monographs 123. Theobald, D. M. In press. Fragmentation by inholdings and exurb an development. Pages:OOO-OOO,in R L, Knight, F. W. Smith, S. W. Buskirk, W.H. Roming, and W.L. Baker, editors. Forest �140 fragmentation in the southern Rocky Mountains. University Press of Boulder, Boulder, CO, USA. Theobald, D. M, and N. T. Hobbs. 1998. Forecasting rural land use change: a comparison of regression- and spatial transition-based models. Geographical and Environmental Modeling 2:57-74. Wallestad, R 1971. Summer movements and habitat use by sage grouse broods in central Montana Journal of Wildlife Management 35:129-136. Wallestad, R, and D. Pyrah. 1974. Movement and nesting of sage grouse hens in central Montana Journal of Wildlife Management 38 :630-63 3. Table 5.1. Amount of area within the current distribution of Gunnison sage grouse in southwestern Colorado classified into each offive human housing densities in 1990 and projected to the year 2020. Housing density Amount (knr') in 1990 Amount (knr') in 2020 Rural « 1 unit per 80 ac) 6,950 4,891 Ranchette « 1 unit per 40 ac) 1,750 2,058 273 1,967 Suburban «1 unit per 2 ac) 74 130 Urban (> 1 unit per 2 ac) 16 16 Exurban « 1 unit per 10 ac) Table 5.2. Amount of area within the. current distribution of sagebrush in southwestern Colorado classified into each of five human housing densities in 1990 and projected to the year 2020. Housing density Rural « 1 unit per 80 ac) « Ranchette Exurban « 1 unit per 40 ac) 1 unit per 10 ac) Suburban «1 unit per 2 ac) Urban (> 1 unit per 2 ac) Amount (krrr') in 1990 Amount (km") in 2020 30,797 24,269 3,712 6,347 2,580 6,371 320 347 38 113 �141 [:~ ,". .c, ~ : •.;.;;.,t.~ t. ~ .\ I ~\ _ __ ..__ ) L- -_ __ < Figure 5.1. Historic (top) and current (bottom) distribution of sage grouse and Gunnison sage grouse (lower left) in Colorado. �142 Figure 5.2. Distribution of 8 populations of Gunnison sage grouse in southwestern Colorado. Polygons with the same color represent the same population. �143 Figure 5.3. Location of cities and towns in southwestern Colorado. �144 N paved road sage grouse population size BIll D 0-10 11-100 •• 101-175 •• 176-:DOO Figure 5.4. Distribution of Gunnison sage grouse in southwestern differences in population size. Colorado color coded to show �145 Npavedroad average # of alleles/locus _ no data MI!!!!IIIH 1- 1.75 _1.75-2.5 •• 2.5-3.75 Figure 5.5. Genetic diversity (average number of alleles per locus) of four populations of Gunnison sage grouse in southwestern Colorado (Chapters Two, Three). �146 IV paved road probability of occupancy _ 0.013-0.237 D 0.237 - 0.577 _ 0.577 - 0.853 _ 0.853 - 0.999 .Figure 5.6. Probability of occupancy of all sagebrush patches in southwestern Colorado (calculated using model averaging procedure in Chapter One). �147 /\I paved road % annual loss of sagebrush habitat 0-0.33 0.34 - 0.68 0.68 - 0.98 0.98 -1.9 no data Figure 5.7. Annual loss (%) of sagebrush habitat (as measured in Chapter Four) in southwestern Colorado. �148 IV paved road priorities for mitigation _ 0-0.25 [=:=I 025 - 0.5 _ 0.5-0.75 _ 0.75-1 c:==I No Data Figure 5.8. Priorities for land protection and mitigation in southwestern Colorado (based on size of the existing sage grouse population, distance to the nearest population, and probability of occupancy). �149 /'V paved roa d patch suitability I I 0.081 - 0.311 0.311 - 0.54 _ 0.54-0.77 _ 0.77-1 _ I occupied I areas data outside sagebrush range Figure 5.9. Priorities for sage grouse reintroduction into unoccupied areas of southwestern Colorado. Priorities are based on a combination of distance to the nearest population and probability of occupancy. �150 paved sage 1990 roa d grouse population density Rural«1 in sage unitperllO Ranchette «1 « Exurban Surburban Urban distribution (>1 unltper40 1 unit «1 per areas ae) 10 a e ) unitper2 unitper2 grouse a e) ae) ae) Figure 5.10. Human housing density (1990) in areas currently occupied by Gunnison sage grouse in southwestern Colorado. �151 \ ~ paved &alle 2020 IIrou&e population _ I road density Rural«1 I distribution in sage grouse areas unitper80ac) Ranchette «1 unitper40 _ Exurb an «1 _ Su rb urba n « 1 unit _ Urban ac) un it per 10 ac) (>1 unit per 2 ac) per 2 ac) Figure 5.11. Human housing density projected to 2020 in areas currently occupied by Gunnison sage grouse. �152 paved ~ I 1990 road sa lie brush d Istrlb utlon pop u Ia tio n den s ity in sag e b ru s hare ! R u ra I « 1 unit Ranchette ex «1 a° per unltper40 u rb an « 1 unit per S u rb u rb an « 1 unit U r ban (,.. 1 unit per as a c) ac) 10 a c) per 2 a c) 2 ac ) Figure 5.12. Human housing density (I 990) in sagebrush areas in southwestern Colorado. �153 paved S8 2020 roa d geb rush population R ura I dlstrib uuo density « Ex u rb an in sagebrush areas 1 unit per 8 0 a c) Ranchette Surburban n «1 « unltper 1 unit «1 per unit U rb a n (> 1 u nit per 40 8C) 10 a c) per2 ae ) 2 ac) Figure 5.13. Human housing density projected to 2020 in sagebrush areas in southwestern Colorado. �154 paved Priority ~ L:] roa d fortranslocation no aenetlc _ 0-0.3 0.3-0.75 _ 0.75-1 c::J Outside to ln c r e a s e diversity datil sage grouse ran ge Figure 5.14. Priority of populations into which sage grouse from the Gunnison Basin should be transloscated. Low values (yellow) receive the highest priority. �155 DISCUSSION Through the course of this dissertation research, much was learned about conservation and management of Gunnison sage grouse (Centrocercus minimus), research and methodology in general, and limitations associated with research. In these final pages, I comment on what was learned, how my work could have been improved, and what future research might follow. From the habitat-based model that I developed in Chapter One, I learned that the model which best described the data included the variables distance to the nearest paved road from the patch centroid, and patch area The major limitation of this model was small sample size (25 patches sampled). With a much larger sample size, many more candidate models with more variables could have been considered. Sample size was small because of the criterion I used for choosing patches to sample (had to have supported populations in the past 20 years) and the time constraint of only one field season. If this criterion was relaxed (although the number of historically occupied patches is limited) and more time was allotted for sampling, more patches (from the sagebrush coverage in Chapter Five) could be visited and sample size could be raised somewhat. In Chapter Two I found that the small-bodied Gunnison sage grouse were distinct from the large-bodied sage grouse (c. urophasianus) and that the small-bodied birds had less genetic diversity and gene flow relative to the large-bodied birds. The main limitation in this study was the small number of microsatellite loci (four) used. With only four loci examined, it becomes more difficult to characterize the uncertainty of the relationship among populations as bootstrap analyses on the neighbor-joining trees are not reliable with such few loci. Further, the data analyses associated with microsatellite markers are not well developed. It has been suggested that microsatellites follow a stepwise mutation model rather than the typical infinite alleles model of mutation. Published genetic distances based on the stepwise mutation model produced spurious results with my data and thus were not used in this dissertation. More research on these models of mutation need to be conducted and better genetic distance measures based on the stepwise mutation model need to be developed to improve the analysis of microsatellite data Future additions to my genetic study might include developing primers specifically for sage grouse to increase the number of loci and also to obtain samples from areas without adequate representation (such as Pinon Mesa and Poncha Pass). Genetic samples from the Poncha Pass population would be particularly interesting because it is an extremely small population which was a transplant from the Gunnison Basin a known number of years ago. This would allow us to look at isolation and genetic drift in this population. The management implications of my genetic study are discussed in Chapter Three. I noted the low genetic diversity in the small-bodied birds, particularly within the population near Dove Creek. I suggested translocating females from the Gunnison Basin population to at least the Dove Creek population to increase its genetic diversity. Again, data used in this study would be improved by obtaining more microsatellite loci which would allow for a better understanding of the relationship among the small-bodied populations. Additionally, future research should focus on monitoring survival and reproductive success of Dove Creek birds before and after any transplant to assess any effects (positive or negative). Genes associated with immunity to disease such as the Major Histocompatibility Complex should be examined in these birds to see if the genetic diversity of this gene is low which might have severe implications should a disease outbreak occur. Analysis of aerial photography in Chapter Four showed a 20% loss and substantial fragmentation of sagebrush-dominated habitat between the mid-50's and the mid-90's. The Gunnison Basin had a lower loss rate than all other strata examined. The design and analysis of this study were sound and likely do not need to be improved. I ground truthed approximately 20 % of the photos and am confident that classification errors were minimal. Errors associated with the zoom transfer scope, scanning, and importing the data into Photoshop were not quantified. To quantify these errors, multiple �156 people would have to be used to assess variability among people and associated errors. I did not have this option for this study, but future studies could incorporate multiple people repeating the same techniques to quantify this error. This has been done by the National Wetlands Institute and was found to be a minimal source of error in their study. The analysis of fragmentation could likely have been documented better by exainining all plots and reporting the fate of each sagebrush polygon. However, this would be extremely time intensive and would not likely add much to the analysis. It would be interesting to look at photos ·from the same plots in another 10 - 20 years. I believe that much of the loss in the past has been the result of conversion of sagebrush into farmland, but that this trend might change given current human influx into Colorado. I predict that future loss and fragmentation will be due to human development rather than farming and ranching. In Chapter Five, I developed a GIS-based model to assess potential conservation strategies for Gunnison sage grouse. It is a course scale model based on data from the Colorado GAP Project. Finer scale data could be used, yet it is prohibitive now due to the enormous size of the computer files containing this type of data Currently, this model contains information on sage grouse populations (size and genetic diversity), information on sagebrush patches in southwestern Colorado, and information on human population densities. Additional information which could be-incorporated into this model include data on whether land is public or private, more complete genetic data, population data on Gunnison sage grouse (such as survival and reproduction), and habitat quality data on sagebrush patches. Predications from this model could be made and field tested (e.g., patch suitability). Overall, I believe that the information in this dissertation has improved the knowledge base of certain aspects of Gunnison sage grouse and that it can be incorporated into conservation plans for this species. This species will likely be petitioned for listing as a threatened or endangered species. My habitat-based model (Chapter One) and GIS-based final model (Chapter Five) can be used to address habitat issues and to determine areas to protect and areas for reintroduction. Data on habitat loss (Chapter Four) will help assess past causes of extirpation and may be used to assess how human population growth may affect Gunnison sage grouse in the future. The genetic data (Chapters Two, Three) support the distinction of Gunnison sage grouse as a species and hopefully can be used to manage against the loss of genetic diversity. A conservation plan that integrates the information from this dissertation and from other studies, should be completed and implemented to assure the persistence of this species. �157 Colorado Division of Wildlife Wildlife Research Report April 1999 JOB PROGRESS REPORT Smteof ~C~o~lo~r=ad~o~ Project: W..:...:.....-~16:::o...7:....-=R::......_ _ Work Plan _8 _ _ Avian Research Job _6_ Job Title: _--"'D::...:e::....:v'-"e::..!:lo~p=m.:..:e::..:n~t o><-'f,_,a:..C=on:.::s:<,::e:,:..rv.:..,:a:::;ti:..::::o.:.:,n ..••. P..:.:lan=..:....:D""'o.:....r '""'L""es:::.:::s:..::::er:....P~rat::.:· e"--.:::ch,,::i.:::ck~e:<:;n""'s~in..:....::::CO!:Oo::..!:lo:.:..ra=d~ _ no..:..:; Period Covered: Author: Personnel: 0 1 January through 31 December 1998 Kenneth M. Giesen Kenneth M. Giesen, Ed Gorman, Judy Sheppard, Jeff Yost, Colorado Division of Wildlife ABSTRACf The Lesser prairie-chicken (Tympanuchus pallidicinctus) was petitioned for listing under the Federal Endangered Species Act in 1996. As a result, the Lesser Prairie-chicken Interstate Working Group (LPCIWG) was formed in 1997 to address the conservation needs of this species, and several meetings were held in 1998 to address the petition and to develop a range-wide Conservation Plan. Increased knowledge of population size and distribution, and development of habitat-based management plans to cooperatively manage sand sagebrush (Artemisia jilifolia) rangelands on public and private lands were identified as priorities in Colorado for this species. Consequently, intensive breeding surveys were conducted in 1998 resulting in 311 males and 18 females being counted on 33 active leks. This is a 30. percent decline since 1988. A cooperative agreement between several private landowners, the Colorado Division of Wildlife, U.S. Forest Service, U.S. Fish and Wildlife Service, and the Natural Resources Conservation Service was completed to manage >23,000 acres for Lesser Prairie-chickens . in southeast Colorado (Baca County). �158 �159 DEVELOPMENT OF A CONSERVATION PLAN FOR LESSER PRAIRIE-CmCKENS IN COLORADO Kenneth M. Giesen INTRODUCTION The lesser prairie-chicken (Tympanuchus pallidicinctus) was petitioned for listing under the Endangered Species Act in 1996. In June 1998 the ruling by the U.S. Fish and Wildlife Service on the petition was "warranted but precluded". A multi-agency committee, the Lesser Prairie-chicken Interstate Working Group (LPCIWG) was established in 1996 to address causes for the declines in .distribution and population size of this species. A review of pertinent literature suggested that the most promising approach to reversing the population declines was to restore and manage habitats used by this species. Priorities for each state included more intensive monitoring of breeding populations and working with land owners and land management agencies to manage or restore habitats, especially nesting and brood-rearing habitats, for the lesser prairie-chicken. P. N. OBJECTIVES The primary objective of this study is to monitor populations of lesser prairie-chickens in Colorado and work cooperatively with other agencies and landowners to develop a conservation plan for this species in Colorado. SEGMENT OBJECTIVES 1. Review literature on lesser prairie-chicken biology and habitat use. 2. Monitor breeding populations of lesser prairie-chickens in Baca, Prowers, and Kiowa counties in southeastern Colorado. . 3. Cooperate with NRCS, U.S. Forest Service, the LPCIWG, CSU Extension Service, other agencies, and private landowners in developing and implementing conservation strategies to benefit the lesser prairie-chicken. 4. Prepare annual progress report. METHODS Intensive ground searches were conducted by 16-18 biologists from 6-9 April following a training session on detection of leks and males by sight and sound. Each person was assigned 4 sections of rangeland each morning to search, and was provided a map on which all recently active and historical lek locations were plotted. Observers were instructed to search from all access roads and also walk the interior of all sections, while listening and scanning for lesser prairie-chicken lek activity. Searches began prior to sunrise and were completed by 10:00 each morning. Other searches occurred prior to sunset when observers scouted habitats to be searched on subsequent mornings. Following the intensive searches, additional counts of males and females were made in April by a trained part-time employee. �160 RESULTS AND DISCUSSION The intensive survey of lesser prairie-chicken habitats in Baca, Prowers, and Kiowa counties resulted in 302 total birds being counted on 33 active leks. There were 198 birds counted on 26 leks in Baca County (including 113 males, 11 females), 92 total birds in Prowers County on 5 active leks (including 81 males and 7 females), but no birds were observed on several historical leks in Kiowa County. However, two new leks were located in Cheyenne County with a total of 12 birds counted. While the number of leks and total birds counted is higher than in 1997 (I06 birds on 16 leks), the populations is substantially lower from high counts in 1988 (448 birds on 35 leks). The Lesser Prairie-chicken Interstate Working Group and various subcommittees held several meetings to address issues concerning the decline in numbers and distribution of lesser prairie-chickens and draft a regional conservation strategy. This conservation strategy is expected to be completed and distributed to interested agencies and individuals in 1999. Within Colorado, a cooperative agreement involving habitat management and Lesser Prairie-chicken conservation on >23,000 acres of public and private lands was drafted. Implementation of this agreement will be dependent upon funding from several agencies and private landowners. Annual meetings will be scheduled to access Lesser Prairie-chicken habitat and habitat goals .. Prepared by: ~~ Kenneth M Giesen �161 Colorado Division of Wildlife Wildlife Research Report April 1999 JOB PROGRESS REPORT State of: C=o=l=o.:..:ra=d=o Project: W-'-'--"""16::::...;7'--~R'- Work Plan 13 Job _ Avian Research _ 11 Job Title: Evaluation of Columbian Sharp-tailed Grouse Reintroduction Colorado Period Covered: Author: 0 1 January through 31 December Opportunities in Western 1998 Richard W. Hoffinan Personnel: Mike Bauman, Jennifer Boisvert. Jim Haskins, Jim Hicks, Richard Hoffinan, Mike Middleton, and Libbie Miller, Colorado Division of Wildlife; John Monarch, Monarch and Associates; Bonnie Postovit, Powder River Eagle Studies Inc. ABSTRACT Efforts for this reporting period focused on reviewing literature, conducting lek surveys, conducting meetings, gathering information to assist in the preparation of a conservation plan, administering the Grouse Habitat Improvement Program (GHJP), and preparing a study plan to evaluate Columbian sharp-tailed grouse (Tympanuchus phasianellus columbianus) use of Conservation Reserve and mine reclamation lands. Three public informational meetings were held in northwest Colorado in 1998. These meetings resulted in the formation of the Northwest Colorado Columbian Sharp-tailed Grouse Working Group. The group has made progresss in terms of developing a mission statement and identifying 36 issues that may potentially impact sharptails. GHJP dollars were used to plant 1,334 acres with enhanced seed mixtures of grasses and forbs to benefit sharp-tailed grouse. Twenty-three of 24 known lek sites in Moffat County and 84 of94 known leks in Routt County were checked in 1998; 12 and 56 were classified as active, respectively. Twenty-eight new leks were found in 1998. The revised database contains 146 known active and inactive lek sites in Moffat (rr = 39) and Routt (rr = 107) counties. Ninety-two percent of the leks occur on private land, 43 and 23% occur on CRP and mine reclamation lands, respectively. No leks were found west of Colorado Highway 13 north of Craig. No surveys were conducted in Rio Blanco County. �162 �163 EVALUATION OF COLUMBIAN SHARP-TAILED REINTRODUCTION OPPORTUNITIES IN WESTERN COLORADO Richard W. Hoffinan INTRODUCTION Use of common names and misidentification of blue grouse mendragapus obscurus) and sage grouse (Centrocercus urophasianus) by early explorers have made it difficult to ascertain the precise distribution of Columbian sharp-tailed grouse in Colorado (Rogers 1969, Giesen and Braun 1993). However, historical records suggest this subspecies may have occurred in at least 22 counties in western Colorado (Bailey and Niedrach 1965, Rogers 1969). Recent surveys indicate viable populations are restricted to Moffat, Routt, and Rio Blanco counties, with possible remnant populations in Mesa and Montrose counties (Giesen and Braun 1993). Similar reductions in the distribution of Columbian sharp-tailed grouse have occurred throughout western North America (Miller and Graul 1980). This decrease in distribution resulted in designation as a Category 2 species (D. S; Dep. Inter. 1989). Factors responsible for the reduction in distribution include conversion of native rangeland to cropland, excessive grazing by livestock, vegetative succession due to fire suppression, herbicide treatments, mineral exploitation, and urban development (Meints et al. 1992, Giesen and Connelly 1993). These factors have had the most pronounced impact on nesting, brood rearing, and winter cover through loss of native grasses and deciduous shrubs (Giesen and Braun 1993). Cover types used by Columbian sharp-tailed grouse tend to be structurally and vegetatively diverse with an extensive deciduous shrub component (Meints et al. 1992, Giesen and Connelly 1993). In Colorado, Columbian sharp-tailed grouse occur in mountain shrub communities interspersed with grasslands, small aspen ~opulus tremuloides) stands, and riparian zones (Giesen 1987). Serviceberry (Amelanchier spp.) is an essential element of these communities and usually grows in association with one or more of the following deciduous shrubs: Gambel oak (Quercus gambelii), common chokecherry ~runus virginiana), snowberry (Symporicarpos spp.), and sagebrush (Artemisia spp.) (Giesen 1987). Wheat is the primary agricultural crop within the range of sharptails in western Colorado. Wheatfields may be used during late summer and fall after harvest. These fields are usually snow-covered and unavailable during winter. Much of what is known about Columbian sharp-tailed grouse in western Colorado has resulted from studies in the northwest portion of the state (Dargan et al. 1942, Rogers 1969, Giesen 1987). Little is known about sharp-tailed grouse in southwestern Colorado other than they once occurred there and may still exist in low densities on the north end of the Uncompahgre Plateau (Rogers 1969, Giesen 1985). It has been 10 years since the last intensive effort to conduct lek surveys for Columbian sharptailed grouse in western Colorado. Another intensive effort is needed because changes in land use practices have occurred since then including implementation of the Conservation Reserve Program (CRP), additional mining and development activities, and alteration of grazing practices. Perhaps the most important action in the last 10 years affecting the need for current populationand distribution data has been the petition to list Columbian sharp-tailed grouse as "threatened" or "endangered" in the lower 48 conterminous United States pursuant to the Federal Endangered Species Act (Carlton 1995). This action is of special significance in Colorado because Idaho, Utah, and Colorado are the only states that allow hunting of Columbian sharp-tailed grouse and that still have adequate populations to provide transplant stock for future restoration programs. �164 Opportunities for management of sharptails in western Colorado may be limited because much of the occupied habitat occurs on private lands. The most extensive areas of public lands within the historic distribution of Columbian sharp-tailed grouse are in southwest Colorado. The last confirmed sighting of sharptails on these lands was in 1985 (Giesen 1985). Before a reintroduction program can be implemented, current habitat conditions and status of sharptails on these lands must be evaluated and management strategies formulated based on the outcome of the evaluation. It is likely that any effort to restore sharptails in western Colorado will require a commensurate effort to restore and protect habitat. P. N. OBJECTIVES Objectives of this project are to (1) form a sharp-tailed grouse working group with broad citizen, community, and agency representation, and in cooperation with this group, prepare a conservation plan for Columbian sharp-tailed grouse in Colorado, (2) conduct intensive lek surveys of Columbian sharptailed grouse in northwest Colorado, (3) ascertain presence or absence of sharptails in historic range in southwest Colorado, (4) identify potential reintroduction sites within the historic range of Columbian sharp-tailed grouse, (5) evaluate existing habitat conditions on these sites, and (6) cooperate with other western states in preparing conservation strategies for Columbian sharp-tailed grouse, . SEGMENT OBJECTIVES 1. 2. 3. 4. 5. 6. 7. Review literature pertinent to the objectives of this study. FOrni working group and conduct regularly scheduled meetings to develop management strategies and prepare conservation plan. Prepare conservation plan in collaboration with working group. Prepare and monitor contracts for sharp-tailed grouse habitat improvement projects. Prepare study plan to evaluate use of Conservation Reserve Lands as breeding, nesting, and brood rearing habitat for Columbian sharp-tailed grouse in northwestern Colorado. Conduct leks surveys in Moffat, Routt, and Rio Blanco counties. Compile data, analyze results, and prepare progress report. RESULTS AND DISCUSSION Segment Objective 1 - Literature on all aspects of the biology and ecology of sharp-tailed grouse was reviewed including 8 Master's Theses and 2 Ph.D. Dissertations obtained through interlibrary loan. Literature searches were conducted through Current Contents and the Fish and Wildlife Reference Service. Efforts were made to review draft and final conservation plans and strategies prepared by other states and to talk with the people involved in preparing these documents. Efforts also were made to review all documents pertaining to the petition to list the Columbian Sharp-tailed grouse as threatened or endangered. Segment Objectives 2-3 - Three public informational meetings were held in northwest Colorado in 1998. The purpose of these meetings was to inform the public about the status, distribution, and biology of Columbian sharp-tailed grouse, introduce them to some of the issues related to management of this grouse, and ascertain their interest in forming a working group to develop a conservation plan. Thirty-six people attended the meetings. Everyone agreed we should move forward with preparation of a conservation plan. There was general agreement we should form one working group that includes representatives from all counties (Moffat, Routt, and Rio Blanco) within the current range of sharptails in northwest Colorado. A mailing list of 230 potential stakeholders was developed with assistance �165 from local personnel from the Colorado Division of Wildlife (CDOW). U.S. Forest Service (USFS). Bureau of Land Management (BLM). and Natural Resource Conservation Service (NRCS). Everyone on the list was notified about the formation of the working group and invited to participate. The inaugural meeting was held in January 1999 at which time a regular meeting schedule of the last Tuesday of every month was established. The group has made progress in terms of developing a mission statement and identifying the issues that need to be addressed in the conservation plan. Following is the mission statement agreed upon by the group: To conserve and enhance Columbian sharp-tailed grouse populations and habitats in northwest Colorado in ways· that are compatible with existing and future land uses thereby insuring the opportunity for people to enjoy this wildlife resource in perpetuity. Following are the issues identified by the group: . Agriculture Management Pollution Herbicides (shrub control) Degradation of riparian zones Weed Control Loss of Topsoil Loss ofCRP Poor quality CRP Grazing (timing. duration, intensity) Insecticides Invasion noxious/exotic plants Irrigation practices Conversion of native mountain shrub communities Increased numbers of elk Range expansion/reintroduction Inadequate inventory data Private land access Poaching Lack of extension information Poor historical information Lek harassment due to research/monitoring Protection of other species (i.e .• sage grouse) Development Recreation Power lines Subdivisions Private property rights Roads Land zoning Mining/energy development Increased human activity Hunting Off road vehicles Lek harassment by wildlife viewers Biology Habitat Fire suppression Habitat fragmentation Pinyon/juniper invasion Lack of vegetative diversity Predators When discussions started on the issues. it became evident that not all key stakeholders were present. The group did not feel comfortable discussing issues that did not pertain to them without having someone in attendance that was directly affected by the issue. The issue discussions were halted. The group decided to use the list of issues to identify additional stakeholders. The group further decided to develop a schedule for discussion of the different issues. This schedule will outline what issues will be discussed at each meeting. This way stakeholders that have a vested interest in any particular issue know what meeting they need to attend to voice their concerns. �166 Segment Objective 4 - Using Great Outdoors Colorado funds obtained from lottery sales, the CDOW has been cost-sharing with selected landowners to plant enhanced mixtures of grasses and forbs to benefit sharp-tailed grouse. In 1998, 1,224 acres of CRP lands and 110 acres of non-CRP ground were planted with the enhanced seed mixtures at a cost of $14,940.00 to the Division. This program will be expanded in 1999 to include establishment of shrub thickets and treatment of 1,500 acres of CRP and 500 acres of non-CRP ground .. Cost of establishing the shrub thickets will be shared with the NRCS through their Wildlife Habitat Improvement Program (WIllP). Depending on the site, the thickets will encompass 0.25 to 0.50 acres and include a mixture of serviceberry, chokecherry, hawthorn (Crataegus spp.), and possibly skunkbush sumac (£hus trilobata). The thickets will be located within or immediately adjacent to CRP fields. The thickets will be fenced to protect against browsing by big game and domestic livestock. Due to the high cost of seed, especially for native grasses and forbs, the enhanced mixtures planted for grouse have been less than ideal. In an effort to obtain more desired seed mixtures, proposals have been submitted to two local Habitat Partnership Program (HPP) committees requesting funding to purchase bulk quantities of high quality native forage (i.e., bluebunch wheatgrass, Idaho fescue, Sherman big bluegrass, and basin wild rye) for planting new CRP and re-seeding existing CRP. The HPP program, also administered by the CDOW, returns a portion of the money generated from big game license sales back to the local communities for use in alleviating game damage. The proposals were prepared with the objective of creating a diverse mix of grasses and forbs, including legumes, that provide quality forage for big game and nesting and brood-rearing habitat for sharp-tailed grouse. The mix would be planted on CRP ground only in an effort to attract big game away from pastures used by domestic livestock and onto CRP, which cannot be grazed by domestic livestock. At least 25% of the mixture must contain a sod-forming grass to qualify for the CRP program. Thus, the mixture that has been proposed to the HPP committees includes 20% alfalfa (Ladak and Ranger), 25% meadow brome (qualifies as sod-former), 10% tall wheatgrass, 10% bluebunch wheatgrass, 10% Sherman big bluegrass, 10% Idaho fescue, 10% cicer milkvetch or sainfoin, and 5% flax. The mixture will cost approximately $32.00/acre to plant, not including labor (landowner contribution). Segment Objective 5 - A study plan entitled "Ecology of Columbian Sharp-tailed Grouse Breeding in Conservation Reserve and Post-Act Mine Reclamation Lands in Northwestern Colorado" has been prepared and submitted for internal and external peer review. Data collection is schedule to begin in April 1999. Segment Objective 6 - Traditionally, lek counts were designed to provide information on average number of males per lek and average number of birds per lek. These estimates, when collected consistently over long periods, were presumed to provide trends in population size. However, Kobriger (1975) concluded that lek counts have little value in measuring population size or documenting population trends of sharp-tailed grouse because of inconsistent lek attendance patterns within and among years. Beck and Braun (1980) likewise concluded that high variation in attendance patterns by males at leks seriously limits the utility of lek counts as a population index for sage grouse. Cannon and Knopf (1981) recommended replacing lek counts with lek surveys (i.e., number of active leks) as a trend index to prairie grouse populations. Their recommendation was based on the observation that when populations increase, males respond by forming more leks instead of increasing the average number of males on each lek. These findings suggest that lek surveys should include two components: (1) surveys of known lek sites to ascertain status (active or inactive), and (2) searches for new leks. If lek counts are conducted, the results should be interpreted with caution (Rippin and Boag 1974, Emmons and Braun 1984). �167 Lek surveys were conducted between 1 April and 6 June and consisted of the following: (1) surveys of known lek sites to ascertain status (active or inactive), (2) searches for new leks, and (3) counts of the total birds per lek, and if possible, the number of males per lek. Accurate counts were not always possible. Birds were frequently obscured by vegetation or there was no vantage point from which to observe the entire lek. In such cases, a flush count was obtained. Also, due to the vast area that needed to be searched and the large number of leks that needed to be surveyed, most leks were checked only once. In 1997, searches for new leks and surveys of known leks were conducted primarily in Routt County (excluding the extreme northern and southern parts) and a small portion of east-central Moffat County. In 1998, searches for new leks were focused in eastern Moffat County and southern and north-central Routt County, whereas, surveys of known leks were conducted throughout Routt and eastern Moffat counties. Lek surveys also were conducted by Area 10 and 6 personnel and by private consultants working for Peabody and Colowyo Coal companies. This report summarizes all surveys and counts conducted in 1997 and 1998. Data collected in 1998 were used to revise the survey results for 1997. This report contains the most current information on the location and status of all known sharp-tailed grouse leks in northwest Colorado and should be used as the basis for future surveys. The original WRIS (Wildlife Resource Inventory System) database contained 77 sharp-tailed grouse lek locations for Routt (n =57) and Moffat counties (n = 20). This database was revised in 1997 to reflect more current information. The following duplicate listings (more than one name for the same lek) were deleted from the database: Gray's Divide - same as Soash Missy - same as Fish Creek Wiseman's 2 - same as Long Gulch RCR27-9SE - same as Annan's 1 and, RCR27-9SENW - same as Annan's 1. In addition, there were two leks named Cottonwood Creek of which one was not a valid site, and thus, was deleted from the database. The valid Cottonwood Creek lek was renamed Dry Fork Elkhead to more accurately reflect its location. Five known leks were added to the database including three leks (Hocket, Buck Mountain 1, and Wilderness Ranch) found in 1996, an historic lek site (Morapas Gas Field) reported by Rogers (1969), and Slater Park 1, a known site that was periodically checked by USFS personnel. These sites were never entered into the original database. The revised database contained 76 leks of which 54 were in Routt County and 22 in Moffat County. Forty-five of the 54 known lek sites in Routt County and nine of22lek checked in 1997; 30 and 6 sites, respectively, were classified as active known leks, at least 33, and possibly 40, new leks were found in 1997 County. Counts were obtained for 43 of the 94 leks checked in 1997. 13.1 and males per lek averaged 11.4 (Table 3). sites in Moffat County were (Table 1). In addition to the (Table 2). All were in Routt Total birds per lek averaged �· 168 The database was revised again after the 1997 field season. The 40 new leks (Table 2) found in 1997, plus the following 4 leks found by consultants working for Colowyo and Peabody Coal companies were added to the database: Annan's 3 - found in 1993 Morgan - found in 1997 Seneca Mine 1 - found in 1995 and, Wilson - found in 1994. Also, Salt Creek leks 1 and 2 were combined into one lek and renamed George's Gulch and Elk Mountain 4 (also called Clark's Pasture) was deleted from the database because it was not a separate lek, but instead was a replacement lek for Elk Mountain 3. The Salt Creek sites were abandoned in favor of a single, large lek that formed in a nearby CRP field. Birds at Elk Mountain 3 relocated to the site identified as Elk Mountain 4 when the sagebrush flat where they displayed was plowed and reseeded to grasses for pasture. The revised database now included 118 leks of which 94 were in Routt County and 24 in Moffat County. In 1998, we checked 23 of24 known lek sites in Moffat County and 12 were classified as active (Table 4). This included the 13 sites not checked in 1997. The only known Moffat County lek not checked in 1998 was Elkhead Road 4. This lek was checked and classified as inactive in 1997. We found 15 new leks in Moffat County in 1998 (Table 5). Trapper Mine 1, which was found on mine reclamation land, was initially classified as a new lek. However, this lek was within 200 m of the historic Wingate lek. No effort had been made to check the Wingate lek for 10+ years because it was assumed the sharptails had abandoned the area due to the mining activity. Although the birds were probably displaced temporarily, they reoccupied the site after it was reclaimed. Based on the size of the lek (23 males), it had probably been established since at least 1995. Seventeen years passed from when the area was mined (1978) and 11 years passed from when reclamation was completed (1984) to when sharptails reoccupied the area Another lek (Trapper Mine 2) was located about 2 km east of Trapper Mine 1 (Wingate). Both the Trapper Mine leks are within 1,500 m of an active pit. In 1998, we checked 84 of the 94 known lek sites in Routt County and 56 were classified as active (Table 4). This included the 8 sites not checked in 1997. We suspect that some leks were mistakenly classified as inactive because they were checked late in the season or during inclement weather when birds were absent. We found 13 new leks in Routt County in 1998 (Table 5). Total counts were obtained for 85 leks in 1998 (Table 6). Birds per lek averaged 14.4 (range The average number of males per lek was 12.7 (range = 2-44,!! = 72). = 2-52). �169 Table 1. 1997 Columbian sharp-tailed grouse lek surveys - status of previously documented leks. Active - Routt Inactive - Routt Active - Moffat Annan's 1* Annan's 2* Barnes Dry Elkhead Ridge* Dry Fork Elkhead * Eckman Park 1 (Energy Fuels) database) Elkhead Road 3* Elk Mountain 2 (Sam's)* Elk Mountain 4 (Clark's Pasture)** Foidel Creek* George's Gulch ( Salt Creek 1&2)*** Hayden Divide* Heidel Hinkle Hocket (not in database) Horton Knoll 1 Maneotis McKinney Ranch* _ Morgan Creek Mud Springs* Rock Creek 2 Sage Creek* Smiths* Soash (Gray's Divide) Twentymile Twentymile Cliffs 4 (Scott) Woods Yellowjacket Road (Sidney Peak)* Yellowjacket 2 (Willie Ranch) 80 Road California Park Road 1* California Park Road 2 Dry Gulch 2* Dry Gulch 3 Elk Mountain 1* Buck Mountain 1 (not in database) Little Buck 1 Little Buck 2 Long Gulch (Wiseman's 2) Villards Wilderness Ranch (not in Elk Mountain 3 * Elk River Cemetery* Gillilands* Green Acres Hicks Milner Robinson Sherrod-Sandelin Yellowjacket 1* Not Checked - Routt California Park County Airport Dinwiddie Fish Creek (Missy) Five Pines Mesa Horton Knoll 2 Rock Creek 1* Slater Park 1 (not in database) Wymans * historic lek site ** possible replacement lek for Elk Mountain 3 *** two leks combined into 1 Inactive - Moffat Elkhead Road 1* Elkhead Road 2* Elkhead Road 4 Not Checked - Moffat Baker's Peak 1 Cedar Hill Gulch Fly Creek Fortification Rocks Des Dome (Stinking Gulch)* McInturf Mesa Morapas* (not in database) Nolands Pelleys* Schneiders* Taylors* Wingate* Wiseman's 1* �170 Table 2. 1997 Columbian sharp-tailed LekName County Bloomquist Calf Creek Deep Creek Dry Creek Eckman Park 2 Eckman Park 3 Eckman Park 4 Eckman Park 5 Eckman Park 6 Elk Creek 1 Elkhead Finger Rock Gnat Hill Hightail Hillberry Hoffman Homestead Homestead Ditch Little Hunter Middle Creek Morning Crow North Giant Penny Pleasant Valley RCR33b Ricks Rogers Saddle Mountain 1 Saddle Mountain 2 Shivers Smuin Gulch Gravel Pit Smuin Gulch Oil WeIll Stokes Gulch Twenty-mile Cliffs 1 Twenty-mile Cliffs 2 Twenty-mile Cliffs 3 Twenty-mile Cliffs 5 Turner Creek Warrick Pasture Wolf Mountain Ranch Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt grouse lek surveys USGS Quad Wolf Mountain Quaker Mountain Clark BlacktailMountain Rattlesnake Butte Rattlesnake Butte Rattlesnake Butte Rattlesnake Butte Rattlesnake Butte Milner Quaker Mountain Quaker Mountain Breeze Mountain Rattlesnake Butte Mount Harris Cow Creek Rattlesnake Butte Cow Creek Clark Cow Creek Mad Creek Mad. Creek Rock Springs Blacktail Mountain Cow Creek Rattlesnake Butte Rattlesnake Butte Cow Creek Cow Creek Rattlesnake Butte Hayden Hayden Breeze Mountain Rattlesnake Butte Rattlesnake Butte Rattlesnake Butte Rattlesnake Butte Wolf Mountain Hooker Mountain Hooker Mountain - new lek locations. Legal 7N86W4SW 8N88W14NE 8N85W30SW 5N84W32NW 4N86W18NW 4N86W18NE 4N86W18SE 4N86W8SE 4N86W33SW 6N86W28NE 8N88W13NE 8N88WliSW 6N89W20SE 4N87W12NE 5N87W6SW 5N86W12SW 4N87W12SW 5N86W24NE 8N85W19NW 5N86W13 SE 7N85W8NW 7N85W6NW 8N88W29SE 4N84W9NW 6N85W30SW 5N86W26NW 5N87W36SW 6N85W19NW 6N86W23SE 4N87W11SE 6N89W21NE 6N89W23SE 5N89W3NW 5N86W30SE 5N87W25SE 5N86W19SW 5N86W20SE 7N86W5SW 7N88WliSW 6N87W4SE UTMX UTMY 327400 4495050 311850 4502800 333550 4498950 344050 4467900 322500 4465700 323250 4465600 323200 4464600 325300 4466200 326150 4467450 327500 4479850 313600 4502100 311050 4503150 297200 4481150 320900 4467100 313700 4476400 330900 4474250 320350 4466450 332150 4471750 333650 4500500 331900 4472650 334650 4494550 333250 4495550 307200 4498650 345350 4466650 333250 4478950 329500 4470050 321250 4467800 332750 4481650 332600 4480900 319750 4466350 298750 4482050 301800 4481500 298850 4477200 323600 4468600 322350 4469500 322950 4470700 325850 4470600 325250 ·4495650 311250 4493900 318350 4485350 Comments riot sure if lek not sure if lek not sure if lek not sure if lek not sure if lek not sure if lek satellite 20mi#2 satellite 20mi#4 not sure if lek �171 Table 3. 1997 Columbian LekName Annan's 2 Bloomquist Calf Creek Deep Creek Dry Creek Dry Elkhead Ridge Eckman Park 1 Eckman Park 2 Eckman Park 3 Eckman Park 4 Eckman Park 5 Eckman Park 6 Elk Creek 1 Elkhead Elkhead Road 3 Elk Mountain 2 Elk Mountain 4 Foidel Creek George's Gulch Hightail Hillberry Hinkle Hoffman Homestead Homestead Ditch Maneotis McKinney Ranch North Giant RCR33b Ricks Rogers Saddle Mountain 1 Shivers Smuin Gulch Gravel Pit Smuin Gulch Oil Well 1 Twentymile Cliffs 1 Twentymile Cliffs 2 Twentymile Cliffs 3 Twentymile Cliffs 4 Twentymile Cliffs 5 Turner Creek Woods Yellowjacket Road Total Leks Counted Total Birds Counted Average BirdslLek (SD) sharp-tailed Males grouse lek counts. Total Birds 16 9 6 9 4 17 18 17 10 10 11 12 6 11 13 10 5 11 7 15 7 19 44 10 15 5 6 14 10 3 10 4 8 20 14 3 43 562 13.1 (9.1) 16 9 12 6 10 4 20 21 20 10 10 13 15 10 8 6 35 11 24 15 11 5 11 6 7 15 10 19 53 10 16 5 6 14 10 3 11 4 10 25 16 17 3 36 409 11.4 (7.2) Lek Status historic new new new. new historic known new new new new new new new historic historic historic? historic historic new new known new new new known historic new new new new new new new new new new. new known new new known historic Comments Energy Fuels minimum count minimum count minimum count minimum count Clark's Pasture Salt Creek 1 and 2 may be satellite lek may be satellite lek Scott minimum count Sidney Peak �172 Table 4. 1998 Columbian sharp-tailed grouse lek surveys - status of previously documented leks. Active - Routt 80 Road Annan's 1* Annan's 2* Annan's 3 Barnes Bloomquist California Park 1 Dry Creek Dry Fork Elkhead * Eckman Park 1 (Energy Fuels) Eckman Park 2 Eckman Park 3 Eckman Park 4 Eckman Park 6 Elk Creek 1 Elkhead Road 3* Elk Mountain 2 (Sams)* Elk Mt 3 (Elk Mt 4/Clark's Pasture)* Fish Creek (Missy) Foidel Creek* George's Gulch (Salt Creek 1&2)* Hayden Divide* Heidel Hightail Hillberry Hinkle Hocket Hoffman Homestead Homestead Ditch Horton Knoll 1 Maneotis Morgan Creek Reservior Mud Springs* North Giant RCR33b Ricks:" Rock Creek 2 Seneca Mine 1 Slater Park 1 Smiths * Smuin Gulch Gravel Pit Smuin Gulch Oil WeIll Soash Stokes Gulch TumerCreek Twentymile Twentymile Cliffs 1 * historic lek site Active - Routt Twentymile Cliffs 2 Twentymile Cliffs 3 Twentymile Cliffs 4 (Scott) Twentymile Cliffs 5 Warrick Pasture Woods Wymans Yellowjacket Rd (Sidney Peak)* Inactive - Routt California Park Road 1* California Park Road 2 County Airport Deep Creek Dinwiddie Dry Gulch 2* Dry Gulch 3 Eckman Park 5 Elk River Cemetery* Five Pines Mesa Gillilands· Gnat Hill GreenAcres Hicks Horton Knoll 2 McKinney Ranch= Milner Middle Creek Morning Crow Penny Pleasant Valley Robinson Rogers Rock Creek 1· Sage Creek" Sherrod-Sandelin Shivers Yellowjacket 1· Not Checked - Routt Calf Creek Dry Elkhead Ridge" Elkhead Elk Mountain 1· Finger Rock Little Hunter Saddle Mountain 1 and 2 Wolf Mountain Ranch Yellowjacket 2 (Willie Ranch) Active - Moffat Buck Mountain 1 Fly Creek Iles Dome (Stinking Gulch)* Little Buck 1 Little Buck 2 Long Gulch (Wiseman's 2) Morapas Gas Field" Morgan Nolands Trapper Mine 1 (Wingate)" Wilderness Ranch 1 Wilson Inactive - Moffat Baker's Peak Cedar Hill Gulch Elkhead Road 1* Elkhead Road 2* Fortification Rocks McInturf Mesa Pelleys· Schneiders= Taylors" Wiseman's 1· Villards Not Checked - Moffat Elkhead Road 4 �173 Table 5. 1998 Columbian sharp-tailed grouse lek surveys - new lek locations. LekName County USGS Quad Big Elk 1 Big Elk 2 Buck Mountain 2 Buck Mountain 3 Buck Mountain 4 Bum California Park 2 California Park 3 Cole Gulch 1 Cole Gulch 2 Cull Reservior Dresher Reservior Earle Edna Mine Elk Creek 2 MCR18 Pinnacle Mountain 1 Pinnacle Mountain 2 Pinnacle Mountain'S Pondella Ranch Postovit Seneca Mine 2 Smuin Gulch Oil Well 2 Straight Gulch Trapper Mine 2 William's Park Windemere Yampa Routt Routt Moffat Moffat Moffat Moffat Routt Routt Moffat Moffat Moffat Moffat Routt Routt Routt Moffat Moffat Moffat Moffat Moffat Routt Routt Routt Moffat Moffat Routt Routt Routt Hayden Hayden Slide Mountain Slide Mountain Slide Mountain Easton Gulch Quaker Mountain Bears Ears McInturf Mesa McInturf Mesa Freeman Reservior Breeze Mountain Hayden Oak Creek Milner Slide Mountain Slide Mountain McInturf Mesa McInturf Mesa Fortification Hayden Milner Hayden Easton Gulch Castor Gulch Dunkley Mad Creek Yampa Legal U1MX 5N88W7NE 305200 5N88W7SE 304450 8N89WliNE 302650 8N89W2NW 301400 8N89WllSW 301250 4N94W27SE .251900 9N87W16SE 318700 9N87WI0NW 319450 8N89W19SE 295850 8N89W21SW 298500 10N90W9NW 289650 6N90WliSE 292500 6N89W36SE 302950 333150 4N85W7SE 6N86W21SE 327600 8N89W15SW 300100 9N89W34NE 300550 9N89W32NW 297100 8N89W9NW 298350 10N90W5SE 288650 5N88W17NE 306650 5N87W12NW 321550 6N89W28NE 298600 4N94W27NW 250400 5N91W3NE 280950 4N87W19NW 312250 7N86W36SE 332500 2N85W16SE 336150 U1MY Comments 4475500 4475150 4504800 4505750 4503300 4463450 4511800 4514350 4500900 4500200 4523800 4484650 not sure if lek 4478050 4466000 4480650 not sure if lek 4501550 4507850 4507500 4504600 near historic Pelly lek 4524800 4474350 4474550 4479900 4464050 4477950 4464100 4487150 4444500 �174 Table 6. 1998 Columbian LekName 80 Road Annan's 1 Annan's 2 Annan's 3 Big Elk 1 Big Elk 2 Bloomquist Buck Mountain 1 Buck Mountain 2 Buck Mountain 3 Buck Mountain 4 Burn California Park 1 California Park 2 California Park 3 Cole Gulch 1 Cole Gulch 2 Cull Reservior Dresher Reservior Dry Creek Dry Fork Elkhead Earle Eckman Park 1 Eckman Park 2 Eckman Park 3 Eckman Park 4 Eckman Park 6 Edna Mine Elk Creek 1 Elk Mountain 2 Elk Mountain 3 Fish Creek Fly Creek George's Gulch. Hayden Di.vide .. Heidel Hightail Hillberry Hinkle Hocket Hoffman Homestead Ditch Horton Knoll 1 IlesDome Little Buck 1 Little Buck 2 sharp-tailed Males 8 22 14 18 8 11 14 8 3 2 19 2 10 4. 4 13 18 2 17 30 12 17 11 8 13 9 9 31 25 17 8 22 6 8 18 7 3 7 grouse lek counts. Total Birds 10 18 27 17 18 8 14 20 9 3 3 21 2 11 4 6 15 22 2 17 34 12 19 17 11 8 13 12 9 16 42 15 12 22 25 17 8 22 6 12 29 7 11 6 15 8 Lek Status Comments known historic historic known new new? known known new new new new known new new new new new new known known new known known known known known new known historic known known known historic historicknown known known known known known known known historic known known previously inactive Rocky Creek, Energy Fuels Eckman Park Reclamation Aspen Island Lonebush Missy replaced Salt Creek leks 1&2 moved, HG Reclamation �175 Table 6 (continued). 1998 Columbian sharp-tailed grouse lek counts. LekName Long Gulch Maneotis MCR18 Morapas Gas Field Morgan Morgan Creek Reservior Mud Springs Nolands North Giant Pinnacle Mountain 1 Pinnacle Mountain 2 Pinnacle Mountain 3 Pondella Ranch Postovit RCR33b Ricks Rock Creek 2 Seneca Mine 1 Seneca Mine 2 Slater Park I Smiths Soash Smuin Gulch Gravel Pit Smuin Gulch Oil WeIll Smuin Gulch Oil Well 2 Stokes Gulch Straight Gulch Trapper Mine 1 Trapper Mine 2 Turner Creek Twentymile Cliffs 2 Twentymile Cliffs 4 Twentymile Cliffs 5 Wilderness Ranch 1 William's Park Wilson Woods Yampa Yellow jacket Road Total Leks Counted Total Birds Counted Average BirdslLek (SD) Males 2 7 4 44 30 20 7 9 9 4 10 10 38 10 12 15 12 7 6 15 10 10 21 14 23 6 15 24 9 14 5 24 5 4 Total Birds 2 7 4 21 52 30 20 7 17 13 10 4 10 14 44 10 12 17 14 10 11 8 15 10 10 30 17 23 7 15 24 9 17 7 7 28 6 5 4 72 85 913 1226 12.7 (8.6) 14.4 (9.3) Lek Status Comments known known new historic known known historic known known new new new new new known known known known new known historic known known known new known new new new known known known known known new known known new historic Wiseman's 2 lek has moved Moffat Routt, minimum count minimum count minimum count Wingate minimum count Scott Sidney Peak �176 We created a new database of all (n = 146) known active and inactive lek sites in Moffat (n = 39) and Routt (n = 107) counties (Appendix A). This database includes all leks listed in the original WRIS database, previously documented leks that were never entered into the WRIS database, and new leks located in 1997 and 1998. Ninety-two percent of the leks occurred on private land; 43 and 23 % of the active leks were found on Conservation Reserve Program and mine reclamation lands, respectively. Rogers (1969) reported finding 21 leks in Routt County and 10 leks in Moffat County. These are identified as historic sites in the database. Rogers (1969) reported that Elkhead Road 3 was in Moffat County, whereas, it is actually in Routt County. Two other sites (Wingate and Cottonwood Creek) identified as historic leks have been renamed (Trapper Mine 1 and Dry Fork Elkhead, respectively) to better reflect where they are located. Of the 31 historic sites surveyed in 1997 and 1998, 17 were still active. Although sharptails are known to occur west of Colorado 13 north of Craig (Baggs Highway), no leks were found in this area during 1998. Fortification Rocks, the only lek in the database located west of the Baggs Highway, was checked twice, but no birds were observed. No effort was made to search for leks west of Colorado 13 south of Craig to Axial Basin. There are no known leks in this area, but sharptails may occur there. The only leks known to occur west of Colorado 13 south of Craig were found by John Monarch on Colowyo property south of Axial Basin. Leks were found up to 13 km west of Axial. No surveys were conducted in Rio Blanco County. Sharptails have been observed in north-central Rio Blanco County, but no leks have been documented to date. The nearest active lek (Morapas Gas Field) was in Moffat County within 1.4 km of the county line. The nearest known lek was Taylors, an historic site identified by Rogers (1969). This lek was within 0.5 km of Rio Blanco County. It was checked twice, but no birds were observed. Five Pines Mesa, the only known lek in southern Routt County between Oak Creek and Toponas, was inactive. However, another new lek was located in this area about 1.5 km southwest of Yampa Recommendations 1. The following leks are inactive and can be deleted from the database: California Park Road 1 California Park Road 2 County Airport Dry GUlch 2 Dry Gulch 3 Elkhead Road 1 Elkhead Road 2 Elk Mountain 1 Elk River Cemetery Five Pines Mesa 2. Gillilands GreenAcres Hicks Horton Knoll 2 Milner Robinson Schneiders Sherrod-Sadelin Taylors Information about inactive leks should be maintained in a separate database in case the sites become active in the future. �177 3. Lek searches should be intensified within a 1 km radius of inactive leks to insure the lek has not relocated to a new site. Any active lek found within 1 km of an inactive lek should be classified as a known lek that has shifted location. If the known lek is active and another lek is found within 1 km, it can be classified as a new lek provided it is at least 0.5 km from the known lek and has four or more males on the lek. Otherwise, it should be classified as a satellite lek. 4. Need to verify if the following sites are valid lek locations: Penny Pleasant Valley Gnat Hill Middle Creek Dresher Reservoir Morning Crow Saddle Mountain 2 Wolf Mountain Ranch Elk Creek 2 From 1 to 5 males were observed displaying at Gnat Hill, Penny, Pleasant Valley, Middle Creek, and Morning Crow in 1997, but no birds were observed at these sites in 1998. All were new leks in 1997. Saddle Mountain 2 and Wolf Mountain Ranch were not checked in 1998, but they were classified as questionable new lek sites in 1997 due to their proximity to other leks. DresherReservoir and Elk Creek 2, which were found in 1998, also were classified as questionable lek sites, due to their proximity to other leks. 5. The primary goal should be to check all leks each year to determine if they are active or inactive. This may not be possible due to manpower constraints. Thus, any lek not checked in a given year should be given priority the next year. The following leks were not checked in 1998 and therefore should be checked in 1999: Calf Creek Elkhead Finger Rock Little Hunter Saddle Mountain 1 Wolf Mountain Ranch Dry Elkhead Ridge Elk Mountain 1 Foidel Creek Elkhead Road 4 Saddle Mountain 2 Yellowjacket 2 6. Lek surveys are often conducted by District Wildlife Managers in conjunction withArea Biologists; therefore, leks should be identified according to the DWM District where they are located (Appendix B). The DWMs should receive a list of alilek sites within their district each year. The biologist and DWM should meet to decide who will survey/count the leks within that district. 7. The secondary objective should be to obtain counts for at least 40 leks each year. These should be identified as the count leks. The initial sample should be randomly selected from the most current list of active leks (Appendix A). If a count lek becomes inactive, it should be replaced with another lek randomly-selected from the list of known active leks (excluding the count leks). 8. Counts should be conducted during the peak of breeding activity, which in most years occurs between 20 April and 3 May in northwest Colorado. Adjust the timing one week earlier following mild winters and one week later following severe winters. Conduct a minimum of two and preferably three counts per lek. Conduct the counts from Y2 hour before sunrise to 2 hours after sunrise. Attempt to count on mornings with no precipitation and with wind speeds <15 mph. �178 9. The preferred method to conduct the count is to find a suitable vantage point from which the entire lek can be observed. However, this is not always possible. The alternative method is to flush the birds off the lek and count birds flying away. Be sure to walk through the lek several times to ensure you flushed all the birds. It is not necessary, and frequently is not possible, to distinguish males from females The objective is to obtain a total count of birds on the lek. Promptly leave the lek after conducting the count. 10. Use the standardized form included in this report (Appendix C) to record alilek count and survey data Fill out the form each time you visit a lek site regardless of whether you observe birds or not. . 11. Leks checked late in the morning, late in the season, or during inclement weather may be mistakenly classified as inactive. Whenever birds are not observed at a known lek site, it is best to search the site for signs of activity (i.e., droppings, tracks, feathers, matted vegetation). Leks that may have been rnisclassified as inactive in 1998 include Deep Creek, Dinwiddie, Eckman Park 5, McKinney Ranch, Rogers, Sage Creek, and Shivers. Efforts should be made to confirm the status of these leks in 1999. 12. Efforts to locate new leks should be focused in north-central Rio Blanco County, southern Routt County between Oak Creek and Toponas, and west of Highway 13 north and south of Craig. Any leks located in these areas will help more clearly define the distribution of sharptails in northwest Colorado. LITERATURE Bailey, A. M., and R J. Niedrach. CO. 454pp. CITED 1965. Birds of Colorado, Vol. 1. Denver Mus. Nat. Hist., Denver, Beck, T. D. 1, and C. E. Braun. The strutting ground count: variation, traditionalism, management needs. Proc. West. Assoc. Fish and Wildl. Agencies 60:558-566. Cannon, R W., and F. L. Knopf 1981. Lek numbers as a trend index to prairie grouse populations. J. Wildl. Manage. 45:776-778. Carlton. J. C .. 1995. Petition for a rule to list the Columbian sharp-tailed grouse, Tympanuchus phasianellus columbianus, as "threatened" or "endangered" in the conterminous United States under the Endangered Species Act, 16 U.S.C. Sec. 1531 et seq. (1973) as amended. Biodiversity Legal Foundation, Boulder, CO. 52pp. Dargan, L. M., H. R Shepherd, and R N. Randall. 1942. Data on sharp-tailed grouse in Moffat and Routt counties. Colorado Game, Fish, and Parks Dep, Sage Grouse Survey, Vol 4, Denver. 28pp. Emmons, S. R, and C. E. Braun. 48:1023-1028. 1984. Lek Attendance of male sage grouse. J. Wildl. Manage. �179 Giesen, K. M. 1985. Inventory of Columbian sharp-tailed grouse in western Colorado. Colorado Div. Wildl., Unpubl. Rep, Fort Collins. 6pp. Giesen, K. M. 1987. Population characteristics and habitat use by Columbian sharp-tailed grouse in northwest Colorado. Pages 251-279 in Wildlife Res. Rep., Part 2. Colorado Div. Wildl., Fed Aid Proj. W-152-R, Apr. 1987. Giesen, K. M, and C. E. Braun. 1993. Status and distribution of Columbian sharp-tailed grouse in Colorado. Prairie Nat. 25:237-242. Giesen, K. M., and 1. W. Coruielly. 1993. Guidelines for management of Columbian sharp-tailed grouse habitats. Wildl. Soc. Bull. 21 :325-333. Kobriger, G. D. 1975. Correlation of sharp-tailed grouse population parameters. North Dakota Outdoors 25(5): 10-13. Meints, D. R, J. W. Connelly, K. P. Reese, A. R Sands, and T. P. Hemker. 1992. Habitat suitability index procedure for Columbian sharp-tailed grouse. Univ. Idaho For., Wildl., and Range Exp. Stn. Bull. 55: 27pp. . ., Miller, G. C., and W. D. Graul. 1980. Status of sharp-tailed grouse in North America Pages 18-28 in P. A. Vohs and F. L. Knopf, eds. Proceedings of the prairie grouse symposium. Oklahoma State Univ., Stillwater. Rippin, A. B., and D. A. Boag. 1974. Recruitment to populations of male sharp-tailed grouse. 1. Wildl. Manage. 38:616-621. Robel, R J., J. N. Briggs, A. D. Dayton, and L. C. Hulbert. 1970. Relationships between visual obstruction and weight of grassland vegetation. J. Range Manage. 23:295-297. Rogers, G. E. 1969. The sharp-tailed grouse in Colorado. Colorado Division Game, Fish, and Parks Tech. Publ. 23. 94pp. U. S. Department of the Interior. 1989. Endangered and threatened wildlife and plants; annual notice of review; proposed rules. Fed. Register 54:560. Prepared by : ~~!lfFLSSRIV �•.... APPENDIX A - Columbian sharp-tailed grouse lek locations in Colorado. LekName 80 Road Annan's Twenty Mile 1 Annan's Twenty Mile 2 Annan's Twenty Mile 3 Baker's Peak Barnes Big Elk 1 Big Elk 2 Bloomquist Buck Mountain 1 Buck Mountain 2 ' Buck Mountain 3 Buck Mountain 4 Bum Calf Creek California Park 1 California Park 2 California Park 3 California Park Road 1 California Park Road 2 Cedar Hill Gulch Cole Gulch 1 Cole Gulch 2 County Airport Cull Reservoir Deep Creek Dinwiddie Dresher Reservoir Dry Creek Dry Elkhead Ridge Dry Fork Elkhead Dry Gulch 2 Dry Gulch 3 Earle Eckman Park 1 Eckman Park 2 Eckman Park 3 Eckman Park 4 County USGS Quad Routt Routt Routt Routt Moffat Routt Routt Routt Routt ' Moffat Moffat Moffat Moffat Moffat Routt Routt Routt Routt Routt Routt Moffat Moffat Moffat Routt Moffat Routt Routt Moffat Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt HookerMt MOWlt Harris MOWlt Harris MOWlt Harris Baker's Peak Hayden , 'Hayden ' Hayden WolfMt Slide Mt SlideMt Slide Mt SlideMt Easton Gulch QuakerMt Bears Ears QuakerMt Bears Ears HookerMt HookerMt McInturf Mesa McInturf Mesa McInturf Mesa Rocky Peak Freeman Reservoir Clark Hayden BreezeMt Blacktail Mt Rock Springs Gulch QuakerMt Mad Creek Mad Creek Hayden Rattlesnake' Butte Rattlesnake Butte Rattlesnake Butte Rattlesnake Butte Legal UTMY 7N88W24NW 5N87W9SE 5N87W5SE 5N87W9NW IIN90WI6NE 5N88W18SE 5N88W7NE 5N88W7SE 7N86W4SW 9N89W36SE 8N89W11NE 8N89W2NW 8N89W11SW 4N94W27SE 8N88W14NE 9N87W9SW 9N87W16SE 9N87W10NW 7N88W12NE 7N88W23SE 8N89W6NE 8N89W19SE 8N89W21SW 6N85W1NW 10N90W9NW 8N85W30SW 5N89W14NE 6N90W11SE 5N84W32NW 7N88W4SW 8N88W24SE 7N85W21NW 7N85W16SE 6N89W36SE 4N88W18SW 4N86W18NW 4N86W18NE 4N86W18SE 312350 317800 316450 317100 290200 305050 305200 304450 327400 304250 302650 301400 301250. 251900 311850 318100 318700 319450 313600 311800 296150 295850 298500 341850 289650 333550 301500 292500 344050 308150 313350 336300 337400 302950 321900 322500 323250 323200 00 UTMX 4491600 4474600 4476000 4474950 4532200 4473350 4475500 4475150 4495050 4506900 4504800 4505750 4503300 4463450 4502800 4513150 4511800 4514350 4494200 4490800 4506400 4500900 4500200 4486000 4523800 4498950 4473600 4484650 4467900 4495150 4500050 4491150 4491600 4478050 4464550 ' 4465700 4465600 4464600 Type· K H H K K K N N N K N N N N N K N, N H K K N N K N N K N N H H H K N K N N N Status" I A A N N A N N A A N N N N A N N N I I N N N N N A N N A A A I I N A A A A A A A A I I A A A A A A A A N A A A I I I A A I A I I A A N A I I A A A A A Comments/other names moved to west side RCR 80 birds dance next to RCR 27 not in original database need to verify status birds possibly moved to Big Elk 2, verify status found in 1998 found in 1998 found in 1997 not in original database found in 1998 found in 1998 found in 1998 found in 1998 found in 1997 found in 1998 found in 1998 found new lek (warrick) 2 km WSW found in 1998. found in 1998 lost to airport expansion, no new lek found in 1998 found in 1997, birds seen in area in 98 found in 1998 found in 1997 active in 1997 Cottonwood Creek found in 1998 old EnerFuels lek that relocatedlRocky Creek found 1997/Eckman lek found 1997IReclamation lek found 19971Aspen Island �APPENDIX A (continued) - Columbian sharp-tailed grouse lek locations in Colorado. LekName Eckman Park 5 Eckman Park 6 Edna Mine Elk Creek 1 Elk Creek 2 Elkhead Elkhead Road 1 Elkhead Road 2 Elkhead Road 3 Elkhead Road 4 Elk Mountain 1 Elk Mountain 2 Elk Mountain 3 Elk River Cemetery Finger Rock Fish Creek Five Pines Mesa Fly Creek Foidel Creek Fortification Rocks George's Gulch Gillilands Gnat Hill Green Acres Hayden Divide Heidel Hicks Hightail Hillberry Hinkle Hocket Hoffman Homestead Homestead Ditch Horton Knoll 1 Horton Knoll 2 llesDome Little Buck 1 Little Buck 2 County Routt Routt Routt Routt Routt Routt Moffat Moffat Routt Moffat Routt Routt Routt Routt Routt Routt Routt Moffat Routt Moffat Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Moffat Moffat Moffat USGS Quad Legal UTMY Rattlesnake Butte Rattlesnake Butte Oak Creek Milner Milner QuakerMt Ralph White Lake Slide Mt Slide Mt Slide Mt WolfMt Mad Creek WolfMt Mad Creek QuakerMt Milner Trapper Fly Creek Rattlesnake Butte Fortification Mad Creek Rock Springs Gulch Breeze Basin Oak Creek Hayden Gulch Hayden Oak Creek Rattlesnake Butte MOWlt Harris Rattlesnake Butte MOWlt Harris Cow Creek Rattlesnake Butte Cow Creek Rock Springs Gulch HookerMt Monument Butte CraigNE McInturf Mesa 4N86W8SE 4N86W33SW 4N85W7SE 6N86W28NE 6N86W21SE 8N88W13NE 7N89W16NW 8N89W26NE 8N88W18SE 8N89W24SE 7N86W14SW 7N86W2SE 7N86WIONE 7N85W29NE 8N88WliSW 6N86W35SE IN85W2SE 12N89W25SE 5N87W36NE 9N91W4NW 8N86W26SE 7N88W19SE 6N89W20SE 5N86W25SE 5N88W30SE 6N86W24SW 5N86W25SW 4N87W12NE 5N87W6SW 5N86W27SW 6N87W31NE 5N86W12SW 4N87W12SW 5N86W24NE 8N88W33NE 8N88W26NW 4N92W35SW 9N90WI7NE 9N90W26NW 325300 326150 333150 327500 327600 313600 297950 302250 305850 304200 329900 330900 329100 335900 311050 330400 339350 304950 322500 279400 331250 305250 297200 331800 304800 302900 330900 320900 313700 328150 314900 330900 320350 332150 309100 311200 271550 288800 292000 UTMX Type" Status" 4466200 4467450 4466000 4479850 4480650 4502100 4493350 4499900 4501850 4500500 4491900 4494800 4494300 4489550 4503150 4476950 4438150 4437000 4468550· 4516200 4497950 4490800 4481150 4468950 4470450 4481250 4469300 4467100 4476400 4469050 4478950 4474250 4466450 4471750 4498300 4499200 4461250 4513300 4509750 N N N N N N H H H K H H H H N K K K H K H H N K H K K N N K K N N N K K H K K A A N A N A I I A I I A A I A N N N A N A I A I A A I A A A A A A A A N N A A I A A A A N I I A N N A A I N A I A N I A I I I A A I A A A A A A A A I A A A Comments/other names found 1997/Redeye found 19971Lonebush lek found in 1998 found in 1997 found in 1998, not sue if valid lek found in 1997 displaced by reservoir construction lek site moved from historic site inactive in 1997 Sams moved from historic sitelElk Mt 4, Clark's Pasture found in 1997 lek has moved from original site! Missy on RouttIMoffat Co line active 1997IPinnacie Peak possibly displaced by fire replaced Salt Creek 1 and 2 possibly displaced by subdivision found 1997/not sure if valid lek moved across road! HG Reclamation found in 1997 found in 1997 not in found found found original database in 1997 in 1997 in 1997 lek has moved! Stinking Gulch need better location .... 00 .... �..... APPENDIX A (continued) - Columbian sharp-tailed grouse lek locations in Colorado. LekName Little Hunter Long Gulch Maneotis Mcinturf Mesa McKinney Ranch MCR18 Middle Creek Milner Morapas Gas Field Morgan Morgan Creek Reservoir MomingCrow Mud Springs Nolands North Giant Pelleys Penny Pinnacle Mountain 1 Pinnacle Mountain 2 Pinnacle Mountain 3 Pleasant Valley Pondella Ranch Postovit RCR 33b Rick's Robinson Rock Creek 1 Rock Creek 2 Rogers Saddle Mountain 1 Saddle Mountain 2 Sage Creek Schneiders Seneca Mine 1 Seneca Mine 2 Sherrod-Sadelin Shivers Slater Park 1 County Routt Moffat Routt Moffat Routt Moffat Routt Routt Moffat Moffat Routt Routt Routt Moffat Routt Moffat Routt Moffat Moffat Moffat Routt Moffat Routt Routt Routt Routt Routt Routt Routt Routt Routt Routt Moffat Routt Routt Routt Routt Routt USGS Quad Clark Slide Mt Oak Creek Mcinturf Mesa QuakerMt Slide Mt Cow Creek Cow Creek Thornburgh Easton Gulch HookerMt Mad Creek Rock Springs BreezeMt Mad Creek Mcinturf Mesa Rock Springs. Slide Mt Mcinturf Me;S8 Mcinturf Mesa Blacktail Mt . Fortification Hayden Cow Creek . Rattlesnake Butte Mad Creek . Rock Springs Rock Springs Rattlesnake Butte Cow Creek Cow Creek. Mount Harris Monument Butte Milner Milner Mad Creek Rattlesnake Butte Bears Ears Peak Legal UTMY 8N85W19NW 8N89W24SW 5N86W36SW 9N89W21SW 8N88W14SW 8N89W15SW 5N86W13SE 6N86W26NE 3N91W8SE 4N94W15SW 8N87W32SW 7N85W8NW 7N89W22SE 6N90W15SW 7N85W6NW 8N89W8SE 8N88W29SE 9N89W34NE 9N89W32NW 8N89W9NW 4N84W9NW 10N90W5SE 5N88WI7NE 6N85W30SW 5N86W26NW 7N85W29NE 7N88W31NE 7N89W26NE 5N87W36SW 6N85W19NW 6N86W24SE 6N88W36NE 4N91W31SE 5N87W2NW 5N87W12NW 7N85W20SE 4N87W11SE 10N87W4NE 333650 302900 330750 298750 311400 300100 331900 329950 276050 250900 316050 334650 300800 289950 333250 297500 307200 300550 297100 298350 345350 288650 306650 333250 329500 335650 304950 302000 321250 332750 332600 313150 275000 320500 321550 335900 319750 318600 00 UTMX Type- Status" 97 98 4500500 4500340 4467700 4509650 4501550 4501550 4472650 4479800 4457350 4466750 4497000 4494550 4490600 4482900 4495550 4503850 4498650 4507850 ·4507500 4504600 4466650 4524800 4474350 4478950 4470050 4489700 4488350 4489950 4467800 4481650 4480900 4478300 4461400 4476900 4474550 4490200 4466350 4525550 N K K K H .N N K H N K N H K N H N N N N N N N N N K H K N N N H H K N K N K A A A N A N A I N A A A A N A N A N N N A N N A A I N A A A A A N N N I A N N A A I I A I I A A A I A A A I I A A A I A A A A I I A I N N I I A A I I A Comments/other names found in 1997 Wiseman's 2 moved west across drainage birds seen nearby in 1998 birds seen nearby in 1998 but not on lek found in 1998 found 1997, not sure if valid lek searched for in 1997 and 1998 lek moved north, not in original database found 1997 but not reported found 1997, not sure if valid lek lek has shifted locations lek has shifted location found in 1997 possibly replaced by Pinnacle Mt 3 found 1997, not sure if valid lek found in 1998 found in 1998 found in 1998 found 1997, not sure if valid lek found in 1998 found in 1998 found in 1997 found in 1997 found 1997, probably active in 98, checked late found in 1997, need to verify status found in 1997, need to verify status need to verify status/location site was checked 3 times in 1998 found in 1996, not in original database found in 1998 inactive for several years found 1997, probably active in 98, checked late not in original database �APPENDIX A (continued) - Columbian sharp-tailed grouse lek locations in Colorado. LekName Smiths Smuin Gulch Gravel Pit Smuin Gulch Oil WeIll Smuin Gulch Oil Well 2 Soash Stokes Gulch Straight Gulch Taylors Trapper Mine 1 Trapper Mine 2 Turner Creek Twentymile Twentymile Cliffs 1 Twentymile Cliffs 2 Twentymile Cliffs 3 Twentymile Cliffs 4 Twentymile Cliffs 5 Villards Warrick Pasture Wilderness Ranch William's Park Wilson Windemere Wiseman's I Wolf Mountain Ranch Woods Wymans Yampa Yellowjacket Road YeIlowjacket 1 Yellowjacket 2 County USGS Quad Legal UTMY Routt Routt Routt Routt Routt Routt Moffat Moffat Moffat Moffat Routt Routt Routt Routt Routt Routt Routt . Moffat Routt Moffat Routt Moffat Routt Moffat Routt Routt Routt Routt Routt Routt Routt Rock Spring's Gulch Hayden Hayden Hayden Mad Creek Hayden Easton Gulch Sleepy Cat Castor Gulch Castor Gulch Wolf Mountain Mount Harris Rattlesnake Butte Rattlesnake Butte Rattlesnake Butte Rattlesnake Butte Rattlesnake Butte Freeman Reservoir Hooker Mountain Baker's Peak Dunkley Axial Mad Creek Rock Springs Gulch Hooker Mountain Mad Creek Hayden Gulch Yampa Blacktail Oak Creek Oak Creek 7N88W29SE 6N89W21NE 6N89W23SE 6N89W28NE 7N86W25SW 6N89W34SE 4N94W27NW 3N91W14SW 6N91W33SW 5N91W3NE 7N86W5SW 5N87W22NW 5N86W30SE 5N87W25SE 5N86W19SW 5N86W19SW 5N86W20SE 10N90W33NE 7N88WlISW I1N89W16SW 4N87W19NW 4N93W32SW 7N86W36SE 8N98W25SW 6N87W4SE 7N85W22NW 4N89W22NE 2N85W16SE 5N84W19SE 5N85W36NW 5N85W26NW 306900 298750 301800 298600 331550 299600 ·250400 280350 278150 280950 325250 318700 323600 322350 322950 323350 325850 289800 311250 298900 312250 257300 332550 302900 318350 338050 298850 336150 342750 340300 339350 UTMX Type· Status" 97 98 4489450 4482050 4481500 4479900 4488550 4477700 4464050 4456050 4478300 4477950 4495250 4471900 4468600 4469500 4470700 4470800 4470600 4518050 4493900 4531000 4464100 4461650 4487150 4499000 4485350 4490800 4464500 4444490 4470600 4468400 4469700 H N N N K N N H H N N K N N N K N K N K N K N H N K K N H H K A A A N A A N N N N A A A A A A A A A A N N N N A A N N A I A A A A A A A A I A A A A I A A A A I A A A A A I N A I A A I N Comments/other names found in 1997 found in 1997 found in 1998 found in 1997 found in 1998 checked in 1998 found 1998, replaced historic Wingate lek between 2 active coal pits found 1997 only sign found on lek, need to verify satellite lek ofTwentymile Cliffs 2/Cyprus found in 1997 found 97, moved 98, satellite of 20mi cliffs 4 old Scott Lek found in I 997IHope Lek need to verify status found in 1997 found in 1997 found fall 1997 found 1994, not in original database found 1998, suspected in 1997 found 1997, not sure if valid lek in Campell subdivision checked late in season, need to verify Sidney Peak need to verify/Willie Ranch • H = historic lek site (identified in Rogers, G. E. 1969. The sharp-tailed grouse in Colorado. Colo Div. Game, Fish, and Parks Tech. PubI23.); K new lek site found in 1997 or 1998. b A = active, I = inactive, N = known lek site (found prior to 1997); N = = not checked. •... 00 w �184 APPENDIXB Distribution of Columbian sharp-tailed grouse known lek sites lek by DWM districts, Routt County, northwest Colorado. Steamboat North Steamboat 80 Road Annan's 20-mile 1, 2, and 3 Bames Big Elk 1and 2 Calf Creek California Park 1,2, and 3 California Park Road 1 and 2 Dinwiddie Dry Elkhead Ridge Dry Fork Elkhead Earle Elkhead Elkhead Road 3 Finger Rock Gillilands Gnat Hill Heidel Hayden Divide Hillberry Hocket Horton Knoll 1 and 2 McKinney Ranch Morgan Reservior Mud Springs Penny Postovit Rock Creek 1 and 2 Sage Creek Seneca Mine 1and 2 Slater Park 1 Smiths Smuin Gulch Gravel Pit Smuin Gulch Oil WeIll Smuin Gulch Oil Well 2 Stokes Gulch Twentymile Warrick Pasture William's Park Wolf Mountain Ranch Wymans Bloomquist County Airport Deep Creek Dry Gulch 2 Dry Creek 2 Elk Mt 1, 2, and 3 Elk River Cemetery George's Gulch Little Hunter Morning Crow North Giant Robinson Sherrod-Sadelin Soash Turner Creek Windemere Woods Five Pines Mesa Dry Creek Yampa Eckman Park 1, 2, and 3 Eckman Park 4, 5, and 6 Edna Mine Elk Creek 1and 2 Fish Creek Foidel Creek GreenAcres Hicks Hinkle Hightail Hoffman Homestead Homestead Ditch Maneotis . Middle Creek Milner Pleasant Valley RCR33b Ricks Rogers Saddle Mt 1and 2 Shivers Twentymile Cliffs 1,2, and 3 Twentymile Cliffs 4 and 5 Yellowjacket Road Yellowjacket 1and 2 N=49 N= 19 N=37 Hayden South Yampa N=2 �185 APPENDIX B (continued) Distribution of Columbian sharp-tailed grouse known lek sites lek by DWM districts, Moffat County, northwest Colorado. Craig North Craig South Meeker North Baker's Peak Buck Mt 1, 2, 3, and 4 Cedar Hill Gulch Cole Gulch 1 and 2 Elkhead Road 1 and 2 Elkhead Road 4 Fly Creek Fortification Rocks Little Buck 1 and 2 Long Gulch McInturf Mesa MCR18 Pelleys Pinnacle Mt 1, 2, and 3 Pondella Ranch Villards Wilderness Ranch Wiseman's 1 Cull Reservior Dresher Reservior Des Dome Morapas Gas Field Nolands Schneiders Taylors Trapper Mine 1 and 2 Burn Morgan Straight Gulch Wilson N=26 N=9 N=4 �186 APPENDIXC Instructions for Conducting Grouse Lek Surveys and Counts These instructions and the accompanying form are an attempt to standardize the information we collect. Please follow the steps outlined below. 1. Conduct surveys from 30 minutes before to two hours after sunrise. 2. Timing of breeding activities can vary 2-3 weeks from one year to the next depending on spring conditions. In most years, the best time to conduct surveys is from 10 April to 15 May. Only one visit per lek IS necessary ifbirds are observed on the lek. If birds are not found, then at least one and preferably two more visits should be made to the lek to confirm that it is inactive. Complete a form each time you visit the lek. 3. These instructions are specific for Columbian sharp-tailed grouse, but the form can be used for other species of grouse. Therefore, be sure to record the species being surveyed in the appropriate space. 4. Provide the complete date (mm/dd/yy). 5. Lek names can be confusing. Sometimes the lek name has no relevance to where the lek is located. Over the years, lek names may change or old leks that shift locations are given new names. Therefore, it is important when recording the lek name to be as complete and specific as possible. If the lek has been referred to by other names, also list those names. Be sure to include numbers that are part of the lek name; i.e., Eckman Park #2, Eckman Park #2 ...etc. 6. Give the official name of the district; i.e., Steamboat South, Hayden, Yampa ...etc. 7. For lek status, check either established (previously documented) or new. If the lek is established, check whether it is active or inactive (this is the most important priority for established leks), and list the USGS quad and county in which the lek is located. If the lek is new, provide the UTM coordinates, indicate whether the coordinates were read from a map or determined with a GPS unit, list the USGS quad and county in which the lek is located, and identify if the lek is on public or private land. 8. If the lek is active, attempt to make three counts at five minute intervals. First attempt to count the total birds present on the lek, then classify them as males, females, and unknown birds. It may be difficult to distinguish males from females especially if the males are not displaying. Topographic and vegetative features also may make it difficult to observe and count all birds on the lek. If it is apparent that you will not be able to obtain an accurate count, then flush the birds, record the total count, and check the space that indicates the birds were flushed 9. Provide whatever comments you feel are necessary to interpret the data that was gathered, such as weather conditions, presence of predators, and additional information about the location of the lek. �187 1998 GROUSE LEK SURVEY FORM SPECIES: DATE: _ LEK NAME: OBSERVERS: DWM District: _ _ LEKSTATUS: __ ESTABLISHED (known lek site) NEW ACTIVE INACTIVE UTM Y (Northing) UTM X (Easting) USGS Quad _ . UTM coordinates determined by __ GPS COUNTY PUBLIC __ TOTAL MALES HIGH COUNTS: MALES FEMALES TOTAL BiRDS Map _ LAND STATUS: __ TIME __ PRIVATE TOTAL FEMALES TOTAL UNKNOWN UNKNOWN FLUSH COUNT * --- "The flush count is only necessary when birds are flushed inadvertently and opts to flush the birds to obtain a count. COMMENTS: TOTAL BIRDS or the observer is unable to observe birds on the lek _ �188 �189 Colorado Division of Wildlife Wildlife Research Report April 1999 JOB PROGRESS REPORT State of: Colorado Project: W-167-R Work Plan: .za, :Job Job Title: Avian Research _1_ Avian Research Publications Period Covered: 01 January through 31 December 1998 Author: Clait E. Braun Personnel: Clait E. Braun, lH. Garnmonley, K M. Giesen, Christian A. Hagen, and Sara J. OylerMcCance, Colorado Division of Wildlife ABSTRACT The following articles were published in 1998: Braun, C.E. 1998. Sage grouse declines in western North America: what are the problems? Proc. Western Assoc. Fish and Wildl. Agencies. 78-139-156. Gammonley, J.H., and L.R. Fredrickson. 1998. Breeding duck populations and productivity on montane wetlands in Arizona Southwest. Nat. 43:219-227. Giesen, KM. 1998. Harvest of Columbian sharp-tailed grouse in Colorado: implications for management. Proc. Prairie Grouse Tech. Council. 22: Abstract. ___ " 1998. Lesser prairie-chicken (Tympanuchus pallidicinctus). In The Birds of North America, No. 364 (A. Poole and F. Gill, eds.) The Birds of North America, Inc., Philadelphia, PA. Hagen, c.A., RK Baydack, and C.E. Braun. 1998. Movements of sage grouse in a fragmented landscape of northwestern Colorado. 1Colorado-Wyoming Acad. Sci. 30(1):35. __ -' __ -' and . 1998. Sage grouse habitat use of a fragmented landscape in northwestern Colorado. Proc. Annu. Conf , The Wildl. Soc. 5:91. �190 Oyler-McCance, S.1., N.W. Kahn, C.E. Braun, and T.W. Quinn. 1998. Genetic support for a proposed new species of sage grouse in Colorado. Proc. Annu. Conf, The Wildl. Soc. 5:122. Prepared by Clait E. Braun Avian Research Program Manager �191 Colorado Division of Wildlife Wildlife Research Report April 1999 JOB PROGRESS REPORT State of: Colorado Project W-167-R Upland Bird Research Work Plan: _l§_ : Job _1_ Job Title: Analysis of Upland Bird Population Trends Period Covered: 0 1 January through 31 December 1998 Author: Clait E. Braun Personnel: Clait E. Braun, Kenneth M. Giesen, Richard W. Hoffman, and Thomas E. Remington, Colorado Division of Wildlife ABSTRACT The following reports were published in 1998: Braun, C.E. 1998. Sage grouse harvest report, North Park, 1998. 1998. Sage grouse harvest report, Eagle, 1998. 1998. Sage grouse harvest report, Gunnison Basin, 1998. 1998. Sage grouse harvest report, Lower Moffat and western Routt County, 1998. 1998. Sage grouse harvest report, Middle Park, 1998. 1998. Sage grouse harvest report, Yampa Area, 1998. Hagen, c.A., and C.E. Braun. 1998. Habitat use and seasonal movements of sage grouse in the Piceance Basin, Rio Blanco County, Colorado. Colorado Div. Wildt. Unpubl. Rep., Fort Collins. Hoffman, R W. 1998. Columbian sharp-tailed grouse harvest data, northwest Colorado, 1976-98. ____ . 1998. Columbian sharp-tailed grouse lek surveys and lek counts for northwest Colorado. Unpubl. Rep., Fort Collins. 9pp. �192 1998. Blue grouse wing analyses in northwest Colorado for 1998. 1998. Blue grouse wing analyses in southwest Colorado for 1998. 1998. Evaluation of wild turkey transplant opportunities in Middle Park and at Radium State Wildlife Area Larison, J.R 1998. Multiple-metal stress in an avian herbivore: the reproductive and physiological effects of chronic exposure to toxic metals. Cornell Univ., Unpubl. Rep. 18pp. Prepared by _~~.....;..._~_. _£._. :...:.~=------:-__ Clait E. Braun Avian Program Manager �193 Colorado Division of Wildlife Wildlife Research Report April 1999 JOB PROGRESS REPORT State of: ____,:C=o=lo=r=a=do=--_ Project: Work Plan: Job Title: ___lL: Job _1_ Evaluate Population Trends of Selected Species ofNeotropical Colorado Period Covered: Author: Avian Research W..!..!--~16~7!.....-"'-'R.__ _ 01 January through 31 December Migratory Birds in 1998 Kenneth M. Giesen Personnel: Kenneth M Giesen, Colorado Division of Wildlife; Michael F. Carter and Tony Leukering, Colorado Bird Observatory ABSTRACT Monitoring of selected species of passerine Neotropical migratory birds was initiated in three habitat types in Colorado in 1998 (Spruce-Fir, Ponderosa Pine, Aspen) facilitated by a Division of Wildlife contract with the Colorado Bird Observatory (CBO). Data collected indicated 26 of29 target species in these habitats will be effectively monitored using distance-measured point counts. A study plan to evaluate survey methodology and investigate productivity of selected grassland birds was completed, peer-reviewed, and finalized. Study areas to evaluate distance-measured point counts and transects on eastern Colorado grasslands were selected from suitable habitats on the Comanche and Pawnee National Grasslands. Species selected for intensive evaluation include Cassin's sparrow (Aimophila cassinii) and westernmeadowlark(S(Umella neglecta) on the Comanche N.G., and McCown's longspur (Calcarius mccowni), Chestnut-collared longspur (Calcarius omatus), and western meadowlark on the Pawnee N. G. �194 �195 EVALUATE POPULATION TRENDS OF SELECTED SPECIES OF NEOTROPICAL MIGRATORY BIRDS IN COLORADO Kenneth M Giesen INTRODUCTION There is widespread concern that populations of many species ofNearctic-Neotropical migratory passerine birds have declined in the last 30 years (Robbins et al. 1986, Robbins et al. 1992). Although US. Fish and Wildlife Breeding Bird Survey (BBS) data monitors populations for many species, other species are not sampled adequately because of low densities, geographic distribution, or access. Many Colorado passerines are monitored annually with BBS routes, but other species are not represented or sampled in numbers too small to ascertain population status and trend, even if one assumes BBS indices reflect actual population trends (Colorado Bird Observatory 1997). There is no statewide program for monitoring population status of most Colorado avian species, and little specific information on nest success or other measures of reproductive performance for those species apparently declining (e.g., grassland birds). A program was developed cooperatively between the Colorado Division of Wildlife, US. Forest Service, Bureau of Land Managment, and the Colorado Bird Observatory to monitor Colorado's breeding birds using a system of distance-measured point counts (Buckland et al. 1993) stratified by habitats found in Colorado. This program (Colorado Birds Monitored 2001) is designed to be able to detect a 2: 3 percent population change with a statistical significance of 0.1 and a power of 0.8 for target species in 13 Colorado habitats. Because many grassland avian species are reported to have undergone the largest population declines in the last 3 decades, additional efforts will be focused on evaluation of inventory methodology for selected grassland bird species and examination of their reproductive performance (i.e., nest success, parasitism rates, fledging success). P.N. OBJECTIVES The primary objectives of this study are to evaluate and implement a statistically reliable population monitoring program for passerine birds in Colorado, and to investigate population monitoring protocols and nesting success for selected species of grassland birds. SEGMENT OBJECTIVES 1. Review literature appropriate to monitoring Neotropical migratory birds and literature on ecology and biology of selected avian species. 2. Develop a study plan to monitor and evaluate population trends for selected species of Neotropical migratory birds. 3. Select protocols and study sites to monitor selected Neotropical migratory bird species. 4. Cooperate with the Colorado Bird Observatory, US.Geological Survey -Biological Resources Division, U.S. Forest Service, Bureau of Land Management, National Park Service, and other entities to coordinate and evaluate statewide monitoring efforts. �196 5. Prepare annual report. METIIODS Literature concerning monitoring of avian populations and detecting population trends was reviewed and discussed with personnel of the Colorado Bird Observatory (CBO) and other agencies involved with bird monitoring in Colorado. A distance-based point count methodology (Buckland et al. 1993) was used to design a program for population monitoring of avian species in Colorado's 13 major habitats. Study sites for comparison of distance-measured point counts and transects were located on the Pawnee and Comanche National Grasslands. and 4 grassland species were selected for intensive surveys of their population densities and nesting success. RESULTS AND DISCUSSION The Colorado Bird Observatory (CBO), in cooperation with many partners, initiated the initial year of distance-based point count monitoring in aspen, ponderosa pine, and spruce-fir habitats. The report of CBO's results is attached (Appendix A). Preliminary information suggests that 26 of29 species using these three habitats will be able to be effectively monitored using distance-based point counts. Further, the number of years to detect a trend in populations will be fewer (avg.= 6.86 years) than when using standard analysis (> 19 years). Study sites for intensive comparison and evaluation of distance-based point counts and transects were located on the Comanche and Pawnee National Grasslands because they provided necessary access and had known populations of grassland birds thought to be declining based on BBS data. The species selected for both inventory and nest monitoring include the western meadowlark, because of its abundance and conspicuous behavior in both areas, Cassin's Sparrow on the Comanche National Grassland, and chestnut-collared longspur and McCown's longspur on the Pawnee National Grassland. Field work is scheduled to begin in 1999. LITERATURE CITED Buckland, S. T., K. R Anderson, K. P. Burnham, and J. L. Laake. 1993. Distance sampling: estimating abundance of biological populations. Chapman & Hall, New York 446 pp. Colorado Bird Observatory. 1997. 1996 reference guide to the monitoring and conservation status of Colorado's breeding birds. Colorado Bird Observatory, Colorado Div.Wildl., Great Outdoor Colorado Trust Fund, and partners, Denver. Robbins, C. S., D. Bystrak, and P. H. Geissler. 1986. The breeding bird survey: its first fifteen years, 1965-1979. U.S. Dep. Inter., Fish and Wildlife Servo Resour. Pub. 157. 196 pp. Robbins, C. S.; 1. R Sauer, and B. G. Peterjohn. 1992. Population trends and management opportunities for neotropical migrants. Pages 17-23 in D. M. Finch and P. W. Stangel (eds). Status and management ofNeotropical migratory birds. U.S. Dep.Agric., Forest Serv., Gen. Tech. Rep. RM-229. Prepared by: ·~-If~ Kenneth M. Giesen �197 COLORADO BIRDS MONITORED BY 2001: RESULTS OF POINT TRANSECTS IN THREE COLORADO HABITATS wrm AN APPENDIX OF RESULTS OF SPECIAL SPECIES MONITORING Michael Carter Tony Leukering ABSTRACT In 1998, Colorado Bird Observatory, with many partners, initiated the pilot year of count-based monitoring as outlined in the plan Colorado Birds Monitored by 2001 (CBM 2001)(Carter and Leukering 1998 and updates). The CBM 2001 plan set the goal of being able to detect a ~3.0% population change with statistical significance of 0.1 and power of 0.8 for species dependent (target species) on 13 Colorado habitats. For a test of methodology, we randomly established 30 point transects within 30 randomly-selected stands in three habitats (Aspen, Ponderosa Pine, and Spruce-Fir) within the state. 'We used roads as access points to transects but each transect headed in a random direction from its road access point. Each transect consisted of 15 five-minute counts with the sum of detections for each species for each transect treated as a replicate. We analyzed these data using distance-sampling theory via the program DISTANCE (Laake et al. 1994) and a "standard analysis" using unlimited detection radii. Results using both methods were then used to model monitoring efficiency using the program MONITOR (pWRC 1998). Data analyzed via the program DISTANCE exhibited significantly lower variance (vs. the standard analysis) thus resulting in fewer number of years to meet our monitoring goals (5<=6.86yrs., range=6-9 yrs.). Assuming the density-based variance we obtained will be approximately stable through time, we will be able to effectively monitor 26 of the 29 target species in the three habitats in time periods that are reasonable, i.e. we will start seeing results for most species before a decade has passed. The unmonitored species (purple Martin in Aspen and Sharp-shinned Hawk and Boreal Owl in Spruce-Fir) may never be monitorable using countbased methods during the breeding season and have been addressed under the special-species techniques within CBM 2001. The species that are monitorable under this plan using count-based methods are (habitats are A=Aspen, P=Ponderosa Pine, and S=Spruce-Fir): Broad-tailed Hummingbird (A), Red-naped Sapsucker (A), Hail)' Woodpecker (P), Three-toed Woodpecker (S), Olive-sided Flycatcher (S), Western Wood-Pewee (A), Hammond's Flycatcher (S), Warbling Vireo (A), Gray Jay (S), Clark's Nutcracker (S), Tree Swallow (A), Violet-green Swallow (A), Mountain Chickadee (S), Red-breasted Nuthatch (S), Pygmy Nuthatch (P), Golden-crowned Kinglet (S), Rubycrowned Kinglet (S), Mountain Bluebird (A), Western Bluebird (P), Hermit Thrush (S), Grace's Warbler (P), Chipping Sparrow (P), Pine Grosbeak (S), Cassin's Finch (S), Red CrossbiU(S), and Pine Siskin (S). We expect that the annual costs of monitoring birds in these habitats through this plan will not exceed $10,000 per habitat per year (1998 dollars) for the first decade. INTRODUCTION Conservation and management of Colorado's birds depend on adequate monitoring information. The Breeding Bird Survey (BBS) has accumulated a 33-year data set, but the routes in Colorado effectively monitor <20% of the state's breeding bird species (BBS web site 1998). In addition, data that corroborates the BBS data set are entirely lacking across the range of that project. Monitoring information is required by legislative and land/wildlife management agency mandates, as well as a host of long-range plans, Forest plans, ecoregional plans, preserve management plans, etc. From 'a global �198 biodiversity perspective, Colorado hosts some species at high abundances, thus Colorado has high responsibility for those species (partners in Flight 1998). In cooperation with the agencies/organizations charged with protecting and managing Colorado's birds, Colorado Bird Observatory (CBO) has developed and proposed a program of bird monitoring for the state, Colorado Birds Monitored by 2001 (CBM 2001)(Carter and Leukering 1998), in which every interested agency/organization contributes and benefits. The first phase of this plan calls for establishing a statewide, statistically-robust program of randomly-selected point-count transects in each of 13 habitats. With funding from the Great Outdoors Colorado Trust Fund through the Colorado Division of Wildlife and the U.S.D.A. Forest Service, CBO established the transects in three habitats in 1998. In developing CBM 2001, we defined suites of species that are restricted to or that are found at their highest abundances in each of the 16 defined habitats; the species in each group were termed "target species" and were considered indicators of that habitat. We here slightly redefine "target species" as those species that achieve their highest abundance in the state in that habitat and which are common enough for us to be able to monitor their populations, detecting trends of ~3.0o/olyr (positive or negative) with statistical significance of 0.1 and power of 80%. The plan also developed a variation on sampling methodology by combining aspects of point-count and transect methodologies for the point transect protocol used herein. METHODS We established transects of 15 point counts in each of30 randomly-selected stands in each of three habitats: Aspen, Ponderosa Pine, and Spruce-Fir. Using USDA Forest Service GIS data, we numbered all publicly-owned stands of the habitats in Colorado (n=478 in Aspen (2,248,526 acres), 521 in Ponderosa Pine (1,672,074 acres), and 490 (4,185,844 acres) in Spruce-Fir) and then randomly selected 53 to 59 from each habitat. We then randomly selected 30 of those in which we established point transects. In a few instances, selected stands were not the indicated habitat or access across private land was. denied, so we discarded them and randomly selected a replacement from the original set of randomly-selected stands. Each transect was conducted by one observer using protocol established by Leukering (1998). The observer located the selected stand on the ground and ran the transect along a randomly-selected bearing. It was usually impossible to run the entire transect along the random bearing, as stand boundaries, property boundaries, and physical obstructions forced turns in the transect direction. When this happened, the observer randomly turned right or left perpendicular to the random bearing, subsequently alternating perpendicular directions if additional turns were necessary. In some stands, the narrowness of the stands predicated the location and bearing of the transects. . Transects consisted of 15 5-minute point counts spaced at 250-m intervals along a line. We considered the intervals between points as legs of a true transect. At the individual points, we recorded the distance to each bird detected. Along the transect legs, we recorded only individuals of a short list of the habitat's target species whose population densities are relatively low (thus, poorly-recorded on point counts) and estimated distance to each. However, the protocol allowed recording individual birds on both a point and a transect leg, thus eliminating the possibility of analyzing pooled point and transect data In future years we will record individuals of low-density target species on either a point or a transect leg, not both, with points having priority over transect legs. �199 Observers recorded weather data (sky condition--cloud cover and precipitation, wind--Beaufort scale, and temperature) and the time at the start and end of each transect. At each point, the observer recorded whether the point was within 100m of a road. Also at each point, he/she recorded the specific habitat and seral stage (1-5 scale; Buttery and Gillam 1983) of each of the two predominant habitats around the point (often there was only one habitat present). Upon arriving at a point, the observer recorded habitat data, then conducted the point count. Though we anticipated using the data from the transect legs, for virtually all target species, the point data was robust enough to stand alone. We analyzed the point data grouped by transect, with transects as replicates, and developed species means and various descriptive statistics. DISTANCE Program We used program DISTANCE (Laake et al. 1994) to analyze distance-estimate data; in this report, all references to density estimates are values provided by DISTANCE from our data The notation, concepts, and analysis methods of the program were developed in Buckland et al. (1993). The program can analyze several forms of distance sampling data, fitting a detection curve to the data set to be analyzed. The program avoids some serious biases inherent in traditional analysis of point-count data (e.g. detectability among habitats or years), but comes with three assumptions: all birds at distance 0 are detected; distances of birds close to the point are measured accurately, and birds do not move in response to the observer's presence. We considered well-sampled those species for which DISTANCE provided a model that met three criteria: coefficient of variation (CV) of <50%, <3 parameters included in the detection curve function, and total variance reasonably balanced between the variance caused by sample size and that caused by the detection probability (ratio from :::2:1 to :::1:2; D. Anderson pers. corrim.). For those species for which unlimited distance did not meet all three criteria, we truncated the data sets for individual species at various distances (using cut points developed by DISTANCE) and reran DISTANCE. We used two programs, TRENDS (Gerrodette 1993) and MONITOR (patuxent Wildlife Research Center 1998), to model monitoring efficacy. The output provided is the number of years required to detect a given trend with certain assumptions (trend of ~3.0% (both positive and negative) with statistical significance of 0.1 and power of 80%, assuming 30 transects run annually). To develop the indices, these programs require input ofCVs, for which we used the CVs developed both by "standard analysis" of unlimited-distance radii and density-estimate CVs provided by DISTANCE. The two programs (TRENDS and MONITOR) differed by a factor of 2 or more in these indices. Since we are unsure of the reason( s) for the difference, we report the results from both. RESULTS We recorded a total of 11,408 birds of 104 species on 89 transects (one Spruce-Fir transect's data were lost) (Table 1; all scientific names are presented in Appendix A). Of these, 10,431 non-flyover individuals of 95 species were recorded on the 1335 point counts, with only 21 points having no birds (8 in Aspen, 1 in Ponderosa Pine, and 12 in Spruce-Fir). Species totals on the 89 transects ranged from 1 for many species to 1059 Warbling Vireos (almost 10% of all birds recorded; Table 1). Using unlimited-radius detections, we obtained CVs of under 150% for 12 species and under 100% for seven species, with the lowest CV being 46% for Y ellow-rumped Warbler. �200 The distance-estimate data provided CV s of under 100%, in at least one habitat, for all species with sample size of >5 in at least one habitat (except for Common Nighthawk; total n=59 species; Table 2). Of those 58 species with CVs <100%,53 had CVs under 50% in at least one habitat. For 21 species with CVs of under 50% in at least one habitat, we obtained robust results (well-balanced variance sources and<3 parameters in the detection-curve model that incorporated the complete data sets) (Table 2). By truncating outliers at various distances for individual species, we attempted to optimize CV s, decrease the number of parameters included in the models, and to balance the two sources of variance: sample size and probability of detection. We truncated data for 36 species in the three habitats. These were primarily target species, but included some species that were recorded in their highest numbers in habitats for which they were not targets. For a few species, low sample sizes precluded truncation (e.g. Red Crossbill). We truncated the data at various distances and report those results by habitat (Tables 3-5). Finally, using 1998 data as a baseline, we provide estimates of the number of years required to monitor various target species (Table 6). DISCUSSION The high number of species with low CV s (Table 2) indicates that point transects will be adequate for detecting population trends in the studied habitats and, by extension, all habitats in Colorado. This is particularly so with the use of detection-curve models developed by DISTANCE (Laake et al. 1994). The use of DISTANCE is particularly important, as the results provided by density estimates are much more powerful than those obtained by considering unlimited-radius detections alone (see Table 7 for the comparison in Spruce-Fir). In fact, the power of DISTANCE enabled us to tighten up the thresholds for effective monitoring. We also increased the size of the population change (from 2.5% to 3.0%) that we wished to detect to align with currently-accepted thresholds (e.g. Patuxent Wildlife Research Center 1998). . There were only three target species in the three habitats that we either did not record or did not record in numbers high enough to analyze: Purple Martin in Aspen (one individual counted) and Sharpshinned Hawk and Boreal Owl in Spruce-Fir (none of either counted). We achieved sample size with all Ponderosa Pine target species, including the limited-range Grace' s Warbler. We will need to establish species-specific inventory methods for Purple Martin and Sharp-shinned Hawk and nocturnal transects may result in useful data for Boreal Owl. A few target species with low sample sizes had surprisingly robust CV s and/or fairly balanced sources of variance when distance estimates were analyzed (Table 2). All other species for which we obtained low sample sizes are species that are either much more common in other habitats or widespread habitat generalists for which we will need to analyze data from all habitats to monitor. Of the 59 species recorded in sample sizes large enough to analyze (>5), we discuss below those species that were target species in one of the three habitats in which we worked, though we do discuss some non-target species. Trend detection using unlimited-radius detections will, for most species, not be possible in the term of 30 years that was set in CBM 2001. Additionally, program TRENDS does not permit input ofCVs >99%. Therefore, unless otherwise noted in species accounts below, trend-detection timetables are generated from density-based analysis from DISTANCE. Please refer to Table 6 for trend-detection estimates using unlimited distances. When perusing the results presented in the various tables and the species accounts below, it is important to keep in mind that most individual transects traversed multiple habitats due to the interdigitation of high-elevation habitats in mountainous Colorado. While we attempted to run transects in �201 target habitats, GIS systems are often unable to distinguish various coniferous habitats from each other (Lodgepole Pine vs. Spruce-Fir or Mixed Conifer vs. both Ponderosa Pine and Spruce-Fir). In addition, Aspen is regularly mixed in with all coniferous habitats in the state, particularly from the Ponderosa Pine elevations upward. Therefore, many individual points of transects fell in habitats that were not targeted by those transects, thus increasing variance in the data for all habitats. It is also important to know that we are unsure which is better when faced with a decision between an unbalanced model with <3 parameters and a balanced model with three parameters; we chose the former. In the accounts below, the reference Andrews and Righter (1992) is abbreviated to A&R In addition, "Aspen" (capitalized) refers to the habitat and "aspen" (not capitalized) refers to individual trees. Band-tailed Pigeon-We recorded only nine individuals of this inhabitant of Ponderosa Pine and Gambel Oak, despite being fairly common to common in southwestern and south-central forests (A&R). It is a relatively quiet species and despite its size, can be difficult to detect (Leukering pers. obs.). All individuals recorded were found on southwestern Colorado transects as the randomization process selected few Ponderosa Pine transects in south-central Colorado. Common Nighthawk-We were surprised to record 11 of this species which we designated as requiring species-focused effort (CBM 2001). The most interesting aspect of this species' data was the result from DISTANCE. Common Nighthawk is a species that, though fairly common (A&R); is usually only detected during morning hours by walking very close to, thus flushing, individuals roosting on the ground. This fact was detected by DISTANCE as the variance due to detectability accounted for 99% of the total variance with only 1% due to encounter rate. Broad-tailed Hummingbird-We detected this Aspen target species more often on Ponderosa Pine transects than on Aspen transects (95 vs. 68; Table 2). Variance was balanced In both habitats, but in Aspen the detection curve required four parameters. Truncating the Aspen data set at 52 m provided a better model. Williamson's Sapsucker-Despite the good sample size for this species in Ponderosa Pine (n=46), the sources of variance were highly skewed and truncation of data failed to reconcile this problem. We are unsure of the reasons for our inability to find a good model for this species and await further years' data for definition of the problem. Our experience with this species in north-central Colorado suggests that, though it prefers Ponderosa Pine forest, it usually nests in Aspens as that tree species provides more cavity-excavation sites than does any other tree species (Leukering pers. obs.) Thus, though we were surprised that data from Aspen provided a better model for trend detection in this species, we were not surprised at the number encountered there. Red-naped Sapsucker-The robust data for this species in Aspen required no truncation to produce a solid model. In Ponderosa Pine, despite a reasonable sample size, DISTANCE good detection curve. was unable to fit a Hairy Woodpecker-We designated this species a target in Mixed Conifer (CBM 2001), but we obtained reasonable sample sizes for the species in all three habitats this year, with highest numbers in Aspen (n=33). Though Hairy Woodpecker is more of a forest generalist than is Williamson's Sapsucker, like the latter species, it often nests in Aspens, though, in the mountains, preferring coniferous forests (Leukering pers. obs.). This partly explains the seeming preference for Aspen in our data In Aspen, truncating the data set at 118 m produced the best of three well-balanced models. �202 Three-toed Woodpecker-We only recorded six individuals on point counts of this very quiet species whose population is naturally of very low density in unburned forest. Interestingly, despite the tiny sample size, the CV of the density estimate was very low (16.7%), though, all of the variance was due to the low sample size (Table 3). In addition, we counted seven Three-toeds on transect legs (though, with some duplication with the point counts). Thus, we believe that we will develop useful data on this species in future years with a slight change in survey protocol. Also of importance is that we recorded ten unidentified woodpeckers on point counts, all of which were drumming birds thought by the observers to be either Hairy or Three-toed woodpeckers. Since drums of the two species are very similar, unseen drumming woodpeckers are best left unidentified, though Stark et al. (1998) suggest that syntopic woodpecker species have distinctive drums. The CV from DISTANCE would result in trend detection in 6 to 15 years (Table 4). Northern Flicker-We recorded this species, a forest generalist which we designated as a target species in Lowland Riparian (CBM 2001), in good numbers in both Aspen and Ponderosa Pine (n=91 and 45, respectively). As in other woodpecker species, the high number of individuals in Aspen is partly due to Aspen being such a good provider of cavity-excavation sites. Detection-curve models were unbalanced in both Aspen and Ponderosa Pine, but truncation at 164 m in Ponderosa Pine produced a good model. Olive-sided Flycatcher-This is a relatively low-density species with specific structural habitat requirements that are consistent across habitats (Hutto 1995, Leukering pers. obs.). Olive-sideds usually occur in sites with particular combinations of snags, forest, open areas, and water. These situations occur at most montane elevations in Colorado, so the species is distributed across elevations. In 1998, we recorded the species in all three habitats, with the largest numbers occurring in Aspen, despite its designation as a target in Spruce-Fir (in which we recorded the fewest individuals). Though the complete Aspen data set for this species provided a good model, we will analyze the data for this species across all habitats in future years. Western Wood-Pewee-We found this Aspen target species in highest numbers in Aspen (n=164), but also had significant numbers in Ponderosa Pine (n= 102). However, the presence of Aspen interdigitated with most coniferous habitats in Colorado probably accounts for a large proportion of pewees recorded on Ponderosa Pine transects. The model produced by the complete Aspen data set was balanced but required three parameters. By truncating the data at 105 m, we were able to obtain a model with only one parameter, but one which was unbalanced. Hammond's Flycatcher-A&R state that Hammond's Flycatcher is primarily an inhabitant of "mature, closed-canopy spruce-fir forests ...", but also say that in "some areas it may occur in greater numbers in ponderosa pine forests than in other habitats (J. Sedgwick, pers. comm.)" and that many other forest types are selected by this species. We found Hammond's Flycatchers in decreasing numbers across habitats from Aspen to Ponderosa Pine to Spruce-Fir (n=25, 20, and 14, respectively), despite being designated a target species of the latter. This agrees with our experience with this species in northcentral Colorado (Leukering pers. obs.). We did not obtain a good model for Hammond's in Aspen (Table 3), but did so in both Ponderosa Pine and Spruce-Fir. One confounding factor is the difficulty that many observers have in distinguishing this species from the very similar Dusky Flycatcher, although we believe that the distribution in our results of the two species within the three habitats suggest that there were few, if any, mis-identifications in our data set. �203 Dusky Flycatcher-We recorded a large number of Duskies in Ponderosa Pine (n=120), though we designated this a target species in Mountain Shrub land (CBM 2001). This is readily explained in that this species is a shrub inhabitant and that many Ponderosa Pine forests in Colorado have a robust oak understory in which this species achieves amazingly high densities (Hutchings et al. 1998, Leukering pers. obs.). In Ponderosa Pine, the complete data set and all truncations produced unbalanced models (Table 4), though truncation at 84 m produced a model with only one parameter that was nearly balanced. It will be interesting to see Dusky Flycatcher data from the Mountain Shrub land transects when those are initiated in 1999. Plumbeous Vireo-Though we designated this species a target in Pifion-Juniper (CBM 2001), it is somewhat of a low- to mid-elevation forest generalist, breeding also in Lowland Riparian in western Colorado (A&R, Leukering pers. obs.) and in Aspen. In Pinon-Juniper, it is most common in denser high-elevation forest (A&R, S. Hutchings pers. comm.) and the distribution of transects in that habitat will have an affect on the detection rate. Should lower-elevation transects predominate among the Pifion-Juniper transects, then the Ponderosa Pine transects may produce higher detection rates for this species. With that in mind, the complete data set in Ponderosa Pine in 1998 (n=43) produced a balanced model, but one with three parameters. By truncating the data set at 86 m, we obtained a good model (Table 4). Warbling Vireo-As we expected, this was the most abundant bird on transects in anyone habitat and in all habitats combined (Table 2), being recorded at rates of 1.84 birds per point in Aspen and 0.81 birds per point overall. Despite this abundance, we did not obtain a good model in Aspen; models for all truncation distances were unbalanced (Table 3). However, the Spruce-Fir data set produced a good model. Gray Jay- This species' data was the most difficult to analyze as it is a species with low-density populations (Leukering pers. obs.) for which we obtained data indicating high density (density of 26.60/ha in Aspen). However, Gray Jays are exceedingly curious and tame and regularly inspect humans and other disturbances in their vicinity. Thus, we often recorded the species at very close range as individuals or groups came toward the observers before being detected, which violates one of the assumptions of DISTANCE. Because of the species' often quiet nature, we believe that our data are skewed to short distances. We will have to solve the observer-attraction problem before we are comfortable with any results. Steller's Jay- This is a coniferous-forest generalist that we designated a target in Mixed Conifer (CBM 2001). We obtained good models in all three habitats with the complete data sets. This bodes extremely well for monitoring this species and anticipate initiating transects in Mixed Conifer in 1999. Clark's Nutcracker-Nutcrackers are difficult to obtain robust data on due to their low-density populations and predilection for habitats at or near timberline. Thus, we were surprised with the data that we obtained in Ponderosa Pine (n=60) and in Spruce-Fir (n=61); it is a target species in the latter. However, the Spruce-Fir model required three parameters, thus we opted to reanalyze the data set. By truncating the data set at 292 m, we obtained a balanced model with only one parameter. The Ponderosa Pine model was suitable using the complete data set. Tree Swallow-This is a species that nests primarily in Aspen forest, but one which readily adapts to other habitats given the presence of suitable cavity sites (A&R). We were surprised at the low sample �204 size we obtained in Aspen (n=30), particularly since the complete data set model required five parameters. By truncating the Aspen data set at 98 In, we obtained a robust model. Violet-green Swallow-CBM 2001 designated this a target of Aspen, though it is also a very common nester in CliffIRock. This affinity for the latter habitat means that it can be found breeding in many other habitat types, as CliffIRock is present within many forest habitats. In addition, the presence of Aspen intermingled with Ponderosa Pine helps to explain the small difference in sample size that Aspen (n=91) obtained over Ponderosa Pine (n=81). The complete data sets for both habitats produced balanced models, but that for Ponderosa Pine required three parameters while that for Aspen required four. Truncating the Aspen data at 171 m produced an unbalanced model with only one parameter. A truncation at 73 m produced a model with two parameters that wasjust shy of balancing the variance sources (32.8% vs. the minimum of33.3%). Mountain Chickadee-This species is a widespread inhabitant of coniferous forests throughout the state and densities undoubtedly vary by habitat. CBM 2001 designated this a Spruce-Fir bird and we recorded the highest number of individuals in that habitat (n=250). We recorded large sample sizes in all habitats, but only in Spruce-Fir did we obtain a robust model from the complete data set. Red-breasted Nuthatch-This species is a conifer generalist, however, one with a predilection in Colorado for Spruce-Fir and Lodgepole Pine (A&R). We counted more of this species in Aspen than we did in Spruce-Fir (19 vs. 18), but, as with many woodpeckers (above), this species readily nests in aspens (Leukering pers. obs.) among coniferous forest. In both Aspen and Spruce-Fir, we obtained good models with the complete data sets. However, we recorded more Red-breasted Nuthatches in Ponderosa Pine (n=44) than we did in the other two habitats combined. We did not obtain a balanced model from the Ponderosa Pine data set, with or without truncation, all being skewed to encounter rate (Table 4). This species is dependent upon varying conifer seed crops (A&R) and, thus, like other nomadic species (e.g. many cardueline finches, which, see below), it varies in abundance temporally, spatially, and in regard to what conifer species it exploits. This will make it difficult to monitor this species' population in a short time period and/or in only one habitat. Further years' data may determine whether we continue to consider this species a target of Spruce-Fir or not. White-breasted Nuthatch-CBM 2001 designated this a target of Mixed Conifer. We obtained a good sample size in Ponderosa Pine and recorded it in all three habitats that we studied in 1998. The complete Ponderosa Pine data set produced a balanced model, but one with three parameters. Truncating this data set at 107 m produced a robust model. It will be interesting to see the results of the Mixed Conifer transects when we initiate them in 1999, because in 1998, all three nuthatch species were recorded in highest numbers in Ponderosa Pine, despite CBM 2001 designating the three species as target species in three different habitats (see Pygmy Nuthatch, below). Pygmy Nuthatch-This species is strongly associated with Ponderosa Pine in Colorado, so much so, that its range in the state closely parallels the range of Ponderosa in the state (A&R). In fact, we did not record Pygmy Nuthatches in any other habitat in 1998. The complete data set (n=87) produced a balanced model, but one with three parameters. By truncating distances at 82 m, we obtained a barelybalanced model. Brown Creeper-Though in CBM 2001 this is a Mixed Conifer target species, our experience with it in Spruce-Fir, particularly its selection of the oldest seral stages in that habitat (Carter and Gillihan in �205 press), suggested that we analyze Brown Creeper data in that habitat. The Spruce-Fir data set (n=29) provided a slightly-unbalanced model; truncating the data at 47 m produced a robust model (Table 5). Golden-crowned Kinglet-This species, though difficult to detect beyond 35 meters, was recorded at fairly long distances a few times, thus requiring a model with three parameters to fit the detection curve. Truncating the data at 81 m produced a much more parsimonious model. Ruby-crowned Kinglet-This species is one of the ubiquitous species of Spruce-Fir forest in Colorado and our sample size reflected that (the 314 detections was the second-highest total of detections for any Spruce-Fir target species). We tried two truncation distances one of which came close to balancing variance sources, but added a second parameter to the detection-curve model. The other truncation (248 m) made a slight improvement in the model over the non-truncated data set and kept the number of parameters at one, so we selected that one as the best model (Table 5). Western Bluebird-We only recorded this species in Ponderosa Pine, the habitat in which it is a target. This species seems to occur in denser forests than does Mountain Bluebird (below). Whether this is an actual preference or the result of competition with the habitat-generalist Mountain has yet-to be worked out. The complete data set for this species provided a robust model. Mountain Bluebird-Though CBM 2001 designated this an Aspen target species, we counted more Mountain Bluebirds in Ponderosa Pine. This species is actually a habitat generalist, breeding in many habitats that provide cavities and fairly open conditions (A&R, Letikering pers. obs.). The complete Aspen data set (n=30) provided a robust model, but the Ponderosa Pine data set (n=43) required truncation to produce such a model. Three different truncation distances only slightly improved the balance, but the truncation at 177 mjust managed to get the model balanced (Table 4). Townsend 's Solitaire-This, a Cliff/Rock species according to CBM 2001, is a conifer generalist with a limiting structure requirement of rocky slopes or embankments for nesting (A&R). As we will not be conducting Cliff/Rock transects, we will need to track this species' trends in all habitats in which it is relatively numerous. However, the bulk of our detections of Solitaires (103 of 137) was in Ponderosa Pine in which the complete data set provided a balanced model, but one requiring three parameters. We truncated the data at two long distances, one of which produced a one-parameter model that was not balanced, this despite a large sample size. Hermit Thrush-Though oflow density, this species' loud voice permitted us to record it for the largest number of detections for any Spruce-Fir target species (Table 2). We detected 342 in Spruce-Fir, the habitat in which it is a target. Density estimation with the complete data set produced a very low CV and very balanced sources of variance, but required a whopping 6 parameters in the model. We tried various truncation distances and selected a distance of 189 m as the best model as it produced a low CV with only one parameter. However, this was at the expense of balanced variance sources (Table 3). The truncation at 147 m almost produced a balanced model. American Robin-We recorded this habitat generalist in large sample sizes in all habitats, but with the 330 in Ponderosa Pine just besting the Aspen total of326 (Table 2). Of the complete data sets, Aspen provided the only robust model, though the Ponderosa Pine model only missed being robust by requiring three parameters. Truncating the Ponderosa Pine data did not produce a better model. American Robin will probably be our most general of habitat generalists and we will need to analyze all habitats in which it breeds to develop a statewide trend for the species. �206 Yellow-rumped Warbler-This, the second-most numerous species on the transects (Table 2), is a coniferous forest generalist, though more common in forests with a Fir component (CBM 2001 allocated it to Mixed Conifer), that we recorded in highest numbers in Spruce-Fir. The complete . Spruce-Fir data set produced a robust model, with none of four different truncation distances producing a better model. Grace 's Warbler-This species is limited in Colorado to Ponderosa Pine forests in the southwestern comer of the state (A&R). Since it is so limited in distribution in the state, we were pleasantly surprised at the reasonably-good sample size, which, in its complete form (n=20), provided a robust model. Western Tanager-CBM 2001 records this species as a target in Mixed Conifer. However, we counted a large number of tanagers (213) in Ponderosa Pine and await initiation of Mixed Conifer transects to compare sample sizes. Unfortunately, the complete Ponderosa data set produced an unbalanced model with four parameters. Truncating the data at 132 m resulted in a good model. Green-tailed Towhee-This is a Sage Shrub land target species (CBM 2001) that we recorded in large numbers in Ponderosa Pine (n= 197) due to the strong oak: or sage understory of many stands. The complete data set in Ponderosa produced an unbalanced model that we could not correct by truncation. Interestingly, the complete Aspen data set (with a much lower sample size) did produce a robust model. Chipping Sparrow-We recorded this Ponderosa Pine target species (CBM 2001) in all three habitats in at least reasonable numbers, though among complete data sets, only Spruce-Fir produced a robust model. By truncating distances in Ponderosa Pine, we obtained two robust models, with the one we selected producing a lower CV (Table 4). Lincoln's Sparrow-Though we designated this a target species of High-elevation Riparian (CBM 2001), Lincoln's Sparrow also occupies moist, grassy slopes in Aspen forest, thus accounting for the large number detected on the Aspen transects (n= 119). The complete data set produced a poor model. Truncation did not produce a balanced model, but did provide a model with only one parameter (Table 3). White-crowned Sparrow-This is a target species of High-elevation Riparian, though it achieves highest densities in krummholz Spruce-Fir (T. Leukering, M. Carter pers. obs.). However, we felt that we would not get enough transects in that aspect of Spruce-Fir so selected another habitat in which it is common. Since High-elevation Riparian runs through all other high-elevation habitats, we recorded White-crowns in good numbers in both Aspen and Spruce-Fir. However, due to these birds being in riparian areas in those two habitats, variance sources were very unbalanced in both. We did not attempt truncation in either habitat and await initiation of High-elevation Riparian transects to fully analyze data for this species. Dark-eyedJunco-We have allocated this species to Mixed Conifer (CBM 2001), though it is a forest generalist requiring a particular structure and which we recorded most often in Aspen (n=330). Numerous truncation attempts in Aspen (Table 3) and in Spruce-Fir (Leukering and Carter 1998) did not produce a model better than that produced by the complete data set in either habitat. �207 Cardueline finches: special cases in bird-monitoring-The Cardueline finches (subfamily Carduelinae) are, for the most part, nomadic or semi-nomadic due to their dependence (during at least parts of every year) on highly variable food sources, often conifer seeds. The Red-breasted Nuthatch (see above) has a similar lifestyle for the same reason. This nomadism makes monitoring populations very difficult, except in very long time periods. We anticipate that densities and CV s in this group will vary widely across the history of this project. Whether we will be successful in monitoring these species awaits the compilation of many years of data Pine Grosbeak-Despite this bird's name, it is most common in Spruce-Fir forest, virtually throughout its holarctic breeding range. It is a tame species that we often recorded at very close range on the transects. We tried four truncation distances in Spruce-Fir to provide a better model than that produced by the complete data set. Two truncation distances (89 m and 118 m) produced robust models, but the model from the shorter truncation required only one parameter (vs. 2 for 118 m; Table 5). Cassin's Finch-Despite the low sample size for this species in Spruce-Fir (the habitat in which it is a target), the CV of 40.3% is quite good, particularly when combined with a good model. . Red Crossbill-This is a very enigmatic species and is probably the most highly-nomadic species in Colorado. Tc'compound the problem, recent research on Red Crossbill suggests that the various taxonomic units of that species, each of which is tied to a particular conifer species, may all be good species (Groth 1993). Thus, our anticipated highly-variable results across years will be made even more variable when considering individual taxa In Colorado, two forms are regular breeders, a Ponderosa Pine specialist and a Lodgepole Pine specialist (C. Benkman, pers. comm., Leukering pers. obs.). At least two other forms have occurred in the state and could breed. Since vocalizations are the primary field-detectable differences between forms (Groth 1993) and since most birders and ornithologists have not yet leamed how to separate these various forms (T. Leukering, pers. obs.), data on occurrence of the various forms in the state are generally unavailable (A&R predates general knowledge of this information). Analysis of results for this species should more correctly be performed on flocks, not on individuals, as this species was almost always recorded in flocks, thus clumping distance estimates. Because we did not record flock composition in the field, we are unable to perform a by-flock analysis and we advise that the results for this species in Table 3 be viewed accordingly. One last confounding factor in monitoring Red Crossbills is that this species can breed at any time of year (often in mid-winter), thus our transects performed in June will probably not record this species during their breeding season in most years. Complete data sets in Ponderosa Pine and Spruce-Fir produced poor models and sample sizes were too small to attempt any truncation. Pine Siskin-This is the most common of the carduelines in Colorado and the one most likely to be found in a given place from year to year. However, previous banding work performed by CBO in the Arapaho National Forest (Leukering 1996, CBO unpubl. data) has shown that though this species may be common at one site year after year, very few individuals are site-faithful. The 1998 transect data for this species are difficult to analyze due to the high proportion of individuals recorded as flyovers and different interpretations by various observers as to the "countability" of these flyovers (some observers identified all as flyovers, some identified many of them as "on point"). We believe that this is the primary reason behind the poor performance of the large sample (n=212) at compromising between number of parameters and balanced sources of variance in Spruce-Fir. Despite many truncation attempts, we could not produce a balanced model (Table 5). Perhaps; more rigorous training on identifying whether individuals are on point or not will help with this problem. �208 ACKNOWLEDGMENTS Colorado Birds Monitored by 2001 is a large-scale, cooperative effort funded in large part through the Colorado Division of Wildlife by the Great Outdoors Colorado Trust Fund (contracts 2183-98 and 2196-99). In addition, U.S.D.A. Forest Service provided funding to CBO for the project (CCS-09-0098-043) in addition to providing salary and expenses for Chris Schultz (San Juan-Rio Grande National Forests) to work on many aspects of the project, including coordinating the southwestern transects and running most of them. We would like to thank Chris and Bob Tribble, also ofU.S.D.A. Forest Service, for conducting the original stand selection for this project from Forest Service GIS data. We wish to express our gratitude to the field staff that conducted most of the transects: Sue Bonfield, Glenn Giroir, Dave Hallock, Dave Hedeen, Rich Levad, and Chris Wood. We would also like to thank Dr. David Anderson for advice on distance sampling and analysis. LITERATURE CITED Andrews, R. and R Righter. 1992. Colorado Birds: A Reference to Their Distribution and Habitat. Denver Mus. of Natural History, Denver. Breeding Bird Survey. 1998. Breeding Bird Survey home page, v. 1998.10. http://www.mbr-pwrc.usgs.govlbbslbbs.html. Buckland, S.T.,.D.R. Anderson, K.P. Burnham, and J.L. Laake. 1993. Distance sampling: estimating abundance of biological populations. Chapman & Hall, London. Buttery, R L. and B. C. Gillam. 1983. Ecosystem descriptions. Pgs. 43-71 in RL. Hoover and D. L. Wills, eds., Managing Forested Lands for Wildlife. Colo. Div. of Wildlife, in cooperation with USDA Forest Service, Rocky Mtn. Region, Denver. Carter, M. C., and S. W. Gillihan. In press. Influence of stand shape, size, and sera! stage on forest bird communities in Colorado. In R. L. Knight, F. W. Smith, S. W. Buskirk, W. H. Romme, and W. L. Baker, eds. Forest Fragmentation in the Southern Rocky Mountains. University Press of Colorado. Carter, M. and T. Leukering. 1998. Colorado Birds Monitored by 2001: The plan for count-based monitoring. Unpubl. ms. Colorado Bird Observatory, Brighton, CO. Gerrodette, T. 1993. Program TRENDS. Southwest Fisheries Science Center, La Jolla, CA. Groth, J. G. 1993. Evolutionary differentiation in morphology, vocalizations, and allozymes among nomadic sibling species in the North American red crossbill (Loxia curvirostra) complex. Univ. California Publ. Zoology, vol. 127. Hutchings, S., T. Leukering, and M. Carter. 1998. Assessment of breeding bird monitoring station at Bodo State Wildlife Area, La Plata County, Colorado, 1995-1998. Unpubl. report. CBO, Brighton, CO. Hutto, RL. 1995. Composition of bird Communities following stand-replacement fires in northern Rocky Mountain (U.S.A.) conifer forests. Conservation Biology 9:1041-1058. Laake, J. L., S. T. Buckland, D. R Anderson,and K. P. Burnham. 1994. DISTANCE User's Guide, ver. 2.1. Colorado Cooperative Fish & Wildlife Research Unit, Colorado State University, Ft. Collins, co. 84pp. Leukering, T. 1996. Alfred M. Bailey Bird Nesting Area Breeding Bird Monitoring and Research. Unpubl. report. CBO, Brighton, CO. __ . 1998. Point transect protocol. Unpubl. doc. CBO, Brighton, CO. __ and M. Carter. 1998. Colorado Birds Monitored by 2001: Results of the Spruce-Fir Pilot Transects. Unpubl. report. CBO, Brighton, CO. Partners in Flight. 1998. Priority scores for Colorado, v. 1998.10. http://members.aol.comlnocedalcbo/CO.html. Patuxent Wildlife Research Center. 1998. MONITOR, v. 1998.10. (www.im.nbs.gov/pub/sofiwareimonitor). Stark, RD., D. 1. Dodenhoff, and E. V. Johnson. 1998. A quantitative analysis of woodpecker drumming. Condor 100:350-356. �Table 1. Birds detected on point transects in three habitats in Aspen, Ponderosa Pine, and Spruce-Fir in Colorado in summer 1998. Categories are: P=birds detected on point counts, F=birds detected only as flyovers, and T=birds detected on transect legs (see text). Species Turkey Vulture Canada Goose Mallard Osprey Sharp-shinned Hawk Cooper's Hawk Northern Goshawk unidentified accipiter Red-tailed Hawk Golden Eagle American Kestrel Prairie Falcon Blue Grouse Wild Turkey Sora Spotted Sandpiper Conunon Snipe Wilson's Phalarope Band-tailed Pigeon Mourning Dove Great Horned Owl Conunon Nighthawk White-throated Swift Broad-tailed Hununingbird Belted Kingfisher Lewis's Woodpecker Williamson's Sapsucker Red-naped Sapsucker unidentified sapsucker Downy Woodpecker Hairy Woodpecker Three-toed Woodpecker Northern Flicker unidentified woodpecker Olive-sided Flycatcher Western Wood-Pewee Hanunond's Flycatcher Dusky Flycatcher Cordilleran Flycatcher unidentified flycatcher P F Spruce-Fir Ponderosa Pine Aspen T Total 3 0 1 3 1 2 1 0 0 0 1 0 0 5 1 6 1 1 8 2 2 6 0 0 1 0 3 6 2 1 0 0 0 0 2 1 2 0 0 2 68 18 1 87 11 36 0 0 2 10 13 46 1 31 0 1 4 12 5 44 44 2 33 164 24 62 25 0 2 0 0 0 0 0 5 1 5 0 15 1 0 49 5 38 164 39 63 25 Total P F T Total 0 0 0 0 0 2 1 2 1 4 1 1 4 3 0 1 0 1 3 0 1 1 0 0 0 4 0 2 0 1 1 1 13 1 3 1 5 1 1 1 0 1 2 1 0 1 2 0 1 3 1 0 5 26 0 0 0 2 0 1 6 0 1 1 11 28 3 0 0 3 8 3 87 1 0 41 14 6 0 27 1 2 21 0 0 0 0 0 0 0 6 0 1 0 2 14 8 1 3 20 13 7 0 20 5 2 0 0 1 0 6 2 80 9 18 98 15 114 20 1 1 0 0 0 0 0 0 0 14 2 4 0 2 0 0 0 15 5 109 1 2 55 22 7 3 47 0 95 11 22 98 17 114 20 1 20 6 9 10 11 3 14 4 10 0 0 1 8 7 0 2 3 0 3 0 1 28 13 10 12 14 3 17 4 11 P F 0 1 2 0 1 2 2 2 0 2 1 0 0 0 6 1 0 0 4 0 0 T 0 0 0 0 0 0 2 N 0 \C �Table 1. Continued. .- N Ponderosa Aspen Species Ash-throated Flycatcher Plumbeous Vireo Warbling Vireo Gray Jay Steller's Jay Clark's Nutcracker Black-billed Magpie American Crow Common Raven Purple Martin Tree Swallow Violet-green Swallow N. Rough-winged Swallow Cliff Swallow unidentified swallow Black-capped Chickadee Mountain Chickadee Red-breasted Nuthatch White-breasted Nuthatch Pygmy Nuthatch Brown Creeper Rock Wren House Wren American Dipper Golden-crowned Kinglet Ruby-crowned Kinglet Blue-gray Gnatcatcher Western Bluebird Mountain Bluebird Townsend's Solitaire Veery Swainson's Thrush Hermit Thrush American Robin European Starling Orange-crowned Warbler Virginia's Warbler Yellow Warbler Yellow-rurnped Warbler Grace's Warbler MacGillivray's Warbler P F Total 1 828 17 48 19 0 0 0 0 4 0 0 0 1 3 1 828 17 49 26 1 11 1 30 91 0 8 0 4 5 0 0 1 0 0 1 19 2 34 96 6 4 177 19 12 1 5 1 0 0 0 0 0 0 0 0 0 4 1 0 3 7 4 177 23 13 1 8 302 1 10 89 0 0 0 0 0 0 6 0 302 1 16 89 30 8 1 1 0 0 31 9 1 97 326 0 0 2 0 1 0 1 98 328 85 2 7 328 I 0 0 1 0 0 0 0 86 2 7 329 31 0 1 32 T Pine Spruce-Fir P F T Total 1 43 202 0 0 1 0 0 0 133 53 7 13 14 0 9 2 5 18 0 12 0 0 0 1 43 203 0 133 74 9 18 32 10 77 1 0 3 24 0 1 0 0 0 0 13 101 10 1 196 31 69 82 13 13 96 2 0 0 1 0 0 0 0 4 1 5 10 1 0 198 35 70 88 23 14 96 1 92 5 24 42 94 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 93 5 24 42 94 1 112 305 1 38 27 0 4 0 0 0 0 0 0 0 0 112 309 207 20 13 1 0 0 0 0 0 0 P F T Total 28 77 46 61 0 3 0 2 0 10 0 9 28 90 46 72 1 30 0 9 0 0 1 39 5 5 0 10 250 18 5 0 1 0 251 19 5 29 1 0 0 0 0 13 42 3 3 91 314 0 0 0 0 03 2 46 0 5 137 314 15 26 0 0 0 2 15 28 2 342 185 0 0 2 0 0 0 2 342 187 38 27 3 0 0 3 208 20 13 414 1 0 415 5 0 0 5 I �Table 1. Continued. Aspen S~cies Wilson's Warbler Western Tanager Green-tailed Towhee Spotted Towhee Chipping Sparrow Brewer's Sparrow Vesper Sparrow Lark Sparrow Fox Sparrow Song Sparrow Lincoln's Sparrow White-crowned Sparrow Dark-eyed Junco Black-headed Grosbeak Blue Grosbeak Lazuli Bunting Red-winged Blackbird Western Meadowlark Yellow-headed Blackbird Brewer's Blackbird Brown-headed Cowbird Pine Grosbeak Cassin's Finch Red Crossbill White-winged Crossbill Pine Siskin American Goldfinch Evening Grosbeak unidentified finch Totals Ponderosa P F T Total 8 49 44 3 22 0 0 0 0 0 0 0 0 0 0 8 49 44 3 22 4 0 0 4 0 0 0 1 0" 0 0 0 0 0 1 119 41 331 10 1 0 0 1 1 0 0 1 18 5 7 3 2 1 1 14 0 1 0 0 20 7 8 17 121 83 0 204 2 4 0 6 3899 158 86 4143 1 119 41 330 10 P Pine SEruce-Fir F T 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 204 190 38 126 2 30 2 2 3 0 0 0 0 2 3 261 28 2 4 7 3 2 2 47 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 261 28 2 4 7 3 2 2 49 10 11 0 67 0 0 10 78 113 0 9 1 97 3 9 0 0 0 0 0 210 3 18 1 3727 287 129 4143 I 204 190 38 126 2 30 Total P F T 28 21 4 0 0 0 0 0 0 28 21 4 20 0 0 20 2 0 0 2 48 47 248 1 0 0 0 0 0 0 0 0 48 47 248 1 0 59 19 11 0 213 1. 4 0 38 8 100 0 16 1 0 0 0 1 79 20 49 8 313 15 2 4 0 2 0 21 2 2805 187 130 3122 Total N �212 Table 2. Species recorded on point transects in three habitats, Aspen (A), Ponderosa Pine (P), and Spruce-Fir (S), in Colorado, Summer 1998. Only species that were recorded >5 times in at least one habitat are included. Asterisks indicate target species in listed habitat. Density (D) is n/ha; CV (D) is the coefficient of variation of the density; % var (n) is the percentage of the variance due to variation in sample size; P=probability of detection; K=number of transects on which the species was recorded; and m=the number of parameters included in the detection-curve function. Species Habitat Band-tailed Pigeon P* K m n D CV(D) % var (n) P 9 0.60 42.1 100.0 1.000 4 0 Mourning Dove P 26 1.14 36.5 37.4 0.165 11 3 Common Nighthawk P 11 183.35 171.9 1.0 0.221 6 3 Broad-tailed Hummingbird A* P S 68 95 13 12.40 17.33 2.24 24.8 25.3 15.4 41.5 48.6 100.0 0.086 0.021 1.000 24 24 11 4 2 0 Williamson's Sapsucker A S 11 46 8 3.42 6.31 1.60 58.0 82.6 25.0 39.4 7.1 4.4 0.237 0.050 0.154 3 18 5 2 2 2 Red-naped Sapsucker A* P S 36 20 2 3.16 0.80 18.8 33.0 39.2 100.0 0.191 1.000 22 8 1 0 Hairy Woodpecker A P S 33 30 20 3.52 2.09 2.90 25.1 46.2 18.7 55.2 16.5 44.5 0.119 0.155 0.169 14 18 9 3 4 2 Three-toed Woodpecker S· 6 1.89 16.7 100.0 1.000 5 0 Northern Flicker A P S 45 91 9 1.80 1.53 1.66 31.0 19.4 73.5 26.2 23.0 4.1 0.075 0.068 ·0.094 17 28 6 2 2 3 Olive-sided Flycatcher A P S· 33 20 11 0.53 0.68 0.46 19.3 21.5 33.5 48.8 21.6 12.9 0.321 0.379 0.420 15 11 9 2 1 1 Western Wood-Pewee A* P S 164 102 7 3.84 2.44 25.6 24.6 53.5 87.4 0.235 0.133 24 23 3 1 Hammond's A P S· 25 20 14 4.50 6.13 3.97 24.5 27,7 25.3 100.0 52.3 51.1 1.000 0.251 0.349 8 6 6 0 2 1 Dusky Flycatcher A P S 62 120 4 4.89 11.62 42.3 17.8 28.2 75.5 0.362 0.070 17 21 4 4 Cordilleran Flycatcher A P S 25 21 10 3.19 3.02 0.54 65.8 31.6 22.4 8.4 17.2 100.0 0.123 0.123 1.000 13 11 5 4 3 0 Plumbeous Vireo P 43 6.64 26.7 55.7 0.130 11 3 A* P S 828 237 28 23.70 4.70 0.96 9.1 23.1 21.0 88.4 95.0 34.0 0.101 0.099 0.195 30 23 11 1 1 1 Gray Jay A S· Steller's Jay A 17 77 48 140 46 26.60 9.88 1.50 2.85 1.17 64.1 26.2 21.4 19.0 48.0 11.5 31.5 59.5 38.3 47.8 0.003 0.015 0.176 0.071 0.101 7 17 15 27 18 4 3 1 2 2 Warbling Flycatcher Vireo p. P S �213 Table 2. Continued. Species Habitat n D CV(D) % var (n) p K m Clark's Nutcracker A P S* 19 60 61 2.63 0.51 1.02 90.5 21.6 19.2 17.4 41.2 52.4 0.076 0.117 0.045 6 20 15 2 2 3 Black-billed P 10 0.59 59.7 28.0 0.270 5 2 American Crow P S 13 1 0.51 54.2 55.7 0.160 4 2 Common A P S 10 13 30 1.20 0.14 1.09 59.4 91.5 49.5 28.3 5.8 54.4 0.059 0.116 0.017 5 6 5 4 2 2 A* 30 12 2.18 3.06 32.2 47.1 54.5 20.3 0.130 0.276 11 5 5 2 91 81 5 5.02 4.34 26.9 24.8 52.1 63.8 0.082 0.053 17 18 4 3 Magpie Raven Tree Swallow P Violet-green Swallow A* P S Mountain Chickadee A P S* 177 216 250 10.37 7.20 9.63 27.2 151.7 15.7 42.1 0.8 64.3 0.052 0.050 0.059 24 27 28 3 4 2 Red-breasted A P S* 19 44 18 1.24 1.20 2.74 23.7 38.8 26.8 49.2 80.3 58.9 0.241 0.241 0.299 13 10 . 7 2 1 2 A P S 12 71 5 3.20 2.74 50.3 21.6 25.1 46.8 0.288 0.107 8 24 2 3 Pygmy Nuthatch P* 87 8.19 37.2 52.7 0.144 16 3 Brown Creeper A P S 6 18 29 5.53 1.91 7.36 84.9 17.4 19.1 100.0 100.0 67.6 0.088 1.000 0.244 6 12 17 3 0 2 Rock Wren P 14 0.58 34.6 69.8 0.370 7 1 House Wren A P S 302 107 3 20.42 4.62 17.7 21.0 51.5 63.8 0.077 0.094 24 23 4 3 A S* 12 91 7.64 9.72 80.2 19.0 4.9 53.5 0.033 0.047 8 24 2 3 A P S* 89 107 314 0.90 5.20 5.44 28.6 44.4 11.8 27.2 30.8 87.3 0.264 0.074 0.090 27 16 27 3 3 1 P* 27 6.76 38.3 61.9 0.159 8 2 Mountain Bluebird A* P S 30 43 15 3.64 5.11 3.94 36.1 26.9 20.0 45.4 69.9 100.0 0.109 0.029 1.000 10 8 4 2 2 0 Townsend's A P S 8 103 26 0.86 1.53 0.47 37.7 25.4 34.3 44.0 64.3 75.4 0.260 0.103 0.163 6 23 11 2 3 2 Hermit Thrush A P S* 97 118 342 0.83 1.61 3.53 15.2 28.1 14.6 64.4 77.9 51.0 0.135 0.056 0.062 24 22 28 1 4 6 American A P S 326 330 185 9.37 8.45 7.54 13.1 17.4 24.0 62.7 54.3 35.8 0.063 0.048 0.048 28 29 26 2 3 3 Nuthatch White-breasted Nuthatch Golden-crowned Ruby-crowned Western Kinglet Kinglet Bluebird Solitaire Robin �214 Table 2. Continued. Species Habitat n D CV (D) % var (n) P K m 3 1 Orange-crowned Warbler A P S 85 38 3 6.56 2.90 35.6 30.3 77.4 68.5 0.160 0.285 16 10 Virginia's Warbler P 27 3.24 25.4 38.2 0.271 10 Yellow Warbler A 7 1.70 62.8 82.9 0.381 3 1 Yellow-rumped Warbler A P S 328 225 414 8.97 7.08 12.16 15.0 36.1 11.7 79.0 25.7 59.9 0.146 0.089 0.081 29 26 29 2 5 2 Grace's Warbler P* 20 3.36 45.7 62.2 0.494 6 MacGillivray's Warbler A P S 31 13 5 7.25 2.29 34.4 27.7 61.5 63.3 0.495 0.356 11 6 1 1 Wilson's Warbler A S 8 28 0.80 2.45 25.0 46.0 100.0 73.3 1.000 0.228 5 7 0 2 Western Tanager A P S 49 213 21 2.60 5.62 1.51 26.2 19.0 68.0 74.6 70.8 14.9 0.152 0.105 0.131 16 27 7 1 4 2 Green-tailed Towhee A P S 44 197 4 1.62 5.43 26.3 23.4 53.7 81.9 0.213 0.056 19 23 2 2 Spotted Towhee P 38 4.73 49.5 83.2 0.163 8 2 Chipping Sparrow A P* S 22 126 20 0.85 ·3.58 2.89 22.3 17.9 63.1 81.5 73.2 38.6 0.242 0.122 0.132 10 23 6 2 3 2 Lincoln's Sparrow A S 119 48 8.89 4.33 28.2 33.5 74.8 56.4 0.104 0.067 16 13 3 3 White-crowned Sparrow A S 41 47 2.50 13.94 23.0 191.7 79.9 1.9 0.291 0.024 7 11 2 4 Dark-eyed Junco A P S 330 282 248 23.07 14.11 6.73 21.6 59.2 9.2 31.6 5.5 82.8 0.059 0.024 0.088 30 30 28 3 4 1 Black-headed Grosbeak A P S 10 28 1 0.79 1.05 56.7 43.9 12~5 60.4 0.164 0.059 8 9 2 4 Brown-headed Cowbird A P 18 49 3.48 5.91 29.2 28.1 66.6 47.5 0.291 0.045 7 14 2 2 Pine Grosbeak A S* 6 59 0.36 4.73 19.3 27.3 100.0 32.4 1.000 0.101 4 18 0 2 Cassin's Finch P S* 7 19 0.37 0.79 66.3 40.3 4.6 66.6 0.174 0.267 6 7 2 1 Red Crossbill P S* 14 11 1.39 0.81 73.0 61.9 44.1 12.5 0.157 0.164 5 6 4 2 Pine Siskin A P S· 121 125 212 6.17 7.63 6.96 21.6 17.3 16.8 74.6 84.2 94.1 0.029 0.069 0.083 24 22 2 2 4 1 Evening Grosbeak P S 9 15 5.79 0.83 74.3 55.9 26.8 100.0 0.293 1.000 3 4 2 0 �215 Table 3. Analysis of various truncation distances for selected species in Aspen (target species and nontarget species found in highest abundance in Aspen). See Table 2 for column heading explanations. Check-marks indicate those truncations considered the best for optimization ofCVs, # of parameters, and variance due to sample size. Truncation distance (m) Species n 0 CV(%) % var (n) m Best model 29 52 52 63 68 16.14 17.60 12.40 41.7 22.1 24.8 18.5 54.7 41.5 3 1 4 Hairy Woodpecker 80 97 118 30 31 32 33 3.98 3.88 3.56 3.52 29.2 27.3 24.8 25.1 47.3 51.6 56.1 55.2 1 1 1 3 Western Wood-Pewee 105 136 154 161 3.60 3.82 20.4 19.3 96.3 97.8 1 1 164 3.84 25.6 53.5 3 14 25 4.88 4.50 23.5 24.5 100.0 100.0 0 0 ./ 540 758 826 828 23.38 22.27 24.05 23.70 23.7 10.1 9.2 9.1 22.3 82.9 87.4 88.4 3 1 1 1 ./ 70 98 126 20 24 27 30 3.36 2.82 2.80 2.18 38.3 33.5 41.7 32.2 85.4 60.5 24.9 54.5 1 1 2 5 73 171 64 87 4.83 4.80 43.1 22.4 32.8 75.8 2 1 91 5.02 26.9 52.1 4 Broad-tailed Hammond's Warbling Hummingbird Flycatcher Vireo 61 100 169 Tree Swallow Violet-green 34 Swallow ./ ./ 58 246 19.02 12.0 91.6 1 78 277 18.71 21.4 30.9 3 302 20.42 17.7 51.5 4 66 86 60 71 85 5.50 5.82 6.56 45.7 33.4 35.6 42.4 82.6 77.4 2 1 3 ./ Lincoln's Sparrow 70 98 119 11.69 8.89 24.2 28.2 94.1 74.8 1 3 ./ Dark-eyed Junco 58 97 154 218 290 329 330 30.35 26.48 23.51 23.07 21.6 26.2 21.5 21.6 58.9 24.6 32.0 31.6 3 3 3 3 House Wren Orange-crowned Warbler �216 Table 4. Analysis of various truncation distances for selected species in Ponderosa Pine (target species and non-target species found in highest abundance in Ponderosa Pine). See Table 2 for column heading explanations. Check-marks indicate those truncations con-sidered the best for optimization ofCVs, # of parameters, and variance due to sample size. Truncation distance (m) Species Mourning Dove Williamson's Sapsucker n D CV(%) % var (n) m 168 184 25 25 26 2.04 1.03 1.14 38.1 28.2 36.5 39.0 71.4 37.4 3 1 3 115 39 46 6.74 6.31 223.3 82.6 0.8 7.1 3 2 Northern Flicker 164 290 80 89 91 1.74 1.56 1.53 15.2 19.8 19.4 46.1 23.3 23.0 1 2 2 Dusky Flycatcher 54 84 98 112 120 10.29 11.67 11.62 19.5 20.0 17.8 89.0 71.9 75.5 1 1 4 48 86 29 38 43 6.58 6.42 6.64 34.5 29.1 26.7 70.9 61.7 55.7 1 1 3 123 194 30 43 44 1.25 0.87 1.20 35.5 36.6 38.8 76.5 93.5 80.3 1 1 1 87 107 57 62 71 3.22 3.05 2.74 20.1 21.6 21.6 88.7 63.0 46.8 1 1 3 58 82 76 85 87 8.96 9.82 8.19 33.2 34.2 37.2 94.2 66.5 52.7 1 2 3 136 177 190 42 42 42 43 8.69 8.62 8.68 5.11 28.1 28.2 28.1 26.9 66.9 66.5 66.8 69.9 1 1 1 2 204 240 99 101 103 1.56 1.94 1.53 22.6 23.7 25.4 81.3 74.3 64.3 1 3 3 120 264 274 327 330 10.98 9.08 8.45 14.5 17.8 17.4 71.5 51.5 54.3 4 3 3 Western Tanager 132 148 197 201 213 4.55 4.65 5.62 21.1 17.8 19.0 61.2 83.3 70.8 2 1 4 Green-tailed 156 204 288 191 196 196 197 7.33 6.16 5.14 5.43 22.8 24.6 22.2 23.4 81.3 74.7 92.3 81.9 3 2 1 2 73 101 32 36 38 5.55 4.14 4.73 54.6 46.7 49.5 77.1 96.3 83.2 2 1 2 Plumbeous Vireo Red-breasted Nuthatch White-breasted Nuthatch Pygmy Nuthatch Mountain Bluebird Townsend's American Solitaire Robin Towhee Spotted Towhee Best model ./ ./ ./ ./ ./ ./ ./ ./ ./ ./ �217 Table 4. Continued. Truncation distance (m) Species Chipping Sparrow Black-headed Grosbeak % var (n) m 19.4 17.3 17.9 60.6 65.6 73.2 1 1 3 37.8 47.7 43.9 77.8 48.9 60.4 1 5 4 n D CV(%) 96 128 99 112 126 3.84 3.63 3.58 240 352 27 27 28 1.45 4.01 1.05 Best model .t .t �218 Table 5. Analysis of various truncation distances for selected species in Spruce-Fir (target species and non-target species found in highest abundance in Spruce-Fir). See Table 2 for column heading explanations. Check-marks indicate those truncations considered the best for optimization of Cvs, # of parameters, and variance due to sample size. Species Gray Jay Truncation distance (m) n D CV(%) % var (n) m Best model 63 190 60 73 77 10.75 9.47 9.88 23.9 24.5 26.2 70.5 40.5 31.5 2 2 3 .I 133 292 48 59 61 1.21 1.34 1.02 18.6 21.1 19.2 84.0 50.3 52.4 1 1 3 .I Red-breasted Nuthatch 70 90 95 14 16 16 18 3.33 3.10 2.71 2.74 39.5 35.4 25.4 26.8 22.9 34.3 66.5 58.9 1 1 1 2 Brown Creeper 28 47 21 28 29 10.40 8.07 7.36 30.3 17.3 19.1 29.0 63.2 67.6 1 1 2 81 83 91 12.39 9.72 17.4 19.0 59.4 53.5 1 3 220 248 312 312 314 5.50 5.87 5.44 13.5 12.2 11.8 68.9 84.1 87.3 2 1 1 HermitThrush 147 189 357 259 286 333 342 3.24 3.71 2.96 3.53 23.9 14.0 23.5 14.6 30.2 78.6 20.3 51.0 2 1 5 6 Yellow-rumpedWarbler 86 140 183 194 344 395 405 405 414 14.59 11.41 11.26 12.10 12.16 10.8 15.4 11.2 20.5 11.7 87.8 32.3 64.3 19.3 59.9. 1 3 2 4 2 Pine Grosbeak 74 89 118 141 51 53 57 58 59 3.78 5.56 4.11 5.24 4.73 21.7 24.6 25.2 19.7 27.3 88.9 60.3 43.4 67.7 32.4 1 1 2 3 2 Pine Siskin 124 147 169 180 202 205 208 208 214 28.0 17.1 18.1 18.0 17.0 31.0 83.8 74.3 75.0 94.4 3 1 2 2 1 Clark's Nutcracker Golden-crowned Kinglet Ruby-crowned Kinglet 7.67 9.26 7.66 7.65 6.84 .I .I .I .I �219 Table 6. Number of years required for target trend detection (>3o/olyr. change with statistical significance of 0.1 and power of 80%, assuming 30 transects run annually) for target species in Aspen, Ponderosa Pine, and Spruce-Fir using program MONITOR (pWRC 1998). Asterisks and boldface indicate target species within habitats. Aspen Species Band-tailed Pigeon Broad-tailed Hummingbird Williamson's Sapsucker Red-naped Sapsucker Three-toed Woodpecker Olive-sided Flycatcher Western Wood-Pewee Hammond's Flycatcher Warbling Vireo Gray Jay Clark's Nutcracker Tree Swallow Violet-green Swallow Mountain Chickadee Red-breasted Nuthatch Pygmy Nuthatch Golden-crowned Kinglet Ruby-crowned Kinglet Western Bluebird Mountain Bluebird Hermit Thrush Grace's Warbler Chipping Sparrow Pine Grosbeak Cassin's Finch Red Crossbill Pine Siskin CV(O) #of years 24.8 58.0 18.8 7 * 9 6 * 19.3 25.6 24.5 9.1 64.1 90.5 32.2 26.9 27.2 23.7 6 7 7 5 10 12 7 7 7 7 80.2 28.6 11 7 * * * *. 36.1 15.2 8 * 6 22.3 19.3 6 6 21.6 6 Ponderosa #of years CV(O) 42.1 25.3 82.6 33.0 8 * 7 11 * 8 21.5 24.6 27.7 23.1 6 7 '7 6 21.6 47.1 24.8 151.7 38.8 37.2 6 9 7 14 8 8 * 44.4 38.3 26.9 28.1 45.7 17.9 9 8 * 7 7 9 * 6 * 66.3 73.0 17.3 10 10 6 PineSpruce-Fir #of years CV(O) 15.4 25.0 6 7 16.7 33.5 6 * 8 * 25.3 21.0 26.2 19.2 7 * 6 7 * 6 * 15.7 26.8 6 * 7 * 19.0 11.8 6 * 5 * 20.0 14.6 6 6 * 63.1 27.3 40.3 61.9 16.8 10 7 8 10 6 * * * * �220 Table 7. Comparison, within Spruce-Fir, of coefficients of variation (CV) for results from detections only with those from density estimates, rounded to nearest whole percent. The number of years was calculated from density-derived CVs (Table 2). See Table 4 for more details on truncation results. Unlimited-radius Detections Species Three-toed Woodpecker Olive-sided Flycatcher Hammond's Flycatcher Gray Jay Clark's Nutcracker Mountain Chickadee Red-breasted Nuthatch Golden-crowned Kinglet Ruby-crowned Kinglet Hermit Thrush Pine Grosbeak Cassin's Finch Red Crossbill Pine Siskin CV(%) 242 167 200 121 123 70 201 93 66 62 118 250 215 102 Density estimates (all data) CV(%) 17 # of years to detect trend using TRENDS1 MONITOR2 15 25 24 19 26 19 20 16 16 27 14 34 19 20 16 21 'Gerrodette (1993). 2Patuxent Wildlife Research Center (1998). 7 7 6 6 7 6 29 15 27 20 40 26 7 5 7 8 62 17 35 15 ·6 14 19.64 StD Variance 6 8 5.76 33.17 9 6.79 1.05 1.10 �221 Appendix A. Species recorded on 89 point transects in Colorado, Summer 1998. Species Turkey Vulture Canada Goose Mallard Osprey Sharp-shinned Hawk Cooper's Hawk Northern Goshawk Red-tailed Hawk Golden Eagle American Kestrel Prairie Falcon Blue Grouse Wild Turkey Sora Spotted Sandpiper Common Snipe Wilson's Phalarope Band-tailed Pigeon Mourning Dove Great Horned Owl Common Nighthawk White-throated Swift Broad-tailed Hummingbird Belted Kingfisher Lewis's Woodpecker Red-naped Sapsucker Williamson's Sapsucker Downy Woodpecker Hairy Woodpecker Three-toed Woodpecker Northern Flicker Olive-sided Flycatcher Western Wood-Pewee Hammond's Flycatcher Dusky Flycatcher Cordilleran Flycatcher Ash-throated Flycatcher Plumbeous Vireo Warbling Vireo Gray Jay Steller's Jay Clark's Nutcracker Black-billed Magpie American Crow Common Raven Purple Martin Tree Swallow Violet-green Swallow Northern Rough-winged Swallow Cliff Swallow Black-capped Chickadee Mountain Chickadee Red-breasted Nuthatch White-breasted Nuthatch Pygmy' Nuthatch Brown Creeper Rock Wren House Wren American Dipper Golden-crowned Kinglet Ruby-crowned Kinglet Scientific name Cathartes aura Branta canadensis Anas platyrhynchos Pandion haliaetus Accipiter striatus Accipiter cooperii Accipiter gentilis Buteo jamaicensis Aquila chrysaetos Falco sparverius Falco mexicanus Dendragapus obscurus Meleagris gallopavo Porzana carolina Actitis macularia Gallinago gallinago Phalaropus tricolor Columba fasciata Zenaida macroura Bubo virginianus Chorde/7es minor Aeronautes saxatalis Selasphorus platycercus Ceryle alcyon Melanerpes lewis Sphyrapicus nuchalis Sphyrapicus thyroideus Picoides pubescens Picoides villosus Picoides tridactyfus Colaptes auratus Contopus cooperi Contopus sordidulus Empidonax hammondii Empidonax oberholseri Empidonax occidentalis Myiarchus cinerascens Vireo plumbeous Vireo gilvus Perisoreus canadensis Cyanocitta stelleri Nucifraga columbiana Pica pica Corvus brachyrhynchos Corvus corax Progne subis Tachycineta bicolor Tachycineta thalassina Stelgidopteryx serripennis Petrochelidon pyrrhonota Poecile atricapillus Poecile gambelii Sitta canadensis Sitta carolinensis Sitta pygmaea Certhia americana Salpinctes obsoletus Troglodytes aedon Cindus mexican us Regulus satrapa Regulus calendula �222 Appendix A. Continued. Species Scientific name Blue-gray Gnatcatcher Western Bluebird Mountain Bluebird Townsend's Solitaire Veery Swainson's Thrush Hermit Thrush American Robin European Starling Orange-crowned Warbler Virginia's Warbler Yellow Warbler Magnolia Warbler Yellow-rumped Warbler. Grace's Warbler MacGillivray's Warbler Wilson's Warbler Western Tanager Green-tailed Towhee Spotted Towhee Chipping Sparrow Brewer's Sparrow Vesper Sparrow Lark Sparrow Fox Sparrow Song Sparrow Lincoln's Sparrow White-crowned Sparrow Dark-eyed Junco Black-headed Grosbeak Blue Grosbeak. Lazuli Bunting Red-winged Blackbird Western Meadowlark Yellow-headed Blackbird Brewer's Blackbird Brown-headed Cowbird Pine Grosbeak Cassin's Finch Red Crossbill White-winged Crossbill Pine Siskin American Goldfinch Evening Grosbeak Polioptila ceetulee Sialia mexican a Sialia cutrucokies Myadestes townsendi Catharus fuscescens Catharus ustulatus Catharus guttatus Turdus migratorius Sturn us vulgaris Vermivora celata Vermivora virginiae Dendroica petechia Dendroica magnolia Dendroica coronata Dendroica graciae Oporornis tolmiei Wi/sonia pusilla Piranga /udoviciana Pipilo ch/orurus Pipilo macu/atus Spizella passerina Spizella breweri Pooecetes gramineus Chondestes grammacus Passerella iliaca Melospiza me/odia Melospiza lincolnii Zonotrichia /eucophrys Junco hyemalis Pheucticus melanocephalus Guiraca caerulea Passerina amoena Agelaius phoeniceus Sturn ella neglecta Xanthocephalus xanthocephalus Euphagus cyanocephalus Molothrus ater Pinicola enucleator Carpodacus cassinii Loxia curvirostra Loxia leucoptera . Carduelis pinus Carduelis tristis Coccothraustes vespertinus �223 Appendix B. Colorado special species list, with results from 1998 monitoring efforts. AI 4 4 4 4 3 3 3 3. 3 3 3 3 3 3 3 3 3 3 3 3 3 3 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 Species Common Nighthawk Common Poorwill Cordilleran Flycatcher American Dipper Western Grebe Clark's Grebe American White Pelican Great Blue Heron White-faced Ibis Northern Goshawk Golden Eagle Prairie Falcon Spotted Sandpiper Black Tern Flammulated Owl Western Screech-Owl Great Horned Owl Northern Pygmy-Owl Long-eared Owl Boreal Owl Northern Saw-whet Owl Black Swift Eared Grebe Double-crested Cormorant Great Egret Snowy Egret Cattle Egret Green Heron Black-crowned Night-Heron Osprey Mississippi Kite Black Rail Sora Black-necked Stilt Willet California Gull Forster's Tern Eurasian Collared-Dove Black-billed Cuckoo Barn Owl Eastern Screech-Owl Spotted Owl Black Phoebe Eastern Phoebe Scissor-tailed Flycatcher Bell's Vireo . Purple Martin Bank Swallow # Adults 843 1678 507 # Juveniles 61 70 188 # Nests 26 2 8 69 2585 78 37 1186 11 3 pairs + 2 72 7 males 72 97 pairs 93 3 pairs 4 pairs + 1 32 1 2 6 16 2 �224 AppendixB. Continued. AI 2 2 2 2 I # Adults Species American Redstart Ovenbird 3 males+3 pr 4 males Bobolink Scott's Oriole I pair Least Bittern Little Blue Heron Yellow-crowned Night-Heron Harlequin Duck Broad-winged Hawk Merlin Marbled Godwit Magnificent Hummingbird Acorn Woodpecker I pair Least Flycatcher Vermilion Flycatcher Red-eyed Vireo " Carolina Wren Bendire's Thrasher Golden-winged Warbler Lucy's Warbler Chestnut-sided Bay-breasted Warbler 3 males Warbler Northern Waterthrush Hooded Warbler 1 I pair Hepatic Tanager Northern Cardinal 6 males Eastern Meadowlark Field Sparrow White-winged 14 Crossbill 28 # Juveniles # Nests �225 Colorado Division of Wildlife Wildlife Research Report April 1999 JOB PROGRESS REPORT Stmeof ~C~o~lo~ra~d~o~ _ Avian Research Project: W'-'---'-1:,.;:6:...,:.7.....:-R:..:_ _ WorkPlan_12_ : Job_l_ Job Title: ~Id:::e::..:n.:.:.tify........,D=is.:.:.tn.!.!:·b~u~ti~on~an'""'d~R:::;ep~r~o::::d.::.uc:::.:tr:!.-'. v~e'_"S~t=atu=s ..•• o""-f=M=o"""un=taI:::·n~P=_=lo:...!.v=ers""'_ _ Period Covered: Auilior: Personnel: __ 0 1 January - 31 December ~~~r~ru~d~R~C~r~ru~g 1998 _ Gerald R Craig and Kenneth M. Giesen, Colorado Division of Wildlife ABSTRACT A preliminary study plan was developed, distributed for peer review, and then revised. Changes were incorporated and the study plan was finalized, Prescribed burns on the Pawnee and Comanche National Grasslands were visited and examined in preparation for the study. �226 �227 IDENTIFY DISTRIBUTION AND REPRODUCTIVE STATUS OF MOUNTAIN PLOVERS Gerald R Craig Planning Phase P. N. OBJECTIVES The objectives of the initial planning were to (1) review literature pertinent to monitoring status and trends of mountain plovers (Charadrius montanus), (2) evaluate existing efforts which monitor mountain plovers in Colorado, (3) expand reproductive monitoring efforts to other regions of the state, particularly South Park and portions of southeastern Colorado, and (4) develop a detailed study plan for monitoring mountain plover reproductive status and population trends. RESULTS During this segment, a preliminary research proposal was developed and circulated for peer review. Changes were incorporated and the following P.N. Objectives and Segment Objectives were established for this study. P. N. OBJECTIVES The primary objectives of this study are to (1) develop and evaluate a statewide monitoring program to identify distribution, population trends, and changes in productivity for mountain plover, and (2) investigate contributions of burned grasslands and fallow cultivated fields to plover productivity. Field Phase SEGMENT OBJECTIVES 1. Develop and test a statewide plover population monitoring protocol. Apply and test inventory techniques that permit statistically powerful analysis to assess population densities, productivity and trends. The actual census technique will be developed in consultation with Dr. David Anderson of the Colorado Cooperative Wildlife Research Unit. DWMs and Area Biologists will be encouraged to visit and inventory those habitats that appear suitable for plover nesting, brood rearing and staging. 2. Monitor plover breeding densities, nest success and chick survival on prescribed burns on the Comanche National Grasslands and evaluate the effectiveness of that potential management technique. Up to 3 designated prescribed burn areas will be inventoried a year prior to the burn, the year of the burn and a year following the burn. Up to 30 plovers nesting within the study sites will be trapped, banded and equipped with transmitters. Movements of broods will be tracked so that mortality, dispersal and habitat preferences can be documented. 3. Monitor breeding densities, nest success and chick survival on fallow, cultivated fields. Fallow cultivated fields will be surveyed by the investigators, DWMs, and Area Biologists for the presence of nesting plovers. Documented nests will be monitored to document success and productivity. A sample of up to 30 nesting adults will be trapped and treated as described in 2 to monitor the broods' movements and attrition of broods. �228 4. Compile data and prepare annual report. RESULTS Aside from visits to prescribed bums on the Pawnee and Comanche National grasslands, investigations were undertaken during this segment. Prepared By: __ ----l...C..sz_...:.·:.....r:2=--.:.... ...l...fJ...L:hai?-~l,Gerald R Craig----OWildlife Researcher _ no field � https://cpw.cvlcollections.org/files/original/871ededc07089190c3b6d3fdbfdbccbd.pdf de33a4fa906859548ba07a4ad03c5a3a PDF Text Text 1 Colorado Division of Wildlife Wildlife Research Report July 1999 JOB PROGRESS REPORT Cost Center 3430 State of -"C"""o:,:.::lo:<:.r.=ad=o"--_ Mammals Program Project No. W~-1~5:..:::3....::-R~-..;!.1.::.2_,......--Work Package No. _0.=...;6;:..,;:6;.::2 _ Preble's Meadow Jumping Mouse Conservation Task No. ....:.l _ Develop Conservation Plan for Preble's Meadow Jumping Mouse Period Covered: July 1, 1998 - June 30, 1999 Author: Tanya M. Shenk 'Personnel: CSU J. Brinker, J. Eussen, M. Miller, M. Sivert, M. Wild, CD OW, K. Burnham, G. White, ABSTRACT Preliminary analysis of the movement data collected on radio-collared PMJM indicate (1) maximum movements of> 1 mile, (2) greater use of upland habitats than previously assumed, (3) general site fidelity to both daytime nesting sites and nighttime feeding sites, (4) seasonal shifts in movement patterns, (5) use of both perennial and intermittent tributaries adjacent to the capture drainage, and (6) long distance movements in September to new locations, presumably in preparation for hibernation. Mouse locations were categorized by the distance criteria used in the USFWS Proposed 4-D Rule to provide the greatest amount of information to evaluate if the proposed regulations would be sufficient to ensure protection ofPMJM. Given the high proportion of locations within both the proposed buffer zone (150-300 feet) and outside any area of protection (> 300 feet), data from this study do not support the regulations regarding the 300 foot protection zone from the center of the capture stream. Analyses were not completed using the criteria of protection up to 300 feet from the edge of contiguous wetlands because a delineation of contiguous wetlands for any of the three study sites is not yet available. Once these contiguous wetlands are mapped, similar analyses will be conducted using edge of the wetland rather than stream center for perpendicular distance measurements. Eight possible hibernacula were located during this study. Five of the eight mice using these possible hibernacula traveled ~ 90 meters from the center of their typical September night time locations. Vegetation characteristics of the eight sites located during this study are similar to other hibernacula described for PMJM. Natural mortality factors documented during this study included predation by house cats, garter snakes, rattlesnakes, and fox as well as accidents by drowning and road kill. High percentages of arthropods and endogenous fungus were found in the 85 PMJM fecal samples analyzed thus far. The combination of shifts in both general mouse movements, individual mouse movements, and diet provide strong circumstantial evidence that PMJM may be selecting for or require specific seasonal diets. A total of39 sites were surveyed for PMJM in Larimer and Weld Counties, Colorado with a total of 16,671 trap-nights completed. Of the 39 sites survey, Zapus spp. were captured at 21 �2 locations. A total of 71 unique individuals were captured in the 21 locations, with recaptures of 14 of those individuals. All sites where Zapus spp. were captured were located in Larimer County. Three soil samples from 0-12 inches and 12-24 inches were collected at all sites where possible. Soils at some sites were too rocky to collect the 12-24 inch soil samples. Nine fecal samples were collected from traps where Zapus spp. were captured. Initial result indicate that arthropods, fungus, and willows are important dietary components of the Zapus diet. Measurements of habitat characteristics at both the site and landscape levels were completed. Genetic tissue samples were collected from 68 of the 71 jumping mice captured. Analyses of these tissue samples is pending. . A total of 186 individual mice were captured and PIT-tagged at three study sites (73 mice at Maytag Property, 77 mice at PineCliffRanch, and 36 mice at Woodhouse Ranch) during the 1998 field season. These mice were captured over three different trapping sessions with an effort of 17.330 trapnights. Genetic tissue samples were collected for at least 30 individual mice at each of the three study sites. Density estimates were expected to increase as the summer progressed to account for the birth pulses in late June, and late July-August. This was the trend observed at Maytag. PMJM densities at Woodhouse Ranch increased from June to July but did not increase further during the September trapping session. Mean PMJM density over all sites and sessions was 40.5 (range 16.9-79.0) mice per . kilometer of stream stretch. Defining summer as June 1 - October 5, over-summer survival was estimated as 0.36 (se = 0.056) over all three study sites. Temporary emigration and immigration rates were estimated but both had extremely high variances associated with those estimates. Thus, these estimates cannot be used, with any confidence, to provide information on movements of mice into and out of these populations. Very little new information was gained during this study on reproductive parameters ofPMJM. Most adult mice captured exhibited evidence of active reproductive behavior, either pregnancy, lactation, or enlarged genitalia. However, we were unable to locate any breeding nests, Juvenile mice were not captured during the June trapping session. Juvenile mice were captured at all three sites during both the July and September trapping sessions. Preliminary results from the demography, movement and distribution studies ofPMJM have . suggested the need to continue with research currently being addressed in these three studies. However, the preliminary results also suggest further research be conducted on (1) use of upland habitat by PMJM, (2) refining water requirements ofPMJM (i.e., do they require stream habitat or are wetland areas sufficient), (3) range-wide distributional boundaries (e.g., elevation restrictions), and (4) investigating areas of potential sympatry or hibridization with Z. princeps (western jumping mouse). The demography and movement studies will be modified during the summer 1999 field season to further investigate upland use and water requirements. This report includes results from only the first year of a multi-year project to follow individually marked PMJM through time. Collection of more data, and more years of data, will improve our ability to evaluate demographic parameters, movement patterns, and habitat use and evaluate how they vary across space and time. . �3 PREBLES'S MEADOW JUMPING MOUSE CONSERVATION TanyaM. Shenk P.N. OBJECTIVE Develop and implement a conservation plan for Preble's meadow jumping mouse in Colorado. SEGMENT OBJECTIVES 1. Evaluate movement data of Preble's meadow jumping mouse as it relates to landscape features at two study sites. 2. Conduct presence/absence surveys for Preble's meadow jumping mouse at 30 sites in Larimer and Weld Counties, Colorado. 3. Refine and prioritize needed research components to develop sound strategies for conservation of Preble's meadow jumping mouse. 4. Analyze data and prepare a Federal Aid Job Progress Report. Status Report 1, Evaluate movement data of Preble's meadow jumping mouse as it relates to landscape features at two study sites. Movement data ofPMJM were collected on radio-collared mice to evaluate the effects of landscape features on movement patterns. Preliminary analysis of the movement data is presented as a draft manuscript, Movement patterns of Preble's meadow jumping mouse (Zapus hudsonius preblei) as they vary across time and space, in Appendix A. In summary, movement data collected on radio-collared PMJM indicate (1) maximum movements of> 1 mile, (2) greater use of upland habitats than previously assumed, (3) general site fidelity to both daytime nesting sites and nighttime feeding sites, (4) seasonal shifts in movement patterns, (5) use of both perennial and intermittent tributaries adjacent to the capture drainage, and (6) long distance movements in September to new locations, presumably in preparation for hibernation. 2. Conduct presence/absence surveys for Preble's meadow jumping mouse at 30 sites in Larimer and Weld Counties, Colorado. a. Presence/absence surveys Preliminary analysis of the PMJM survey data is presented as a draft manuscript, Habitat use and distribution of Preble's meadow jumping mouse (Zapus hudsonius preblei) in Larimer and Weld Counties, Colorado, in Appendix B. In summary, a total of39 sites were surveyed for PMJM in Larimer and Weld Counties, Colorado with a total of 16,671 trap-nights completed. Of the 39 sites survey, Zapus spp. were �4 captured at 21 locations. A total of 71 unique individuals were captured in the 21 locations, with recaptures of 14 of those individuals. All sites where Zapus spp. were captured were located in Larimer County. Three soil samples from 0-12 inches and 12-24 inches were collected at all sites where possible. Soils at some sites were too rocky to collect the 12-24 inch soil samples. Nine fecal samples were collected from traps where Zapus spp. were captured. Initial result indicate that arthropods, fungus, and willows are important dietary components of the Zapus diet. Measurements of habitat characteristics at both the site and landscape levels were completed. Genetic tissue samples were collected from 68 of the 71 jumping mice captured. <. b. Abundance, over-summer survival, over-winter, annual survival estimates and population age and sex structure of Preble's meadow jumping mouse populations at two study sites. Preliminary analysis of the demography study ofPMJM is presented as a draft manuscript, Temporal and spatial variation in the demography of Preble's meadow jumping mouse (Zapus hudsonius preblei) , in Appendix C. In summary, a total of 186 individual mice were captured and PIT-tagged at three study sites. Mean PMJM density over all sites and sessions was 40.5 (range 16.9-79.0) mice per kilometer of stream stretch. Defining summer as June 1 - October 5, over-summer survival was estimated as 0.36 (se = 0.056) over all three study sites. Temporary emigration and immigration rates were estimated but both had extremely high variances associated with those estimates. Thus, these estimates cannot be used, with any confidence, to provide information on movements of mice into and out of these populations. Very little new information was gained during this study on reproductive parameters of PMJM. Most adult mice captured exhibited evidence of active reproductive behavior, either pregnancy, lactation, or enlarged genitalia. However, we were unable to locate any breeding nests. Juvenile mice were not captured during the June trapping session. Juvenile mice were captured at all three sites during both the July and September trapping sessions, indicating two birth pulses. 4. Refine and prioritize needed research components to develop sound strategies for conservation of Preble's meadow jumping mouse. Preliminary results from the demography, movement and distribution studies ofPMJM have suggested the need to continue with research currently being addressed in these three studies. However, the preliminary results also suggest further research be conducted on (1) use of upland habitat by PMJM, (2) refining water requirements ofPMJM (i.e., do they require stream habitat or are wetland areas sufficient), (3) range-wide distributional boundaries (e.g., elevation restrictions), and (4) investigating areas of potential sympatry or hibridization with Z. princeps (western jumping mouse). The demography and movement studies will be modified during the summer 1999 field season to further investigate upland use and water requirements. 5. Analyze data and prepare a Federal Aid Job Progress Report. A Federal Aid Job Progress Report was submitted to the CDOW on August 16, 1999. �5 ApPENDIX A MOVEMENT PATTERNS OF PREBLE'S MEADOW JUMPING MOUSE (Zapus hudsonius preblei) As THEY VARY ACROSS TIME AND SPACE Tanya Shenk and Maile M. Sivert Abstract Preliminary analysis of the movement data collected on radio-collared Preble's meadow jumping mice (PMJM) indicate (1) maximum movements of> 1 mile, (2) greater use of upland habitats than previously assumed, (3) general site fidelity to both daytime nesting sites and nighttime feeding sites, (4) seasonal shifts in movement patterns, (5) use of both perennial and intermittent tributaries adjacent to the capture drainage, and (6) long distance movements in September to new locations, presumably in preparation for hibernation. Mouse locations were categorized by the distance criteria used in the USFWS Proposed 4-D Rule to provide the greatest amount of information to evaluate if the proposed regulations would be sufficient to ensure protection of PMJM. Given the high proportion of locations within both the proposed buffer zone (150-300 feet) and outside any area of protection (> 300 feet), data from this study do not support the regulations regarding the 300 foot protection zone from the center of the capture stream. Analyses were not completed using the criteria of protection up to 300 feet from the edge of contiguous wetlands because a delineation of contiguous wetlands for any of the three study sites is not yet available. Once these contiguous wetlands are mapped, similar analyses will be conducted using edge of the wetland rather than stream center for perpendicular distance measurements. Eight possible hibernacula were located during this study. Five of the eight mice using these possible hibernacula traveled ~ 90 meters from the center of their typical September night time locations. Vegetation characteristics of the eight sites located during this study are similar to other hibernacula described for PMJM. Natural mortality factors documented during this study included predation by house cats, garter snakes, rattlesnakes, and fox as well as accidents by drowning and road kill. High percentages of arthropods and endogenous fungus were found in the 85 PMlM fecal samples analyzed thus far. The combination of shifts in both general mouse movements, individual mouse movements, and diet provide strong circumstantial evidence that PMlM may be selecting for or require specific seasonal diets. These data are limited to the first year of a multi-year study and thus annual variation in mouse movement patterns cannot yet be addressed. Continuation of this study, at least through the 1999 field season will provide information on annual variation in PMlM movement patterns. This is an interim progress report. Further analyses will be conducted on these data and data collected during the 1999 field season. Introduction Movement and dispersal pattern information will be key to any conservation strategy designed for Preble's meadow jumping mouse (PMlM). Documenting daily and seasonal movement patterns of PMlM will provide information on habitats used by the mice and on the relative configuration and juxtaposition of these habitats. Configuration of habitats include vegetation and size of the areas used ... Juxtaposition includes relative locations of different habitats used such as nest sites, feeding areas, and movement corridors connecting those areas. Key dispersal factors to document for PMlM include (1) which segment of the population disperses, (2) when do they disperse, (3) through what habitat do they disperse, (4) how far will individuals disperse (i.e., what is the maximum distance that separates adjacent populations) and (5) how critical is dispersal (both into and out of a population) to the persistence of a given population. �6 Areas of suitable habitat must provide requirements to survive throughout the life cycle. These requirements must provide necessities for both the active period and hibernation periods. During the active period suitable habitat must provide requirements for daily survival, reproductive activities (breeding, nesting, and rearing of young to independence), and dispersal. The hibernation period requires sufficient food supplies to assure fat storage prior to hibernation and suitable hibernacula Habitat providing all seasonal and life cycle requirements mayor may not occur in a single contiguous area. If not in a contiguous area, habitat patches must occur in a mosaic of usable areas where suitable .corridors exist for seasonal movement among sites. The habitat matrix within the range of Z. h. preble; is mixed grasslands adjacent to the Colorado Front Range along the Piedmont and along the base of the Laramie Mountains in Wyoming and extends to the Colorado plains. Within this matrix, PMlM occur along stream drainages that contain patches of suitable vegetation. Suitable habitat appears to have at least two major components. The first component is a supply of open water, at least in part of the active season (M. Bakeman, C. Meaney, personal communication). Secondly, areas where PMlM has been found have dense cover (M. Bakeman, C. Meaney, personal communication). Based on studies of Z. h. preble; and Z. hudsonius elsewhere, Z. h. preblei apparently occurs mostly in undergrowth consisting of grasses, forbs, or both in open wet meadows and riparian corridors, or where tall shrubs and low trees form an overstory and provide adequate cover (Armstrong et al. 1997). Meadow jumping mice are widespread in abandoned grassy fields, but are often more abundant in thick vegetation along ponds, streams, and marshes or in rank herbaceous vegetation of wooded areas (Whitaker 1963). The mouse does not appear to have an affinity toward any single plant species but instead favors sites that are structurally diverse and provide adequate cover and food throughout its life cycle. PMlM are typically not found in upland areas away from riparian habitats but are most often captured where either ground water becomes visible as either seep springs or as main water channels (M. Bakeman.T. Ryon, personal communication) suggesting a dependence on open water, at least during their active periods. PMlM have been trapped in natural riparian areas as well as areas altered by anthropogenic influence including ditches and wetlands adjacent to interstate highways, cement-lined ditches with tall cover, ditches along driveways and moderate road use, and moderate cattle grazing (M. Bakeman, personal communication). The only currently available data on dispersal and/or movement for Z. h. preble; are from marked mice at Rocky Flats Environmental Technology Site (T. Ryon unpublished data). Two mice, an adult female and an adult male, were observed approximately 1.6 kilometers from previous locations (incidences occurred separately). Each of the locations were in the same drainage (Woman Creek). Movement of Z. h. preblei on Woman Creek at Rocky Flats Technology Site suggests mice move along corridors of shrub cover, generally Salix exigua (PTI 1996a, T. Ryon, unpublished data), suggesting dispersal habitat is similar to habitats used for other activities. IfPMJM occurs as metapopulations in the classical sense of a set of local populations linked by infrequent dispersal then habitat includes not just one area of suitable habitation but also areas suitable for nearby mouse populations. These suitable areas must also be linked by dispersal habitat. If the mice are dependent on dense riparian habitat for dispersal as well as for areas to reproduce, persistence of discrete populations would require a mosaic of suitable discrete riparian patches interconnected with dispersal corridors of similarly dense riparian vegetation. If mouse populations function in a source (populations where growth rate ~ 1) and sink (those populations where growth rate < 1, maintained through immigration) system, it will be critical to identify and protect those populations serving as sources. Thus, for a source-sink population critical habitat will include those areas that support source population, dispersal habitat to sink areas will be less critical. Iflocal mouse populations are functionally discrete, such a mosaic of interconnected areas of suitable habitat would provide a buffer e, �7 for local, and source, populations against deleterious stochastic events by providing the opportunity for local population failures to be 'rescued' by immigration from other populations. To begin to understand PMJM movement, dispersal, and habitat use we monitored radiocollared mice at three different study areas. Study areas were selected to cover a variety of habitat configurations to evaluate spatial variation in movement patterns ofPMJM. To address temporal variation in PMJM movement, dispersal, and habitat use we willcontinue this study for at least one more field season at the same three field sites. This is an interim report, after one field season of data collection. Study sites To evaluate spatial differences in movements ofPMJM, the three sites selected provided a variety of habitat matrices available to the mouse. The first site selected, the Maytag Property, has one primary water source available to the mouse. This water source is East Plum Creek. Therefore, we predicted mice would restrict their movements to up and down this single drainage. The second site, PineCliffRanch, has both a tributary (Garber Creek) and a main stem drainage (West Plum Creek). This provided the opportunity to investigate whether PMJM will use upland areas to move from one drainage to another or if they are restricted to only moving along riparian corridors. The third study site, Woodhouse Ranch, provided an area with a tributary (Indian Creek) and a series of ponds and irrigation ditches within 0.5 kilometers of the drainage. The configuration at Woodhouse Ranch provided an even greater opportunity to investigate how much the mice use upland areas or if they restrict their movements strictly to riparian corridors. By replicating the same methodologies at each of these three unique habitat matrices we can begin to estimate spatial variation in nightly and seasonal movements ofPMJM. Objectives This PMJM movemerit study is designed to describe nightly and seasonal movement patterns ofPMJM and to describe habitats used by PMJM. These movement patterns will be described for three different study areas to evaluate spatial variation, and over two different years at the same three study areas to evaluate temporal variation. Specific objectives include: 1. Describe nightly movements ofPMJM. Evaluate difference in nightly movements as they relate to sex, age, and habitat available to the animals. l. Describe 30-day (or life of the radio transmitter) interval movements ofPMJM. Evaluate difference in 30-day (or life of the radio transmitter) interval movements as they relate to sex, age, and habitat available to the animals. 2. Describe seasonal movements ofPMJM. Evaluate difference in seasonal movements as they relate to sex, age, and habitat available to the animals. 3. Describe habitats where mice occur: movement corridors, end point descriptions (i.e., movement from what to what), and landscape features (connectivity with other riparian areas and other habitats used). 4. Estimate the mean amount of time PMJM spend in each available habitat. 5. Evaluate spatial and temporal variation in movement patterns ofPMJM. The objective of the habitat use study is to identify and refine habitat requirements of Z. h. preblei, including hibernation sites, and to determine if they influenced any of the demographic parameters that will be estimated in this and complementary studies (see Shenk and Sivert 1999). �8 Methods Movement data were documented primarily from locations of radio-collared mice from each of the three study populations. To place radio transmitters on the animals, mice were captured in Sherman live traps. Mice weighing> 18 grams were fitted with either :MD-2C, I-gram radio transmitters supplied by Holohill Systems Ltd. (used successfully on PMlM by R Schorr, personal communication) or I-gram radio transmitters supplied by Advanced Telemetry Systems (ATS). Three trapping sessions were conducted during the following weeks: June 2-9, 1998; July 21Aug 6, 1998; and September 8-15, 1998. Trappers were advised to follow the Center for Disease Control's Hantavirus instructions and recommendations when dealing with rodents. Due to the nocturnal nature ofPMlM, traps were set between 19:00hrs and 21:OOhrsand checked as early as possible in the morning beginning at 5:00hrs (to reduce stress and the potential for predation on trapped animals). Time required to complete the trap lines varied depending on how many animals were caught. Small mammal Sherman live traps (folding and non-folding) were used to conduct the trapping sessions. Traps were set in two parallel lines of trap stations (1 trap per station) on either side of the drainage. Trap stations were 5 meters apart for a total of 250 meters; the parallel transects were 10 meters apart unless extent of habitat, terrain topography, or stream hydrology do not allow. Location of transects were recorded to the nearest 5 meters in UTM coordinates using a Trimble Geo-Explorer GPS. A small (~1 inch) ball of polyester quilt material was placed in each trap as nestinglbedding material. Baiting material was Manna Pro Sweet 3-way Livestock feed which contains no animal matter; ingredients include flaked barley, flaked corn, flaked oats, and cane molasses. Peanut butter was used to stick the bait to the trap. Traps were checked by two surveyors. All animal captures were recorded. If an animal was captured in a trap, a ziplock plastic bag was placed over the end of the trap. The trap was opened allowing the animal to fall into the plastic bag. The animal was identified to species while in the plastic bag. If the animal was not a PMlM, identification of the animal was recorded and the animal set free. Each captured PMlM was checked to detect the presence/absence of a PIT-tag andlor radio-collar. If a PIT -tag or radio-collar was detected and the mouse had been captured at that same site within the same trapping session, identification of the mouse was recorded and the mouse was released. If the animal was a PMJM and no PIT-tag or radio collar was detected the PMJM was anesthetized for further processing, as follows. All PMlM were PIT-tagged for a complementary demography study on PMJM at these same three study sites (see Shenk and Sivert 1999 for details). If the PMlM weighed> 18 g a radio-collar was also put on the animal. Each PMJM was weighed while in the bag, the weight recorded in grams using a Pesola spring balance. Sex, age (juvenile, adult), and reproductive condition (pregnant, lactating or non-breeding if it is female; for males the position of the testes indicated if in breeding status or non-breeding status) was noted. Each PMJM was measured (total length, length of body, length of hind foot - heel to distal end of claws) in millimeters. Mice were placed in a plexiglass photobox to reduce handling stress while taking the photograph. If the mouse had been captured previously as noted by the detection of a PITtag, no photograph was taken. If there were feces in the trap where a PMJM was captured, the fecal material was collected in a plastic bag labeled with date, location of the site, and animal PIT-tag number. Fecal samples were kept cool until returned to the CDOW office where they were frozen. Salt was added to each specimen bag for preservation. Once the field season was over all fecal samples were analyzed by the Composition Analysis Laboratory, Inc, 622 Y2 Whedbee, Fort Collins, CO for content. All trap mortalities were recorded on a 'Trap Mortality' data form which included information on species, potential duration of time spent in the trap, and any information available to help determine cause of death. All animals found dead were double-bagged in a plastic bag and placed in a cooler with ice. Specimens were frozen as soon as possible and deposited in the CDOW freezer at the office . " .. : �9 in Fort Collins. A museum card was completed and attached to each PMlM specimen and the specimen and card were given to Cheri Jones, Curator of Mammalogy at the Denver Museum of Natural History, for study skins and tissue storage. If an animal was severely injured (e.g., severed limb, large lacerations) it was euthanized by soaking a cotton ball in Metofane (methoxyflurane) and placing the cotton ball and the mouse in a ziplock bag until the animal stopped breathing. If an animal appeared to be only slightly injured (e.g., broken tail, small laceration) the animal was released to the wild. If an animal appeared to be coldstressed attempts were made to warm it by holding it in the surveyors hands and/or against their body. If the animal appeared to be heat stressed, isopropyl alcohol was applied with a cotton swab to the ears, arm pits, and feet to cool it down. During the last trapping session homeopathic first aid treatments were used in an attempt to minimize stress and improve recovery of each PMlM. Homeopathic first aid kits and instruction was provided by WildAgain Wildlife Rehabilitation, Evergreen, Colorado. PIT-tagging and radio-collaring Each PMlM was individually marked with a Passive Integrated Transponder (PIT-tag). PITtags are electromagnetic, glass-encased tags that emit a passive signal (125 kHz) that can be decoded by a portable reader. Destron-Fearing PIT-tags were used on all mice. We used portable readers from Biomark to read the PIT-tags. All mice were anesthetized to PIT-tag them. Anesthesia procedures followed protocols successfully used on PMlM before (R. Schorr personal communication). Mice were anesthetized by placing them and a cotton ball with 1 rnl of Metofane (methoxyflurane) into a sealed ziplock bag (to keep Metofane fumes in bag). The bag was then lain aside to minimize stress for the mouse and the time the mouse struggles in the bag. The less agitated the mouse is in the bag the less time required for the metofane to take affect. The mouse was observed at all times to make sure the mouse did not position itself such that the cotton ball was in direct contact with its face, which might cause the mouse to have a severe reaction to the anesthesia. After the animal stopped moving, the handlers waited one minute before removing the mouse from the bag. The animal typically remained anesthetized for 2-3 minutes once outside the bag. Each PMlM was PIT-tagged using a protocol successfully used by C. Meaney (personal communication). Each PMJM had a PIT-tag inserted above the shoulder blades by lifting the skin on its back and inserting the individually sterilized needle with the PIT-tag under their skin and injecting the tag. Verification of the PIT-tag identification number was made before and after insertion into the mouse by running the PIT-tag scanner across it. The skin behind the opening was then pinched to prevent emergence of the tag. PIT-tag identification number was recorded on the field data form. Each radio transmitter was checked to ensure proper functioning before collaring the animal with the transmitter by tuning the receiver to the transmitter frequency to make sure there was a signal. The radio antenna was fed through a small piece of plastic tubing and then back through the radio to make a collar of the plastic coated antenna. The loop made by the antenna was kept large enough to fit over the mouse's head. Once over the neck of the animal, the antenna was pulled to reduce collar size, making sure rubber tubing was on the dorsal side of the neck. The purpose of the rubber tubing was to prevent damage to the collar and prevent the person attaching the collar from cinching the collar too tight. The collar was tested to be sure it could rotate freely around the animals' neck, but was not loose enough to be removed. The metal crimp was flattened to hold the antenna in the desired collar position. Approximately two inches of antenna was left to extend out behind the animal. If, at any time during the handling of the PMlM the animal appeared to be severely stressed (dramatic changes in heartbeat, respiration and responsiveness or gums turning blue) first aid was administered and, once recovered, released without further processing. �10 Movement data Once transmitters were in place, locations of individual mice were made nightly. Because very little is known about the movements ofPMlM pilot protocols were established and then modified. Final protocols included tracking up to eight individual mice per person per night. A minimum of six locations per individual mouse per night were made. Each individual mouse was tracked a minimum ~ of three nights per week. The six locations per night per individual were scattered throughout the night to maximize information learned about nightly movements (i.e., eight locations taken within a single hour contains less information than eight locations taken one each hour). Each location was determined by locating the mouse and recording the position as a file in a Trimble Geo-Explorer GPS. Each GPS location (file) was differentially corrected for 2-5 meter accuracy. Daytime locations were taken as time permitted. To describe nightly movements the following parameters were estimated: (1) minimum and maximum distances moved each night animal was followed, (2) mean minimum and maximum distances moved each night for all mice, (3) minimum and maximum distances moved away from the stream center each night by individual mice, and (4) mean minimum and maximum distances moved away from the stream center each night for all mice. To describe 30-day (or life of the radio transmitter) movements the following parameters were estimated:(I) minimum and maximum distances moved each 30-day (or life of the radio transmitter) interval animal was followed, (2) mean minimum and maximum distances moved each 30-day (or life of the radio transmitter) for all mice, (3) minimum and maximum distances moved away from the stream center each 30-day (or life of the radio transmitter) interval by individual mice, (4) mean minimum and maximum distances moved away from the stream center each 30-day (or life of the radio transmitter) interval for all mice. Mortality factors for any mouse found dead was recorded Both site specific and landscape features were recorded at each of the three population study sites to document the extent of spatial variation. These quantitative measures of spatial variation were used in the analyses to determine if they influenced any of the movement parameters estimated in this study. The following habitat characteristics were measured and recorded for each site. Site characteristics Cover was measured at 20 random locations along the 250 meter sampled stream stretch. Mean cover and standard error of the mean was estimated. Cover was estimated using a vegetation profile board (Nudds 1977) that allowed for an assessment of visual obstruction in 0.5 meter vertical intervals above ground. The board was 2.0 meters high and 30.48 cm wide. The board was marked in alternate black and white colors at 0.5 meter intervals. Horizontal cover was assessed in each interval by viewing the board from 15 meters away in a randomly chosen direction. The percentage of each interval concealed by vegetation was recorded as 0-20, 21-40, 41-60, 61-80, and 81-100% estimated concealment. Vegetation species composition and richness was estimated using the Modified-Whittaker nested vegetation sampling method (Stohlgren et al. 1995). A modified Whittaker plot is 20-meters x 50-meters with 101m2 and 210m2 subplots arranged systematically around the perimeter and 1 100m2 subplot centered in the inside of the 20 meter by 50 meter plot. Species composition and percent cover of each species was recorded for each subplot. Soil samples were taken at each site for a hydrometer textural analysis (percent sand, silt and clay). Samples were collected using a soil probe. Samples were taken from 0-12 inches, and 12+ to 24 inches. The 0-12 inch sample was taken first, removing the probe and soil. Samples were placed in a labeled ziplock bag. The probe was then placed in the same hole for the 12+ - 24 inch probe with that soil sample being placed in a separately labeled ziplock bag. The hydrometer textural analysis was conducted at the Colorado State University Soils Laboratory. ~ '. .' �11 Mean stream width was estimated by measuring stream width at 30 locations along the entire site sampled. The 30 locations were stratified equidistant from each other to cover the entire stream stretch in increments of site stream length/30. Each nest site was described including distance from the stream and vegetation. Landscape characteristics The following landscape habitat characteristics were measured using either GIS, topographic maps, or aerial infrared photography. 1. Distance to nearest human habitation and human disturbance. 2. Connectivity of sites to other streams. 3. Total length of stream stretch at each site and total length of stream stretch with suitable habitat at each site. Hibernation site measurements Hibemation sites were identified by following radio-collared mice in late September and early October. Once a radio-signal remained stationary for 2-10 nights we assumed we located a hibernaculum. The following measurements were taken at each hibernation site: 1. 2. 3. 4. Distance ofhibernaculum to stream. Vertical elevation gain from stream to hibernaculum. A soil sample was taken as described above. Vegetation species richness within a 5-m radius. Density of vegetation was measured as described above. Results PIT-tagging and radio-tracking effort A total of 186 individual mice were captured and PIT-tagged at the three study sites (Table 1: 73 mice at Maytag Property, 77 mice at PineCliffRanch, and 36 mice at Woodhouse Ranch). These mice were captured over three different trapping session with an effort of 17,330 trap-nights. A total of 125 radio-collars were put on mice over the three trapping sessions (Table 1: 48 at Maytag property, 47 at PineCliffRanch, and 30 at Woodhouse Ranch). A total of62 females and 63 males were radiocollared over the field season. Initial protocols called for using triangulation methods (simultaneous compass bearing readings from three different locations) to determine mouse locations. Triangulation was used during the month of June. Locations were estimated in late June from these data and found to have large variances associated with most locations. Field protocols were changed for the latter two tracking sessions to walk-up locations. A walk-up location required technicians to use the signal strength indicator on the receiver to determine when they were within 2-7 meters of the mouse. GPS location files were collected each time a mouse was located. File name was recorded as well as the UTM coordinates displayed on the GPS. These files were later differentially corrected to provide a positional accuracy of 2-5 meters. Due to the large variances on the mouse locations collected during June these locations are not reported here. Further analyses will be conducted on these data to see if any information on PMJM movements can be gained. The walk-up method was first evaluated to ensure we were not causing the mice to move in response to us. Given the consistency of individual mouse locations throughout the night and on a day to day basis we felt the method was being employed in such a way that we were not affecting mouse movement patterns. However, it should be noted that T. Ryon (personal communication) observed mice moving in response to him when he tried to conduct similar walk-up locations. �12 Daily movements Percent of PMlM locations, as determined from radio-tracking, at each of three distance categories were calculated for each study site and the latter two radio-tracking sessions (Table 2). The majority of locations at Maytag, PineCliff, and Woodhouse were within 46m of stream center for both tracking periods (July-August and September-October). For all areas and tracking periods except Maytag during the last tracking session, 90% of PMlM movements were within 91m of either side of the center of the capture drainage. Mouse movements at Maytag during September-October resulted in 32.6% of the locations being> 91m from the stream center. Analyses were also conducted on mice captured on PineCliffRanch separating mice by drainage of initial capture. These analyses were conducted to address questions that might arise when connecting drainagesoccur in areas known to have PMlM, but only one drainage has been surveyed. Mice initially captured on Garber Creek showed similar movement patterns as far as distance from center of capture drainage (i.e., Garber Creek) as mice from both Maytag and Woodhouse (Table 2). Mice initially captured on West Plum Creek showed movements more often> 91m from center of the capture drainage (Table 2). Seasonal movement The distribution of mouse locations at Maytag Property for the July-August tracking session is different from the distribution ofPMlM locations for the September-October tracking session. The pattern shifts from heavy use along East Plum Creek to the north of the trapline to a more concentrated distribution either side of the trap line. The more concentrated use of the areas east and west of the trapline also resulted in locations being further from the center of the creek. The northern end of the trapline is largely vegetated by willows with the remainder of the trapline more diversely vegetated. Mouse movements were more concentrated along the drainages at PineCliff Ranch during the September-October tracking sessions than during the July-August tracking sessions. Vegetation along both West Plum Creek and Garber Creek along the trapline was primarily willow. At Woodhouse Ranch, the distribution of mouse locations also shifted between tracking sessions. Mouse movements were more concentrated along the southern half of the trapline in September-October as compared to the more even distribution of mouse locations recorded for JulyAugust. The northern end of the trap line is dominated by willows, the middle and southern end of the trapline is in more diverse riparian habitat. Eight mice were captured, radio-collared, and tracked for two tracking sessions, one mouse was captured, radio-collared, and tracked for all three tracking sessions (Table 3). Other mice were captured during more than one trapping session but radio collars were not replaced either because all the radio collars had been put on other mice or because the physical condition of the animal was such that we decided not to replace the radio-collar. Such conditions included mice who had significant hair worn off the neck from the previous radio collar. In no incidence did we find open wounds caused by radio collars. Of the nine mice radio-collared for more than one session, four provided information to evaluate seasonal changes in areas and habitats use between the July-August tracking period and the September-October tracking period. One mouse, a male from Maytag was observed only once during the September-October tracking session, providing no information concerning seasonal changes in movement patterns. Movement data from June have not been fully analyzed and are not included in this report. Thus, changes in areas used by individual mice between June and either the July-August or September-October tracking session are not summarized here. A female at Maytag exhibited a seasonal movement shift, small sample size prohibited any evaluation of the movement patterns of the Maytag male (n = 1 in September-October). There does not appear to be any shift in areas used by the male tracked during both latter sessions at Pine Cliff Ranch, .. " �13 however, sample sizes were small (n = 17, 14). Two females at Woodhouse Ranch exhibited a shift in areas used between July-August and September-October. However, caution should be used in interpreting these results because of low sample sizes in the latter tracking session (n = 14,6). Nest sites Numerous daytime nest sites were located at all thee study sites. Nest sites were made of tightly woven vegetation, located on the ground. Nest material included leaves, grass, and small sticks. There was only one small entrance hole on each of the nests found. When observing mice in their nests the mice would peer out of the hole and remain motionless as long as we were present. No breeding nests were located, however there were several locations at each of the study sites where adult females returned to repeatedly. Each of these sites were in patches of extremely dense vegetation or underneath a large downed log. We did not disturb these areas because we did not want to cause failure of the nest. Hibemacula Eight potential hibernacula were located, at least one at each study site (Table 4). Mortality factors Mortalities factors of radio-collared mice included predation by rattlesnake, garter snake, fox, and house cat (Table 5). Four probable predations were also noted and identified by finding tightly crimped (i.e., no possibility of the mouse having slipped the collar over its head) radio-collars lying on the ground. Two accidental deaths were documented, a road kill and drowning. Mortality factors also included trapping and/or handling mortalities and unknown causes. Fecal analysis The amount of bait found in each of the fecal samples was quantified as either 100%, abundant « 10%) or 0%. The following summaries are made after eliminating all samples of 100% bait, and then only classifying the fecal sample contents other than bait. Fecal analyses indicate a seasonal shift in diets. The most common item found in the fecal samples during the June trapping session at all three sites were arthropods. This was true for three of the five June samples collected at Maytag (Table 6), eight of ten June samples at PineCliff (Table 7), and four of five June samples collected at Woodhouse (Table 8). Other common items included endogenous fungus (1) and seed (1) at Maytag; endogenous fungus (2) at PineCIiff,; and Poa (1) at Woodhouse. During the July 21-August 4 trapping session the most common items in the fecal samples at Maytag were arthropod (2), endogenous fungus (1), pollen (1), and Carex (1). During the same trapping period, the most common items in five of eight samples collected at PineCliff were endogenous fungus, the other three samples having a majority of arthropod (1), moss (1), and pollen (1). The two samples from Woodhouse during this trapping session were composed primarily of either mushroom or seed. On August 13, nine samples were collected during an extra trapping session (on the same trapline as the other trapping sessions occur) with the most common items in the fecal samples being moss (4), endogenous fungus (3) or pollen (2). All three samples collected from PMJM captured at a back drainage on August 23rd at Woodhouse Ranch were primarily fungus. The September trapping session at Maytag yielded samples with majority fecal contents of arthropod (2), moss (2), pollen (2), endogenous fungus (1), and seed (1). Ten samples from PineCliff during September had majority fecal contents of arthropod (3), endogenous fungus (3), and seed (3). Eight of nine samples taken from Woodhouse contained primarily arthropods, with one a trace of seed. (> 70%), trace �14 Discussion Preliminary analysis of the movement data collected on radio-collared Preble's meadow jumping mice (PMJM) indicate (1) maximum movements of> 1 mile> (2) greater use of upland habitats than previously assumed, (3) general site fidelity to both daytime nesting sites and nighttime feeding sites, (4) seasonal shifts in movement patterns, (5) use of both perennial and intermittent tributaries adjacent to the capture drainage, and (6) long distance <movements in September to new locations, presumably in preparation for hibernation. Initial evaluation ofPMJM movement data focused on maximum distances mice moved perpendicular to the capture drainage. This focus resulted from criteria used in the USFWS Draft 4-D Ruling (1998) to provide special regulations for the mouse. The 4-D Rule defines a Mouse Protection Area (MPA) as "extending 300 feet on each side of the stream measured from the centerline, or 300 feet from the exterior boundary of any contiguous wetlands, whichever is further." The 4-D Rule also states "the basis for the 300-foot standard is that mice have been documented to regularly move up to 150 feet from streams and wetlands. The remaining ISO-foot zone serves as a buffer zone to avoid disturbance ofPMJM habitat associated with human activities. We believe that this zone will encompass the normal home range of the Preble's and will provide an adequate buffer from adjoining development. " Mouse locations were categorized by the distance criteria used in the 4-D Rule to provide the greatest amount of information to evaluate if the proposed regulations would be sufficient to ensure protection ofPMJM. Given the high proportion of locations within both the buffer zone (150-300 feet) and outside any area of protection (> 300 feet), data from this study do not support the regulations regarding the 300 foot buffer from the center of the capture stream. Analyses were not completed using the criteria of protection up to 300 feet from the edge of contiguous wetlands because a delineation of contiguous wetlands for any of the three study sites is not yet available. Once these contiguous wetlands are mapped, similar analyses will be conducted using edge of the wetland rather than stream center for perpendicular distance measurements. To address the issue of whether or not connected but untrapped tributaries should be included in the designation ofMPA's, the more detailed analyses of mouse movements at PineCliffRanch were conducted. The issue of whether or not Garber Creek, a tributary of West Plum Creek, would be included in that particular MP A is not at issue since both the main drainage, West Plum Creek and the tributary, Garber Creek have been trapped in that area with mice captured on both drainages. Rather, the analysis was conducted to test how well the proposed regulation to protect only 300 foot from the center of the capture stream would have 'encompassed the normal home range of the Preble's and will provide an adequate buffer from adjoining development." It is clear from the results that the regulation would have failed to accomplish its objective, with up to 47% of the mouse locations falling beyond the 300 foot demarcation when only the capture drainage was considered. Use of tributaries was also an issue at the Maytag Property. Two mice moved away from East Plum Creek in September and focused their movements ~300 meters from their previous locations, up a dry drainage dominated by upland grasses and gambel oak. These two mice continued to use this new area and were not observed again near East Plum Creek for at least two weeks. Normal radiofailure (i.e., batteries failed after ~4 weeks) after this time did not allow us to determine if the mice ever returned to East Plum Creek or if they hibernated in their new location. No tributaries were available to mice at Woodhouse Ranch. In summary, movement patterns of mice at the Maytag Property suggest use of tributaries, and at PineCliffRanch extensive use of a tributary. Jumping mice of the genus Zapus are true hibernators, spending much of their lives in hibernation. Meadow jumping mice spend approximately 7 months (~210 days) per year in hibernation (Quimby 1951) whereas estimates for Z. princeps indicate that some populations (e.g., in the western mountains of Utah) spend up to 300 days per year in hibernation (Cranford 1983). �15 Jumping mice hibernate in underground burrows (Quimby 1951, Whitaker 1963). They are excellent burrowers and create their own hibernacula Meadow jumping mice are generally solitary hibernators, however, there have been occurrences of more than one mouse found in a single hibernaculum. Eight possible hibernacula were located during this study. The greatest perpendicular distance from the center of the main drainage at the site (East Plum Creek for Maytag Property, West Plum Creek for PineCliffRanch, and Indian Creek for Woodhouse Ranch) was 341 meters. However, if tributaries were considered, greatest distance from center of the nearest drainage to any of the eight sites. was 78 meters. Five of the eight mice using these possible hibernacula traveled ~ 90 meters from the center of their typical September night time locations. One mouse, a female at Maytag moved 750 meters to a possible hibernacula Vegetation characteristics of the eight sites located during this study (Table 4) are similar to other hibernacula described for PMJM. One confirmed hibernaculum, located on Rocky Flats Environmental Technology Site, used by Z. h. preblei has been located (Armstrong et al. 1997). This site was 9m above a creek bed (Walnut Creek); it had a thick cover of chokecherry (Prunus virginiana) and snowberry (Symphoricarpos spp.), the mouse was found in a leaf litter nest 30cm beneath the ground in coarse textured soil (Armstrong et al. 1997). Four possible hibernacula were located by tracking radio-telemetered mice at the U. S. Air Force Academy in fall 1997. These sites are located 7, 12,29, and 31m from a creek bed (R. Schorr, unpublished data). There was no consistency among sites in aspect. Three sites were in vegetation dominated by coyote willow (Salix exigua), one site was in vegetation dominated by snowberry and mullein (Verbascum thapsus). However, all four hibernacula appeared to be below coyote willows. The eight sites located during this study and the four U. S. Air Force Academy sites were not disturbed to protect any hibernating mice and therefore are only possible hibernacula because there is no confirmation a mouse actually hibernated there. Confirmation of a true hibernaculum cannot be made until a chamber, or nest is located. The other explanation for these collar locations might be either locations of radios discarded by the mice or dead mice carried underground by a predator. Location or more hibernation sites was limited primarily by normal battery failure of the radio transmitters before mice went into hibernation. Prior to the 1998 field season, natural mortality factors reported for Z. hudsonius included only insufficient fat storage prior to hibernation (Whitaker 1963), predation (Whitaker 1963, Poly and Boucher 1997, R. Schorr unpublished data) and cannibalism (Sheldon 1934). Other assumed natural mortality factors for Z. h. preblei included starvation, exposure, and disease. This study further reports confirmation of predation by house cats, garter snakes, rattlesnakes, and fox. Other documented natural mortality factors now include accidents by drowning and road kill. The high percentage of arthropods and endogenous fungus found in the fecal samples provides a new aspect to evaluating PMJM habitat requirements. When considering habitats used by PMJM, or in trying to predict habitats that might be suitable for the subspecies, consideration should be given as to whether those habitats could support the arthropods and fungus apparently being selected for by the mouse. Arthropods are a food source high in protein and fat which would benefit mice emerging from hibernation and mice preparing for hibernation. Whitaker (1963) reported a 67% loss of individuals over hibernation and that average body mass of individuals emerging from hibernation was greater than the average for mice entering hibernation. Because no mice are known to store food in their hibernacula, this indicates that the lighter individuals died during hibernation and only those entering with higher masses survived. All the energy they use during hibernation and the periodic arousals (the energetically most expensive part of hibernation) must be the fat they carry into hibernation (B. Wunder, personal communication). The ability to put on sufficient fat for overwinter survival during hibernation is a critical factor in the life history of these mice. Thus, appropriate and sufficient food sources must be available to the mouse to meet these nutritional requirements. �16 The apparent seasonal shift in mouse movements between the July-August and September tracking sessions may be a result of diet switching. The broader diets suggested by the fecal analyses from samples collected during July and August possibly represent both a wider availability of suitable foods and the ability ofPMlM to exploit these resources. It might also suggest a need to exploit these other resources to provide the mouse with necessary food requirements for breeding. Because mice tracked during each session were not generally the same mice there is a possibility these apparent movement shifts might only be different areas used by different mice. The probability of this being the case is small for two reasons. The first is the low density of mice in the stream (Shenk and Sivert 1999). Given that, the proportion of mice followed during each session is a high proportion of the mice in the population. Thus, each subset of mice should be representative of the population. Secondly, evidence from the four mice followed during both the latter tracking sessions also exhibited movement shifts between sessions. The combination of shifts in both general mouse movements, individual mouse movements, and diet provide strong circumstantial evidence that PMlM may be selecting for or require specific seasonal diets. If this is the case, all these requirements must be considered and provided for to ensure conservation of the subspecies. A detailed literature search and further studies need to be conducted to investigate the possible implications of these dietary patterns. These data are limited to the first year of a multi-year study and thus annual variation in mouse movement patterns cannot yet be addressed. Continuation of this study, at least through the 1999 field season will provide information on annual variation in PMlM movement patterns. This study looked at PMJM movements at only three sites. These sites were selected based on known presence ofPMlM. Other sites, perhaps because of different habitat configurations or sites of poorer quality may require mice to move further from the creek or may allow mice to remain closer to the creek in order to obtain all life requirements. However, these study sites were specifically selected to address spatial variation in movement patterns ofPMlM due to different spatial configuration and juxtaposition of habitats. And although all study sites were different they could all be considered within the general description of what is consider typical habitat for PMlM. It should also be noted that we generally trapped along the creek. Thus, mice captured and subsequently radio-collared and followed were those mice that use the area near the most prominent drainage. If there are mice that do not regularly use the main channel and thus were not available for capture there would be a bias in the data towards fewer observations 300 feet away from the creek. To test for such a bias, further research needs to be conducted trying to capture mice further from the creek and compare their movements and locations of those movements in relation to distance from the creek. We plan to test for this bias during the 1999 field season at Maytag Property and Woodhouse Ranch. This is an interim progress report of a multi-year study. Further analyses will be conducted on these data and data collected during the 1999 field season. Literature Cited Armstrong, D. M., M. E. Bakeman, A. Deans, C. A. Meaney, and T. R. Ryon. 1997. Conclusions and recommendations in: Report on habitat findings on the Preble's meadow jumping mouse. Edited by M. E. Bakeman. Report to USFWS and Colorado Division of Wildlife. Bailey, B. 1929. Mammals of Sherburne County, Minnesota. Journal ofMammalogy 10:153-164. Bailey, V. 1923. Mammals of the District of Columbia Proceedings of the Biological Society. Washington 36:103-138. Cook, T. D. and D. C. Campbell. 1979. Quasi-experimentation: design and analysis issues for field settings. Houghton-Mifflin, Boston. �17 Cormack, R M. 1964. Estimates of survival from the sightings of marked animals. Biometrika 51:429-438. ERO Resources. 1995. Environmental review of South Boulder Creek Management Area Prepared for City of Boulder Real Estate/Open Space Department. Prepared by ERO Resources Crp., Denver, Colorado in association with Stoecker Ecological Consultants, Boulder, Colorado. Fitzgerald, J. P., C. A. Meaney, and D. M. Armstrong. 1994. Mammals of Colorado. Denver Museum of Natural History, University Press of Colorado. Niwot, Colorado. Hafner, D. J., K. E. Petersen, and T. L. Yates. 1981. Evolutionary relationships of jumping mice (genus Zapus) of the southwestern United States. Journal ofMammalogy 62:501-512. Hall, E. R 1981. The mammals of North America. John Wiley and Sons, Inc., New York, New York, 2 volumes. Hamilton, W. J., Jr. 1935. Habits of jumping mice. American Midland Naturalist 16:187-200. Jolly, G. M. 1965. Explicit estimates from capture-recapture data with both death and immigration stochastic model. Biometrika 52:225-247. Jones, C. A. 1996. Mammals of the James John and Lake Dorothey State Wildlife Areas. Final Report, submitted to the Colorado Division of Wildlife and Colorado Natural Areas Program. Kendall, W. L., and J. D. Nichols. 1995. On the use of secondary capture-recapture samples to estimate temporary emigration and breeding proportions. Journal of Applied Statistics.22:751762. Kendall, W. L., K. H. Pollock, and C. Brownie. 1995. A likelihood-based approach to capturerecapture estimation of demographic parameters under the robust design. Biometrics 51:293308. Kendall, W. L., J. D. Nichols, and J. E. Hines. 1997. Estimating temporary emigration using capturerecapture data with Pollock's robust design. Ecology 78:563-578. Krutzsch, P. H. 1954. North American jumping mice (genus Zapus). University of Kansas Publications, Museum of Natural History 7:349-472. Meaney, C. A., N. W. Clippinger, A. Deans, and M. OShea-Stone. 1996. Second year survey for Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. Report prepared for the Colorado Division of Wildlife. Meaney, C. A., A. Deans, N. W. Clippinger, M. Rider, N. Daly, and M. O'Shea-Stone. 1997. Third year survey for Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. Report prepared for the Colorado Division of Wildlife. Nudds, T. D. 1977. Quantifying the vegetative structure of wildlife cover. Wildlife Society Bulletin 5:113-117. Poly, W. J., and C. E. Boucher. 1997. Record ofa creek chub preying on ajumping mouse in Bruffey Creek, West Virginia. Brimleyana 24: 29-32. PTI Environmental Services. 1996a. Preble's Meadow Jumping Mouse Study at Rocky Flats Environmental Technology Site, Annual Report 1996. Final. Rocky Flats Environmental Technology Site, Golden, Colorado. PTI Environmental Services. 1996b. Preble's Meadow Jumping Mouse Study at Rocky Flats Environmental Technology Site, Spring 1996. Final. Rocky Flats Environmental Technology Site, Golden, Colorado. Quimby, D. C. 1951. The life history and ecology of the jumping mouse, Zapus hudsonius. Ecological Monographs 21:61-95. Riggs, L. A., J. M. Dempey, and C. Orrego. 1997. Evaluating distinctness and evolutionary significance of Preble's meadow jumping mouse: Phylogeography of mitochondrial DNA noncoding region variation. Final Report for the Colorado Division of Wildlife. Denver, Colorado. �18 Seber, G. A. F. 1965. A note on the multiple recapture census. Biometrika 52:249-259. Sheldon, C. 1934. Studies on the life histories of Zapus and Napaeozapus in Nova Scotia Journal of Mammalogy 15:290-300. Shenk, T. M. 1998. Conservation assessment and preliminary conservation strategy for Preble's meadow jumping mouse (Zapus hudsonius preblei). Colorado Division of Wildlife FY 199798 Armual Report. Shenk, T. M. and M. M. Sivert. 1999. Temporal and spatial variation in the demography of Preble's meadow jumping mouse (Zapus hudsonius preblei). Colorado Division of Wildlife JanuaryMarch 1999 Quarterly Report. Stohlgren,T. J., M. B. Falkner, and L. D. Schell. 1995. A Modified-Whittaker nested vegetation sampling method. Vegetatio 117:113-121. USFWS. 1997. Interim survey guidelines for Preble's meadow jumping mouse. USFWS. Denver, Colorado. Whitaker, J. 0., Jr. 1963. A study of the meadow jumping mouse, Zapus hudsonius (Zimmerman), in cental N ew York. Ecological Monographs 33:3. e . Table 1. Number of Preble's meadow jumping mice captured and radio-collared at each study site during each trapping session. # Site Trapping session Trap nights female Maytag June 1949 Maytag July Maytag CaQturedt # Radio-collared male (%) total female male total 9 (64) 5 (36) 14 9 5 14 2370 15(54) I3 (46) 28 11 5 16 September 3256 18(43) 24 (57) 42 10 8 18 PineC1ifT June 1095 6 (32) 13(68) 19 5 I3 18 PineC1ifT July 1603 8 (50) 8 (50) 16 6 7 13 PineClifT September 1544 13(28) 33(72) 46 4 12 16 Woodhouse June 1525 4 (57) 3 (43) 7 2 3 5 Woodhouse July 2420 9 (64) 5(36) 14 8 4 12 Woodhouse September 1568 8 (44) 10(56) 18 7 6 I3 17330 90 114 204 62 63 125 TOTALS (%) t SomecapturedanimalswerePIT-taggedduringprevioustrappingsessions �19 Table 2. Percent locations of Preble's meadow jumping mice at each of three distance categories for each study site and radio-tracking session. PineCliffRanch locations are also divided into percent locations for each distance category from center of the stream of initial capture for mice captured on either West Plum Creek or Garber Creek. % locations at each distance from center of main Site Tracking period N Maytag Jul-Aug 1998 Maytag <46m (150 feet) 46 - 91 m 150-300 feet >91 m (300 feet) 901 75.6 21.0 3.4 Sep-Oct 1998 900 47.8 19.6 32.6 PineClifTRanch Jul-Aug 1998 302 86.1 11.9 2.0 PineClifTRanch Sep-Oct 1998 520 81.2 9.2 9.6 Woodhouse Ranch Jul-Aug 1998 377 91.5 8.5 0.0 Woodhouse Ranch Sep-Oct 1998 533 67.6 32.1 0.4 Pine Cliff Ranch Garber Creek Jul-Aug 1998 210 68.1 10.5 21.4 Pine Cliff Ranch Garber Creek Sep-Oct1998 499 75.6 9.8 14.6 Pine ClifTRanch West Plum Creek Jul-Aug 1998 92 32.6 20.7 46.7 Pine ClifTRanch West Plum Creek Sep-Oct 1998 21 52.4 0.0 47.6 Table 3. Preble's meadow jumping mice radio-collared and tracked (X) for more than one tracking session. Location of capture, sex, identification, and number of locations (n) for each mouse during each tracking session is noted. Tracking Session I Site Sex PIT-tag # Maytag F Maytag June' July-Aug n Sep-Oct 4140371625 X X 25 * F 4141435234 X Maytag F 4140753620 X Maytag F 41412B4A2D X Maytag M 4141241836 Woodhouse F 41413E610B Woodhouse F 41413E7807 PineClifT F 4141125066 PineClifT M 41413D4C04 Locational data from June is not included in this report. * Mouse captured but radio-collar not replaced. X X n X 52 X 82 103 X 105 X 93 X X 78 X 14 X 66 X 5 X 13 X 18 X 15 * �20 Table 4. Possible hibemacula for Preble's meadow jumping mouse at three study areas in Douglas County, Colorado. Sex, beginning date of stationary signal, duration of stationary signal, distances from the main drainage (East Plum Creek at Maytag Property, West Plum Creek at PineCliffRanch, Indian Creek at Woodhouse), nearest tributary, and center of night time locations are presented. Primary vegetation at the site is also noted. or Distance (m) from: Site Sex Date Days Main Drainage Tributary Center Night Location Vegetation Woodhouse M 9122 12 40 N/A 91 skunkbush, sumac, ha~orne,chokecherry Woodhouse M 9127 9 33 N/A 67 clematis vine, snowberry Woodhouse M 9/29 5 23 N/A 116 narrowleaf cottonwood, Bromus spp., chokecherry PineCliff M 9117 5 210 10 29 Salix spp. PineCliff M 9/23 11 35 40 90 Salix spp., alysiwn PineCliff M 9/23 11 120 35 70 Salix spp., grass PineCliff M 9/25 9 100 78 72 snowberry, Salix spp., alysium, thistle PineCliff F 9125 9 70 105 105 snowberry, Salix spp. PineCliff M 9/25 9 341 0 Salix spp., cottonwood, snowberry Maytag F 9123 14 120 750 gambel oak, chokecherry, alysiwn N/A �Table 5. Known and probable mortalities of Preble's meadow jumping mice from Maytag Property, PineCliffRanch, and Woodhouse Ranch the 1998 field season (June l-October 31). Probable cause of death, date radio-collars located, description of how what was found, sex of the animal: and disQosition of sQecimen noted. collected during Site where found Mortality May tag Date How obtained Sex Current location of specimen handling stress June 8 died during processing of the mouse, never came out of anesthesia female DMNH May tag handling stress July 27 died from anesthesia shock female DMNH May tag trap mortality June 10 trap mortality male DMNH Maytag trap mortality: heat stress Sept. 10 trap mortality, probably over-heated female DMNH Maytag predation July 2 tracked radio-collar, found collar in fox scat female no body, scat at CDOW .May tag predation Sept. 21 radio-collar tracked to a rattlesnake, waited and picked up snake scat with the radiocollar in it female no body, scat at CDOW Maytag road kill Sept. 15 tracked radio-collar; found highly decomposed, unsalvageable body covered with ants, side of road-probably road kill male no body Maytag unknown Aug. 1 tracked radio-collar; found highly decomposed, unsalvageable body male no body PineCliff possible predation Aug. 6 possible predation; crimp still tightly on collar when retrieved female no body PineCliff possible predation Aug. 12 tightly crimped collar found on ground, no part of mouse detected; suspect predation since movement detected earlier female no body PineCliff possible predation Sept. 13 possible predation; crimp still tightly on collar when retrieved male no body PineCliff drowning Aug. 6 radio-tracked collar; found mouse in water female D.MN":I Woodhouse handling stress June 5 died during handling while trying to readjust radio-collar male DMNH radio-collar: suffocation/starvation Aug. 6 radio-tracked collar, found dead mouse; collar may have been too tight female DMNH Woodhouse predation Aug. 11· radio-collar found in snake scat female no body, scat at CDOW Woodhouse predation Sept. 23 radio-collar tracked to a garter snake, waited and found collar in snake scat female no body, scat at CDOW Woodhouse predation Sept. 29 radio-collar tracked to a house cat, waited and found collar in house cat scat female no body, scat at CDOW Woodhouse possible predation Aug. I tightly crimped radio-collar found in exposed area female nobody Woodhouse unknown Aug. 7 radio-tracked collar, found dead mouse (skeleton) on shrub female DMNH Woodhouse unknown Sept. IS radio-tracked collar and found dead mouse male DMNH . Woodhouse factor .- N �N Table 6. Percent contents of fecal samples collected from traps where Preble's meadow jumping mice were captured. All samples were collected from Maytag Property during the 1998 field season. Several samples are from the same animal captured on different dates. Amount of bait in each sample was quannneu as IVV/,o f\ ror aounaam \? IV/,o or me Sample}, 1 ror trace amoum \~ IV/,o or me Sample}, or V/'o or me sam pie was oan, endogenous LesVerAgro- Equiunknown mushID Date Bait arthropod fungus seed moss loollen Poa Helianthus Salix ouerella bascum Carex pvron setum flower forb room 41405E402 Jun3 A 76 414125476E Jun 3 64 '4140271625 Jun8 T T T ,414119674D Jun II A 91 41407E5E35 Ju121 0 16 4141106B52 Ju121 100 414128794B Jul28 100 4141382614 Jun4 24 7 92 100 A 82 18 4141407328 Jul28 T 2 30 4141360708 Ju129 A 47 41411E6879 Aug 3 T 4 96 41412B3D24 Aug13 0 3 16 41413A026A Aug 13 A 17 UNKNOWN Aug 13 A 25 4140371625 Aug 13 A 100 414020547B Aug 13 7 100 6 Aug 13 414125476 Sep 7 A 98 2 . 4140753620 Sep 9 A 4 43 414164363C Sep 9 T 41412CC443 Sep 10 A 41417CIE42 Sep 10 T 60 2 46 3 25 11 12 6 3 67 11 4140173A6D .4141241836 12 46 29 T T T T T Aug 13 4 9 Jul28 Aug 13 16 I 4 Jul28 Aug 13 13 96 41417AI807 4141187C76 23 .' 4 . 4141297FOF 41412B4A2D N 16 59 5 1'. 100 89 20 2 5 5 84 30 19 16 21 41411E6879 Sep II A 73 41412B4A2D Sep 12 A 20 4140754707 _Sep 12 T 24 4 84 14 90 II 78 65 5 5 27 80 7 35 34 ------- I �Table 7. Percent contents offecal samples collected from traps where Preble's meadow jumping mice were captured. All samples were collected from Pinef'liff'Ranch during the 1998 field season. Several samples are from the same animal captured on different dates. Amount of bait in each sample was quantified as 100%, 'A' for abundant (> 70% of the sample 'T' for trace amount « 10% of the sample), or 0% of the sample was bait. ID UNKNOWN 4140761237 41411A3D4B 4140755157 UNKNOWN UNKNOWN 4140755157 UNKNOWN UNKNOWN 41411AII14 4141714141 41413D4C04 4141505F54 AI4125D66 413F6F7EOA 4140784BI4 41411AECI2 41412F2433 41412F5E45 41413C604F .414176064A 4141656001 414138666D 41415B4748 41412F2F62 4141370528 4141350155 41415E434A 4141491F02 4140502204 Date Bait arthropod Jun 3 100 A Jun4 0 96 4 Jun4 A Jun4 97 A Jun4 T A Jun4 A 88 Jun4 8 T Jun 7 T T Jun 8 T A Jun 8 T 88 Jul II A 4 Jul21 0 13 Jul21 0 Jul21 100 Jul21 T 8 Jul21 T 100 Juln Jul22 A Jul24 A Jul24 41 A Sep 9 T 3 Sep9 T 3 Sep II A Sep 11 T 14 Sep 14 A 96 Sep 14 T 22 Sep 15 A 83 Sep 15 A T Sep 16 A Sep 16 T 17 endogenous fungus seed moss Bromus Poa Chenopodium seed grass Eleagnus Helianthus lily pollen Salsola seed composite Festuca 4 96 .' 3 T 6 6 88 T 4 6 0 6 96 2 37 85 63 92 8 8 84 7 8 100 6 4 75 ~ 100 100 18 90 86 41 3 74 6 3 12 5 ---- ----- ----- --------- --- 83 - 10-). w �,Table 8. Percent contents of fecal samples collected from traps where Preble's meadow jumping mice were captured. All samples were collected from Woodhouse Ranch during the 1998 field season. Several samples are from the same animal captured on different dates. Amount of bait in , each sample was quantified as 100%, 'A' for abundant (> 70% of the sample), 'T'for trace amount « 10% of the sample), or 0% of the sample .was bait PIT-tag # Date Bait arthropod 41407B2575 Jun 3 A 100 41413E610B Jun 3 A 100 41413E610B Jun4 A 100 41407B2575 Jun4 T 100 414147354F Jun4 T 24 UNKNOWN Jul27 T 41413E610B Aug 4 T 4141076B41 Aug 4 100 4141250646 Aug 23 0 12 41404A7D58 Aug 23 T 4 AI4128F2607 Aug 23 0 41415D402E Aug 23 100 41406AI049 Sep 9 A 100 4141313E6F Sep 9 A 74 4141350C2A Sep 10 100 '414133085 Sep 10 A UNKNOWN Sep 11 0 100 414173462B Sep II T 70 4143P5772B Sep II 0 45 4141611413 Sep 12 0 98 41414A6B6F Sep 12 0 88 4140241303 Sep 12 0 88 endogenous fungus mushroom Poa seed Agropyron Sporobolus seed Chenopodium composite Equisetum flower 76 50 50 23 77 88 96 89 11 26 f' T 30 33 15 7 2 12 2 10 N ~ �25 ApPENDIXB HABITAT USE AND DISTRIBUTION OF PREBLE'S MEADOW JUMPING MOUSE (Zapus hudsonius preblei) IN LARIMER AND WELD COUNTIES, COLORADO Tanya M. Shenk and James T. Eussen Abstract A total of39 sites were surveyed for Preble's meadow jumping mouse in Larimer and Weld Counties, Colorado with a total of 16,671 trap-nights completed. Of the 39 sites survey, Zapus spp. were captured at 21 locations. A total of 71 unique individuals were captured in the 21 locations, with recaptures of 14 of those individuals. All sites where Zapus spp. were captured were located in Larimer County. Three soil samples from 0-12 inches and 12-24 inches were collected at all sites where possible. Soils at some sites were too rocky to collect the 12-24 inch soil samples. Nine fecal samples were collected from traps where Zapus spp. were captured. Initial result indicate that arthropods, fungus, and willows are important dietary components of the Zapus diet. Two trap mortalities of Zapus spp. occurred. Both specimens were saved and delivered to Cheri Jones of the Denver Museum of Natural History on October 26, 1998. Measurements of habitat characteristics at both the site and landscape levels were completed. Genetic tissue samples were collected from 68 of the 71 jumping mice captured. Analyses of these tissue samples is pending. Introduction The meadow jumping mouse (Z hudsonius) is broadly distributed across North America from the Atlantic to Pacific coasts, extending south into the United States to Alabama and Georgia and west across the Great Plains to the base of the Rocky Mountains. In general, it is a common inhabitant of moist, grassy and herbaceous fields. Eleven living subspecies have been described (Whitaker 1972). Rather et al. (1981) describe a twelfth subspecies, Z. h. luteus. Z. h. preblei occurs only in eastern Colorado and southeastern Wyoming (Krutzsch 1954, Long 1965, Armstrong 1972). From its limited ecological and geographic distribution, Fitzgerald et al. (1994) suggest it is an Ice Age relict, once widespread in tallgrass prairie across the eastern plains of Colorado but now restricted to scattered localities on the Colorado Piedmont. Similar relict populations of meadow jumping mice in the White Mountains of Arizona and the Sacramento Mountains and Rio Grande Valley of New Mexico are described as the subspecies Z. h. luteus (Rather et al. 1981). On May 12, 1998 the U. S. Fish and Wildlife Service (USFWS) published a final rule in the Federal Register (63 FR 26517) to list Preble's meadow jumping mouse (Zapus hudsonius preblei) as 'threatened' under the Federal Endangered Species Act (ESA) of 1973, as amended. Scarcity of suitable habitat presumably limits current distribution of Preble's meadow jumping mouse (PMJM) and thus, maintenance of quality habitat has been identified by the USFWS (63 FR 26517) as the principal conservation goal. Although Meaney et al. (1997) reported an improved ability to recognize suitable habitat for PMJM, a more refined and complete definition of potentially suitable habitat for the mouse does not exist. Because the protection of potentially suitable habitat for PMlM may occur under the ESA, as well as protection of known locations where the subspecies occurs, the definition of potentially suitable habitat, as determined by the USFWS, will directly influence site specific regulatory procedures. Ideally, a definition of potentially suitable habitat for the subspecies would identify areas where the PMlM could survive and reproduce in sufficient numbers to sustain populations throughout its range of natural variability over an extended length of time. During the active period of the life cycle of the mouse, suitable habitat must provide requirements for daily survival, reproductive activities (breeding, nestin?, and rearing of young to independence), and dispersal. The hibernation period �26 requires habitat with sufficient food supplies to assure fat storage prior to hibernation and hibernacula sites. Habitat providing all seasonal and life cycle requirements mayor may not occur in a single contiguous area If not in a contiguous area, habitat patches must occur in a mosaic of usable areas where suitable corridors exist for seasonal movement among sites. Because very little is known about the ecological requirements of the subspecies, potentially suitable habitat is currently defined only as areas of well-developed, dense herbaceous vegetation consisting 'of a variety of grasses, forbs and thick shrubs in close proximity to open water. This definition is vague and possibly incomplete. Numerous surveys have been conducted since 1990 to establish the current distribution of PMlM. However, very few surveys for determining the presence or absence ofPMlM have been conducted in Larimer or Weld Counties, Colorado. Of the surveys conducted up to 1997, only three sites yielded captures ofPMlM in either Larimer or Weld Counties. In Larimer County, two sites (Lone Pine and Rabbit Creeks) yielded captures of PMlM, based on field identification and supported by genetic analyses conducted to date (Riggs 1998). One mouse was captured on Lone Tree Creek in Weld County that was identified as Z. h. preblei in the field but was genetically found to be more. closely allied with the western jumping mouse, Z. princeps (Riggs 1998). Therefore, very little information is known about habitat use or current distribution of the subspecies in these two counties. Both Larimer and Weld Counties include habitat that is currently perceived to be outside the ecological limits of PMlM. Larimer County provides the opportunity to explore elevational limitations, currently thought to be 7400 feet (2260m) (USFWS 1997); Weld County extends beyond the currently believed eastern boundary of the subspecies. Both of these boundaries will be challenged by selecting survey sites within the two counties beyond the perceived ecological limits of the subspecies. Genetic tissue samples will be collected on all PMlM captured to assist identification through DNA analysis, particularly for those animals captured outside the currently perceived ecological limits, and to provide information for future studies to better define the relationships among different populations of Z. h. preblei, other subspecies of Z. hudsonius and other species of Zapus. Quantifying and comparing both landscape and site specific characteristics of habitats where PMlM are found and not found in these two counties would further our understanding of habitat use by the subspecies, particularly in the northern half of its probable range. Any new locations where PMlM are found during this study would contribute to the range-wide distribution map currently available for the subspecies. Repeated annual visits to these same randomly selected sites would also provide information on population persistence at a given site. Such a monitoring scheme would be necessary for evaluating the continued status of the mouse at each site, providing further information as to the suitability of the habitat for long-term persistence of the population. Thus, the primary objective of this study is to quantify and define habitats used and identify ecological limits of the subspecies in Larimer and Weld Counties, Colorado. Such information could be used by the USFWS to further refine the current definition of potentially suitable habitat for PMlM. Objectives Conservation of Z. h. preblei should require maintaining populations of the subspecies throughout the range of its natural variation and to try to identify ecological limits for the subspecies. Specific objectives for the distributional component of this study are to (1) clarify the distribution of the subspecies in Larimer and Weld Counties, Colorado, and (2) define and quantify the amount of suitable habitat within Larimer and Weld Counties. The objective of the habitat study is to identify and refine ecological requirements of Z. h. preblei in Larimer and Weld Counties, Colorado. Because of the uncertainty of the identification of the mouse captured in Weld County, Colorado, and because currently perceived ecological boundaries of PMlM will be challenged in the survey effort, the objective of the genetic component of this study is to genetically identify all Zapus spp captured during this study. Therefore, genetic tissue samples will be . collected from each jumping mouse captured to confirm identification of the animal. These genetic �27 tissue samples could also provide further data to better define the relationships of different populations of Z. h. preblei as well as to other subspecies of Z. hudsonius and other species of Zapus through molecular systematic relationships and to link these genetic relationships to systematic studies of Z. hudsonius. Specific objectives for this study on PMlM include: . 1. Define and quantify habitat characteristics at sites where PMlM was found and sites where PMlM was not found. 2. Identify elevational and eastern boundary limitations for PMlM in Larimer and Weld Counties, Colorado. 3. Describe the distribution ofPMJM in Larimer and Weld Counties, Colorado. 4. Select sites and establish monitoring protocols for determining long-term trends in populations of PMlM. 5. Collect genetic tissue samples to assure identification of jumping mice captured during this study and for future genetic analyses to better define the. relationship of populations of Z. h. preblei to each other, other subspecies of Z. hudsonius and other species of Zapus. Study sites Trapping surveys will be conducted at a minimum of30 sites selected at random from the sampling frame developed for Larimer and Weld Counties, Colorado. No survey site will occur at elevations greater than 7600 feet and not east of the UTM Easting Coordinate of 602000. The molecular systematic studies will be conducted in genetic laboratories using the genetic tissue samples collected in the field (ear punches). Methods For simplicity, the methods described below assume all jumping mice captured are PMlM. To meet these objectives, this study included (1) constructing a sampling frame of potentially suitable habitat within the area of interest (Larimer and Weld Counties, Colorado) and (2) conducting trapping surveys to determine presence or absence at a minimum of30 sites randomly selected from the sampling frame. Specific approaches to each of these tasks follow. Sampling frame A sampling frame was developed to delineate potentially suitable habitat for PMlM from unsuitable habitat. To produce a preliminary map displaying existing potential habitat for PMlM would require at least three primary layers of ecological information. Two initial layers, hydrology and vegetative cover would provide a map displaying all areas where these two ecological requirements of PMlM co-occur. The hydrology layer must be in enough detail to provide locations of intermittent and small order streams as well as identify water source (e.g., irrigation ditches, stream, seep). Degree of density of vegetative cover is sufficient for initial mapping efforts. Demarcation of shrub, trees and grasses will suffice in these initial efforts rather than species identification. However, not all combinations of vegetative cover and water provide sufficient habitat to support PMJM. Thus, mapping potential habitat as described above would produce a map displaying far more potential habitat than exists. For example, an extremely critical ecological requirement for the survival of the mouse, that to date we have very little information for, is potential hibernacula sites. As a possible index to hibernacula habitat, a third GIS layer, indicating the presence of alluvial deposits may provide some insight in to hibernation requirements. Areas where alluvial deposits co-occur with the presence of water and vegetative cover may provide a more realistic map of potentially suitable habitat for the mouse. Our ability to survey a site selected at random would be restricted to those areas where we are . able to obtain permission to access the land. We shouldbe ableto survey any randomly selected site �28 on public lands and thus our inference should be unrestricted. Such readily available access would probably not occur on privately owned or leased lands, thus, restricting inferences made on private lands. By stratifying the sampling frame, this lack of access on private lands would not restrict inferences that can be made on public lands. Inference on private lands will depend on the percent of access we get from private landowners. The sampling frame was developed from the 1:24,000 schle Hydrographic GIS layer developed by the CDOW (Reese Tietje, unpublished data). From this sampling frame a three-stage sampling design was used to select 40 random sites (to ensure meeting our objective of a minimum of 30) for conducting PMlM surveys. Primary sampling units were 3rd order streams. Of the 150 3rd order streams that exist within the sampling frame, 40 were selected on public lands, 40 on private lands. The secondary sampling unit was the stream stretch selected within the primary (3rd order stream) unit. The tertiary sampling unit was the actual sites along the stream stretch selected at the secondary sampling stage that were sampled. Each of the primary, secondary, and tertiary sampling units were selected using simple random sampling. If the secondary sampling unit did not yield potentially suitable habitat (i.e., no dense vegetation) secondary sampling units continued to be selected at random until one yielded potentially suitable habitat. Determination of suitable habitat was made by field site visits. Total stream length of all the secondary sampling units were recorded as well as total stream length containing potentially suitable habitat. From these measurements an estimate of the probability of potentially suitable habitat occurring within the primary sampling unit can be made for each area surveyed. A mean estimate, over all sites surveyed, of the probability of a primary sampling unit having potentially suitable habitat was made. Because of the random selection of primary sampling units, inference can be made as to the probability of potentially suitable habitat existing on all3rd order streams within the sampling frame. By recording presence or absence ofPMlM within the tertiary sampling units we can estimate the probability of PMlM occurring within potentially suitable habitat. Because of the random selection of sampling units in each of the two strata (public and private lands) we can then estimate the probability ofPMlM occurring in potentially suitable habitat within both public and private lands. Trapping surveys Trapping surveys to determine presence/absence ofPMlM were primarily conducted following protocols defined by the USFWS (1997). Additional guidelines have also been developed to accommodate the needs of this project. Trapping surveys were conducted within the period from June 1 to September 15, 1998. These dates correspond to dates when PMlM is known to be out of hibernation. Trappers wereadvised to follow the Center for Disease Control's Hantavirus instructions and recommendations when dealing with rodents. Due to the nocturnal nature ofPMlM, traps were set between 19:00hrs and 21 :OOhrs and checked as early as possible in the morning beginning at 5:00hrs to reduce stress and the potential for predation on trapped animals. Time required to complete the trap lines varied depending on how many animals were caught. Small mammal Sherman live traps (folding and non-folding) were used to conduct the trapping sessions. Traps were set in two parallel lines of trap stations (1 trap per station) on either side of the drainage. Trap stations were 5 meters apart for a total of 250 meters; the parallel transects were 10 meters apart unless extent of habitat, terrain topography, or stream hydrology did not allow. Location of transects were recorded on field data sheets and 7.50 topographic maps. A small (-1 inch) ball of polyester quilt was placed in each trap as nestinglbedding material. Baiting material was Manna Pro Sweet 3-way Livestock feed which contains no animal matter; ingredients include flaked barley, flaked com, flaked oats, and cane molasses. Peanut butter was used to stick the bait to the trap. �29 Traps were checked by two surveyors. All animal captures were recorded. If an animal had been captured in a trap, a ziplock plastic bag was placed over the end of the trap. The trap was opened allowing the animal to fall into the plastic bag. The animal was identified to species while in the plastic bag. If the animal was not a PMJM, identification of the animal was recorded and the animal set free without further handling. Each captured PMJM was checked to detect the presence/absence of a PITtag (See below). If a PIT-tag was detected and the mouse had been captured at that same site within the same trapping session, identification of the mouse was recorded and the mouse was released. If the animal was a PMJM and no PIT -tag was detected the PMJM was anesthetized for further processing, as follows. All PMlM were PIT-tagged. Each PMJM was weighed while in the bag, recording the weight in grams using a Pesola spring balance. Sex, age (juvenile, adult), and reproductive condition (pregnant, lactating or nonbreeding if it was female; for males the position of the testes indicated if in breeding status or nonbreeding status) was noted. Each PMJM was measured (total length, length of body, length of hind foot - heel to distal end of claws) in millimeters. Capture of every PMJM was documented by taking a photograph of the mouse on first capture. Mice were placed in a plexiglass photobox to reduce handling stress while taking the photograph. If the mouse had been captured previously as noted by the detection of a PIT -tag, no photograph was taken. If there were feces in the trap where a PMJM was captured, the fecal material was collected in a plastic bag labeled with date, location of the site, and animal PIT-tag number. Fecal samples were kept cool until returned to the CDOW office where they were frozen. Salt was added to each specimen bag for preservation. Once the field season was over all fecal samples were analyzed by the Composition Analysis Laboratory, Inc, 622 Y2 Whedbee, Fort Collins, CO for content. All trap mortalities were recorded on a 'Trap Mortality' data form which included information on species, potential duration of time spent in the trap, and any information available to help determine cause of death. All animals found dead were double-bagged in a plastic bag and placed in a cooler with ice. Specimens were frozen as soon as possible and deposited in the CD OW freezer at the office in Fort Collins. A museum card was completed and attached to each PMJM specimen and the specimen and card were given to Cheri Jones, Curator of Mammalogy at the Denver Museum of Natural History, for study skins and tissue storage. If an animal was severely injured (e.g., severed limb, large lacerations) it was euthanized by soaking a cotton ball in Metofane (methoxyflurane) and placing the cotton ball and the mouse in a ziplock bag until the animal stopped breathing. If an animal appeared to be only slightly injured (e.g., broken tail, small laceration) the animal was released to the wild. If an animal appeared to be coldstressed attempts were made to warm it by holding it in the surveyors hands and/or against their body. If the animal appeared to be heat stressed, isopropyl alcohol was applied with a cotton swab to the ears, arm pits, and feet to cool it down. PIT-tagging Each PMJM was individually marked with a Passive Integrated Transponder (PIT -tag). PITtags are electromagnetic, glass-encased tags that emit a passive signal (125 kHz) that can be decoded by a portable reader. Destron-Fearing PIT-tags were used on all mice. We used portable readers from Biomark to read the PIT-tags. All mice were anesthetized to PIT-tag them. Anesthesia procedures followed protocols successfully used on PMJM before (R. Schorr personal communication). Mice were anesthetized by placing them and a cotton ball with 1 ml of Me tofane (methoxyflurane) into a sealed ziplock bag (to keep Metofane fumes in bag). The bag was then lain aside to minimize stress for the mouse and the time the mouse struggles in the bag. The less agitated the mouse is in the bag the less time required for the metofane to take affect. The mouse was observed at all times to make sure the mouse did not .. position itself such that the cotton ball was in direct contact with its face, which might cause the mouse o· • " •• �30 to have a severe reaction to the anesthesia After the animal stopped moving, the handlers waited one minute before removing the mouse from the bag. The animal typically remained anesthetized for 2-3 minutes once outside the bag. Each PMJM was PIT -tagged using a protocol successfully used by C. Meaney (personal communication). Each PMJM had a PIT -tag inserted above the shoulder blades by lifting the skin on its back and inserting the individually sterilized needle with the PIT-tag under their skin and injecting the tag. Verification of the PIT -tag identification number was made before and after insertion into the mouse by running the PIT -tag scanner across it. The skin behind the opening was then pinched to prevent emergence of the tag. PIT-tag identification number was recorded on the field data form. If, at any time during the handling of the PMJM the animal appeared to be severely stressed (dramatic changes in heartbeat, respiration and responsiveness or gums turning blue) first aid was administered and, once recovered, released without further processing. Absence ofPMJM at a site was defined as no captures ofPMJM from a minimum of four consecutive nights and 400 trap nights (a trap night is defined as the sum total number of traps available each night). Presence ofPMJM at a site was defined as at least one capture ofPMJM at the site. However, to accommodate sample size requirements for the monitoring and genetics study (see below) trapping efforts continued until at least 10 individuals were tagged with PIT-tags. Habitat Use The general approach was to measure both site specific and landscape features of at least 30 sites selected at random to be surveyed for the presence ofPMJM. A comparison of sites where PMJM were found and were not found was then made in an attempt to refme our current understanding of the ecological requirements of the subspecies. The following habitat characteristics were measured and recorded for each site. Site characteristics Cover was measured at 20 random locations along the 250 meter sampled stream stretch. Mean cover and standard error of the mean will be estimated. Cover was estimated using a vegetation profile board (N udds 1977) that allows for an asse~sment of visual obstruction in 0.5 meter vertical intervals above ground. The board was 2.0 meters high and 30.48 ern wide. The board was marked in alternate black and white colors at 0.25 (0.50) meter intervals. Horizontal cover was assessed in each interval by viewing the board from 15 meters away in a randomly chosen direction. The percentage of each interval concealed by vegetation was recorded as 0-20, 21-40, 41-60, 61-80, and 81-100% estimated concealment. Vegetation species composition and richness was estimated using the Modified-Whittaker nested vegetation sampling method (Stohlgren et al. 1995). A modified Whittaker plot is 20-meters x 50-meters with 101m2 and 210m2 subplots arranged systematically around the perimeter and 1 100m2 subplot centered in the inside of the 20 meter by 50 meter plot. Species composition and percent cover (optical estimate) of each species was recorded for each subplot. Soil samples were taken randomly at each site for a hydrometer textural analysis (percent sand, silt and clay). Samples were collected using an AMS 24" soil recovery probe. Samples were taken from 0-12 inches, and 12+ to 24 inches. The 0-12 inch sample was taken first, removing the probe and soil. Samples were placed in a labeled zip lock bag. The probe was then placed in the same hole for the 12+ - 24 inch probe, with that soil sample being placed in a separately labeled zip lock bag. The hydrometer textural analysis will be conducted at the CSU Soils Laboratory. Mean stream width was estimated by measuring stream width at 30 locations along the entire site sampled. The 30 locations were stratified equidistant from each other to cover the entire stream stretch in increments of site stream lengthl30. : .--_ �31 The above listed parameter estimates will be compared for sites where PMlM were. captured and sites where PMlM were not captured. Landscape characteristics The following landscape habitat characteristics will be measured and compared for sites where PMlM were captured and sites where PMlM were not captured using either GIS, topographic maps, or aerial infrared photography. 1. Distance to nearest human habitation and human disturbance. 2. Connectivity of sites to other streams from 1:24,000 topographic maps. 3. Total length of stream stretch at each site and total length of stream stretch with suitable habitat at each site. Molecular Systematics Genetic tissue samples were collected from all PMJM captured during the study. To date, positive identification of the individual to subspecies cannot be made using only genetic information. However, results of the DNA analysis combined with the morphometric data that will also be collected (e.g., body length, tail length) and the photograph of the individual should provide sufficient information to support the field identification to subspecies. As established by the USFWS (1997) Survey Guidelines, presence ofPMJM is established when only one individual is captured. Thus, we could stop our trapping efforts after the first PMJM capture. However, because we would also like to provide sufficient genetic data to more fully explore the relationships of different populations of Z. h. preblei we will attempt to collect genetic tissue samples from a minimum of 10 individuals per successful survey site. A sample of at least 10 individuals from a population will provide enough information to document the genetic variation within the population (T. Quinn, personal communication). Genetic tissue sampling Genetic tissue samples were collected from every PMJM captured at each survey site. The following protocol was used based on previous success with PMJM (M. Bakeman, C. Meaney, T. Ryon, R. Schorr, personal communication). A fresh pair of clean latex gloves were used to handle each mouse. With a clean ear punch tool, one tissue plug was obtained from each mouse ear. If there was excessive bleeding, gentle pressure was applied to the injured area for approximately one minute. Each ear plug was placed in a vial of 95% ethanol. Both the vial and the corresponding record on the data sheet were labeled with a unique identifier as follows. A seven-place alpha-numeric code was composed as follows: a three-letter designator for the survey location (e.g., MYT = Maytag Property); a number beginning with two digits indicating the year (e.g., 98); followed by 2 digits specifying the individual trapped, numbered sequentially. For example: MYT9801 = first animal sampled at Maytag Property in 1998. After returning from the field, all samples were put in a cool place or refrigerated until delivery to the CDOW office at 317 West Prospect, Fort Collins, CO where they are being held in the freezer for future analyses. Ear punch tools were cleaned after each use by immersing them in a 10% bleach solution for a few minutes, rinsing them thoroughly with clean water, and then dried thoroughly to prevent rusting. Dull ear punch were discarded and replaced with new sharp punches to ensure the quickest, cleanest cut possible. Results Trapping results Thirty-nine sites were surveyed for PMJM in Larimer and Weld Counties, Colorado (Fig. 1), with a total survey effort of 16,671 trap-nights. Of the 39 sites survey, Zapus spp. were captured at 21 survey sites (Table 1). A total of 71unique.individuals were captured in the 21 locations, with �32 recaptures of 14 of those individuals. All sites where jumping mice were captured were located in Larimer County. Two trap mortalities of Zapus spp. occurred. Both specimens were saved and delivered to Cheri Jones of the Denver Museum of Natural History on October 26, 1998. Numbers of other small mammal species captures was recorded for each site surveyed (Table < 2). Nine fecal samples were collected from traps where Zapus spp. were captured. Results of eight samples indicate the jumping mice at the capture sites in Larimer County feed on arthropods and fungus which occurred in 50% (4) of the samples, followed by willows, pollen, and moss occurring in 38% (3), 25% (2) and 25% (2) of samples respectively. Other food items found included, moss, horsetail, unknown flowers, and unknown seeds (Table 3). Habitat characteristics Measurements of habitat characteristics at both the site and landscape levels were completed. No single habitat feature could be identified as unique to either sites where jumping mice were captured or sites where jumping mice were not captured. Kentucky blue grass (Poa pratensis), snowberry (Symphoricarpos spp), several species of willow (Salix spp.), and Mountain Alder were most abundant in areas occupied by Zapus spp. Horsetail (Equisetum spp), wild rose (Rosa spp), and cottonwood were also common. Dominant vegetation at survey sites with negative trapping results for jumping mice included wheat grass (Agropyron spp), Canary reed grass (Phalaris angustifolia), thistle, and willows (Table 4). Russian olive (Elaeaqnus angustifoliat occurred in 6 (33%) of the areas where no jumping mice were found, while absent in areas where jumping mice were captured. Jumping mice were captured in vegetation adjacent to large rivers, small streams, ponds, ditches, and seeps. Mean stream width varied considerably for both sites where jumping mice were captured (0.3-24.5m) and where no jumping mice were captured (0.0-20m)(Table 4). No jumping mice were found in locations surveyed east of Interstate 25(125) during 4,716 nights of trapping effort. Habitat features along certain stretches of Willow Creek within the Pawnee National Grasslands and near the town of Briggsdale are similar to areas where jumping mice were found west ofI25. However, these sites are extremely isolated and the surrounding landscape is in either rangeland or agriculture production. From late June through late August, 22 locations in Larimer County were surveyed. Zapus spp were found in 20 of these 22 sites. The exceptions being the effluent below Seaman Reservoir on the Poudre River and Soldier Canyon west of Horsetooth Reservoir. Below Seaman Reservoir just prior to and during trapping efforts, livestock (primarily cattle) were allowed to graze along this section of the Poudre River. Habitat composition along Soldier Canyon is similar to that of Arthur's Rock -3/4 miles to the south (occupied by Zapus spp). However water is permanent along the Arthur's Rock drainage but flows intermittently through Soldiers Canyon. Only one jumping mouse was captured west ofI25 after August 24 during 5,382 nights of trapping effort at nine different sites. All locations trapped after August 24 were within drainages known to be occupied by Zapus spp., including four sites along the Big Thompson River (this study), two along the Poudre River (this study), and one each along Rabbit Creek (Meaney et al. 1997), Bull Creek (this study) and Lone Tree Creek (Meaney et al. 1997). The surrounding areas that were trapped along Bull Creek and Lone Tree Creek had been extensively grazed, leaving only isolated patches of vegetation over most of the site, however all other locations were similar in habitat composition to known PMIM locations. Three soil samples from 0-12 inches and 12-24 inches were collected at all sites where possible. Soils at some sites were too rocky to collect the 12-24 inch soil samples. A total of 104012inch samples and 46 12-24 inch soil samples were delivered on October 21, 1998 to the Colorado State University-Soil Composition lab for soil textural analyses .. To date these analyses have not been. completed. '.' . �33 Genetic sampling Genetic tissue samples were collected from 68 of the 71 jumping mice captured. these tissue samples is pending. Analyses of Discussion The habitat matrix within the range of Z. h. preblei is mixed grasslands adjacent to the Colorado Front Range along the Piedmont and along the base of the Laramie Mountains in Wyoming and extends to the Colorado plains. Within this matrix, PMlM occur along stream drainages that contain patches of suitable vegetation. Suitable habitat appears to have at least two major components. The first component is a supply of open water, at least in part of the active season (M. Bakeman, C. Meaney, personal communication). Secondly, areas where PMlM has been found have dense cover (M. Bakeman, C. Meaney, personal communication). Based on studies of Z. h. preblei and Z. hudsonius elsewhere, Z. h. preblei apparently occurs mostly in undergrowth consisting of grasses, forbs, or both in open wet meadows and riparian corridors, or where tall shrubs and low trees form an overstory and provide adequate cover (Armstrong et al. 1997). Meadow jumping mice are widespread in abandoned grassy fields, but are often more abundant in thick vegetation along ponds, streams, and marshes or in rank herbaceous vegetation of wooded areas (Whitaker 1963). Vegetation at sites where Zapus spp were captured in this study were structurally diverse. No single plant species dominated areas occupied or unoccupied by Zapus spp. Such diversity in a multistory cover has been reported for the majority of other sites where Z. h. preblei have been found (Armstrong et al. 1997, Meaney et al. 1997). Vegetation composition of the dense cover varied considerably and included both native and non-native species as was found by other studies (Meaney et al. 1996, 1997, Armstrong et al. 1997, M. Bakeman, personal communication). Results from this study continue to support the habitat descriptions provided by Armstrong et al. (1997) which follows: The herbaceous understory is primarily grasses or forbs or a mixture of the two. Few of the sites, however, are dominated by fewer than two understory species. The tall shrub canopy at most sites is willow (Salix spp.), although scrub oak (Quercus gambelli), birch (Betula spp.), and alder (Alnus spp.) occur in sites south of the Palmer Divide Ponderosa pine (pinus ponderosa) is the most common tree at higher elevations. The mouse appears to tolerate weedy or exotic species in areas that are structurally diverse and species rich; nearly every successful site contained Canada thistle (Cirsium arvense). Thus, the mouse does not appear to have an affinity toward any single plant species but instead favors sites that are structurally diverse and provide adequate cover and food throughout its life cycle. PMlM have been trapped in natural riparian areas as well as areas altered by anthropogenic influence including ditches and wetlands adjacent to interstate highways, cement-lined ditches with tall cover, ditches along driveways and moderate road use, and moderate cattle grazing (M. Bakeman, personal communication). Open water was present at all sites surveyed in this study, with stream width varying across locations with and without jumping mice. Jumping mice were captured in 21 new locations in both the Poudre River and the Big Thompson River drainages in Larimer County. No jumping mice were captured at any survey site located in Weld County, however there may still be some sites within and adjacent to the Pawnee National Grasslands that may support PMlM (M. Ball, personal communication). Fish Creek, north of Bull Creek is the only location trapped after the 24th of August where jumping mice occurred. Poor trap success in late summer especially in areas along the Big Thompson and Poudre Rivers, including locations near Cherokee Park SW A, may be influenced in part by jumping mice entering hibernation. Elevation at these sites ranged from 5800 ft along the Poudre River to 7600 ft along Bull Creek north of Cherokee Park, much higher then the maximum elevation in Douglas (Shenk.and Sivert 199.9,<1,1999b.) and Boulder Counties. (Meaneyet al, 1999) whereno .. , reduction in trap su.cces~ could be detected. ..... . . .. . . .. " '.' . �34 Food habit studies using scat analysis indicate jumping mice captured in Larimer County fed on several different food types. This is similar to Shenk and Sivert (1999b) in Douglas County, Colorado who also reported several food items in Z. h. preblei scat. It appears jumping mice captured in Larimer County fed on willows, with willow present in 38% of the scats examined. This is in contrast to Shenk and Sivert (1999b) who report virtually no willows in the scat examined from 85 fecal samples collected at three sites in Douglas County. PMlM are only one component of the small mammal community inhabiting areas where they were captured. These mice were more often found at sites with high species richness and small mammal abundance (Table 2). The highest occurrences of Mus musculus captures occurred on sites where jumping mice were not found. This is similar to results summarized in Shenk (1998). The presence of large numbers of house mice might suggest degradation of habitat for Z. h. preblei in those areas or possibly competition between the two species (Ryon 1996). Jumping mice were also not found in areas where large numbers of captures of deer mice (> 115) occurred. These preliminary analyses have treated all the jumping mice captured the same. However, the truth may be they are all PMJM, all western jumping mice, or a mixture of the two. Once the genetic study is completed and species identifications confirmed these analyses will need to be reevaluated in light of that information. If some of the mice captured are identified as western jumping mice it would be most beneficial to compare areas of occupancy by this species to areas occupied by PMlM and to areas potentially occupied by both. According to Fitzgerald et al. (1994) the distributions of Z. princeps and Z. hudsonius do overlap. The area of overlap occurs in eastern Wyoming. The distribution of Z. hudsonius in Colorado is now known to be larger than shown in Fitzgerald et al. (1994). The boundaries are currently as far south as Las Animas County, based on genetically identified specimens of Z. h. preblei (Riggs 1998). Captures of Z. princeps and Z. hudsonius (as identified in the museum) have occurred as close as eight miles of one another within the same drainage (Armstrong 1972). Z. princeps were reported captured in 1981 (Olson and Knopf 1988) at the Lone Pine site in Larimer County, Colorado, where Z. h. preblei were captured by Menaey et al. (1997). Because neither specimens or genetic samples were taken in the 1981 study, identification of those mice will remain in question. The discrepancy may be explained by field misidentification or Meaney et al. (1997) also suggest this discrepancy might be explained by displacement of Z. princeps with z. h. preblei sometime in the sixteen intervening years between trapping efforts. Although assumed to have different ecological requirements, genetic evidence presented by Riggs (1998) suggests further investigation of possible distributional overlap between Z. h. preblei and Z. princeps. These preliminary results from this study suggests further research should be conducted to continue evaluating range-wide distributional boundaries (e.g., elevation restrictions). Further studies should also be conducted to investigate areas of potential sympatry or hibridization ofPMlM with Z. princeps (western jumping mouse). G Literature Cited Armstrong, D. M. 1972. Distribution of mammals in Colorado. University of Kansas, Museum of Natural History Monograph 3:1-415. Armstrong, D. M., M. E. Bakeman, A. Deans, C. A. Meaney, and T. R. Ryon. 1997. Conclusions and recommendations in: Report on habitat findings on the Preble's meadow jumping mouse. Edited by M. E. Bakeman. Report to USFWS and Colorado Division of Wildlife. EG&G. 1993. Report of Findings: 2nd Year Survey for the Preble's meadow jumping mouse. Prepared by Stoecker Environmental Consultants for ESCO Associates, Inc., Rocky Flats Environmental Technology Site, Jefferson County, Colorado Fitzgerald, J. P., C. A. Meaney, and D. M. Armstrong. 1994. Mammals of Colorado. Denver Museum of Natural History, University Press of Colorado. Niwot, Colorado. Hafner, D. l,K, E. Petersen, .and T. L. Yates. 1981.. Evolutionary relationships of jumping mice (genus Zapus) of the southwestern United States. Journal of'Mamrnalogy 6~:501-512 .. �35 Hall, E. R. 1981. The mammals of North America John Wiley and Sons, Inc., New York, New York, 2 volumes. Jones, C. A 1996. Mammals of the James John and Lake Dorothey State Wildlife Areas. Final Report, submitted to the Colorado Division of Wildlife and Colorado Natural Areas Program. Krutzsch, P. H. 1954. North American jumping mice (genus Zapus). University of Kansas ~ Publications, Museum of Natural History 7:349-472. Levins, R. 1970. Extinction. Lectures in Mathematical Life Sciences 2:75-107. Long, C. A 1965. The mammals of Wyoming. University of Kansas Publications, Museum of Natural History, 14:493-758. Meaney, C. A, N. W. Clippinger, A Deans, and M. OShea-Stone. 1996. Second year survey for Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. Report prepared for the Colorado Division of Wildlife. Meaney, C. A, A Deans, N. W. Clippinger, M. Rider, N. Daly, and M. O'Shea-Storie. 1997. Third year survey for Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. Report prepared for the Colorado Division of Wildlife. Nudds, T. D. 1977. Quantifying the vegetative structure of wildlife cover. Wildlife Society Bulletin 5:113-117. Olson, T. E., and F. L. Knopf. 1988. Patterns of relative diversity within riparian small mammal communities, Platte River watershed, Colorado. Pg. 379-386 in: Proceedings of the symposium: Management of amphibians, reptiles and small mammals in North America. Flagstaff, Arizona U. S. Forest Service, General Technical Report RM-166. PTI Environmental Services. 1996a Preble's Meadow Jumping Mouse Study at Rocky Flats Environmental Technology Site, Annual Report 1996. Final. Rocky Flats Environmental Technology Site, Golden, Colorado. . Quimby, D. C. 1951. The life history and ecology of the jumping mouse, Zapus hudsonius. Ecological Monographs 21:61-95. Riggs, L. A, J M. Dempey, and C. Orrego. 1997. Evaluating distinctness and evolutionary significance of Preble's meadow jumping mouse: Phylogeography of mitochondrial DNA noncoding region variation. Final Report for the Colorado Division of Wildlife. Denver, Colorado. Ryon, T. R. 1996. Evaluation of historical capture sites of the Preble's meadow jumping mouse in Colorado, final report. MS Thesis. University of Denver, Denver, Colorado. Shenk, T. M. 1998. Conservation assessment and preliminary conservation strategy for Preble's meadow jumping mouse (Zapus hudsonius preblei). Colorado Division of Wildlife FY 1997-98 Annual Report. Shenk, T. M. and M. M. Sivert. 1999. Temporal and spatial variation in the demography of Preble's meadow jumping mouse (Zapus hudsonius preblei). Colorado Division of Wildlife JanuaryMarch 1999 Quarterly Report. Shenk, T. M. and Sivert M. 1999. Movement patterns of Preble's meadow jumping mouse (Zapus hudsonius preblei) as they vary across time and space. Colorado Division of Wildlife JanuaryMarch 1999 Quarterly Report. Stohlgren,T. J, M. B. Falkner, and L. D. Schell. 1995. A Modified-Whittaker nested vegetation sampling method. Vegetatio 117:113-121. USFWS. 1997a Proposal to list the Preble's meadow jumping mouse as an endangered species. USFWS 50 CFR part 17. USFWS. 1997b. Interim survey guidelines for Preble's meadow jumping mouse. USFWS. Denver, Colorado. Whitaker, J. 0., Jr. 1963. A study of the meadow jumping mouse, Zapus hudsonius (Zimmerman), in cental New York. Ecological Monographs 33:3. ...... Whitaker, 1, 0., Jr. ·}972. Zapushudsonius. Mammalian Species.ll.l-Z. -, '.' �36 Table 1. Survey site locations and trapping results for presence of jumping mice (Zapus Larimer and Weld Counties, Colorado. Positive identification to species is pending. spp.) in Drainage Dates trapped UTM coordinates Elevation Number of Zap us spp.found Weights (grams) General Description of Location Arthur's Rock 6/23/986/26/98 E 485250 N 4490700 5520 feet 3 total 3 females 2i,23.5 unnamed drainage to north of Arthur's Rock drainage Bull Creek 6/28/98- E458594 N 4525510 7480 feet 711/98 13 total 6 females 7 males 20,21.5,28,22, 26.6, 19,23,22, 25,20.5,23, 22, 23 west of Prairie Divide Road 7nt987110/98 E485875 N 4498154 5120 feet 2 total 1 male 1 female 18.5,19 near Watson Lake, northwest of Laporte 7nt98- E 467500 N 4502140 6320 feet 7/9/98 12 total 8 females 4 males 21.5,25, 19,20, 24,21,19,21, 23.5,21,23 drainage runs along west side of Stove Prairie Road into the Poudre River Meadow Creek 7110/987113/98 E467149 N 4524532 6680 feet 3 total 3 females 17,22.5,25.5 along Cherokee Park Road YOWlg Gulch 7112/987115/98 E470750 N 4502380 E470650 N 4503800 6240 feet 1 total 1 female 2 total 2 female 22 tributary ofPoudre River E450611 N 4505834 7000 feet 2 total I female I male 24,24.8 7115/98 tributary ofPoudre River N. Fork Poudre River 7/207/22/98 E 468100 N 4527485 6440 feet 2 total 1 female 1 male 20,22.5 Cherokee Park, drainage above Halligan Reservoir Pendergrass Creek 7/267/29/98 E461380 N 4501300 7000 feet 1 total 1 male 28 tributary ofS. Fork of the Poudre River Buck Gulch 8/3-4/98 E463575 N 4502360 6400 feet 2 total 2 females 10,23.5 tributary ofPoudre River just east of Big Narrows Lakey Canyon 8/3-4/98 E466550 N 4491360 7120 feet '" total 3 females I male 28.5,22, 30, 13 tributary of Buckhorn Creek Little Bear Gulch 8/5-7/98 E 475650 N 4483745 6600 feet 2 total 2 female 29,16 Buckhorn Mtn. area Bear Gulch 8/6-7/98 E472500 N 4483550 7200 feet I total 1 female 19.5 tributary of Buckhorn Creek N. Fork, Big Thompson 8/9-11198 E 462950 N 4478090 7080 feet 1 total 1 female 24.5 by Glen Haven N. Fork Poudre River 8/9-11/98 E 478692 N 4516030 5900 feet 7 total 3 female 3 male ,1 escaped 22,17,5,17.5,18, 17,28, Robert's property Poudre River Skin Gulch Sevenmile Creek 7/12/98- 5840 feet .. 33,23.5 ~ : .:. ,.' �37 Drainage Dates trapped UTM coordinates Elevation Number of Zap us spp.found Weights (grams) General Description of Location North Poudre Canal 8117-18/98 E478000 N 4518600 6000 feet 4 total 4 male 23,19,13,18 irrigation ditch and pond - Knight's property NorthFork ofPoudre 8118-19/98 E472935 N 4523400 6100 feet 1 total 1 female 28.5 Phantom Canyon Little Thompson River 8/19-20/98 E468500 N 4459400 6600 feet 1 total 1 male 17.5 northwest of Pinewood Sprngs, offHwy 36 CedarCreek 8/21-22/98 E 477097 N 4477899 6080 feet 5 total 3 female 2 male 20,28, 18.5,33,15 Sheep Creek 8/21-22/98 E 470451 N 4490818 6700 feet I total I male 18.5 tributary of Buckhorn Creek Fish Creek drainage 9/9-10/98 E462800 N 4537100 7400 feet 1 total I female 18.5 tributary ofFish Creek Geary Creek 6/2-6/5/98 E 547250 N 4522450 5160 feet 0 Geary Creek, Pawnee National Grasslands Windsor Ditch 6n-6110/98 E 502350 N 4506750 5180 feet 0 Windsor ditch, Cobb LakeSWA Willow Creek 61116114/98 E 537260 N 4521900 5200 feet 0 Willow Creek, Pawnee National Grasslands Willow Creek 61126115/98 E 545310 N 4516687 5060 feet 0 Willow Creek, Pawnee National Grasslands 6116- E 555711 N 4499260 4820 feet 0 Crow Creek, Pawnee National Grasslands E 522550 N 4523050 5450 feet 0 Owl Creek, Pawnee National Grasslands Crow Creek 6119198 Owl Creek 6123- 6/26/98 '" Soldier Canyon 6/28-7/1/98 E 484350 N 4492950 5520-5600 feet 0 Soldier Canyon drainage, Lory State Park Seaman Reservoir 7/237/26/98 E 480100 N 4505680 5400 feet 0 efiluentofSeaman Reservoir, North Fork of the Poudre River South Fork Rabbit Creek 8/248/27/98 E 470150 N 4515100 6334 feet 0 South Fork Rabbit Creek, Cherokee Park SWA Banner Lakes 8/248/26/98 E 537850 N4435190 5030 feet 0 Banner Lakes SWA Big Thompson River 8/27-30/98 E497210 N 4470130 4870 feet 0 Big Thompson River at Simpson Ponds SWA South Side Ditch 9/1-4/98 E 484350 N 4473690 5100 feet 0 South Side Ditch, Big Thompson Canyon " "" �38 Drainage Dates trapped UTM coordinates Elevation Number of Zap us spp.found Big Thompson River 911-4/98 E467150 N 4473120 6800 feet 0 Long Gulch 911-9/4/98 E465850 N 4473450 7100 feet 0 Long Gulch, Big Thompson drainage LoneTree Creek 9/9-9112/98 E N E N 506850 4536000 506320 4535400 5900 feet 0 Long Tree Creek, Meadow Springs Ranch Bull Creek 91129/15/98 E462150 N 4534150 7600 feet 0 Bull Creek South Fork Poudre River 9/139/16/98 E462250 N 4503650 6400 feet 0 South Fork Poudre River Poudre River 91139116/98 E470955 N 4504350 5800 feet 0 Poudre River Weights (grams) General Description of Location Big Thompson River t • �·Table 2. Numbers of other small mammal captures at the 39 sites surveyed for Preble's meadow jumping mouse in Larimer and Weld Countiesz Colorado. SQecies codes are listed below the table. Mumu Other Drainage PMJM SosQ Sys12 Tas12 Tahu Chhi Res12 Perna Petr PesQ Neme Mis12 Arthur's Rock · Bull Creek '.Poudre River at Watson Lake • Skin Gulch Meadow Creek Youn Gulch Seven Mile Creek North Fork Poudre River · . Pender~ass Creek · Buck Gulch · Bear Gulch " North Fork Big ThomQson · North Fork ofPoudre River North Poudre Canal North Fork ofPoudre River · Little ThomQson River CedarCreek Shee Creek .: Fish Creek drainage Geary Creek Windsor Ditch · Willow Creek Willow Creek Crow Creek Owl Creek . Soldier Canyon Seamen Reservoir South Fork Rabbit Creek · B8IU1erLakes 2 3 13 2 12 3 3 2 2 1 1 5 2 7 I 25 3 3 27 34 43 79 106 113 114 11 22 6 5 14 8 4 1 2 2 2 11 5 8 22 2 7 4 2 2 I I 5 I I 0 0 0 0 3 65 15 5 2 2 4 3 ___ 5 I 0 0 0 0 0 0 3 5 9 4 1 1 1 15 46 15 6 23 8 2 I 5 2 15 3 37 16 1 6 8 1 43 14 8 9 211 176 110 32 39 2 6 2 2 8 8 30 174 12 1 93 13 1 3 1 13 4 28 166 VJ \0 6 �~ Drainage PMJM Big Thompson River SospSysQ 0 0 0 0 0 Big Thompson River Long Gulch Lone Tree Creek Chhi Resp 2 :' Sosp = Sorex spp. • Chhi = Chaetodipus hispidus Petr = Peromyscus truei Mumu = Mus musculus Pesp Mumu 10 25 3 8 16 31 1 13 11 104 67 5 1 22 14 4 1 4 3 Neme 186 83 174 1 1 0 Poudre River Perna· Petr 28 0 0 Bull Creek . South Fork Poudre River Tasp _Tahu Misp Other 81 9 72 Tasp = Tamiasciurus hudsonicus Perna = Peromyscus maniculatus Neme= Neotoma mexicana Sysp = Sylvilagus spp. Resp = Reithrodontomys spp . Pesp = Peromyscus spp. Misp = Microtis spp. "Table 3. Percent contents offecal samples collected from traps where Preble's meadow jumping mice were captured. All samples were .collected during the 1998 field season. Amount of bait in each sample was quantified as 100%, 'A' for abundant (> 70% of the sample), 'T' for trace amount « 10% of the sample), or 0% of the sample was bait. endogenous Salix seed Location Date bait arthroEods moss Eollen Equisetum flower Salsola funBuS PIT-ta~ ID# 41412PH3D , 41405A3326 41405C6818 41414A5459 Juvenile Juvenile 41413A7757 ... 4141310654 Juvenile Seven Mile Creek July 14 Seven Mile Creek North Fork Poudre North Fork Poudre Sheep Creek July July Aug Aug 18 21 II 22 Buck Gulch Buck Gulch Aug 4 Aug 4 Bear Gulch Fish Creek Aug 7 Sept 10 T T A T A A T A 0 89 II 53 2 9 89 41 6 78 15 100 T 7 9 91 7 7 100 79 7 �41 Table 4. Mean stream width and dominant understory, shrub, and overstory vegetation at each of the sites surveyed for Preble's meadow jumping in Larimer and Weld County, Colorado during 1998. Mean Stream Dominant Overstory widt~,<m) Drainage+ PMJM Arthur's Rock y Bull Creek y Poudre R. at Watson Lake y y Skin Gulch Meadow Creek Young Gulch Seven Mile Creek Y Y 0.97 1.06 3.38 1.14 Y Y Lakey Canyon Y Little Bear Gulch Y Y Y North Fork Big Thompson North Fork ofPoudre River Y Y North Poudre Canal CedarCreek Y Y Y Sheep Creek Y Fish Creek drainage Y Geary Creek N Windsor Ditch N Willow Creek N Willow Creek N Crow Creek N Owl Creek N North Fork ofPoudre River Little Thompson River Soldier Canyon N Seamen Reservoir N South Fork Rabbit Creek 5.15 1.22 Y Bear Gulch 0.28 1.01 0.2 0.54 1.77 Pendergrass Buck Gulch 0.77 1.75 24.47 se(X) y North Fork Poudre River Creek X 8.30 1.72 0.52 0.84 0.46 0.53 6.93 3.03 0.53 0.26 0.18 0.26 1.9 1.73 12.50 11.97 6.97 1.24 8.30 2.93 2.07 2.58 0.49 1.43 l.82 1.1 0.4 2.12 0.00 7.07 9.20 8.00 3.37 2.40 l.20 9.87 GrasslForb Shrub BluegrassIHorsctail Wild Plum Elm Bluegrass Willow Mountain Alder Willow Cottonwood Alta Fescue Bluegrass/Canada Thistle Tree Snowberryl Chock Cherry Cottonwood Bluegrass/Canada thistle Snowberry Mountain alder Bluegrass Willow Ponderosa pine Bluegrass Wax current Juniper Bluegrass! wheatgrass Willow Mountain alder Aspen! maple Black-eyed susan Snowbeny Wax current Spruce Columbine Juniper Aster Wild rose! wild grape Bluegrass Wild raspberry Aspen Bluegrass! yarrow Willow Mountain alder Bluegrass Snowberry Mountain alder Bluegrass!Canada thistle Willow Cottonwood Bluegrass Willow Cottonwood Bluegrass! Canada thistle Snowberry Mountain alder Canada thistle Willow Cottonwood Bluegrass Willow Ponderosa pine Blue grama Wild rose Mountain alder Canada thistle Willow Ponderosa pine 0.91 5.74 2.41 Wheatgrass (Absent) Cottonwood Wheatgrass Willow (Absent) Wheatgrass (Absent) (Absent) 1.78 0.89 Canada Thistle Wax Current (Absent) Canada Thistle Wax Current Elm 0.39 Cone Flower Willow Russian Olive 0.42 0.79 2.41 Bluegrass! Dandelion Willow Cottonwood Cheatgrass Willow Ponderosa pine �• II • • Zapus spp. Found Site Trapped No Zap us spp. Found D County Line NStreams IV Roads Figure 1. 1998 survey site locations and trapping results for presence of Preble's meadow jumping mouse (PMJM) in Larimer and Weld Counties, Colorado. Locations where PMJM were found are indicated by a red circle, locations where PMJM were not found are indicated by a green circle. ~ !-oJ �43 ApPENDIXC TEMPORAL MOUSE AND SPATIAL VARIATION IN THE DEMOGRAPHY OF PREBLE'S MEADOW JUMPING (Zapus hudsonius preblei) , \ Tariya M. Shenk and Maile M. Sivert Abstract A total of 186 individual mice were captured and PIT-tagged at three study sites (73 mice at Maytag Property, 77 mice at PineCliffRanch, and 36 mice at Woodhouse Ranch). These mice were captured over three different trapping sessions with an effort of 17,330 trap-nights. Genetic tissue samples were collected for at least 30 individual mice at each of the three study sites. Density estimates were expected to increase as the summer progressed to account for the birth pulses in late June, and late July-August. This was the trend observed' at Maytag. PMJM densities at Woodhouse Ranch increased from June to July but did not increase further during the September trapping session. Mean PMJM density over all sites and sessions was 40.5 (range 16.9-79.0) mice per kilometer of stream stretch. Highest densities occurred at PineCliff, where the vegetation is primarily willow and both a mainstem and tributary were used by mice. Lowest densities occurred at Woodhouse Ranch where the riparian vegetation has fewer willow but was dense with other riparian vegetation. Vegetation at Maytag provided some areas of dense willow with the remaining areas being of moderate density of riparian vegetation. The lower density ofPMJM reported for Woodhouse Ranch might be explained by the composition and densities of other small mammals at that site. Woodhouse Ranch had the highest captures of both house mice and voles of the three sites studied. Defining summer as June 1 - October 5, over-summer survival was estimated as 0.36 (se = 0.056) over all three study sites. Temporary emigration and immigration rates were estimated but both had extremely high variances associated with those estimates. Thus, these estimates cannot be used, with any confidence, to provide information on movements of mice into and out of these populations. Very little new information was gained during this study on reproductive parameters ofPMJM. Most adult mice captured exhibited evidence of active reproductive behavior, either pregnancy, lactation, or enlarged genitalia. However, we were unable to locate any breeding nests. Juvenile mice were not captured during the June trapping session. Juvenile mice were captured at all three sites during both the July and September trapping sessions. This report includes results from only the first year of a multi-year project to follow individually marked PMJM through time. Collection of more data, and more years of data, will improve our ability to evaluate demographic parameters and estimates of how they vary across space and time. Introduction On May 12, 1998 the U.S. Fish and Wildlife Service (USFWS) published a final rule in the Federal Register (63 FR 26517) to list Preble's meadow jumping mouse (Zapus hudsonius preblei) as 'threatened' under the Federal Endangered Species Act (ESA) of 1973, as amended. Recovery goals for Preble's meadow jumping mouse (PMJM) should work towards the sustainability, protection, and restoration of Z. h. preblei populations and habitats on both private and public lands to provide the spatial, genetic, and demographic structure needed to promote long-term species viability and provide species management flexibility. Recovery efforts for the subspecies will be most effective if reliable information is available on the basic ecology of the subspecies and this information used to design recovery efforts such as Habitat Conservation Plans. A review of studies conducted on PMJM shows that there is insufficient information to fully address defining range-wide ecological requirements, .'. limiting factors, limits of species tolerance, .01: population status (Shenk 1998). Most work to date has ..... �44 focused on geographic distribution (presence or absence of Z. h. preblei), taxonomy, and habitat descriptions of sites where mice have and have not been captured. For PMlM in particular, information on dispersal, habitat use, and population dynamics is most needed to identify minimal ecological requirements of the subspecies. Monitoring, by way of estimating demographic parameters (such as survival, abundance, and reproduction), of a single population over time provides an OPPOl.1Unityto document demography of a species, estimate temporal fluctuations in demography, and gain insights into the temporal variation inherent in demographic parameters. Monitoring of multiple populations over time and over multiple geographic locations provides the opportunity to gain further understanding of population processes and detaches time effects from spatial effects. Population monitoring activities do not constitute scientific experiments, in the spirit of manipulation of salient ecological variables, however, replication of monitoring activities for natural populations over long periods of time and in diverse geographic locations can lead to insights into population processes (Cook and Campbell 1979). These insights can then be translated into hypotheses useful for predicting changes in population demography resulting from either natural perturbations (e.g., flooding events) or anthropogenic modifications (e.g., gravel mining). Experimentation would then be required to test these hypotheses and establish cause and effect. The primary objective of this study is to investigate spatial and temporal variation in the demography ofPMJM. Demographic parameters to be estimated include survival, reproduction, temporary emigration, immigration, population structure, and density. These demographic parameters will be estimated from individually marked animals from geographically distinct populations. To evaluate spatial differences in the demography ofPMlM, populations were selected from three sites that provided a variety of habitat matrices available to the mouse. Multiple years of Conducting the study at those same sites will provide an estimate of the temporal variation in demography ofPMlM. Study sites Demography ofPMlM was evaluated for three populations. All three populations selected were located in areas where PMJM had previously been found. To evaluate spatial differences in the demography ofPMJM, populations were selected on sites that provided a variety of habitat matrices available to the mouse. The first site selected, Colorado Division of Wildlife (CDOW) Maytag Property, has one primary water source available to the mouse. This water source is East Plum Creek. The second site, Colorado Open Lands PineCliffRanch, provides both a tributary (Garber Creek) and a main stem drainage (West Plum Creek). The third study site, CDOW Woodhouse Ranch provides an area containing a tributary (Indian Creek) and a series of ponds and irrigation ditches scattered throughout the property. Objectives Objectives of the demography study are to: 1. Estimate abundance and density of PMJM for each of three study populations. 2. Estimate over-summer, over-winter, and annual survival ofPMJM at each of three study populations. 3. Estimate temporary emigration of PMJM at each of three study populations. 4. Estimate immigration of marked PMlM back into each of three study populations. 5. Estimate reproduction ofPMlM at-each of three study populations. 6. Evaluate the affect of weight, sex, age, abundance (i.e., density dependent response), and habitat features such as stream reach, vegetation composition and density on survival, reproduction, abundance, temporary emigration, and immigration of marked animals back into three study populations of PMJM. 7, Estimate ..age and, sex ratios Of PMJM at each of three study populations .. : .. , �45 Methods Trapping Three trapping sessions were conducted during the following weeks: June 2-9, 1998; July 21August 5, 1998; and September 8-15, 1998. Trappers were advised to follow the Center for Disease Control's Hantavirus instructions and recommendations when dealing with rodents. Due to the nocturnal nature ofPMJM, traps were set between 19:00hrs and'21 :OOhrsand checked as early as possible in the morning beginning at 5:00hrs to reduce stress and the potential for predation on trapped animals. Time required to complete the traplines varied depending on how many animals were caught. Small mammal Sherman live traps (folding and non-folding) were used to conduct the trapping sessions. Traps were set in two parallel lines of trap stations (1 trap per station) on either side of the drainage. Trap stations were 5 meters apart for a total of 250 meters; the parallel transects were 10 meters apart unless extent of habitat, terrain topography, or stream hydrology did not allow. Location of transects were recorded on field data sheets. Each trap location was also recorded to the nearest 5 meters in UTM coordinates using a Trimble Geo-Explorer GPS. A small (-1 inch) ball of polyester quilt was placed in each trap as nesting/bedding material. Baiting material was Manna Pro Sweet 3-way Livestock feed which contains no animal matter; ingredients include flaked barley, flaked corn, flaked oats, and cane molasses. Peanut butter was used to stick the bait to the trap. Traps were checked by two surveyors. All animal captures were recorded. If an animal had been captured in a trap, a ziplock plastic bag was placed over the end of the trap .. The trap was opened allowing the animal to fall into the plastic bag. The animal was identified to species while in the plastic bag. If the animal was not a PMIM, identification of the animal was recorded and the animal set free without further handling. Each captured PMIM was checked to detect the presence/absence of a PITtag and/or radio-collar. If a PIT-tag or radio-collar was detected and the mouse had been captured at that same site within the same trapping session, identification of the mouse was recorded and the mouse was released. If the animalwas aPMJM and no PIT-tag or radio collar was detected the PMIM was anesthetized for further processing, as follows. All PMIM were PIT -tagged. If the PMIM weighed> 18 g a radio-collar was also put on the animal for movement data collection (see Shenk and Sivert 1999). Each PMIM was weighed while in the bag, recording the weight in grams using a Pesola spring balance. Sex, age (juvenile, adult), and reproductive condition (pregnant, lactating or nonbreeding if it was female; for males the position of the testes indicated if in breeding status or nonbreeding status) was noted. Each PMIM was measured (total length, length of body, length of hind foot - heel to distal end of claws) in millimeters. Capture of every PMIM was documented by taking a photograph of the mouse on first capture. Mice were placed in a plexiglass photobox to reduce handling stress to take a photograph. If the mouse had been captured previously as noted by the detection of a PIT-tag, no photograph was taken. If there were feces in the trap where a PMJM was captured, the fecal material was collected in a plastic bag labeled with date, location of the site, and animal PIT-tag number. Fecal samples were kept cool until returned to the CDOW office where they were frozen. Salt was added to each specimen bag for preservation. Once the field season was over all fecal samples were analyzed by the Composition Analysis Laboratory, Inc, 622 Y2 Whedbee, Fort Collins, CO for content. These analyses were for a complementary study on food habits and movements ofPMIM at the same three study areas (see Shenk and Sivert 1999 for details). All trap mortalities were recorded on a 'Trap Mortality' data form which included information on species, potential duration of time spent in the trap, and any information available to help determine cause of death. All animals found dead were double-bagged in a plastic bag and placed in a cooler with ice. Specimens were frozen as soon as possible and deposited in the CDOW freezer at the office in Fort CoIlins.. A museum card was.completed and attached-to each PMJM specimen and the . . . _". �46 specimen and card were given to Cheri Jones, Curator ofMammalogy at the Denver Museum of Natural History, for study skins and tissue storage. If an animal was severely injured (e.g., severed limb, large lacerations) it was euthanized by soaking a cotton ball in Metofane (methoxyflurane) and placing the cotton ball and the mouse in a ziplock bag until the animal stopped breathing. If an animal appeared to be only slightly injured (e.g., broken tail, small laceration) the animal was released to the wild.~ If an animal appeared to be coldstressed attempts were made to warm it by holding it in the surveyors hands and/or against their body. If the animal appeared to be heat stressed, isopropyl alcohol was applied with a cotton swab to the ears, arm pits, and feet to cool it down. During the last trapping session homeopathic first aid treatments were used in an attempt to minimize stress and improve recovery of each PMJM. Homeopathic first aid kits and instruction was provided by WildAgain Wildlife Rehabilitation, Evergreen, Colorado. PIT-tagging Each PMJM was individually marked with a Passive Integrated Transponder (PIT -tag). PITtags are electromagnetic, glass-encased tags that emit a passive signal (125 kHz) that can be decoded by a portable reader. Destron-Fearing PIT-tags were used on all mice. We used portable readers from .Biomark to read the PIT-tags. All mice were anesthetized to PIT -tag them. Anesthesia procedures followed protocols successfully used on PMJM before (R. Schorr personal communication). Mice were anesthetized by placing them and a cotton ball with 1 ml of Me tofane (methoxyflurane) into a sealed ziplock bag (to keep Metofane fumes in bag). The bag was then lain aside to minimize stress for the mouse and the time the mouse struggles in the bag. The less agitated the mouse is in the bag the less time required for the metofane to take affect. The mouse was observed at all times to make sure the mouse did not position itself such that the cotton ball was in direct contact with its face, which might cause the mouse to have a severe reaction to the anesthesia After the animal stopped moving, the handlers waited one minute before removing the mouse from the bag. The animal typically remained anesthetized for 2-3 minutes once outside the bag. Each PMJM was PIT -tagged using a protocol successfully used by C. Meaney (personal communication). Each PMJM had a PIT-tag inserted above the shoulder blades by lifting the skin on its back and inserting the individually sterilized needle with the PIT -tag under their skin and injecting the tag. Verification of the PIT-tag identification number was made before and after insertion into the mouse by running the PIT -tag scanner across it. The skin behind the opening was then pinched to prevent emergence of the tag. PIT-tag identification number was recorded on the field data form. If, at any time during the handling of the PMJM the animal appeared to be severely stressed (dramatic changes in heartbeat, respiration and responsiveness or gums turning blue) first aid was administered and, once recovered, released without further processing. Genetic tissue sampling '. . . Genetic tissue samples were collected from the first 30 PMJM captured at each study site. The following protocol was used based on previous success with PMJM (M. Bakeman, C. Meaney, T. Ryon, R. Schorr, personal communication). A fresh pair of clean latex gloves were used to handle each mouse. With a clean ear punch tool, one tissue plug was obtained from each mouse ear. If there was excessive bleeding, gentle pressure was applied to the injured area for approximately one minute. Each ear plug was placed in a vial of 95% ethanol. Both the vial and the corresponding record on the data sheet were labeled with a unique identifier as follows. A seven-place alpha-numeric code was composed as follows: a three-letter designator for the survey location (e.g., MYT = Maytag Property); a number beginning with two digits indicating the year.Ie.g., 98); followed by 2· digits specifying the individual trapped, numbered .. sequentially, FoI' example: MYT980J ~. first ~i~aI sampled ,at Maytag Propertyin J 9~.8. �After returning from the field, all samples were put in a cool place or refrigerated until delivery to the CDOW office at 317 West Prospect, Fort Collins, CO where they are being held in the freezer for future analyses. Ear punch tools were cleaned after each use by immersing them in a 10% bleach solution for a few minutes, rinsing them thoroughly with clean water, and then dried thoroughly to prevent rusting. Dull ear punch were discarded and replaced with new sharp punches to ensure the quickest, cleanest cut possible. " Demographic parameter estimation Mark-recapture estimation techniques were used to estimate abundance, over-summer survival, over-winter survival, temporary emigration, and immigration of marked animals back in to the three study populations of PMJM. Abundance: Abundance was estimated using Pollock's Robust Design (see Kendall et al. 1997, 1995 and Kendall and Nichols 1995 for detail) in Program MARK (White and Burnham 1999). The robust design is a combination of the Cormack-Jolly-Seber (CJS)(Cormack 1964, Jolly 1965, Seber 1965) live recapture model and the closed capture models. The key difference from the CJS model is that instead of just one capture occasion between survival intervals multiple capture occasions are used. These occasions are close together in time allowing the assumption that no mortality or emigration occurs during these short time intervals. The closely spaced encounter occasions are termed "trapping sessions" and each trapping session is viewed as a closed capture survey whereby abundance can be estimated. Three 7-day trapping sessions were conducted during the following weeks: June 2-9, 1998; July 21-28, 1998; and September 8-15, 1998. Abundance estimates were calculated for each trapping session. Survival: By using the estimate of the probability that an animal is captured at least once from the trapping sessions designed to estimate abundance, survival between the longer intervals was estimated. Analyses were conducted to estimate over-summer survival, and survival from June-July and August-September. A fourth 7-day trapping session to be conducted June 1-8, 1999 will provide information to estimate over-hibernation and annual survival. Temporary emigration and immigration: The longer intervals between trapping sessions also allows estimation of temporary emigration from the trapping area, and immigration of marked animals back to the trapping area using Pollock's Robust Design. Reproduction: Reproductive parameters were estimated by evaluating the reproductive status and age of captured PMJM. Results Trapping and PIT-tagging Effort A total of 186 individual PMJM were captured from the three study areas during the three 1998 trapping sessions -(Table 1: 73 at Maytag Property, 77 at PineCliffRanch, and 36 at Woodhouse Ranch). These mice were captured over three different trapping sessions with an effort of 17,330 trapnights. Every PMJM captured was PIT-tagged, providing each mouse with a unique, permanent, lifetime identification marker. Other small mammals captured during the trapping sessions included Peromyscus spp., Mexican wood rat (Neotoma mexicana), house mouse (Mus musculus), vole (Microtis spp.), western harvest mouse (Reithrodontomys megalotis), hispid pocket mouse (Chaetodipus hispidus), and shrew (Sorex spp.). House mice were captured at all three study areas during either June or September (Tables 2,3). Species and relative numbers of captures within each of those species were similar for Maytag Property and PineCliffRanch (Tables 2,3). Species richness and relative number of captures between these sites were also similar in June and September with the exception of an increase in vole captures at both sites in September. Most of these common species were also captured at _Woodhouse Ranch during both }un_~and September (Tables 2,~): ..HC)."",ey~r,. hispid pocket mice !3I1~ ".: "." �48 Mexican woodrats were also captured at Woodhouse Ranch and not at either of the other two sites. Numbers of house mice and vole captures at Woodhouse Ranch were greater than at either Maytag or PineCliff during June and September, with numbers of captures for both species increasing in the September trapping session (Tables 2,3). Age and Sex Ratios As expected, no juvenile PMlM were captured during the June trapping session as it was too soon after hibernation for any litters to have been born and/or if born the juveniles would still be restricted to their nests. Juvenile mice were captured during both the July and September trapping sessions at all three sites. Juveniles as small as 109 were captured. Sex ratios may be biased towards males, however further analyses need to be completed to confirm this hypothesis (Table I). Genetic tissue sampling Genetic tissue samples were collected from 40 individual mice at Maytag, 30 mice at PineCliff Ranch, and 32 mice at Woodhouse Ranch. Additional samples were collected from four mice captured at a back drainage on the Woodhouse Ranch. Samples are currently being stored in the cooler at the CDOW Research Center in Fort Collins, Colorado. A Requestfor Proposals was submitted through the Colorado Department of Natural Resources. A committee is currently evaluating the three proposal that were submitted. Samples from these three populations will be used in studies to evaluate genetic uniqueness ofPMlM populations. Abundance and Density Mark-recapture analysis methodologies were used to estimate abundance at each of the three study sites for each of the three trapping sessions (Table 4). Abundance estimates and length of the .trapline were used to estimate an unadjusted density of mice per kilometer of stream stretch. Mice that typically don't use the area of stream stretch where the traps were placed could be trapped because they were attracted to the area by the artificial food source. Including these mice in the density estimates for the stream stretch covered by the trapline would artificially increase density estimates. Therefore, an adjustment was made to the density estimates. Locations of each individual mouse were classified as either within the stream stretch that was trapped or above or below the stream stretch where traps were placed. If greater than 50% oflocations of an individual mouse were off the trap line the mouse was categorized as an 'off-trapline' mouse; if less than or 50% of the locations were within the trapline the mouse was categorized as an 'on-trapline' mouse. The percent of off-line mice for each trapping session and each site was calculated and a mean adjustment parameter (P) estimated. This adjustment parameter is sensitive to the length of the trapline. The shorter the trapline the greater the adjustment would have to be. Regression models were run and AlC used to select the best regression model to estimate p by trapline length. These adjustment parameters are simply an estimated percent of mice that should be considered resident mice along the stream stretch where traps were placed. The density estimates were then decreased to percentage p. Use of the adjustment parameter provided a more realistic estimate of density of mice per kilometer of stream stretch. Without the adjustment densities would be inflated. Survival Survival rate was estimated using the mark-recapture estimator for Pollock's Robust Design in Program MARK. The best fitting model combined data from all three study sites over all three trapping sessions. Survival was estimated as 0.75 (se = 0.033) for a five week period. Extrapolating this survival rate across the summer results in an over-summer (June 1- October 5, 18 weeks) survival rate of 0.355 (se = 0.056). No estimates of over-winter survival can be made until trapping sessions .occur in Sillllmer.l9.99., ,. -;' ;. . . .' ' " '_.. .' . '., :... . .~." . �49 Temporary emigration and immigration Temporary emigration rate and immigration was estimated using the mark-recapture estimator for Pollock's Robust Design in Program MARK. The best fitting model combined data from all three study sites over all three trapping sessions for these two parameters. Temporary emigration was estimated as 0.64 (se = 2.21), immigration was estimated as 0.45 (se = 89.94). It; Reproduction No estimates of reproduction could be made. We located what we believed to be nest sites at several locations. We believe these sites to be nest locations with young because we followed radiocollared adult females (see Shenk and Sivert 1999) to these sites on numerous morning during breeding season. However, all sites were in very thick vegetation and we were not able to visually observe a nest with young. We felt it to be in the best interest of the mouse to not disturb the areas which might possibly cause failure of the nest. Discussion Information on the population dynamics ofPMJM is necessary to determine which areas support viable populations. To begin to evaluate the viability of a population information on key demographic parameters must be obtained. Conducting studies on individually marked animals provides the greatest insight on the demography of a population. In general, estimated sex ratios from this study are comparable to those found by Armstrong et al. (1997) who reported an overall sex ratio for all captured PMJM of 51.6 males: 48.4 females; approximately 86.0% of captures were identified as adults. There is a possible male sex bias at PineCliffRanch. However, small sample sizes and possible trapping biases by sex may explain the discrepancy. . Density estimates were expected to increase as the summer progressed to account for the birth pulses in late June, and late July-August. This was the trend observed at Maytag. PMJM densities at Woodhouse Ranch increased from June to July but did not increase further during the September trapping session. Mean PMJM density over all sites and sessions was 40.5 (range 16.9-79.0) mice per kilometer of stream stretch. Highest densities occurred at PineCliff, where the vegetation is primarily willow and both a mainstem and tributary were used by mice. Lowest densities occurred at Woodhouse Ranch where the riparian vegetation has fewer willow but was dense with other riparian vegetation. Vegetation at Maytag provided some areas of dense willow with the remaining areas being of moderate density of riparian vegetation. The lower density ofPMJM reported for Woodhouse Ranch might be explained by the composition and densities of other small mammals at that site. Woodhouse Ranch had the highest captures of both house mice and voles of the three sites studied. Defining summer as June 1 - October 5, over-summer survival was estimated as 0.36 (se = 0.056) over all three study sites. Meaney et al. (1999) report a one-month summer survival rate of 78%. Extrapolating Meaney et al.'s (1999) estimate over our summer period (- four months) would result in a similar over-summer survival rate estimate of 36%. Prior to studies conducted in 1998 no information existed on survival rates for populations of Z. h. preblei although Whitaker (1963) reported a 67% loss of Z. hudsonius over hibernation. Temporary emigration and immigration rates were estimated but both had extremely high variances associated with those estimates. Thus these estimates cannot be used, with any confidence, to provide information on movements of mice into and out of these populations. Very little new information was gained during this study on reproductive parameters ofPMJM. Most adult mice captured exhibited evidence of active reproductive behavior, either pregnancy, lactation, or enlarged genitalia. However, we were unable to locate any breeding nests. Juvenile mice were not captured during the June trapping session. Juvenile mice were captured at all three sites - .. ';" .,._, during.both theJuly andSeptember trapping sessions, ..Given that breeding peaks .appear to,qc_<;(tu: in '. . . . '. .. '. " . ....•... ", . �50 early to mid-June and August with a possible third litter in September (Whitaker 1963) this was not unexpected and agrees with previous observations (Meaney et al. 1996, 1997, PTI 1996a, M. Bakeman unpublished data, T. Ryon unpublished data). This report includes results from only the first year of a multi-year project to follow individually marked PMlM through time. Collection of more data, and more years of data, will improve our ability to evaluate demographic parameters and estimates of how they vary across space and time. . :: ".": .'. Literature Cited Armstrong, D. M., M. E. Bakeman, A Deans, C. A Meaney, and T. R. Ryon. 1997. Conclusions and recommendations in: Report on habitat findings on the Preble's meadow jumping mouse. Edited by M. E. Bakeman. Report to USFWS and Colorado Division of Wildlife. Bailey, B. 1929. Mammals of Sherburne County, Minnesota Journal ofMammalogy 10:153-164. Bailey, V. 1923. Mammals of the District of Columbia Proceedings of the Biological Society. Washington 36:103-138. Cook, T. D. and D. C. Campbell. 1979. Quasi-experimentation: design and analysis issues for field settings. Houghton-Mifflin, Boston. Cormack, R. M. 1964. Estimates of survival from the sightings of marked animals. Biometrika 51:429-438. ERO Resources. 1995. Environmental review of South Boulder Creek Management Area Prepared for City of Boulder Real Estate/Open Space Department. Prepared by ERO Resources Crp., Denver, Colorado in association with Stoecker Ecological Consultants, Boulder, Colorado. Fitzgerald.J. P., C. A Meaney, and D. M. Armstrong. 1994. Mammals of Colorado. Denver Museum of Natural History, University Press of Colorado. Niwot, Colorado. Hafner, D. 1., K. E. Petersen, and T. L. Yates; 1981. Evolutionary relationships of jumping mice (genus Zapus) of the southwestern United States. Journal ofMammalogy 62:501-512. Hall, E. R. 1981. The mammals of North America. John Wiley and Sons, Inc., New York, New York, 2 volumes. Hamilton, W. 1., Jr. 1935. Habits of jumping mice. American Midland Naturalist 16:187-200. Jolly, G. M. 1965. Explicit estimates from capture-recapture data with both death and immigration stochastic model. Biometrika 52:225-247. Jones, C. A 1996. Mammals of the James John and Lake Dorothey State Wildlife Areas. Final Report, submitted to the Colorado Division of Wildlife and Colorado Natural Areas Program. Kendall, W. L., and 1. D. Nichols. 1995. On the use of secondary capture-recapture samples to estimate temporary emigration and breeding proportions. Journal of Applied Statistics.22:751762. Kendall, W. L., K. H. Pollock, and C. Brownie. 1995. A likelihood-based approach to capturerecapture estimation of demographic parameters under the robust design. Biometrics 51:293308. Kendall, W. L., 1. D. Nichols, and 1. E. Hines. 1997. Estimating temporary emigration using capturerecapture data with Pollock's robust design. Ecology 78:563-578. Krutzsch, P. H. 1954. North American jumping mice (genus Zapus). University of Kansas Publications, Museum of Natural History 7:349-472. Meaney, C. A, N. W. Clippinger, A Deans, and M. OShea-Stone. 1996. Second year survey for Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. Report prepared for the Colorado Division of Wildlife. Meaney, C. A, A. Deans, N. W. Clippinger, M. Rider, N. Daly, and M. O'Shea-Stone. 1997. Third year survey for Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. , ... '. Report prepared for. the Colorado Division of W.ildlife_....•..: .. "..' .... .'.... :,. " '.' .. ' ." . " .. ' .. . .. ;' -., '.- :,' : :.' .. . ~:' ' . .' '.' " . �51 Meaney, C. A., A. Ruggles, B. Lubow, N. W. Clippinger, and A. Deans. 1999. Preliminary results: Second year study of the impact of trails on small mammals and population estimates for Preble's meadow jumping mice on City of Boulder Open Space. Report for Greenways Program, City of Boulder Transportation Department, City of Boulder Open Space, and Environmental Protection Agency, Region 8. , Nudds, T. D. 1977. Quantifying the vegetative structure of wildlife cover. Wildlife Society Bulletin 5:113-117. Poly, W. 1., and C. E. Boucher. 1997. Record of a creek chub preying on ajumping mouse in Bruffey Creek, West Virginia Brimleyana 24: 29-32. PTI Environmental Services. 1996a Preble's Meadow Jumping Mouse Study at Rocky Flats Environmental Technology Site, Annual Report 1996. Final. Rocky Flats Environmental Technology Site, Golden, Colorado. PTI Environmental Services. 1996b. Preble's Meadow Jumping Mouse Study at Rocky Flats Environmental Technology Site, Spring 1996. Final. Rocky Flats Environmental Technology Site, Golden, Colorado. Quimby, D. C. 1951. The life history and ecology of the jumping mouse, Zapus hudsonius. Ecological Monographs 21:61-95. Riggs, L. A., 1. M. Dempey, and C. Orrego. 1997. Evaluating distinctness and evolutionary significance of Preble's meadow jumping mouse: Phylogeography of mitochondrial DNA noncoding region variation. Final Report for the Colorado Division of Wildlife. Denver, Colorado. Seber, G. A. F. 1965. A note on the multiple recapture census. Biometrika 52:249-259. Sheldon, C. 1934. Studies on the life histories of Zapus and Napaeozapus in Nova Scotia Journal of Mammalogy 15:290-300. Shenk, T. M. 1998. Conservation assessment and preliminary conservation strategy for Preble's meadow jumping mouse (Zapus hudsonius preblei). Colorado Division of Wildlife FY199798 Annual Report. Shenk, T. M. and Sivert M. 1999. Movement patterns of Preble's meadow jumping mouse (Zapus hudsonius prebleii as they vary across time and space. Colorado Division of Wildlife January - March 1999 Quarterly Report. Stohlgren,T. 1., M. B. Falkner, and L. D. Schell. 1995. A Modified-Whittaker nested vegetation sampling method. Vegetatio 117:113-121. USFWS. 1997. Interim survey guidelines for Preble's meadow jumping mouse. USFWS. Denver, Colorado. Whitaker.T, 0., Jr. 1963. A study of the meadow jumping mouse, Zapus hudsonius (Zimmerman), in cental New York. Ecological Monographs 33:3. • -." ...• ..... ...•. " •. ',~ ,. -'_. !., . ". ~.' ••.• ,': ..". '1 ••.•• .;. • . .,.... '" ~ • ~ . " ",: '" "':' ',,: .:. .... '..... _. :. . .. "', . ,_ ." '/: . ".: . ~. ~. ':'." �52 Table 1. Number of Preble's meadow jumping mice PIT -tagged and captured at each of the three study sites for each of the three trapping sessions. Trapping effort is recorded as number of trap nights for each session at each site. # PIT-tagged # Captured! Site Trapping session Trap nights Female Male Unknown Total Female (%) Male (%) Total Maytag Jun 1949 9 5 0 14 9 (64) 5 (36) 14 Maytag Jul 2370 13 13 0 26 15 (54) 13 (46) 28 Maytag Sep 3256 12 20 33 18(43) 24 (57) 42 PineCliff Jun 1095 6 13 19 6 (32) 13 (68) 19 PineCliff Jul 1603 7 8 15 8 (50) 8 (50) 16 PineCliff Sep 1544 12 31 43 13 (28) 33(72) 46 Woodhouse Jun 1525 4 3 7 4 (57) 3 (43) 7 Woodhouse Jul 2420 8 5 13 9 (64) 5(36) 14 Woodhouse Sep 1568 6 9 16 8 (44) 10 (56) 18 186 90 114 204 t Some o o o o o 17330 77 107 TOTALS 2 captured animals were PIT-tagged during previous trapping sessions Table 2. Number of other small mammal species captured during the June 1998 trapping session at Maytag Property, PineCliffRanch, and Woodhouse Ranch. Peromyscus ...~'. ',. ";" ," Reithrodontomys megalotis Harvest mice Date Site spp. Jun2 Maytag 27 Jun 3 Maytag 42 Jun4 Maytag 48 Jun 5 Maytag 34 4 1 1 Jun 7 Maytag 53 o Chaetodipus hispidus Hispid pocket mice Microtis spp. voles o o o o o 2 Jun8 Maytag 39 2 Jun 11 Maytag 41 2 Jun 12 Maytag 45 2 o 4 Jun 13 Maytag _ 36 o o o o o 3 Pine Cliff Jun 3 Pine ClifT 36 o o Jun 4 Pine ClifT 39 6 Jun 5 Pine ClifT 34 3 Jun 7 Pine ClifT 22 Jun 8 Pine ClifT 35 o 4 o 2 o o 2 Woodhouse 13 Jun 3 Woodhouse 23 o o Jun4 Woodhouse 21 1 Jun 5 Woodhouse 12 4 Jun 7 Woodhouse 13 1 Jun 8 Woodhouse 18 Jun 14 Woodhouse . )tih·15· -. Woodhouse 16 o o 7 o o :.. o o o o 2 o o o o 6 o o o o 3 o 1 10 o 3 . " 0 14 6 _...... 0 ·1 . ...... Mexican woodrat o o o o o o o o o o o o 4 2 3 o o o 5 0 9 Neatoma mexicana 1 o o o 7 3 3 .... 4 o o o 5 ·5 o o o o o o o Zapus hudsonius preblei PMJM o o o o o Jun2 o o o o o o o Jun2 Mus musculus House mice Sorex spp. shrews " . o o o 4 o o o o o o 5 o o o o 3 3 5 o o o o ',_ '", " " 't' �53 Table 3. Number of other small mammal species captured during the September 1998 trapping session at Maytag Property, PineCliffRanch, and Woodhouse Ranch. Reithrodontomys myscus megalotis spp. harvest mice Pero- Date Site Sep 9 Sep 10 Sep 11 Sep 12 Sep 13 Sep 14 Sep 15 Sep 16 Sep9 Sep 10 Sep11 Sep 12 Sep 13 Sep 14 Sep 15 Sep 16 Sep 9 Sep 10 Sep 11 Sep 12 Sep 13 Sep 14 Sep 15 Maytag Maytag Maytag Maytag Maytag Maytag Maytag Maytag Pinecliff Pinecliff Pinecliff Pinecliff Pinecliff Pinecliff Pinecliff Pinecliff Woodhouse Woodhouse Woodhouse Woodhouse Woodhouse Woodhouse Woodhouse 30 39 35 35 31 30 36 31 15 19 16 21 18 17 16 22 15 15 18 18 13 19 17 0 0 4 6 o 3 2 3 o o o o o o o Chaetodipus Microtis Hispid spp. pocket mice voles hispidus o o o o o o o o o o o o 10 9 17 16 2 4 7 7 o 11 o 8 12 16 47 60 70 73 67 80 73 1 4 4 8 o o o o o o o o o o 2 3 Mus Sorex musculus spp, house shrews mice ;..''1 o o o o 2 2 o o o o Zapus hudsonius preblei PMJM Neatoma mexicana Mexican woodrat 12 19 11 o 5 o o 8 o 11 2 9 o o o o o o o 8 8 7 2 6 o o o o 7 12 8 7 14 o 7 15 21 18 13 15 14 2 7 o o o o o o o 13 2 o o o o 11 o 2 o o o o o o 8 2 1 o o 6 3 Table 4. Stream reach abundance eN) and density estimates for Preble's meadow jumping mouse (PMJM) from three sites in Douglas County, Colorado for three trapping sessions. Both adjusted and unadjusted density estimates (PMJM per km of stream stretch) are reported. Adjustments were made to the density estimates to account for the positive bias introduced when animals are attracted to a trap line. 95% Confidence Site .... , . Trapping session Interval Lower Upper Trapline Length p (m) Unadjusted Density (PMJMIkm) Adjusted Density (pM1M/ km) Maytag Jun 18 2.7 15 21 550 32.7 0.80 26.0 Maytag Jul 31 11.4 20 42 608 51.0 0.81 41.4 Maytag Sep 44 1.8 42 46 494 89.1 0.78 69.3 PineCliff Jun 30 5.7 24 36 490 61.2 0.78 47.5 PineCliff Jul 17 0.9 16 18 504 33.7 0.78 26.3 PineCliff Sep 51 3.3 48 54 504 101.2 0.78 79.0 Woodhouse Jun 11 3.8 7 15 510 21.6 0.78 16.9 Woodhouse luI 20 5.2 15 25 516 38.8 0.79 30.4 ~W~ood~~ho~u~se~~~S~e~p~~~2~0~~73~.~I~ __ ~17~~~2~3~ '.. . . ~5~74~__ ~ __ 3~4~.8~ ~0~.8~0~ __ ~2~8~~ �54 �55 Colorado Division of Wildlife Wildlife Research Report July 1999 JOB PROGRESS REPORT State of Colorado Project No. . W-153-R-12 Work Package No. --"0-=.66=3"-Task No. Period Covered: ~1,-- _ _ Cost Center 3430 Mammals Program Species of Special Concern/Species Conservation Kit fox Conservation at Risk July 1, 1998,. June 30, 1999 . Author: T. D. I. Beck Personnel: T. Beck, D. Coven, P. Creeden, V. Graham, M McLain, P. Schnurr, B. Sommerville, L. Willmarth; CDOW ABSTRAcr Biological assessment work was expanded to the Colorado River Valley in this segment. Fiftyfour cameras, triggered by active-infra-red sensors, were deployed throughout a nearly 800 km2 area for 12-30 days each. No photos of kit fox were obtained. Surveys of kit fox dens active in 1998 found only one den active in 1999; however, the pair apparently did not have any young again this year. A combination of GAP mapping and ground mapping was used to estimate the habitat potential for restoration of kit fox in western Colorado. Three distinct areas were identified: the Colorado, Gunnison, and Uncompahgre Valleys. The Colorado valley is isolated from the other two by urban development and irrigated agriculture. The Colorado River Valley is the largest contiguous area (685 knr') and has the highest percentage of public land. Thus, it is likely the best area for kit fox restoration efforts. Successful restoration of kit fox throughout the entire 1310 km2 of suitable kit fox habitat in Colorado could result in kit fox populations varying from 182-728; based on densities reported in the literature. Primary negative factors to be addressed are housing developments, greater isolation of habitats, and red fox pioneering into salt desert shrub communities. �56 �57 KIT FOX (VULPES MACROTIS) STATUS IN COLORADO Thomas D. I. Beck P.N. Objective Develop and implement a conservation strategy to recover and conserve kit fox populations in Colorado. Segment Objectives 1. 2. 3. Conduct the biological assessment work necessary to develop an effective conservation strategy for the conservation of kit fox in western Colorado. Survey for presence of kit fox in Gunnison and 'Uncompahgre Valley areas with infra-red activated cameras. . Capture and radio-collar adult and juvenile kit fox in both the Uncompahgre and Gunnison valley areas. METHODS AND MATERIALS Plans for field work were modified significantly in July in response to the dearth of kit fox activity found in the Gunnison and Uncompahgre Valleys. Additionally, much of the .landscape in the 2 valleys are either marginally suitable or becoming unsuitable through a combination of natural features and human development. Therefore, emphasis was shifted to better map the available habitat and conduct kit fox surveys in the Colorado River Valley. Ground searches for kit fox sign and dens were conducted in the Colorado River Valley on 40 days during September-November 1998. Remote 35mm cameras, activated by active-infra-red sensors, were deployed in areas with fresh kit fox sign, areas with old fox dens, and likely travel areas. The camera system was developed by TrailMaster (Lenexa, KS) and was essentially the same system as used by Beck on black bear (Ursus americanus) studies (Beck 1995). The distance from the camera to the infra-red transmitter was 2 m; camera height above ground was 15-30 ern. A pipe was driven into the ground 10 em in front of the infra-red transmitter and one of several baits (beaver meat, chicken flesh lure) were placed down in the pipe to minimize bird conflict. The recorder unit was programmed so that the infra-red beam had to be broken for 0.15 sec to be recorded (value ofP = 3) and only a 6 sec delay between successive pictures. Color print film, . ASA 400, in rolls of24 and 36 exposures were used. Cameras were left at locations for varying periods of 12-30 days, depending on area, work schedule, and need for cameras in other areas. Observations of an active kit fox den were conducted at dusk on 3 evenings in May 1999, in hopes of documenting presence of pups. Observations were made with binoculars or a night-vision scope at distances of 30 to 100 m. Historic den sites in the Uncompahgre Valley were visited in May 1999 to check for kit fox activity. Maps of areal extent of salt desert shrub communities were obtained from CDOW Habitat Section. Acreage of salt desert shrub and irrigated lands were estimated from the GAP maps. The land ownership of the primary habitat (salt desert shrub) was also estimated from the GAP maps. Onsite field mapping was conducted to better assess the utility of much of the salt desert shrub component on private land since much of this land has been undergoing significant housing development. Also, IS-minute quad maps were used for plotting camera locations and all fresh fox sign. �58 RESULTS AND DISCUSSION Biological Assessment Kit Fox Distribution and Abundance - Analysis of the work reported by Fitzgerald (1996) suggested that summer was the period of lowest trap success, and the majority of search effort had been conducted in the summer. Thus it was deemed prudent to resurvey much of the available kit fox habitat during spring and fall seasons, utilizing a different technology than live-trapping. Ground searches for kit fox dens and spoor were conducted during spring 1998 in the Gunnison and Uncompahgre Valleys. Similar searches were conducted in the Colorado River Valley during September-November 1998. Fifty-four cameras were placed for periods of 12-30 days. Two camera units were stolen. No kit fox were photographed at the 52 sites spread throughout the approximately 800 km2 Colorado River valley. No active fox dens were located during ground searches. A variety of wildlife was photographed: cottontail rabbits (158), jackrabbits (7), antelope ground squirrel (2), coyote (6), badger (6), rock squirrel (6), bobcat (1), chipmunk (1), pronghorn (1), magpie (31), owl (1), unk. birds (3). Frequent rains and cold temperatures resulted in the loss of38 images during the last 2 weeks of work because the plexiglass dust cover frosted over; causing a frosty blur on the photograph. In May, 1999 a dead male kit fox was recovered along a paved road in Colorado National Monument by DWM Paul Creeden. It was freshly killed when retrieved about 0430. The area is in the pinon-juniper vegetation community, approximately 500 m higher than the desert floor. About 3 hours was spent on a ground search in the area looking for any denning activity or other sign but none was found. It was not the normal vegetation or terrain for kit fox. Ail kit fox dens -active in 1998 were checked in May ·1999 for fox activity. The Montrose landfill den again had a pair of kit fox. This pair was observed on 3 evenings in May but there was no indication of pups. Volunteer observers who work at the landfill reported no pup activity during the summer. None of the other known kit fox dens checked had any indication of kit fox activity. Additionally, 3 of the red fox (Vulpes vulpes) dens active in 1998 which were located in the salt desert shrub community were also active in 1999. Prey Abundance - Comparative prey surveys were initially to be conducted in Fall 1998. However, based on the low numbers of kit fox located in Spring 1998 it was decided to expand general surveys to the Colorado River Valley instead. Also, preliminary mapping suggested the Uncompahgre and Gunnison Valleys provided only modest opportunity for kit fox restoration because of limited habitat. In addition, the Colorado Legislature debated a bill in the 1999 session which could have restricted the authority of the CDOW to implement kit fox augmentation. This bill (HB 99-1299) was passed in late April and signed into law in May. However, final wording did not cover the kit fox program (in contrast to original language ). However, it was decided not to do extensive prey base work pending resolution of this issue. Potential Habitat The western valleys of Colorado which historically supported kit fox were divided into 3 distinct valley segments: Uncompahgre, Gunnison, Colorado. The Gunnison Valley area was subdivided into 3 segments because the central segment is a region with basalt cap rock close to the surface and den sites are extremely limited. Even badger diggings are uncommon in this reach; which is also the area where 3 roadkill kit fox have been documented in the past 5 years. �59 Based on the GAP vegetation maps, there is an estimated 547 krrr' of salt desert shrub in the Uncompahgre Valley unit (Fig. 1). This unit also has the highest area of irrigated farm land (528 km'), all of which historically was salt desert shrub. It also has a high proportion of existing salt desert shrub in private land status (45.8%). Ground mapping was conducted to exclude peripheral areas of salt desert shrub as well as areas which have been developed for human housing. Radio telemetry work on kit fox in this valley strongly indicated the kit foxes did not venture into the developed desert areas but remained in a relatively small core area. Thus, the suitable kit fox habitat for the near future is more likely about 140 krrr', This area is separated from the Gunnison Valley habitats by agricultural land, housing developments, a highway, and a river; resulting in a 7-9 km long obstacle course to inhibit dispersal. Based on the range of kit fox densities summarized by White and Garrott (1997) (0.160.7 /km') this area of habitat would likely only support 22-98 kit foxes. The Gunnison Valley area has approximately 609 km2 of salt desert shrub vegetation (Fig. 2). Only 2 kit fox were located in this region in 1998, 5 were trapped here during 1993-1995, and 3 road kills have been documented. The East Gunnison area (179 krrr') only has about 18 km+of irrigated land but does have some residential development and the irrigated land may serve as a barrier to animals moving east of Surface Creek. Thus, suitable kit fox habitat for recovery is closer to 120 km", of which 66% is in public ownership. The Central Gunnison area (250 krrr') has about 10 km2 of irrigated farmland. This is the area of volcanic rock near the surface which appears to severely limit underground dens. Most of this land is in public ownership (87%) and housing development is less a problem here. Thus, about 225 km2 of land is marginally suitable for kit fox recovery. Because of the lack of den sites for escapement, this area will likely always be a mortality sink for neighboring populations of kit fox. A small area near Cheney Reservoir has the more typical soil formations found in the salt desert shrub areas and did support a kit fox family in 1994-95. No kit fox activity has been recorded there since. The area is private land and the current owner has been unsuccessful at attempts to subdivide for housing, primarily because of issues of potable water supply. Unsolicited information from local varmint hunters suggests that red fox have been colonizing this area rapidly during the past 2 years. Earlier work on kit fox surveys in this area did not document red fox but both dens and red foxes were observed here in spring of 1998. The North Gunnison area abuts the Colorado River and Grand Junction urban area to the north. This area is highly impacted by housing development throughout the salt desert shrub type. Based on the GAP maps, there are 180 km2 of salt desert shrub in this unit, of which 48% is private. There is an additional 59 km2 of irrigated farm land. No kit fox have been located in this area by any of our surveys this decade. Suitable area for kit fox restoration in the near future is probably closer to 140 krrr', Total suitable habitat for kit fox in the Gunnison Valley is approximately 385 krrr', It is unlikely this area would support high densities of kit fox because of the denning problem in the Central area Thus, a projection of 50-150 kit fox possible should restoration be successful seems to be the best possible outcome. The housing development in the north coupled with the den problems in the central suggest that the East unit may be the best area for kit fox populations, which could then support 25-85 kit fox. The Colorado River Valley supports the largest intact area of salt desert shrub, about 797 km2 based on the GAP map. Most (85%) is in public ownership. Irrigated lands amount to 319 krrr', mostly in the eastern end (Fig. 3). Ten kit fox were captured here in 1994 and 1995. An active den with a single kit fox was discovered in 1997 near Fruita However, the lack of any photographs of kit fox or sign at historic kit fox dens in 1998 strongly suggests the current density of kit fox in this valley is low. This unit appears to have the lowest probability for loss of kit fox habitat and will likely have at least 685 km2 of suitable habitat for a long period. Based on kit fox densities reported in White and Garrott (1997), this area could potentially support 110-480 kit fox if fully occupied. �60 Capture Efforts It was decided not to trap and collar adult foxes during the whelping season because of the added stress of this activity to the individual foxes (Cypher 1997). The small number of kit foxes present seems to justify minimum intrusion during this period. There have been no juveniles located in 1998 or 1999 to tag. Literature Cited Beck, T. D. I. 1995. Development of black bear inventory techniques. Colo. Div. Wildlife, Fed. Aid. Rep. W-153-R-8. l lpp. Cypher, B. L. 1997. Effects of radiocollars on San Joaquin Kit Foxes. J. Wildl. Manage. 61(4):14121423. Fitzgerald, J. P. 1996. Status and distribution of the kit fox (Vulpes macrotis) in Western Colorado. Colo. Div. Wildlife, Final Fed. Aid. Rept. W-153-R-7. 78 pp. White, P. J. and R A. Garrott. 1997. Factors regulating kit fox populations. Can. J. Zool. 75:19821988. �F Pl (l) Pi ...•. '"t) (l) (JQ ~ ..., "0 3 0 () c::: ::l (l) s- s· (l) (/) ~ o· ::l (l) ...•. ~ (JQ (l) < 0.. (l) ...•. (l) () !:P- CZl (l) ~ ciQ' c:: ..., c 8lC;::l. :r !.!. Ii 110 [ en Co) ::J ~i i"'. ao"t:l "'" ~ ~ (II iil or CD ~ 1= ~~ ~11(ll1I I1-Ii ,,(11 &~~ g~ o ~~(/) Ci)(/)~ iil~C/) I)) CD . CD' is: ::: ~ l8ao ~g'[ ~~.i5' • ~qg o 1;! 00 00 "c: >"' 50 ""' O~m ('» I I~z ~" / I ) ( ) ~ r-, "\ I I 0\ - �N 0'1 A N " ,,'SubUnit Boundaries Q IrrigatedCrop Type _ Sefected Vegetation Types· * Desert Shrub Saltbush Fans & Rats Greasewood Fans & Flats o I 1 2 Kilometers ~ o 1 2 ~ 3 4 Miles ! R GREAT OUTDOOIU Co LORAno • Colorado Division of Wildlife May, 1999 Figure 2. Selected vegetation types in the Gunnison River Valley. DELTA �N A ~-. o Irrigated Crop Type _ Selected Vegetation Types· Desert Shrub Saltbush Fans & Flats Greasewood Fans & Flats \ v---........_\.}""~ o , 2 Kilometers ~ o , i --! 2 GREAT 3 4 Miles R OUTDOORS. Co LORADO ., <,-, J ~ Figure 3. Selected vegetation types in the Colorado River Valley. 0- W �64 �65 Colorado Division of Wildlife Wildlife Research Report July 1999 JOB FINAL REPORT Smteof ~C~o~lo~r~ad~o~ _ Cost Center 3430 Mammals Program Project No. __ --"W_,_....•. 1=53:::....-..:o.;R::....-.o..!12=--_ Work Package No. _0=6=6~3:._ _ Species of Special Concern Task No. ---=2'--_ Lynx ConservationlRecovery Period Covered: July I, 1998 - June 30, 1999 Author: D. F. Reed ABSTRACT Snowshoe hare pellet counts (Krebs plots) were analyzed and a Division report prepared. �66 �67 LYNX CONSERVATIONIRECOVERY DaleF. Reed P.N. OBJECTIVE Task 2 - Develop a conservation strategy to recover and conserve lynx populations in Colorado. SEGMENT OBJECTIVES 1. Analyze data and prepare report. STUDY AREA The study area includes the forests (Aspen, Douglas Fir, Lodgepole Pine, mixed conifer, mixed forest, Ponderosa Pine, and Spruce-fir) and deciduous oak above 7,500 ft throughout the north-south western half of Colorado as reported by Reed (1998). METHODS AND MATERIALS The methods used in assessing potential habitat for lynx (Felis lynx) involved counts of snowshoe hare pellets (Kreb' s plots; Krebs et al. 1987) via randomly selected points ill forest types and deciduous oak as determined by statewide GAP data and as reported by Reed (1998). RESULTS The results are in the report titled "Snowshoe hare density/distribution reintroduced lynx in Colorado" (Reed and Kindler 1999). LITERATURE Krebs, C. L, B. S. Gilbert, S. Boutin, and R. Boonstra. from turd transects. Can. J. Zool. 65:565-567. Reed, D. F. 1998. Lynx conservation/recovery. estimates - potential habitat for CITED 1987. Estimation of snowshoe hare density Colo. Div. Wildl. Res. Rep. July, 107-112pp. Reed, D. F., and l Kindler. 1999. Snowshoe hare density/distribution estimates - potential habitat for reintroduced lynx in Colorado. Fort Collins, CO :Colo. Div. ofWildl. Division report; no. (in review). Prepared by _ Dale F. Reed �68 �69 Colorado Division of Wildlife Wildlife Research Report July 1999 JOB PROGRESS REPORT Smteof ~C~o~lo~r~a~do=_ _ Project No. -'W"'--'-1'-"5~3~-R~-,,!_1 =-2 _ Work Package No ..__ -=-0.=;,;88::;.,;:0;,_ _ Task No. -=1 _ Period Covered: Mammals Research Black-footed Ferret Recovery Monitoring and Managing Disease in Black-footed Ferrets July 1, 1998 - June 30, 1999. Authors: .M. A. Wild and K. T. Castle Personnel: E. Wheeler. ABSTRACT The black-footed ferret is a federally listed endangered species in the United States, BIackfooted ferrets have been extirpated from Colorado, but are scheduled to be reintroduced to the wild at the Little Snake Management Area (LSMA) in Moffat County, Colorado in the near future. Our research can be sub-divided into two broad sections: disease monitoring in the proposed release area and flea control as a tool to manage sylvatic plague in prairie dogs and black-footed ferrets. Disease .monitoring was performed using collection of carnivores (primarily coyotes) from the LSMA in July 1998 and February 1999. Two potentially devastating diseases, canine distemper and plague, are present at LSMA. Prevalence of titers to canine distemper virus (CDV) in coyotes are low (::::10%). Prevalence of titers to plague (Yersina pestis) were found in 60% and 20% of coyotes tested in the summer and winter collections, respectively. Titers were present in both adult and juvenile animals, suggesting ongoing plague activity in some areas. We began investigations into the bioavailability of lufenuron to orally dosed prairie dogs. A test dose of 300 mg/kg body weight was determined and administered to 30 captive prairie dogs. Prairie dogs were divided into two groups (one group torpid, the other non-torpid). Blood samples for lufenuron assay were collected pre-treatment, at 1 wk postdosing, then at 2~wk intervals through week 9 post-dosing. No adverse affects to lufenuron were observed. Lufenuron assay is pending. We also attempted to establish a colony of fleas of the genus Oropsyl/a for artificial infestations on captive prairie dogs. Although we were unsuccessful in establishing self-sustaining colonies of 0. tuberculata (the prairie dog flea) from wild populations, we did establish a thriving Colony of insectary-reared 0. montana (the ground squirrel flea). These fleas will be used in future investigations into the efficacy of lufenuron to control fleas on captive prairie dogs .. BDOW014186 �70 �71 MONITORING AND MANAGING DISEASE IN BLACK-FOOTED FERRETS Margaret A. Wild and Kevin T. Castle P. N. OBJECTIVES 1. Monitor enzootic disease activity threatening survival of black-footed ferrets reintroduced into the LSMA. 2. Develop techniques to manage plague in the LSMA using insect growth regulators applied orally to prairie dogs. SEGMENT OBJECTIVES 1. Determine the level of activity of canine distemper virus in carnivores and plague in prairie dogs (using coyotes as sentinels) in the LSMA by sampling >40 coyotes. ~. 2. Establish a colony of captive white-tailed prairie dogs that can sustain infestations ~eas__of..the genus Oropsylla. 3. Develop techniques to artificially rear fleas of the genus Oropsylla. 4. Determine an effective dose ofpyriproxifen and oflufenuron to control fleas of the genus Oropsylla on prairie dogs. METHODS AND MATERIALS Carnivore Disease Survey Infectious diseases can severely impact the success of black-footed ferret (Mustela nigripesj reintroduction efforts. As part of the black-footed ferret reintroduction protocol, we monitored disease activity in carnivores in the Little Snake Management Area (LSMA), Colorado. Coyotes (Canis latrans) were collected.in July 1998 and February 1999. Post-mortem examination and sample collection was performed as described in the Program Narrative (Wild 1998). Black-footed Ferret Reintroduction We assisted in preparation of the black-footed ferret allocation request submitted to the US Fish and Wildlife Service by the Colorado-Utah black-footed ferret recovery working team in 1999. We prepared a protocol for the care of captive black-footed ferrets at the LSMA (Attachment 1). Consultation on health matters and veterinary care were provided for captive black-footed ferrets. Artificial Rearing of Oropsylla fleas In summer 1998 and spring 1999 we collected Oropsyl/a spp. fleas from prairie dog burrows at Arapaho National Wildlife Refuge. We attempted to maintain and propagate these wild fleas under �72 insectary conditions (Wild 1998). Additionally, we attempted to produce self-sustaining populations of 0. tuberculata within the nest boxes offour quarantined prairie dogs. To enhance flea survival, we placed a wooden box fitted with a screen top into each nest box. The wooden "flea refuge" allowed fleas to jump on or off the animal at will, and kept the prairie dog from stepping on fleas and eggs. We placed "bed-o-cobs" bedding material on the refuge floor in order to provide hiding places for the fleas. Each prairie dog was infested with up to 60 adult fleas (about 3:2 females:males). To count fleas, we anesthetized and groomed each animal periodically, and noted the number of surviving fleas found on the animal and in its nest box. Because of the decline in o. tuberculata availability in the field and the low survivorship and reproduction of that species in the lab, we decided to explore the use of laboratory-reared 0. montana for experimental purposes. 0. montana is generally found on ground squirrels, such as Spennophilus beecheyi (California ground squirrel); it should be able to reproduce when fed on the closely-related prairie dogs. Other researchers have maintained lab colonies of 0. montana for a number of years, and have found that the species readily reproduces when fed in artificial systems, and when fed on captive neonatal or adult rodents. While we would rather utilize true prairie dog fleas, such as O. tuberculata, in our experiments, the use of lab-reared 0 montana offers some advantages over 0. tuberculata: 1) 0. montana is readily available; 2) the temperature and humidity preferences of the species are wellcharacterized, so it reproduces well in an insectary; and 3) lab-reared 0. montana are plague-free. In summer 1999 we acquired a small colony of about 400 O. montana from the Centers for Disease Control (CDC), Fort Collins. These fleas were maintained under insectary conditions (Wild 1998). Flea Control In Prairie Dogs Thirty-three white-tailed prairie dogs (Cynomys leucurus) collected in June 1998 from Arapaho National Wildlife Refuge were maintained at Foothills Wildlife Research Facility (FWRF). Details of animal husbandry and health are reported by Wild (1999). Initially, we planned to evaluate two insect growth regulators in captive prairie dogs: lufenuron and pyriproxyfen. However, after reviewing the literature, contacting product manufacturers, and investigating methods to measure efficacy, we have determined that lufenuron holds the most promise for effective application. Lufenuron is longer lasting than pyriproxyfen (palma et al. 1993, Rink et al. 1994), researchers with Novartis (the manufacturer oflufenuron) are interested in collaboration, and a blood assay is available to measure lufenuron but not pyriproxyfen. We performed two studies to begin evaluating lufenuron in white-tailed prairie dogs. First, a pilot study to determine bioavailabilty oflufenuron administered orally to captive prairie dogs at three dosages: 20 mg/kg, 60 mg/kg, and 300 mg/kg (n = 1 at each dosage). Blood samples were collected for lufenuron assay pre-treatment and 1, 7, 28, 42, and 56 days post-dosing. Lufenuron assay using HPLC was performed by En-Cas Laboratories. Results of this pilot study were used in planning the following study to determined the seurm profile of lufenuron during active and torpid periods in orallydosed white-tailed prairie dogs: Bioavailability of lufenuron administered orally to captive white-tailed prairie dogs (Cynomys leucurus) Kevin T. Castle', Margaret A. Wildl, and S. Craig Parks' 'Colorado Division of Wildlife, 317 W. Prospect, Ft. Collins, CO 80526 2Novartis Animal Health, P. O. Box 26402, Greensboro, NC 27404 �73 Introduction Mortality of black-footed ferrets (Mustela nigripes) from infection with sylvatic plague is extremely high (Williams et al., 1994). Further, mortality from sylvatic plague in prairie dogs (Cynomys spp.) can markedly impact the prey base of black-footed ferrets. Plague has severely hampered reintroduction efforts in other states (p. Marinari, pers. comm.) and preliminary data suggest ...that plague activity is present in some areas of the Little Snake Management Area (Wild, unpub. data) where black-footed ferrets are scheduled to be released in late 1999. Successful reintroduction will likely require management techniques to controlplague, We believe that the proposed research will provide a novel and effective means of controlling plague in prairie dogs and black-footed ferrets. Plague is maintained primarily in rodent populations, where it is typically transmitted via the . bite of an infected flea Fleas of the genera Opisocrostis and Oropsyl/a have been associated with plague transmission in prairie dogs (Ubico et al., 1988). Insecticide dusts have been applied to prairie .. dogs burrows in an attempt to kill fleas and thus control plague. Carbaryl has been the most commonly used insecticide; however, application is labor intensive and activity is short-lived (Barnes, 1993). Permethrin has been shown effective up to 84 days after application (Beard et al., 1992), but the authors warn that application at the recommended dosage rate would be highly laborious. : Recently developed compounds used to control fleas in pet animals offer a promising alternative to insecticide dusts. Although topically applied flea adulticides, such as imidacloprid, are not feasible for use in wildlife, insect growth regulators (IGRs) with ovicidal and/or larvicidal activity could be delivered orally to free-ranging animals via bait. The IGR lufenuron is a benzoylphenylurea derivative which inhibits formation of chitin in the exoskeleton of insects (Cohen, 1987). A single oral dose oflufenuron has been shown effective in controlling the cat flea (Ctenocephalides felis felisy for at least 30 days in treated cats (Blagburn et al., 1994) and dogs (Rink et al., 1994). Davis (1997) reported a significant reduction in fleas on free-ranging ground squirrels (Spermophilus beecheyi) that had been treated with lufenuron. If lufenuron proves efficacious over a long period of time (::::1 mo) when administered orally to prairie dogs, it could be applied over large areas of prairie dog habitat via a treated bait. Prior to management application, lufenuron should be evaluated in a controlled laboratory setting to determine an effective dose, duration of efficacy, product safety, and to formulate an acceptable bait carrier. Ifresults of laboratory tests are positive, lufenuron should be tested in a controlled field study to determine efficacy under natural conditions. Without this evaluation, the management potential ofIGRs for controlling plague and protecting the health of the endangered blackfooted ferret, as well as human health, will be difficult to discern. We have proposed studies to test the efficacy of lufenuron for controlling fleas on artificiallyinfested captive white-tailed prairie dogs (Cynomys leucurus; Wild and Castle, 1998); however, laboratory rearing of fleas has proven difficult thus far. Further, because Oropsyl/a and Opistocrostis fleas are most readily collected in spring and early summer, our lufenuron-efficacy work is limited to the summer months. As an alternative, preliminary, means of lufenuron evaluation, we will conduct a bioavailabiIity study of lufenuron administered orally to captive white-tailed prairie dogs. Prescribed oral doses oflufenuron for domestic cats and dogs are 30 mg/kg and 10 mg/kg body weight, respectively. The concentration of lufenuron in blood of protected animals is somewhat variable, but protection seems to be conferred to dogs and cats when blood levels reach at least 50-100 parts per billion (B. Blagburn, pers. comm.). No data are available regarding the bioavailability of ingested lufenuron in any wild rodent species; however, assay results are pending from a recent pilot study (Castle and Wild, 1998). Based on results from this pilot study and assuming 100 ppb as the efficacious blood level, we will determine the test dose for this experiment. Although the correlation between blood levels oflufenuron and efficacy of flea control in prairie dogs will need to be determined in future experiments, this preliminary work will provide insights into: 1) between-animal variability �74 and 2) features of the decay curve of lufenuron (e.g. peak concentration andduration of levels over 100 ppb) in the blood of prairie dogs. Because white-tailed prairie dogs are spontaneous, obligate hibernators, they exhibit marked changes in activity and physiologic function seasonally (Harlow, 1997). These activity and physiologic changes may influence lufenuron bioavailability. Throughout much of their range, adult white-tailed prairie dogs enter hibernation in August or early September, and juveniles begin hibernation in late September or early October. Before hibernating, they store large amounts of body fat, to provide energy during their winter-long fast (Harlow, 1997). It is possible that as fat is mobilized during hibernation and upon arousal, lipid-soluble materials that were ingested and stored during late summer and fall may also be mobilized. Because lufenuron is highly lipid-soluble (Blagburn et al., 1994), white-tailed prairie dogs fed treated bait prior to hibernation could potentially store the compound over. winter, and therefore be protected during winter and when they arouse in spring. If prairie dogs were thus protected, flea population increases could be pre-empted, and management of plague outbreaks would be enhanced. To test this hypothesis, we will compare blood lufenuron levels in active vs torpid prairie dogs in a controlled laboratory setting. To summarize, the objective of this study is to determine the serum profile oflufenuron during active and torpid periods in orally-dosed white-tailed prairie dogs. These data will be used in the design of future experiments to evaluate lufenuron in captive and free-ranging prairie dogs, and will provide insights on the between-animal and seasonal variation that is to be expected. When combined with efficacy information, blood assay for lufenuron would provide a useful tool for evaluating lufenuron treatment programs involving free-ranging animals. Methods Care and housing oj prairie dog All captive prairie dogs will be pair-housed in cages under the same conditions described in Castle and Wild, 1998. Prairie dog cages (46 x 91 x 46 em) are constructed of wood and wire fencing. Half of the cage serves as a nest box, and the other half serves as a feeding area Two hinged doors allow access to each half of the cage for cleaning or animal handling. The nest box is a 40 x 28 x 23 em plastic container fitted with a wooden lid. The nest box has a 10 ern diameter hole in one wall to provide access to the feeding area; a 10 em diameter piece of PVC pipe joins the nest b.ox to the feeding area The access hole can be covered to confine the animals to either side of the cage as necessary. Our preliminary work shows that defecation and urination in the nest box are minimal, so we do not expect the nest boxes to become soiled during experiments. The floor of the feeding area consists of 1em mesh wire fencing to allow feces, urine, and spilled food to fall through to a newspaper-lined metal pan below. This mesh size prevents the animal's feet from becoming wedged in the floor, but may encourage the animal to spend most of its time in the nest box when not feeding. Newspaper in the metal pan will be changed daily if soiling occurs. Food (Teklad Rodent Blocks) will be provided ad libitum except as noted below. Water will be available at all times. Prairie dog cages will be kept in two similar buildings. Temperature in each building is thermostatically controlled, and artificial lighting is controlled with timers. Health and attitude of prairie dogs will be assessed daily during the experimental period. Body weight will be measured at 1-2 week intervals while each animal is anesthetized for blood collection. �75 Experimental Design Thirty captive white-tailed prairie. dogs (16 males and 14 females) less than one year old will be blocked by sex and randomly divided into two treatment groups (n = 13 each) and two control groups (n = 2 each). Treatment group 1 and one control group (active-group animals) will be housed in a building under conditions that inhibit hibernation (14L:I0D photoperiod plus natural lighting through windows, ambient temperatures above 15°C). Treatment group 2 and a second control group (torpid-group animals) will be housed in a second building under conditions that promote hibernation (1 OL:14D photoperiod, ambient temperature of 5-6 °C). Intermittent fasting of up to 2 d per 7 d period will also be used in some individuals to promote hibernation if temperature and photoperiod changes prove insufficient. Our goal is to expose the two groups to the most extreme conditions known to inhibit or promote torpor. We do not expect food deprivation to harm the torpid-group animals, because food intake between torpor bouts is minimal (Harlow, 1997). We have determined that these experimental conditions allow us to inhibit or promote hibernation in a majority of our white-tailed prairie dogs; however, despite our efforts, some animals housed under hibernation-promoting conditions remain active, while some housed under hibernationinhibiting conditions become torpid. We will therefore assess each animal daily, and record its activity level. Active-group prairie dogs that become torpid will be aroused by gentle shaking and rubbing; torpid-group animals that remain active will not be disturbed. We will assess the duration of torpor in torpid-group animals by placing a small amount of sawdust on the back of each torpid individual; the presence of sawdust on the back at subsequent checks will be interpreted to mean that the individual was torpid the entire period (Harlow, 1997). DOSingMethods Prairie dogs will be fasted for 24 h, then weighed prior to dosing; water will be available during the fast After the fasting period, each individual in a cage will be confined to one-half of the cage by blocking off the nest box access hole, and temporarily removing the nest box. Each animal in treatment groups 1 and 2 will be fed a bolus dose oflufenuron (20-300 mg/kg dose, pending pilot trial results) mixed thoroughly in a highly palatable bait (ground rat chow and molasses). Control animals will receive the bait only. The bait will be offered in each animal's feeding dish. We previously determined that fasted prairie dogs consume up to 2 g of such a bait within about 30 min, and will consume up to 13 g in less than 12 h. We will observe the progress of bait ingestion in each animal to determine when the dose is ingested. We will weigh the baitllufenuron mixture before and after the prairie dogs are dosed, in order to calculate the actual dose ingested. After both animals in a cage have consumed their dose, the nest box will be returned and they will again be pair-housed. Blood Collection We will collect 3 ml of blood from each prairie dog prior to lufenuron dosing, one week after dosing, then at 2 week intervals through week 9 post-dosing, or until torpor bouts are no longer observed. Anesthesia is required for blood collection. We have determined that prairie dogs respond very well to isoflurane anesthesia. Induction and recovery are quick (approximately 3-5 min for each), and we have detected no adverse effects. All prairie dogs will be aroused from torpor prior to anesthesia, in order to assure sufficient blood pressure for blood collection. Prairie dogswill be placed into a plastic chamber for induction at a concentration of 4% isoflurane in oxygen, then anesthesia will be maintained via mask at about 2-2.5% isoflurane. Heart rate and respiration will be monitored during anesthesia. Blood will be collected by jugular venipuncture and placed into a glass vacutainer blood tube (without anticoagulant). Alternatively, if we are unable to collect an adequate blood sample �76 peripherally, we will collect blood from the vena cava or directly from the heart. Although cardiac puncture is considered to be a safe procedure for collection of large volumes of blood from laboratory rodents (CCAC, 1984), due to the increased risks involved with cardiac puncture (5-10% mortality expected; J. Wimsatt, pers. comm.j.'we will use this technique only to obtain critical samples. Serum will be harvested and frozen within 4 h of collection. Serum lufenuron concentration will be determined by HPLC at Novartis Laboratories. Product safety Based on mode of action, chitin inhibitors should be safe in mammals, even at very high doses (Blagburn et al., 1995; T. Miller, pers. comm.). Cats have been fed more than 5 times the prescribed dose oflufenuron without showing adverse side effects (B. Blagburn, pers. comm.). The high dose to which our prairie dogs may be exposed (300 mglkg) is based upon our estimate of the maximum amount of bait a prairie dog might eat in a day in the field (40-60 g) and a reasonable lufenuron concentration in the bait (5 mg/g). We did not detect adverse effects from lufenuron administration during our pilot study, when 3 prairie dogs received doses of20-300 mg/kg. We do not anticipate any prairie dog mortality due to the dosing regime or blood collection. However, if an animal becomes sick or injured, and recovery is not likely, it will be euthanized with an overdose of inhalant anesthetic. A complete post-mortem examination will be performed on any animal that may die during the experimental period. Data Analysis We will compare lufenuron blood concentrations of the torpid- and active-group prairie dogs over time by conducting a profile analysis of the lufenuron decay curves. In addition, we will use analysis of covariance to compare: 1) peak concentrations obtained in each group, and 2) the duration of concentrations above 100 ppb in each group. The number of days an animal was observed to be active will serve as the covariate in our analyses. Group responses will be considered significantly different at p 0.05. Literature Cited Barnes, A. M. 1993. A review of plague and its relevance to prairie dog populations and the blackfooted ferret. Proceedings of the symposium on the management of prairie dog complexes for the reintroduction of the black-footed ferret. J. L. Oldemeyer, D. E. Biggins, B. J. Miller, and R Crete, eds. 96pp-.· Beard, M. L., S. T. Rose, A. M. Barnes, and J. A. Montenieri. 1992. Control of Oropsylla hirsuta, a plague vector, by treatment of prairie dog burrows with 0.5% permethrin dust. Journal of Medical Entomology 29:25-29. Blagburn, B. L., J. L. Vaughan, D. S. Lindsay, and G. L. Tebbitt. 1994. Efficacy dosage titration of lufenuron against developmental stages of fleas (Ctenocephalidesfelisfelis) in cats. American Journal of Veterinary Research 55:98-101. Blagburn, B. L., C. M. Hendrix, J. L. Vaughan, D. S. Lindsay, and S. H. Barnett. 1995. Efficacy of lufenuron against developmental stages offleas (Ctenocephalidesfelisfelis) in dogs housed in simulated home environments. American Journal of Veterinary Research 56:464-467. Castle, K. T. and M. A. Wild. 1998. Safety and efficacy of pyriproxifen and lufenuron for the control of prairie dog fleas: a pilot study. Colorado Division of Wildlife Animal Care and Use Committee Study Plan. Canadian Council on Animal Care (CCAC). 1984. Guide to the care and use of experimental animals, Vol. 2. Ottawa, Ont., Canada. �77 Cohen, E. 1987. Interference with chitin biosynthesis in insects. In Chitin and benzoylphenyl ureas, series entomologica, vol. 38,1. E. Wright and A. Retnakaran (eds), Dr. W. Junk, Publishers, Boston, pp.33-42. Davis, R. M. 1997. Use of an orally administered insect development inhibitor (Iufenuron) as a flea control agent in the California ground squirrel, Spermophilus beecheyi. Fourth International Symposium on Ectoparasites of Pets, pp. 31-35. Harlow, H. 1. 1997. Winter body fat, food consumption, and nonshivering thermogenesis of representative spontaneous and facultative hibernators: the white-tailed prairie dog and blacktailed prairie dog. Journal of Thermal Biology 22:21-30. Hink, W. F., M. Zakson, and S. Barnett. 1994. Evaluation of a single oral dose of lufenuron to control .flea infestations in dogs. American Journal of Veterinary Research 55:822-824. Ubico, S. R., G. O. Maupin, K. A. Fagerstone, and R. G. McLean. 1988. A plague epizootic in the white-tailed prairie dogs (Cynomys Teucurus) of Meeteetse, Wyoming. Journal of Wildlife Diseases 24:399-406. Williams, E. S.; K. Mills, D. R. Kwiatkowski, E. T. Thome, and A. Boerger-Fields, 1994. Plague in a black-footed ferret (Mustela nigripes). Journal of Wildlife Diseases 30:581-585. Wild, M. A. and K. T. Castle 1998. Monitoring and managing disease in black-footed ferrets. Colorado Division of Wildlife Program Narrative, Project # W-153-R. RESULTS AND DISCUSSION Carnivore Disease Survey With the assistance of USDA Wildlife Services and the Bureau of Land Management (BLM) we collected 19 coyotes and 4 badgers from the LSMA between 27-30 July 1998 and 20 coyotes between 11-12 February 1999. Coyotes were collected using a combination of calling and aerial gunning. Death occurred rapidly and the collection technique was adequate, but much less efficient in summer than in the winter months. Coyotes were collected from various locations in the >4700 mf management area; however, we focused on the Powderwash area and near the site of the preconditioning pens. No lesions indicative of active disease were noted on gross examination of carcasses. We found no serologic evidence of exposure to leptospirosis serovars canicola, grippo, hardjo, and pomona; however, low positive titers to serovar ictero were found in nine coyotes (five in July and four in February). No serologic titers to toxoplasmosis were found. Coyotes were not tested for Aleutian Disease (a mustelid disease), but all four badgers were seronegative for the disease. All badgers and 20% of coyotes sampled during the summer had positive titers to tularemia, while 10% of the coyotes sampled in winter were positive. Positive titers could have been associated with predation on rabbits, prairie dogs, or other rodents or exposure to ectoparasites. Duration of titers to tularemia are likely of short duration «6 mo). The impact of tularemia or leptospirosis on black-footed ferrets is not currently known; however, ferrets and humans are susceptible to these diseases. Canine distemper virus (CDV) and plague are serious threats to survival of black-footed ferrets. The prevalence of coyotes with serological titers to CDV was~10%; markedly lower than in previous years (Fig. 1). However, plague activity continues in LSMA as evidenced by titers in sampled coyotes (Fig. 2). Black-footed Ferret Reintroduction Nineteen black-footed ferrets (8 adult females, 5 juvenile females, 6 juvenile males) were received from the National Black-footed Ferret Conservation Center in November 1998 and placed in breeding pens at LSMA. Seventeen ferrets remained in good health with minimal health treatment �78 required. One adult female ferret died of a biliary adenocarcinoma and one juvenile female is missing and presumed dead. Ten female ferrets were paired with males and five of these gave birth. One litter was apparently consumed by the female when <1 day of age. The other four litters yielded 13 pups. Artificial Rearing of Oropsyl/a fleas Although fleas had been abundant in the spring of 1998, in July fleas seemed to disappear from the field. Burrow swabs consistently returned 0, 1, or 2 fleas per burrow. Because most burrows had no fleas, the ratio offleaslburrow dropped to less than 0.5. The cause of decline is unclear. One possibility is that because many burrows were filled in by prairie dogs escaping the summer heat, the nest areas that house a majority of fleas were no longer accessible from the surface. It is also possible that the flea populations experienced a real decrease during the summer, and that decrease was reflected in swabbing success. Our attempts to artificially rear Oropsyl/a fleas during summer 1998 met with limited success. We had successful completion of the life cycle within our rearingjars, but only about 4 adult fleas were produced from eggs laid in captivity. Our overall success was limited by low flea survival during transport to the lab, and the relatively low numbers of fleas available for collection in late summer when our facilities were readied for use (virtually no fleas were available in the field after 30 June). By the end of October 1998, no fleas were alive in our insectary. We resumed field collections of fleas on 5 April 1999, when snow cover had receded and prairie dog burrows were open. We made five more trips to the field through the middle of May. A total of 1645 fleas (906 of these 0. tuberculata) were collected. We successfully increased flea survival during transport from the field to the lab by keeping fleas under cooler and more humid conditions. Most fleas collected were alive when they arrived at the lab. Unfortunately, adult flea mortality in the insectary was high, despite our attempts to keep the fleas well-fed and under optimal temperature and humidity conditions. Reproduction within the insectary was therefore limited. We collected 8 larva from the insectary jars, and placed them into a separate dish containing rearing media; none of the larva pupated. Attempts to produce self-sustaining populations of 0. tuberculata on prairie dogs were also disappointing. While a few fleas survived for 21-24 days on the prairie dogs, it became clear that selfsustaining populations were not possible under our conditions. Our lack of success was most likely due to our inability to properly mimic natural burrow conditions required by fleas (constant temperature and high humidity). We attempted to simulate burrow conditions by attaching insulation to each nest box (to maintain temperature), and by placing a screen-covered petri dish of moist vermiculite into the flea refuge (to increase humidity). We were able to keep temperatures fairly stable, but were not able to maintain high relative humidity within the nest boxes. Survival and reproduction of 0. montana in the insectary has been high. We will use 0. montana in future experiments investigating flea control in captive prairie dogs. Flea Control In Prairie Dogs In the pilot study we observed no adverse affects with any of the lufenuron dosages. Previous researchers have shown lufenuron to be efficacious for controlling fleas in cats and dogs when serum concentration is 50-100 parts per billion (ppb) (C. Parks, Pers. comm.). Our results indicate that, as expected, baseline samples contained no lufenuron, but that one day after dosing, blood concentration was at or above efficacious levels at all treatment levels. After one week, blood concentrations had decreased considerably; serum concentration in the animals dosed with 20 mg/kg and 60 mg/kg had fallen below the efficacious level, while the concentration in the animal fed 300 mg/kg remained well above the efficacious level at 220 ppb. Analyses of remaining blood samples are pending. �79 In January 1999, we performed the experiment titled "Bioavailability oflufenuron administered orally to captive white-tailed prairie dogs". All prairie dogs except for two non-torpid group animals ingested >95% of their bait within 18 h; all remaining bait was removed and weighed after 18 h. We observed all prairie dogs twice daily (8:00 a.rn. and 4:00 p.rn.) during the experiment for overall health and to quantify length of torpor bouts. Animals that were torpid for two consecutive observations were considered to have been torpid the entire time between observations, unless a small amount of sawdust placed on the back of torpid animals had been displaced. Animals that were torpid at one observation time then awake the next were considered to have awakened at the midway point between observations. Torpid-group animals found hibernating were allowed to remain torpid, while non-torpid group animals were awakened if found hibernating. All torpid animals were awakened prior to anesthesia and blood collection. The duration of the study was 1512 hours; torpid group animals were torpid for an average of 406 h (range = 0 - 936 h), while non-torpid group animals averaged 7.5 h torpid (range = 0 - 36 h). We detected no detrimental effects oflufenuron in any prairie dog, nor were any animals harmed by repeated anesthesia and blood collections. HPLC analysis for blood lufenuron concentrations is pending. LITERATURE CITED Rink, W. F., M. Zakson, and S. Barnett. 1994. Evaluation of a single oral dose of lufenuron to control flea infestations in dogs. Am. J. Vet. Res. 55:822-824. Palma, K. G., S. M. Meola, and R W. Meola 1993. Mode of action ofpyriproxifen and methoprene on eggs of Ctenocephalides felis. J. Med. Entomol. 30:421-426. . Wild, M. A. 1998. Monitoring and managing disease in black-footed ferrets. Colorado Div. Wildt. Res. Rep., 0880-1, Jul1997 - Jun 1998, Fort Collins. Wild, M. A. 1999. Animal and pen support services for mammals research. Colorado Div. Wildl. Res. Rep., 8160-1, Jul1998 - Jun 1999, Fort Collins. 25 20 "0 Q.) Eco 15 II) •... Q.) .c 10 E :J Z 5 0 1997-W 1997-S 1998-W 1998-S 1999-W Sample period CI CDV positive IJ CDV negative I Fig. 1. Prevalence of exposure to canine distemper virus (CDV) in coyotes from the Little Snake Management Area, Colorado, from winter 1997 through winter 1999. All positive coyotes were adults with the exception of one juvenile in summer 1998. Data from winter 1997 (age class unknown) provided by M. Albee. �80 A 20 15 L- a> ..c E 10 :::J Z 5 o +-----------,--1997-W 1997-S 1998-W 1998-S 1999-W Sampling period B 20 15 L- a> E :::::I 10 Z 5 1997-W 1997-S 1998-W 1998-S 1999-W Sampling period I II Plague-Pos [3 Plague-Neg I Fig. 2. Prevalence of exposure to plague in coyotes (A = juveniles; B = adults) from the Little Snake Management Area, Colorado, from winter 1997 through winter 1999. Data from winter 1997 (age class unknown) provided by M. Albee. �81 ATTACHMENT 1 HUSBANDRY PROTOCOL FOR CARE OF CAPTIVE BLACK-FOOTED AT THE LITTLE SNAKE MANAGEMENT AREA FERRETS Pen Construction Breeding pens will be constructed over established prairie dogs towns. If an active prairie dog colony is not present, prairie dogs from plague-free areas will be reintroduced to the pen site prior to . construction. Individual pens will be about 1500-5000 ff and constructed in clusters of four (four-plex design). Generally, females will be housed in three of the pens and a male in the fourth pen. Each pen will contain a vault capable of holding a buried insulated wooden nest box (about 12" x 20" x 16"), which will be covered with a roof to provide shade and to keep runoff away from the nest box. In the females' pens, the vault will be contained a sub-enclosure, or whelping pen, about 100 ff in size. A perimeter fence will be constructed around each four-plex or group of'four-plexes. Tubing and traps will be placed at intervals along the fence to protect and capture potential escapees. Additionally, prior to initial introduction of breeding-age ferrets, the burrows will be checked using a "smoke test" and then radio-collared ferrets will be placed in the pens to further check pen security, Site Security Breeding pens will be located in a remote area with access limited to authorized personnel. Visitors will be allowed only on a limited basis and when accompanied bysite personnel. Human activity near the pens will be limited strictly to ferret husbandry and management practices. No. dogs will be allowed in the pen area. Use of motorized vehicles in the pen area will be minimized. The perimeter fence will serve to contain escaped ferrets and as a deterrent to entrance of wild and feral carnivores; however, other means of carnivore control (nonlethal and lethal) may be necessary to protect ferrets. Biosafety protocols will include the use of site specific coveralls and boots by personnel handling ferrets or entering ferret pens. Treatment of shoe soles with a bacteriocidal/virucidal solution (e.g., Roccal-D) may substitute for designated footwear. Personnel will not enter ferret pens or handle ferrets if they are experiencing flu symptoms (fever, chills, body aches, respiratory signs); caretakers with cold symptoms should wear a mask and gloves when working around ferrets. Ideally, caretakers should receive flu shots annually. Only prairie dogs from the immediate vicinity of the pen site or those completing quarantine-will be provided to ferrets. Personnel will not handle carnivores or prairie dogs (except those completing the quarantine period) during the work day prior to entering the pen site unless they shower and change clothes. Pets of site personnel should be vaccinated annually against canine distemper. General Husbandry Ferrets will be checked daily at dawn or dusk by trained caretakers. Caretakers should minimize contact with ferrets, but should observe ferrets directly or with binoculars to assess health status daily. Abnormalities will be noted in animal records and, if necessary, the attending veterinarian notified. If a ferret is not observed (directly or via sign, e.g., consumed food) for 3 consecutive days, a trap (attached to a nest box via tubing) will be placed in the pen. Traps will be checked at intervals appropriate to assure that the ferret is not adversely affected by temperature, weather, or restraint time �82 (generally, every 2-6 hr). Trapped ferrets will be weighed and examined grossly. Additional examination and treatment will be administered if necessary under the direction of the attending veterinarian. Body weights will be obtained monthly (and anytime the ferret is handled for other reasons) to monitor health status and to evaluate feeding programs. Ferrets will be trapped, weighed, briefly examined, and released back to the pen. Transponder function will be checked at least once monthly and transponders will be replaced (with the ferret under anesthesia) if needed. Nest boxes will be provided only during periods when the ferret is confined to the whelping cage, e.g .., the breeding and whelping period or during required confinement to prevent escape or enhance treatment. Nest boxes will be cleaned every other day (excepting during breeding period when male and female are housed together and immediately prior to and following parturition) using pen-specific instruments, bedding, and trash can when kits are present or if a contagious disease is suspected. Wood shavings (not cedar) will be placed about 3-4 inches deep on one side of the nest box and about 1 inch deep of alphi-dry will be placed on the other side. A minimum-maximum thermometer will be placed on the roof of one nest box in each four-plex. Temperatures will be checked during nest box cleaning or daily during periods of extreme weather. Modifications to the nest box insulation may be needed if temperatures in the nest box are extreme. Maintenance diet will include live prairie dogs supplemented with frozen prairie dog and rabbit carcasses, hamsters, and dry ferret food (Totally Ferret). Dry ferret food will be used only as a supplement if insufficient supplies of prairie dogs are available as food. Dry ferret food will not be provided as a sole feed source, but as a supplement to the meat diet. Prairie dogs will be collected, quarantined, and processed as described in the protocol by Marinari and Williams (1998). Rabbits will be procured from breeders and processed as prairie dogs; however, additionally the head and kidneys will be removed. Dry feed will be stored in rodent-proof containers prior to feeding. Whenever possible, live prairie dogs will be provided to ferrets; however, we will avoid offering large, aggressive prairie dogs and any prairie dog surviving> 3 days will be trapped and removed from the ferret pen. These prairie dogs will be euthanized and fed as meat. Female ferrets will be provided with an average of 100 g feed/day and males will receive 125 g feed/d; however, ferrets may not be provided fresh feed daily (e.g., if a 500 g prairie dog is provided). Amount of feed provided to individuals will vary based on assessment of physical condition. Frozen prairie dogs will be thawed about 48 hr in a refrigerator prior to feeding. Feed may at times be provided in a trap (with the doors wired open) for training purposes. Unconsumed meat left in the pen will be discarded after 24 hr. Water will be injected into prairie dog carcasses during the winter or when available water is minimal. A water source will be provided, but may be frozen during the winter. If snow cover is not present, fresh water will be provided daily. Waterers will be cleaned weekly or more frequently if needed. Records will be maintained to document feed offered and estimate feed consumed by ferrets in each pen. During gestation, whelping, and lactation, modifications of the maintenance diet will be required. About 2 wk following breeding, feed will be increased l.33 times the base diet. About 4 wk following breeding, feed will be increased to 1.66 times the base diet. From whelping to weaning, feed will be offered ad libitum. Ideally, only live prairie dogs and hamsters should be offered, but if unfeasible, ferrets will be supplemented with prairie dog or rabbit meat or dry feed initially, then when kits begin to kill prey, only live prairie dogs (about 150 g/ferret) will be offered until release. Breeding Management Adult males and females will be housed in separate pens except during the breeding period. The breeding period will vary annually based on weather, so multiple handling of ferrets may be required to assess reproductive status. Beginning about I February, or when signs of breeding behavior are noted (increased digging, marking, etc), males will be trapped weekly for breeding �83 soundness examination. With ferrets restrained in handling cages, the testes will be measured (width, depth) and tested for firmness. After testes are firm for 1 mo, males will be considered ready for breeding. When males are showing signs of reproductive activity (testes firm and about >20 mm width or depth), females will be trapped for signs of estrous. The females will be restrained in a handling cage and the vulva measured. If the vulva is small «2 mm x 3 mm), the female will be trapped and measured again in about 7 days. If the vulva is beginning to swell, the female will be handled again at 3 day intervals to monitor the trend in change of the vulva Vaginal cytology may be used when the vulva shows a trend toward increasing size (generally to about 4 x 7 mm). When vaginal cytology indicates >90% cornified cells, we will wait 5 days, then introduce the male to the females whelping pen (male and female will be restricted to whelping pen). The male and female will be monitored remotely using a pen camera or by watching from a distance and listening for vocalizations to determine compatibility and breeding activity. Breeding activity is assumed if chuckling is heard, copulation observed, and/or a neck stain is present on the female. In this case, the male will be removed 3 days after introduction. Function of the females transponder will be checked after copulation and replaced if necessary. Ifbreeding activity is not observed, the male may be left with the female up to 7 days; however, if the male and female show no interest in one another (in separate areas of pen, no neck stain), the male may be removed after 24 hr and a new male introduced 1-2 days later. The male will be removed immediately if it shows aggressive behavior that threatens the female. Males will be given 3-4 days rest after each breeding. A male may be brought from a separate fourplex to breed a female if the resident male is incompatible or unavailable. The female will be trapped 7-10 days following suspected breeding for vaginal cytology. If cells remain highly cornified, breeding likely did not occur and the female will be re-paired. Records will be kept on results of breeding soundness exams, vaginal cytology, pairings, and breedings. The female will be restricted to the whelping area beginning 35 days post-breeding. A waterer will be placed in the whelping area and bedding material placed in the nest box. Vitamin K will be supplemented to the females diet at the rate of 5 drops/day from 5 days.prior to whelping to day 35 following whelping. On day 42 post-breeding, we will begin listening daily for sound from the nest box. If sounds are heard, the animals will be left undisturbed for 5 days. On day 5, the nest box will be cleaned using pen-specific bedding, instruments, and trash can. The nest box will be cleaned again on day 10 and the door to the whelping area opened. The nest box will be cleaned at 2 day intervals until the female moves the kits from the nest box. Prairie dogs and hamsters will be the only feed offered until kit release. If females do not whelp a second breeding will be attempted. About 10 days following the calculated whelping date we will again begin monitoring vaginal cytology. When 90% cornified cells are present, a male will be paired with the female as described previously. Kits will be trapped on day 90 for physical examination, marking with transponders, and vaccination against canine distemper virus (if vaccine available). Kits will be trapped subsequently for booster vaccines if needed. Care of Sick Animals Health status offerrets will be checked daily (or as possible). Healthy ferrets should be active, consume feed, have a clean haircoat, and be free of physical abnormalities (lameness, cuts, swellings, etc.). The pen floor should also be examined for abnormal feces, blood (not from prey item), vomit, etc. Any abnormalities will be noted on the daily observation forms and the attending veterinarian notified. If there is any likelihood that the condition could be contagious, the caretaker will disinfect their hands and boots and ideally change coveralls before proceeding to other pens. If an animal is known to be sick, that animal will be cared for last (or by an alternative caretaker) using good sanitation techniques and any equipment used in the pen will be disinfected. Sick animals may be �84 confined to the nest box and whelping pen to aid in care and treatment. Alternatively, critically ill animals may be placed indoors in quarantine for treatment. If coccidiosis is suspected (lack of appetite, mustard-like feces) and the attending veterinarian cannot be reached, treatment can be initiated using 50 mg/kg Albon orally on day 1 followed by 25 mg/kg Albon orally daily for about 7 days. Fresh water must be available at all times when animals are treated with Albon (parenteral fluid therapy may be needed as well). Specimens (feces, discharges) should be obtained opportunistically if possible (using good sanitation practices) and placed in a whirlpack and refrigerated until further instructions are obtained. If a ferret is found dead, it should be double or triple bagged (e.g., in a ziplock or trash bag), labeled with animal ID, pen, and date and refrigerated. The carcass will be boxed in a cooler and submitted via overnight Federal Express to Dr. Beth Williams, Wyoming State Veterinary Laboratory for necropsy. The facility supervisor, attending veterinarian, and BFFCC should be notified for further instructions. Prairie dogs found sick or dead in the pen area should also be collected and submitted for necropsy. Caretakers should wear gloves and practice good sanitation measures whenever dealing With sick or dead animals. Release We will attempt to maintain 20 breeding animals of various ages: ideally, 5 female and 1 male l-year-olds, 5 female and 2 male 2-year-olds, 5 female and 1 male 3-year-old, and 1 male 4-year-old. Animals will be redistributed to pens each fall to maximize diversity in breeding pairs. Older adults and excess young of the year will be released each fall when young are about 20 weeks of age. Prior to release, all ferrets will be fitted with radiocollars. �85 Colorado Division of Wildlife Wildlife Research Report July 1999 JOB FINAL REPORT State of: Colorado Project No. W.J53-R-12 Work Package No . ....;3:::..;0~0::....:1'-Task No. 1 Period Covered: Authors: Personnel: _ Cost Center 3430 Mammals Program Deer Conservation Experimental Deer Inventory July 1, 1998 - June 30, 1999 R B. Gill, and T. M. Pojar G. Schoonveld, S. Steinert, 1. Olterman, C. Wagner, B. Watkins, R Bartmann, G. White ,';' ABSTRACf Over-winter (Nov-May) survival rates were estimated for 3 Data Analysis Units: Red Feather .(D-4), Middle Park (D-9), and Uncompahgre (D-19). Overall 3 study areas, doe survival averaged 91. 7% and fawn survival averaged 79.4%. Population estimates were obtained for D-9 in late January, 1999 by flying square mile quadrats allocated according to a stratified random sampling scheme. Deer density averaged 12.93 deer/km" and projected to a total population of 11,016 deer ± 2,337 (90% CI). Estimates of the ratios of bucks and fawns: 100 does were obtained from helicopter counts conducted in December 1998. The buck:doe ratio was estimated to be 27 bucks: 100 does. The fawn:doe ratio was estimated to be 62 fawns: 100 does. ~~II~ 11~11~~fi~l~ M~~~~il~mll'i~rnr BDOW014555 �98 �87 EXPERIMENTAL DEER INVENTORY R. Bruce Gill and Thomas M. Pojar P.N. OBJECTIVES 1. Develop and test an experimental deer inventory system in 5 DAUs in western Colorado to determine efficacy in monitoring deer populations. 2. Publish results in a peer-reviewed scientific journal. SEGMENT NARRATIVES 1. Estimate the annual doe survival rate in DAUs D-4 (Red Feather), D-9 (Middle Park), and D-I9 (Uncompahgre). 2. Estimate over-winter fawn survival rate in DAUs d-4 (Red Feather), D-9 (Middle Park), and D19 (Uncompahgre). 3. Estimate the December fawn:doe and buckdoe ratios in DAU D-9 (Middle Park). 4. Estimate winter density in DAU D-9 (Middle Park). 5. Develop a spreadsheet model for predicting deer population performance in DAUs D-4 (Red Feather), D-9 (Middle Park), and D-19 (Uncompahgre). 6. Analyze data and prepare an annual Federal Aid Job Progress Report. INTRODUCTION Initial plans called for the development and testing of experimental inventory systems on 5 deer Data Analysis Units (DAUs) across Colorado's important deer herds. Manpower and financial constraints limited the scope to 3 DAUs: Red Feather (D-4), Middle Park (D-9), and Uncompahgre (D-I9). Measurements on each DAU were to continue for a period of3-5 years before deciding whether to expand implementation of the experimental deer inventory system. However, the project's principal investigator, Richard M. Bartmann, retired during the segment and his vacancy remains unfilled. Consequently, it was decided to terminate the R&D component of the inventory system at the end of the current segment. This report, therefore, constitutes the Final Report of activities accomplished under Work Package 3001, Task 1. STUDY AREAS Study aras D-4 and D-19 were described previously (Bartmann and Pojar 1998). Study area D-9 is situated in the headwaters area of the the Colorado River drainage encompassing areas below approximately 2,600 m in elevation in Grand and Summit Counties. Winter range is primarily �88 sagebrush steppe vegetation type characterized by low growing « 1 m) shrubs. Sagebrush species (Artemisia tridentata tridentata and Artemisia tridentata vaseyana) predominate with other shrub species such as rubber rabbitbrush (Chrysothamnus nauseosusy; sticky-flowered rabbitbrush (Chrysothamnus viscidifloris), mountain snowberry (Symphoricarpus oreophilus), bitterbrush (purshia tridentata), serviceberry (Amelanchier alnifolia), and chokecherry (Prunus virginiana)locally abundant (scientific names 'according to Weber 1976). Mule deer winter over approximately 850 km2 below 2600 m in elevation. METIIODS Bartmann and Pojar (1998) thorougly described the methods used in this investigation, so they will not be repeated here. RESULTS Doe Survival Across all 3 study areas, doe survival during winter 1998-99 averaged 91.7% with 121 of 132 does surving from November 1998 through May 31, 1999. This compares to an over-winter survival rate of 90.1 % for adult does throughout the previous winter (Bartmann 1998). No single source of mortality stood out as a primary (Table 1) which was similar to results from 1997-98. Fawn Survival In general, fawn survival throughout winter 1998-99 increased compared to 1997-98. In 199899, 127 of 160 radio-collared fawns survived from November 1, 1998 through May 31, 1999 for an overrall survival rate of 79.4 %. In 1997-98, 61.0% of radio-collared fawns overall survived the winter. Sex and Age Composition Classification counts of mule deer in Middle Park Data Analysis Unit (D-9) were conducted during the period December 22-24, 1998 and included a total of912 deer, of which 15.9% were bucks, 32.0% were fawns, ancl52.1% were does (Table 2). The ratio of bucks per 100 does computed to 30.5 4.8 bucks: 100 does and 61.5 ± 7.5 fawns: 100 does. ± Population Density Middle Park Data Analysis Unit (D-9) deer census quadrats were flown during the fourth week in January, 1998. Counting conditions and deer distribution were both ideal. Overall, 2,415 deer were tallied on 56 sample quadrats. Number of deer/km? averaged 12.93 with a standard error of 4.00. When projected over the entire winter range ofD-9, the deer population was estimated at 11,016 deer ± 2,337 (90% CI) (Table 3). �89 Table 1. Fates of mule deer does and fawns radiocollared in DAUs D-4 (Red Feather), D-9 (Middle Park}, and D-19 (Uncom~ahgre} during the 1998-99 winter. Red Feather Middle Park Uncompahgre Fate Does Fawns Does Fawns Does Total Radiocollared 52 50 40 50 40 60 Survived 45 40 39 44 37 43 Coyote predation 4 0 Mtn. lion predation 2 0 0 0 2 0 3 0 2 2 0 0 0 0 0 0 0 0 0 0 2 Accidents 0 0 0 0 0 Unknown 2 0 0 2 4 10 1 6 3 17 Other predators 0 Road kills 7 0 Hunter kills 2 Starvation 0 7 Total Mortality Fawns "Total does not equal 100% because of rounding error. Table 2. Results of mule deer classification counts in Middle Park Data Analysis Unit (D-9), December 22-24, 1998. Date GMU l-yr 2-yr bucks bucks Adult bucks Total bucks Does Fawns Total 12/23/98 27 6 6 3 15 66 44 125 12/22-24/98 37 27 23 19 69 175 98 342 12/23-24/98 181 6 5 2 13 44 17 74 12/23-24/98 28 7 12 3 22 130 92 244 12/24/98 18 11 10 5 26 60 41 127 57 56 32 145 475 292 -9h Adult bucks: 100 does Total bucks: 100 does Fawns: 100 does Totals D~9 Summary l-yr bucks: 100 does 2-yr bucks:100 does Ratio estimates 12.0 11.8 6.7 30.5 61.5 Lower 90% CI 9.3 9.1 4.8 25.9 54.3 14.8 14.6 8.8 35.5 69.4 Upper 90% CI �90 Table 3. Deer quadrat census results for Middle Park Data Analysis Unit (D-9), January 1998 ... Stratum Strata Deer Counted (km/) Population Estimate N n Total Deer/km? 80.3 28.5 582 20.43 2 222.7 10.4 13 3 189.1 46.6 4 121.7 5 SE Total ±90% 12.34 1640 629 1.25 3.17 280 449 . 730 15.66 7.64 2961 918 15.5 144 9.27 9.84 1128 761 72.5 15.5 378 24.33 26.41 1764 1216 6 88.1 15.5 494 31.79 24.59 2799 1375 7 77.7 12.9 74 5.71 5.40 444 266 Overall 852.1 145.0 2415 12.93 4.00 11016 2337 Spreadsheet Models A 'spreadsheet population model was developed in a Quatro Pro spreadsheet format for Red Feather (D-4), Middle Park (D-9), and Uncompahgre (D-19) Data Analysis Units. Disk copies of spreadsheets for each DAU were sent to Area Biologists responsible for managing each DAU. Copies of the spreadsheet format may be obtained by writing to: Dr. Gary C. White 211 B JVK Wagar Bldg. Department of Fishery and Wildlife Biology Colorado State University Fort Collins, CO 80523 LITERATURE CITED Bartmann, R. M., and T. M. Pojar. 1998. Experimental deer inventory. Colorado Division of Wildlife. Wildlife Research Report, July, 1998, Part 11:135-142. Weber, W. A. 1976. Rocky mountain flora. Colorado Associated University Press. Boulder, CO. Prepared by _ R. Bruce Gill Wildlife Research Leader Thomas M. Pojar Wildlife Researcher �91 Colorado Division of Wildlife Wildlife Research Report July 1999 JOB PROGRESS REPORT State of __,..--!:C~o~lo::.!.r=ad...,o,,-_ Project No. __ -..l..W!--...:..1=.:53:!...-~R~-~12~_ Work Package No. _---"'-30;:....;0::..,:1'-_ Task No. 3 ------~--------- Period Covered: Authors: Personnel: Cost Center 3430 Mammals Program Deer Management Monitoring and Managing Disease in Deer Chronic Wasting July 1, 1998 - June 30, 1999 M. W. Miller, C. T. Larsen, and J. Gross R Kahn. M. Leslie, K. I. O'Rourke, Williams T. R Spraker, E. Wheeler, M. A. Wild, and E. S. ABSTRACf . Deer from throughout Colorado were examined for occurrence of chronic wasting disease using a combination of targeted surveys and harvest/road-kill surveys. Between June 1998 and May 1999, 8 chronic wasting disease (CWD) cases were diagnosed among 25 "suspect" deer submitted from known endemic portions of northeastern Colorado; CWD was not diagnosed in any of 10 additional "suspect" deer submitted from elsewhere in Colorado. All confirmed CWD cases originated in game management units (GMUs) where the disease had been detected previously. We sampled over 1800 harvested or road-killed deer to randomly survey for CWD in select DAUs. Immunohistochemistry (llIC)-based prevalence in Larimer County (5%; 39/780) and South Platte River bottom/plains (1.6%; 5/315) DADs were unchanged from previous years. We detected no evidence ofCWD in any of369 samples from Middle Park, or in any of over 300 samples from other GMUs outside northeastern Colorado. Late doe hunts in Larimer County DAUs provided an additional 212 samples for comparison of CWD prevalence between sexes; based on data compiled over the last 2 years, CWD prevalence does not differ between male and female mule deer (P = 0.72). Since June 1998, CWD affected 8 of 29 adult (z l-yr-old) mule deer in the resident Foothills Wildlife Research Facility herd. No signs of neurological disease were observed in any of the 11 cattle (subjects) or 11 mule deer from the Rocky Mountain Arsenal National Wildlife Refuge (RMANWR)(contact controls) housed with naturally-infected resident deer since July 1997. Using mc, Prpres was detected in brain tissue (medulla oblongata at the obex) from 1 of2 mule deer experimentally inoculated with a 5 g oral dose of brain tissue homogenate from CWD-infected deer 6 or 9 mo earlier. One of two deer examined 16 mo after inoculation (Pl) had lesions of spongiform encephalopathy, and both were mC-positive. Two deer examined 3 rno PI and 2 examined 12 rno PI were negative, as were all 10 uninoculated controls collected from RMANWR at the same intervals. One of the 8 remaining deer inoculated in December 1997 began �92 showing early clinical signs of CWD ~ 15 mo PI; a second began showing early signs 2-3 wks later, and subtle signs were noted in at least 4 of 8 inoculated deer alive 18 mo PI. An epidemic model of was developed to simulate the dynamics of chronic wasting disease in mule deer populations. As seen in earlier modeling exercises, simulations resulted in projections wherein either the disease or the deer population was eliminated; stable coexistence of chronic wasting disease could not be achieved in simulated populations. Productivity (i.e., harvest) was reduced in populations by very low rates of prevalence. Spread of CWD within a simulated population was highly sensitive to transmission rate, and very small decreases in transmission efficiency resulted in notable decreases in prevalence. Simulated test and slaughter programs revealed the importance of initiating control while CWD prevalence was low « 0.05). Low rates of test and slaughter (e.g., < 20% of the infected population) effectively eliminated wasting disease in simulated populations if control measures were initiated while prevalence was low (i.e., 0.01), but the likelihood of control diminished rapidly as disease prevalence increased. Test and slaughter programs will require an effort sustained over many decades to ensure elimination of chronic wasting disease. A statewide plan for monitoring and managing CWD in deer and elk was drafted. The plan has been distributed for internal and limited external review, and will be finalized by 30 September 1999. cwn �93 MONITORING AND MANAGING CHRONIC WASTING DISEASE IN DEER M. W. Miller and C. T. Larsen P. N. OBJECTIVES '"1. Design, conduct, and report results of: a. targeted surveillance to estimate and detect changes in distribution of chronic wasting disease (CWD) in free-ranging deer populations; and b. harvest or road-kill surveys to estimate and detect changes in prevalence of CWD in enzootic deer populations. ,2. Design, conduct, and report results of experimental studies using captive deer naturally or experimentally infected with CWD. AGREEMENT OBJECTIVES 1. Conduct and report results of targeted surveillance to estimate and detect changes in distribution of CWD in free-ranging deer populations statewide. 2. Conduct and report results of harvest surveys to estimate prevalence ofCWD in DAUs D4, D9 (GMUs 18,28,37,371), DI0( +GMU 29), and D44 (+ GMUs 87, 90, 93, arid 95). 3. Continue an experiment evaluating cattle susceptibility to CWD via natural contact exposure. 4. Continue to study pathogenesis ofCWD in mule deer. 5. Develop an epidemic model ofCWD dynamics in deer populations. 6. Observe epidemiology of naturally-occurring CWD in captive deer. MATERIALS AND METHODS Surveillance We monitored deer populations throughout Colorado for occurrence of CWD using a combination of targeted surveillance and harvest or road-kill surveys. These were organized and conducted as follows: �94 Targeted (= clinical disease) surveillance: Deer showing clinical signs consistent with those seen in chronic wasting disease were collected by field personnel statewide and brain tissues examined for evidence of spongiform encephalopathy. The "suspect case" profile was defined as follows: • Species: mule deer white-tailed deer • Age: ::::18 months • Signs: emaciated and abnormal behavior &lor indifference to human activity &lor increased salivation &lor tremor, stumbling, incoordination &lor difficulty or inefficiency in chewing/swallowing increased drinking and urination &lor Where possible, submissions were subjected to complete necropsy; in some situations, only heads were available for examination and sampling. In all cases, histopathology of brain tissue (Williams and Young 1993) was used to diagnose CWO; in some cases, immunohistochemistry (ll:IC) or other ancillary tests were used to confirm or support diagnoses. Harvest surveys: In order to obtain reliable estimates of CWO prevalence that will serve as a basis for monitoring responses to management interventions, we continued conducting harvest surveys on select deer populations. During the 1998-1999 hunting seasons, fresh brain and select lymphatic tissues were collected from deer harvested in enzootic GMUs; deer harvested or culled in other select GMUs throughout Colorado were also sampled as negative controls. Brain tissues were examined at the Colorado State University Diagnostic Laboratory for anti-PrP immunostaining reactions (O'Rourke et al., 1998) and histopathological lesions (Williams and Young, 1993) consistent with CWO infection. Epizootiological Studies Epizootology of naturally-occurring CWO in captive mule deer and white-tailed deer: Naturallyoccurring CWO was a sporadic disease of resident FWRF mule deer prior to 1985 (Williams and Young, 1992), and also has occurred sporadically since 1994; no cases had been observed in resident white-tailed deer since their addition to FWRF in 1993. We maintained and observed 29 ~ l-yr-old mule deer and 5 ~ l-yr-old white-tailed deer in paddocks at CDOW's Foothiils Wildlife Research Facility (FWRF) during May-September 1998. Deer received natural forage, pelleted rations (high energy supplement and "browser" diet), and alfalfa hay; water and mineralized salt were available ad libitum. All deer were evaluated daily for clinical signs of CWO (and other health problems) in conjunction with routine feeding and handling activities. Resident mule deer also represented the source of infection (= "treatment") in an ongoing study of cattle susceptibility to CWO (see below). Cattle susceptibility to CWO (Williams and Miller): We continued monitoring cattle for clinical evidence of natural CWO transmission as part of a coordinated interagency effort to study cattle susceptibility to CWO. Twelve 4-mo-old calves purchased from a private ranch located near Sheridan, WY, were placed in paddocks at the FWRF with naturally-infected captive mule deer in July 1997 (see above for description of resident deer herd). Twelve 4-mo-old mule deer fawns were captured at the Rocky Mountain Arsenal National Wildlife Refuge (RMANWR) in late September and placed in the �95 same paddocks as contact controls for natural transmission (extensive ongoing surveillance of RMANWR deer has continued to confirm that this population is free of CWD). CWD Pathogenesis in mule deer (Williams and Miller): In a separate but related study, 20 additional 6mo-old mule deer fawns were captured at the RMANWR in early December. Each fawn was given a single oral dose of about 5 g of fresh brain tissue homogenate from captive mule deer with clinical CWD. Fawns were then released into a separate paddock physically removed from the contact transmission study. These experimentally-infected fawns serve two purposes: as controls for domestic calves orally inoculated with a single oral dose of about 50 g of this same CWD brain homogenate (Williams et al., 1998), and as subjects of a study on the pathogenesis of CWD in mule deer. We have randomly sacrificed 2 experimentally-infected deer at 3,6,9, 12, and 16 mo after inoculation; we also collected 2 age- and sex-matched control deer from RMANWR at each time step. Complete necropsies were performed and samples collected as described in the original study plan. . Select sections of brain tissue (obex) were examined after staining with hematoxylin and eosin (H&E) or anti-PrP immunostain (F89/160.1. 5; O'Rourke et al., 1998); examination of other tissues is pending. Epidemic modeling (Gross and Miller): We developed a mechanistic model to simulate the dynamics of chronic wasting disease in mule deer populations. The model projected age-specific disease dynamics, changes in population size, and control strategies involving selective removal of infected animals or changes in rates of disease transmission. Model parameters were estimated from observations of infected and uninfected deer in Colorado and Monte Carlo techniques were used to evaluate likely responses. Appendix A provides a more detailed description of our model. CWD Monitoring and Management Plan A statewide plan for monitoring and managing CWD in deer and elk was drafted. RESULTS AND DISCUSSION Surveillance Targeted (= clinical disease) surveillance: Between June 1998 and May 1999, 8 chronic wasting disease (CWD) cases were diagnosed among 25 "suspect" deer submitted from known endemic .portions of northeastern Colorado; CWD was not diagnosed in any of 10 additional "suspect" deer submitted from elsewhere in Colorado. All CWD cases confirmed in 1998-1999 were mule deer, and all originated in game management units (GMUs) where the disease had been detected previously. Harvest surveys: We sampled over 1800 harvested or road-killed deer to randomly survey for CWD in select DADs. ruC-based prevalence in Larimer County (5%; 39/780) and South Platte River bottom/plains (1.6%; 5/315) DADs were unchanged from previous years. We detected no evidence of CWD in any of369 samples from Middle Park, or in any of over 300 samples from other GMUs outside northeastern Colorado. Late doe hunts in Larimer County DADs provided an additional 212 samples for comparison of CWD prevalence between sexes; based on data compiled over the last 2 years, CWD prevalence does not differ between male and female mule deer (P = 0.72). �96 Survey data gathered during 1996-1999 indicate that CWD is generally more prevalent and more widely distributed in mule deer than in white-tailed deer in northeastern Colorado (Fig. 1). However, limited data from white-tailed deer harvested near the mouth of the Big Thompson Canyon west of Loveland suggest CWD could become quite prevalent among white-tailed deer (Fig. IB). In light of this potential and the apparent spread of CWD eastward along the Big Thompson/South Platte River corridor in both mule deer and white-tailed deer (Fig. 1), natural spread and eventual infection of more densely populated white-tailed deer habitats in the midwestern and eastern US should be anticipated in the coming decades unless management interventions are successfully identified and implemented. A.Muledeer Epizootiological Studies Epidemiology of naturally-occurring CWD in captive mule deer and whitetailed deer: Since June 1998, CWD affected 8 of 29 adult (~ l-yr-old) mule deer in the resident Foothills Wildlife Research Facility herd (Fig. 2A). Two other adult mule deer died between June and May; enterotoxemia was diagnosed in one, and pasteurellosis in the other. Although neither showed neuropathology consistent with CWD, 1 had IHC staining consistent with preclinical CWD. Resident mule deer also represented the source of infection (treatment) in an ongoing 10-yr study of cattle susceptibility to CWD (see below). One of 5 white-tailed deer remaining in our resident FWRF herd developed CWD and died in June 1999. In all, CWD has claimed 7 of 11 captive white-tailed deer held at FWRF since 1993 (Fig. 2B). One of the 4 remaining individuals is showing early signs of CWD. I B. White-tailed deer Figure 1. Estimated CWD prevalence (%) among subpopulations of (A) mule deer and (B) white-tailed deer in northeastern Colorado. Data were aggregated via known deer movement patterns. Prevalence estimates are based on IHC reactions only (IHC+ltotal). Evaluation of tonsillar biopsies from captive deer are still pending. Lack of results has precluded evaluation of tonsillar biopsy as a potential antemortem test for CWD in deer. Cattle susceptibility to CWD (Williams and Miller): No signs of neurological disease were observed in any of the II cattle (subjects) or II mule deer from the Rocky Mountain Arsenal National Wildlife Refuge (RMANWR)(contact controls) housed with naturally-infected resident deer since July 1997. Neither of2 contact control deer that died during October-June showed neuropathology consistent with CWD, although both suffered from chronic weight loss; malnutrition probably contributed to at least I of these deaths. Fifteen of the 36 > l-yr-old naturally-exposed resident FWRF deer developed clinical �97 CWD and died or were euthanized during the first 24 mo of this study, thereby ensuring calves and control deer have received considerable exposure to CWD. Similarly, all 11 cattle exposed to a single 50 g oral dose of brain tissue homogenate from CWD-infected deer remained healthy 21 mo postinoculation (through 30 June 1999) (E. S. Williams, pers. comm.), as did all cattle exposed to CWD via intracerebral inoculation 22 mo ago (R. 1. Cutlip, pers. commun.). A. lD j - ".dee, ,._.-. o ,-- •• : : :: ~ 2l :JJ : i 10 :: :: r-': : ::: • : :::: :::: :::: ::: !---:: :: r'1i i~H:: ~~ ~i ~~ :: :: :: !_.-:: :: :: ! !; H ~~ ~. .: :. : 'III' 8. WlitR-t.Ied dHr lD .., i Xl o t 2) z Pathogenesis of CWD in mule deer (Williams and Miller): rcs Using ruc, Prp was detected in brain tissue (medulla oblongata at the obex) from 1 of2 muIe deer experimentally inoculated with CWD 6 or 9 mo earlier. One of two deer examined 16 mo after inocuIation (PI) had lesions of spongiform Figure 2. Chronic wasting encephalopathy, and both were ruC-positive. Two deer disease continued to affect both mule deer and whiteexamined 3 rno PI and 2 examined 12 mo PI were negative, as tailed deer held at FWRF. were all 10 uninoculated controls collected from RMANWR at the same intervals. One of the 8 remaining deer inoculated in December 1997 began showing early clinical signs of CWD ~ 15 mo PI; a second began showing early signs 2-3 wks later, and subtle signs were noted in at least 4 of 8 inocuIated deer alive 18 mo PI. ResuIts from examination of lymphoid tissues are pending. Epidemic modeling (Gross and Miller): Simulations ofCWD dynamics that used a broad range of parameter values resulted in projections in which either the disease or the deer population was eliminated. We did not identify a set of realistic parameters that resulted in stable coexistence of chronic wasting disease in deer populations. Population productivity (i.e., harvest) was reduced in populations by very low rates of prevalence due to the combined effects of a reduction in per-capita production and a decrease in population density. Spread of the disease was highly sensitive to the rate of transmission, and a very small decreases in the efficiency of transmission resuIted in notable decreases in rate of spread of the disease. Simulatedjest and slaughter programs revealed the importance of initiating control activities while prevalence of the disease was low « 0.05). Low rates of test and slaughter (e.g., < 20% of the infected population) effectively eliminated wasting disease in simulated populations if control measures were initiated while prevalence was low (i.e., 0.01), but the likelihood of control diminished rapidly as disease prevalence increased. Test and slaughter programs that examine will require an effort sustained over many decades to ensure elimination of chronic wasting disease. CWD Monitoring and Management Plan The draft monitoring and management plan (Appendix B) has been distributed for internal and limited external review, and will be finalized by 30 September 1999. Surveillance activities for 1998-1999, as well as surveillance planned for 1999-2000, reflect prioroties established in the monitoring portion of this plan. �98 ACKNOWLEDGMENTS . The statewide CWD monitoring and surveillance program described here relies heavily on efforts of dedicated field personnel throughout the Colorado Division of Wildlife, and truly represents a division-wide effort to improve our understanding and management of this important disease problems. In addition to those specifically listed, we collectively thank all of those regional and area biologists, district and area wildlife managers, volunteers, deer hunters, and others who assisted by submitting suspect cases, harvested animals, or road-killed animals throughout the year. LITERATURE CITED K. 1, T. V. Baszler, 1. M. Miller, T. R Spraker, 1 Sadler-Riggleman, and D. P. Knowles. 1998. Monoclonal antibody F89/160.1.5 defines a conserved epitope on the ruminant prion protein. }. Clin. Microbiol. 36:1750-1755. Williams, E. S., and S. Young. 1993. Neuropathology of chronic wasting disease in mule deer (Odocoileus hemionus) and elk (Cervus elaphus nelsoni). Veterinary Pathology 30:36-45. _--" M. W. Miller, T. 1. Kreeger, H. Van Campen, and T. R Spraker. 1998. Susceptibility of cattle O'Rourke, to cervid spongiform encephalopathy. unpublished grant proposal, submitted to USDA, CSREES, National Research Initiative Competitive Grants Program, 88 pp. Prepared by _ Michael W. Miller Wildlife Research Veterinarian �99 Appendix A CWO: Simulating Chronic Wasting Disease User's Manual and Model Documentation Version 1.0 June 1999 John E. Gross Natural Resource Ecology Laboratory Colorado State University Ft. Collins, CO USA 80523-1499 Email: JohnG@NREL.ColoState.edu �100 CWD: Simulating Chronic Wasting Disease Summary CWO is an individual-based model that represents the most important processes determining the dynamics of animal populations over periods of decades or centuries. It simulates reproduction, disease dynamics, and the interaction of the animals with environmental characteristics that can change through time. Each animal in the population is explicitly represented and individual characteristics including age, sex, and disease state are tracked for the life of an individual. The model simulates 1 to 3 disease period, and includes the capacity to model a wide variety of harvest scenarios. Files are written that facilitate analyses of population responses to disease, harvest, or environmental variation, including changes in disease prevalence, disease-caused mortality, harvest, and changes in population age/sex structure. CWO has a flexible structure and there are no inherent practical limitations to the . number of individuals that can be simulated during a run. Practical limits to the number of individuals are set only by the capabilities of the computer on which the model is run. Introduction To make informed decisions on management of large ungulates, biologists need to evaluate the consequences of alternative management actions on population processes. . Management' actions can include hunting by the public or agency personnel, vaccination, selective harvest, habitat modification, or a combination of these actions. The consequences of these decisions depend on a complex of factors, and it is thus difficult to evaluate the relative merits of alternative management plans. CWO was designed to simulate management actions that might be used in attempts to eliminate CWO or to inhibit its spread. The model simulates each individual in the population and explicitly represents reproduction, natural mortality, disease transmission, non-selective harvests (Le., hunting), and selective harvests (e.g., test and slaughter). The use of an individual-based model is particularly appropriate for CWO because stochastic effects are important in small populations, or where a small number of individuals have a large effect on population processes. This situation clearly applies to CWO, where the behavior of a small number of infectious individuals can have a profound influence on long-term dynamics of a much . larger population. �101 An Overview of Model Structure and General Features Animal populations are simulated within a landscape that potentially can consist of a single or multiple patches. Each individual is simulated and has attributes including a sex, age, disease state, and membership in a local population. Mortality and fecundity are stage specific processes, and each individual is assigned to a specific stage based on its age. Mortality is density-independent, and fecundity (recruitment) responds to density only when density exceeds a threshold value. CWO differs from other population simulators by . incorporating disease, modeling both males and females, and by including functions that permit age and sex-specific patterns of removal. Because the model is individual-based, it is appropriate for representing the dynamics of small populations as well as large. In general, the number of patches or individuals that can be simulated are limited only by computer memory and processor speed. In reality, it is much easier to generate complex model dynamics than It is to understand the output, and most practical limitations will result from' an inability to analyze and understand output. It is essential to run many simple simulations to ensure that you have a complete understanding of model behavior before introducing spatial structure or other complexities. i Patch Attributes and Dynamics A patch is an area of the landscape that can support a local population. A Patch has the attributes of area, current quality (see below), maximum quality, mean environmental quality (from 0-1), maximum environmental quality (0-1), deviation from mean quality, and time since disturbance. Overall patch quality is quantified in a single variable ("forage") calculated from a user-defined function that transforms current forage to quality multiplier' (from 0-1; Fig. 2). This "forage" quality multiplier is used by the local population each year to determine the ability of the patch to support a population of animals (see below). Patch dynamics are simulated by predicting "forage" as a function of time since disturbance. The function that determines "forage" is calculated from a linear interpolation between forage and time values that are user-defined. Patch attributes can dramatically affect population processes through densitydependent effects on recruitment. If you wish to remove density-dependent effects from the model and control populations through harvest, set the patch size to a number much larger than mean population size and set animal density (in the *.spc file) to a number greater than one. Recruitment and Death The likelihood of recruitment changes with animal age and varies from year to year. Yearly variation is represented by an "environmental quality" multiplier that modifies the average recruitment rates. Each year, the model selects a random deviate from a Beta distribution that represents the "environmental quality" for the current year. The distribution of "environmental quality" is defined by the mean and standard deviation (entered by the user), and if the standard deviation is set to 0 there is no annual variation in the overall rate of recruitment. Each year the model loops through all females and compares the probability that a female of that age recruits an offspring into the population to a uniform random deviate. If the random deviate is less than the probability, recruitment occurs. Animals are culled from the population at stage-specific rates once per year. Mortality rates are independent of animal density and are subject only to stochasticity due to random events and to acute catastrophes. Stage-specific mortality rates are input by the user, and �102 for each individual, these rates are compared to a uniform random deviate to determine whether that particular individual dies during the year. Animal density dependent can effect population dynamics by reducing the probability of recruiting into the population. There is no effect of density on survivorship. Population density affects recruitment only when it exceeds a density threshold. Above the density threshold, the probability of recruitment declines linearly to 0 at a user-defined upper recruitment bound (Fig. ). In most situations this model is used to simulate, density plays no role in population dynamics because populations are controlled primarily by harvest. Under these conditions, its best to set the patch size sufficiently large to ensure that a density function is never invoked. Animal and Patch Aging The age of both animals and patches are incremented at the end of summer, after dispersal and prior to mating. At this stage, any patch manipulations are input and patch disturbance occurs. New patch statistics are generated for each patch, to be used in the following year's simulation. Data Output Model output is generated at the end of each year. The frequency of model output to the screen is specific by the user, while file output is generated every year. Output files formats are designed for analysis by statistical packages (e.g., SAS, SPSS) and are fixed format. The year is incremented immediately before model output and before patch dynamics are simulated. Thus year 0 output records are initial conditions. Running a Simulation To run a simulation, it is first necessary to build a set of input files that contain the variables and parameters used in the simulation (Table 1). The overall model run (which may consist of a large number separate Monte Carlo simulations) is controlled by a file with the extension .run. This file contains the names of input files that define life history characteristics, patch attributes, disease dynamics, and other run-time information described in Table 1 and below. This section describes the input files necessary to conduct a model run. The appendices to this manual include sample input file of each type. Input Files A simulation is controlled by variables and parameters specified in files that define the characteristics of the. animal species and landscape, and by the a file that contains information specific to a particular run (e.g., the number of years to simulate, names of input files, and what information is to be written to output files). In general, lines in an input files begin with an integer that tells the program what the rest of the line contains. This is followed by a one word descriptor (e.g., a series of characters and/or numbers that contain no spaces) of that line, followed by one or more parameters. It is essential that the one word descriptor has no spaces, and that there is at lease one space between the descriptor and the parameter(s). Input files are summarized in Table 1. Summary of parameters contained in each input file. 1. *.run Simulation run parameter file: Contains the names of files with parameters to be used in a specific set of runs, the number of runs to be simulated, the length of �103 each run, and which output files are to be written. It contains a variable for the frequency of output to the screen and a delay interval so that screen output can be more easily viewed. Screen output is slow; you can dramatically speed model execution by increasing the interval at which output is printed to the screen. This is the"master" file that and it deals with the highest level of model control. Code: Inputs::ReadRUNFile 2. .. *.pch Patch attributes include the size of each patch, maximum quality of the patch (0-1.0), a mean quality and the standard deviation of the mean patch quality, an initial forage (which remains unchanged in the 'absence of disturbance). Specified changes to patch quality and/or size in a given year can be used to simulate prescribed bums or other management protocols. Code: LandScape::ReadPCHFile 3.:''. *.spc Species characteristics: sex ratio at birth (m:f), number of age/stage classes for males and females, ages associated with each class, litter size, stage specific mortality, birth schedule, adult male and female body size, maximum density. The density threshold is estimated from the maximum density of animals, patch quality, and patch area. This file also specifies a function that relates patch quality (summarized into one variable, called "forage", for convenience) to a quality multiplier. Linear interpolation is used to estimate quality for values intermediate to those entered. Code: MammaIData::ReadSPCFile 4. *.pop Population parameters: This file contains the initial 'size and composition (age, sex) of populations. Code: AnimaIPopulation::ReadPOPFileO 5. *.hrv Harvest. File with parameters that determine the rules used to implement harvests. This includes the frequency of harvest (every year, every other year, etc.), threshold population sizes for determining the size of harvest, rules that govern the number of animals harvested, and the age and sex composition of the harvested population. 6. *.dis Disease:. File with parameters that determine disease dynamics. Includes parameters for transmission, and transition from incubating to infected and infected to dead. ' Model Functions and Details This section describes the structure of specific model functions, the implementation functions, and sources of data used to parameterize the model. of model Patch Dynamics Patches are areas that potentially support a local population, and they are characterized by their area (_area) and quality (_qualify, 0-1), constrained to a maximum for each patch (_maxQualify). Patch quality defines the ability of the patch to support animals relative to their maximum density. For example, if the maximum density for a species is 10 and the quality of a particular patch 0.1, that patch �104 Table 1. Input file extensions and the types of data they contain. file name type of data example file parameters *.pch patch data size, maximum quality, time since disturbance *.spc species attributes sex ratio at birth, stage-specific fecundity and mortality, maximum *.pop local population data number of local populations, age and sex of the members in each *.run simulation run control # of years of the simulation, names of input and output files, output *.hrv harvest information frequency, number harvested, and composition of harvest *.dis disease parameters transmission and transition proabilities I will support 1 animal per unit of area (area is generally expressed in krrf). The quality of each patch can be affected by environmental stochasticity, which is characterized by a normal random variate with a specified mean LmeanEnvirQualify, 0:-1) and standard deviation LsdEnvirQualify). The quality of a patch during any given year is calculated from it's current forage (see Landscape, _forage), constrained to a _maxForage level specific to each patch, and the quality associated with a given value of forage: _qualify = f(minLforage,_maxForage)) and the effects of environmental _qualify = minLmaxQualify, stochasticity (if any): _qualify * norma/_randomLmeanEnvirQualify, _sdEnvirQuality). The function that calculates quality as a function of _forage is defined by the user. and linear interpolation is used to evaluate points between those defined in the input file. If current forage is greater than that defined by the user, the _forage value at the maximum _gTSinceDisfurb is used; e.g .• no values are extrapolated beyond the data. These are combined to determine the population size at which reproduction is inhibited, _densifyThresSize, as a function of patch quality and the maximum density of animals that the highest-quality patch could support. _densifyThresSize _qualify. = _maxDensify *_area * 50 100 150 200 250 300 S50 ••00 Forage This design has been implemented to Fig. 2. User-defined function relating quality to forage. reflect the observation that population size may be restricted by forage availability, which changes over time. or other habitat features (perhaps escape terrain) that do not change. The quality of any single patch is therefore constrained by both a "food" -type variable Lforage) and other patch characteristics LmaxQualify). Code: LocaIPop:SetDensThresSizeO; LocalPop:DensThresSizeO �105 Species Characteristics Each species to be modeled must be described by life history characteristics including the number of different stages for males and females LnMaleStages, _nFemaleStages), inclusive ages of each stage LmStageMinAge, _'StageMinAge), maximum number of young LmaxYoung), the probability of having O-_maxYoung (_pYoungm, stage-specific birth and death probabilities (_pRecruitj, _pMMortality;, _pFMortality;. where I is stage). Density Dependence Density dependence influences the probability of dispersal and recruitment of new individuals into the population. The probability of recruiting one or more young is stage-specific and is determined by user-input values. This probability is then modified to reflect the effects of density and environmental stochasticity, To calculate density-dependent effects, it is necessary to specify. a species-specific maximum density ·LmaxDensity, #/unit-area), the density a species achieves in optimal habitat.. The units of patch size and ...;,maxDensity (e.g., ha) must be the same. The density at which recruitment falls to zero is defined by _upperPopBound, a decimal value greater than 1 used to determine the population size at which the probability of recruiting an individual falls to zero in the absence of environmental stochasticity (Figure XX). When the population size in patch I is greater then _pop Ttuessize; the age-specific probability of recruitment (_pRecruif) is modified to reflect the density-dependence: _pRecruitj .' = _pRecruitj ( popSizej l-----------------------------------~--(l+_upperPopBoundj) ) _densThresSizej Effects of Density on Recruitment The probability of recruitment of a new individual can be further influenced by environmental stochasticity, which is constrained to the range of _envirQualMin andt: environmental Quality = maxL envirQualMin, nonnalrandomLmeanEnvirQualify,_ _pRecruitj = _pRecruitj 11-----------------,. l sdEnvirQuality)) *environmental Quality Fig. 3. DensitY effects, where A = _densThresSize Taken together, these functions account for annual.stochastic and B = _densThresSize • _upperPopBound fluctuations in habitat quality that affect "carrying capacity", and yearly fluctuations in recruitment that result from abiotic or biotic sources (weather, predation, etc.). -. Code: AnimaIPopulation::BreedAndRecruitO, LocaIPop::SetDensThresSizeO; LocaIPop::DensThresSizeO Data: MammaIData._envirQuaIMean, _envirQuaISD, _probRecruitD Mortality Natural mortality occurs once per year (Figure 1), and is a stage-specific function. Stage and sexspecific mortality rates are. user-specified parameters in the *.spc file. For each individual, a uniform random deviate (0-1) is compared to the probability of death. Code: AnimaIPopulation::CUII Aging At the end of each time loop, the age of each individuals is incremented by 1 and an appropriate adjustment is made to _tSinceDisfurbance for each patch and local population. Code: AnimaIPopulation:: AgeO; LocaIPop::SAgeO; LandScape:: SetTimeO �106 Model Output Model output is written to standard ASCII text files that are created in the current directory, typically the directory containing the input and executable files. By default, any existing files of with the same name are erased upon opening, except for the error log file. Output file formats are described below. Any errors that occur during the run are printed to the screen and error file, and the run will be aborted when a severe error is detected. Some common errors in input parameter values are automatically corrected (e.g., an out-of-range parameter), in which case a message is written to the error log file informing the user of any corrections. All simulation result files are formatted to facilitate input to programs for further analysis in statistics, graphics, or spreadsheet programs. Variables in output files are either space or comma delimited. File output ceases when an entire population goes extinct, except for data written to the summary output file. Thus if a population goes extinct in year 23 of a 50 year run, files will generally contain 24 years of data (year 0 + 23 years). The summary file will be filled with 0 values for the full length of the run (year 0 + 50 years). All output file names begin with the same prefix, which can be specified in the *.run file or which will default to "cwd". All output file names have an ".asc" extension except the error file. Output Files Produced by Every Run _pavg.as<;: - 1 line for each year of run. Mean and standard deviation of the number of susceptibles, incubating, infectious, and total population size. _ sum.asc - 1 line for. each year of each run. Number of susceptibles, incubating, infectious, and harvested. CWD _err. txt (if an error is detected). Error information Information about any errors detected during a run are printed to this file. Errors are assigned to categories Warning, Error, and Fatal Error. Warning messages signal an error than can be automatically corrected within the program or conditions that are unlikely to affect program results. Errors are more severe, and may be corrections to parameters that are frequently entered incorrectly or a condition within the program that may be irregular. Depending on the compilation, the program may abort upon detecting an error (this is a compiler switch, and cannot be changed by the user). The program always aborts when a Fatal Error is detected. Fatal errors are usually the result of an incorrect input file format, invalid parameter, or insufficient hard disk space or memory. Table 2. Output file names and contents. Files in italics are always produced, others are produced only when specified by a switching variable in the *.run file. Files are comma-delimited. File Columns Information cwd err.txt text Produced or appended to if an error is reported. _age I-many run, year, patch number, males by age (O-Age max), females by age (0Age max). Age max is specified in the *.spc file. 1-5 run, year, patch number, period, disease-caused deaths by sex and age class -hrv l-many run, year, patch, population size category, males by age (O-MaxAge), females by age (O-MaxAge) _pavg l-many year, mean and standard deviation of susceptible, incubating, infectious, and total number of animals _prevAge l-many run, year, patch number, susceptible males by age category, susceptible females by age category, incubating by sex (m.f) and age category, infectious by sex (m, f) and age category sum 1-5 run, year, patch number, number of susceptible, incubating, infectious, and harvested animals ts 1-9 run, year, patch number, test&slaughter index (O=off, I=on), population size, prevalence before t&s, prevalence after t&s, males removed, females removed disfxhs - (produced only when t&s is on) �107 Sample input files and formats General File Format Used by CWD The general format of CWD input files is to begin each line with an integer number that indicates the content of one or more lines of the input file. "99" is always used to indicate a comment and the program disregards all information on a line following the "99". In general, the integer is followed by one word (i.e., a series of characters and/or numbers that contains no spaces) describing the following parameter(s) or variable(s) .. The descriptive word can contain underscores, upper and lower case letters, numbers, or any other characters except blank spaces or tabs. For situations where there are a: variable number of input lines (e.g., the age structure of an initial population), lines may begin with a' data value not preceded by an integer indicating the line content. Sections of input than can include a variable number of lines always end with a delimiter, which is usually a number less than zero. It is essential that these file formats be followed, and users are very strongly encouraged to modify existing input files rather than create a new file. Virtually all errors in model runs can be traced to a mistake in an input file. Many (but not all) common mistakes in input files will be detected by the program and reported to the error file (CWD _err.txt). Model Run Data (*.run) "The *.run file is the first file read by CWD and it determines program operations at the highest levelwhich modules are used, the names of other input files, which files output files are to be written. _A value of 1 or greater turns on optional functions or optional outputs and the value "0" turns these ',' function off. If Random = 0, the same random number seed is used at the beginning of all runs. nlls option is used for debugging and Random should generally be set to "1". A delay can be used if you wish to slow the program in order to more easily view screen output. 99 cwd.run 99 -2 PCHFileName: cwd_inpt.pch 3 SPCFile: cwd_inpt.spc 4 POPFile: cwdjnpt.pop 5 Harvest: 1 cwd_inpt.hrv 6 Disease: 1 cwd_inpt.dis 9 Test&SI: 1 cwdjnpt.tst 99 -11 RunYears: 100 12 Runs: 250 14 ScreenOutputFreq: 50 99 - if random = 0 use same seed for all runs; if random = 1 use a different seed. 15 Random: 1 99 _ all output files will have the following "prefixed" to their names 99 _ E.g., the file beta_2_11_14_age.asc would be produced with this line. In addition, 99 _ any sensitivity variables are added to the file name (see below). 99 -16 OutputFilePrefix: beta_2_11_14 99 -21 PopAgeOut: 1 _age 22 PatchesOut: o _pch �108 23 DoDisOutput: 1 dis 29 doHarvestOutput: 1 harv 9999 -- line 31: on/off prevalence level to begin output, prevalence level to end output 99 so the following results is output when the prevalence level is >= 0.03 and <= 0.50. 991 .01 .50 31 doPrevAgeOutput_startPrev_endPrev: 1 disDths 32 doDisDeathsByPeriodOutput 9999 101 = sensitivity analysis. 99 Variables are repeated for each variable, so the number of 99 values on the line MUST be divisible by 4. 99 var_number smallest_val largest_val step 99 -101 sens vars 123.4.8 .05 Lines 21 -31: Any integer of 1 or greater tufns output on. 31: These files can be very large, thus they are produced only when prevalence in a patch is equal to or within the range of the starting and ending value. Patch Parameters (*.pclz) 99 filename: cwd_21 May.pch 10000 3 PatchArea: 4 MaxQuality: 1 5 MeanEnvQual: 1 6 sdEnvQual: .0 7 maxForage: 100 100 8 initialForage: PatchArea: Area of patch used by the local population. Set to a large value unless you want density.•.dependent population regulation to operate. MaxQuality: This value reflects the maximum density that a particular patch can support, relative to the species-specific maximum density. The species-specific maximum density is specified in the *.spc input file. MeanEnvQual: Mean quality of the patch (must b~ less than MaxQuality). sdEnvQuality: Standard deviation of the mean quality. If set to 0, there will be no annual variation in the value of the density threshold due to differences in patch quality. maxForage: Maximum forage value that will ever be obtained on a specific patch. initialForage: Forage value at start of the run �109 Harvest Populations are normally controlled by harvest of animals. The rate of harvest can vary depending on population size, which can be categorized as small, normal, or large. Input variables are used to determine threshold sizes between these categories. For each population size category, input variables specify age-specific rates of harvest. There are no limits to the number of age categories that can be specified, but if ages within a category overlap, the last value in the input file will be used. If you want harvest to be independent ot age, specify the same rate for all ages. i. Annual rates of harvest are determined each year by selecting the appropriate mean harvest rate from the input file (lines 11-17) and then modifying this rate to account for annual variation. Mean rates for all age groups are multiplied by a random deviate with mean 1 and CV from line 10. Thus annual variation for all age groups are correlated. 99 CWD for mule deer 99 Rule 1 has proportion harvested for low, med, and high pop numbers, by sex and age 1 harv_rule 1 10 CV_on_percent_harv .2 99 _ FEMALES min_age max_age portion_harvested. Ages are inclusive 11 F_Iow 0 20 .0 12 F_med 0 0 .01 12 F_med 1 2 .02 12 F_med 3 20 .07 13 F_hgh 0 0 .03 13 F_hgh 1 2 .05 13 F_hgh 3 20 .10 99 - MALES 15 M low 0 20 .0 16 M med 0 0 .02 16 M med 1 1 .06 16 M_med 2 20 .20 17 M_hgh 0 0 _..03 17 M_hgh 1 1 .09 17 M_hgh 2 20 .27 99 -- repeat lines 21 & 22 for each local population. 21 localPop 1 22 threshold numbers 500 1000 . �110 Test and Slaughter Test and slaughter is implemented by specifying the efficacy of the program for males and females. Efficacy is the product of the portion of the population tested and the likelihood that a positive animal will be detected. Variables in the input file determine whether the program is implemented from the beginning of the program, or after other criteria are met. A test and slaughter program can begin at a specific year of a model run, or after the population has achieved a specified prevalence rate. Efficacy is the product of the proportion of the population tested and ability to detect the disease. The variable on line 3 (periods to detect) is the number of periods of incubation that must elapse before a disease can be detected in an exposed individual. If there were 2 periods per year and this parameter were set to 1, an exposed individual would not be detected until they had incubated for a full period, equivalent to about 6 months. If you assume that rate of exposure is constant, one could interpret this to suggest that the average incubation time before detection would then be about 9 months, or one full period and (on average) half of another period. However, since test and slaughter occurs only once per year (after harvest and the second disease period), individuals infected during disease period 2 (falilwinter) will not be detected until the following year unless periods of detection is set to O. Infectious individuals are always detected. 99 99 3 4 5 6 7 cwd_test_ and_ slaughter.tst periods_of_incubation_before_detection local_pop_tested 1 Efficacy .5 Start_year 10 Start_prevalence .05 3 �III Sensitivity Analysis The model provides the user with very broad options to conduct sensitivity analysis. Todo so, specify on line 1.01in the run file the variable number (from the table below), the initial value, the maximum value, and the step (amount for the initial value to be incremented for each set of runs). Some variables (will in the future) also require that a local population be specified. When a sensitivity analysis is conducted, the names of all output files reflect this by adding the number of the variable and the step (starting with 0) of variable value to the file name. These values are added after the file name prefix specified on line 16 in the run file and before the normal file suffix. For example: lirie 16 in run file: 16 file_name: 24jan_larimer line 101 in run file: 101 sens_vars 121 .01 .10 .01 Which means: do runs with maternal transmission rates from 0.01 to .1 by .01. Ten sets of runswill be simulated, with 10 output files of each type (e.g., _pavg, _sum, etc.). The summary output files, for example, would be: 24jan_larimer_:_121_0_sum.asc 24jan_larimer_121_1_sum.asc 24jan_larimer_121_9_sum.asc If additional sensitivity variables are added, these are added to the file name in the general format: prefix_var#_step_var#_step ... suffix. There are no internal limits to the number of sensitivity variables, but since all levels are fully crossed, you can easily generate hundreds or even thousands of output files. The difficulties in analyzing such results are substantial. �112 Sensitivit~variables available for anal~sis. Class Affected MammalData MammalData MammalData MammalData MammalData Number 101 102 103 104 105 MammalData MammalData MammalData MammalData MammalData MammalData MammalData AnimalPop # values 106 121 122 123 124 125 126 Variable name value or Mult female survival multiplier male survival multiplier recruitment multiplier sex ratio at birth value mean environmental value quality std environmental quality value maternal transmission value prob. infected recruits value period 1 beta value period 2 beta value period 3 beta value beta for all periods value 3 3 3 3 3 3 3 21 periods for test detection value 3 test efficacy mean - both value harvest lower threshold value harvest uQQerthreshold value 3 3 3 LocalPopulation 51 LocalPopulation 61 LocalPoQulation 62 3 3 3 3 3 �113 AppendixB Chronic Wasting Disease in Deer and Elk: DRAFT Colorado Statewide Monitoring and Experimental Management Plan M. W. Miller and R.H. Kahn Terrestrial Wildlife Resources, Colorado Division of Wildlife . Background Chronic wasting disease (CWD) is a transmissible spongiform encephalopathy (TSE) of native deer (Odocoileus spp.) and elk (Cervus elaphus nelsoni) characterized by behavioral changes and progressive loss of body condition that invariably lead to the death of affected animals (Williams and Young 1992). Neither the causative agent nor its mode of transmission have been identified. There are no tests currently available for diagnosing CWD in live animals, and extant postmortem tests require microscopic examination of brain tissue. There are no known treatments for CWD. Previous attempts to eradicate CWD from research facilities failed on at least 2 occasions (Williams and Young 1992; Miller et al. 1998). Although similar in some respects to other TSEs that affect domestic sheep (scrapie) and cattle (bovine spongiform encephalopathy; BSE), existing data indicate CWD cannot be naturally transmitted to domestic livestock, and that scrapie and BSE cannot be transmitted to native cervids. Moreover, available data indicate that CWD does not present a threat to human health. "Chronic wasting disease" was first recognized by biologists in the 1960's as a disease syndrome of captive deer held in wildlife research facilities in Ft. Collins, CO. This disease was subsequently recognized in captive deer, and later in captive elk, in wildlife research facilities near Ft. Collins, Kremmling, and Meeker, CO and Wheatland, WY (Williams and Young 1980, 1982), as well as at in least two zoological collections (Williams and Young, 1992). More recently, CWD has been diagnosed in captive elk residing in one game ranch in Saskatchewan (Spinato, pers. comm.), two ranches in South Dakota (Holland, 1997), one ranch in Nebraska (W. Cunningham, pers. comm.), and one ranch in Oklahoma (L. Detwiler, pers. comm.). Since 1981, over 100 cases of clinical and preclinical CWD also have been diagnosed in free-ranging mule deer (0. hemionus), white-tailed deer (0. virginianus), and elk from northeastern Colorado; most of these diagnoses have been made since 1990 (Spraker et al. 1997; Miller, 1997). At present, the known world-wide distribution of CWD in wild cervids appears to be limited to northeastern Colorado and southeastern Wyoming (Miller, 1997; Williams et al., 1998). Although CWD was first diagnosed in captive cervids, the original source of CWD in either captive cervids or free-ranging cervids is unknown; whether CWD in captive cervids really preceded CWD in wild cervids, or vice versa, is equally uncertain (Spraker et al. 1997). In Colorado, free-ranging CWD cases in deer and elk have originated from throughout the northeastern portion of'the state. Game Management Units (GMUs) yielding infected deer or elk include 7,8,9, 19, 191,20,29,91,93,94,95,951, and 96. Although cases have come from 13 different GMUs, two Data Analysis Units (D4, DI0) have yielded over 85% of the documented cases. Based on targeted surveillance data and select harvest surveys, wild deer or elk populations in other parts of Colorado are probably not infected with CWD (Miller, 1997; 1998). Both targeted and harvest surveys indicate GMUs 9, 191, 19, and 20 are the main foci ofCWD in Colorado; estimated prevalence in all four GMUs is ~3% (Miller, 1997; 1998; M.W. Miller, unpubl. data). Data from harvest surveys indicate that CWD is relatively rare in other parts of northeastern Colorado, probably affecting ~2% of the deer in GMUs 7, 8, 29, 91, 93, 94, and 96 (Fig. 1). Similarly, harvest surveys indicate <1 % of the elk in DAUs E4 and E9 are infected with CWD (Miller, 1997; 1998; M. W. Miller, unpubl. data). For deer populations in the four most heavily infected eastern Larimer County GMUs, no real trend in prevalence (increasing or decreasing) can be discerned from data available to date. In the absence of historical (20-30 years ago) prevalence data or reliable estimates of transmission rates, it is �114 also unclear whether overall incidence ofCWD in northcentral Colorado DAUs is stable or increasing, or whether short-term observations can accurately forecast long-term trends. Based on prevalence estimates and preliminary results of simulation modeling to predict dynamics and impacts of CWD on affected deer populations, it appears CWD was introduced into northern Larimer County over 30 yrs ago (M.W. Miller and C.W. McCarty, unpubl. data). Beyond Larimer County, the South Platte River corridor may be the single most predictable avenue for spread of CWD. Kufeld and Bowden (1995) reported that a proportion of South Platte River bottom deer were highly mobile; observed deer movements included some to and from eastern Larimer County along the Cache la Poudre and South Platte Rivers. These movement patterns provide a plausible mechanism for the apparent emergence of CWD in South Platte River GMUs detected recently through targeted surveillance and harvest surveys. Other likely routes for deer emigrating from Larimer County have not been identified, and may be less predictable. Altitudinal movements of some elk subpopulations in the southern part of Larimer County are also somewhat predictable (Bear, 1982). Although there is potential for elk to cross the Continental Divide through Rocky Mountain National Park, the relatively low prevalence of CWD among elk diminishes the likelihood of its spread via their movements. The significance of CWD and its impacts on native deer or elk populations have not been determined. Preliminary results of simulation modeling suggest that, sustained at 5% prevalence as observed in the four most heavily infected GMUs, CWD could impact wild deer herds and lead to population declines (M.W. Miller and C.W. McCarty, unpubl. data). Data on CWD prevalence in . female deer are particularly critical to predicting potential impacts of disease on long-term population performance and assessing efficacy of management interventions. Preliminary comparisons made in conjunction with 1997 and 1998 surveys showed no differences in CWD prevalence between male and . female mule deer harvested in Larimer County GMUs (M.W. Miller, 1998; unpubl. data); these data were comparable to results of simulation models where male and female deer were assumed to be equally susceptible to CWD. In the absence of data to the contrary, and considering the difficulties inherent in eliminating CWD from captive or wild cervid populations once established, it seems most prudent to assume CWD could adversely affect native deer or elk populations and manage to reduce its occurrence and prevent its further spread in Colorado. Unfortunately, there is considerable uncertainty in how to manage CWD in free-ranging wildlife, or whether such an endeavor could even be successful. Because a more complete understanding of CWD is fundamental to developing comprehensive management programs, the Colorado Division of Wildlife (CD OW) needs to further understanding about CWD and its management through surveillance and experimental management. Data from completed, ongoing, and proposed research, both basic and applied, will serve as the foundation of an adaptive resource management plan for CWD in deer and elk. This plan will provide a mechanism for incorporating new knowledge gained through surveys, modeling, research studies, and management experiments into a continuously evolving management program for CWD. STATEWIDE MONITORING PROGRAM Goals: 1. Provide reliable estimates of CWD distribution and prevalence to serve as a basis for management decisions and public information. 2. Provide information to improve understanding of CWD epidemiology. 3. Continue developing efficient and reliable techniques for detecting and monitoring CWD in freeranging populations. �115 Strategy: Reliable estimates of CWD prevalence in wild deer and elk populations are needed to guide policy decisions and monitor efficacy of management efforts. We will continue to monitor deer and elk populations throughout Colorado for occurrence of chronic wasting disease using a combination of extensive and intensive approaches. These have been organized and conducted as follows: Targeted (= clinical disease) surveillance: Ongoing statewide "targeted" surveillance (i.e., submissions of "suspect" deer & elk) will be used to determine & monitor changes in CWD distribution. Based on·. experiences in Colorado and elsewhere, this appears to be the most sensitive approach for determining CWD distribution & detecting range extensions (Miller, 1997). A program for acquiring, examining, reporting on, and summarizing "suspect" CWD cases occurring throughout Colorado has been in place since 1990 (Miller, 1997). This program has served to increase ongoing surveillance efforts by field personnel statewide by encouraging submission of carcasses from deer or elk showing clinical signs resembling CWD. A formalized process for submitting cases has been developed; this process includes criteria for acceptable submissions, submission forms, handling instructions, and a system for networking submissions within CDOW and among the 3 Colorado State Veterinary Diagnostic Laboratories and the Wyoming State Veterinary Laboratory. Under this program, deer and elk showing clinical signs consistent with those seen in chronic wasting disease are collected by field personnel statewide; a "suspect case" profile has been defined (Table 1). Brain tissues are subsequently examined for evidence ofspongiform encephalopathy. Where possible, whole carcasses will be submitted for complete necropsy; at minimum, heads from suspect animals will be submitted for examination and sampling. Ancillary diagnostics, including histopathology (Williams and Y oung,1993) and immunohistochemistry of brain tissue (Spraker et a1., 1997), will be performed on all suspect cases; other ancillary tests may also be used to confirmor support diagnoses. Preliminary examination and/or test results and final reports will be faxed to CDOWs Wildlife Research Center. Pertinent data from preliminary and final reports will be entered into a permanent database, and copies of reports will be filed as well as sent to appropriate field personnel. Results from targeted surveillance will be used to identify new potential foci of CWD infection for further evaluation via harvest or road-kill surveys. Harvest and road-kill surveys: Surveys of harvested or, in some areas, road-killed deer and elk will be conducted in endemic and high-risk DAUs/GMUs to estimate and monitor CWD prevalence. We will continue conducting annual surveys in DAUs or GMUs where CWD prevalence is >2% ("endemic foci") to obtain reliable-estimates of prevalence to use as a basis for monitoring natural trends, as well as responses to management interventions. Annual surveys will be designed to provide an estimated prevalence within ±2% at a 95% level of confidence for affected populations where true prevalence is 1%. In addition to annual surveys of endemic foci, we will also continue conducting a series of surveys for CWD in other select deer and elk populations throughout Colorado over the next 5 yrs, with a goal of determining statewide distribution of CWD. Our surveillance strategy is as follows: Areas newly identified as infected via targeted surveillance (e.g., GMU 29, South Platte River corridor GMUs) will be surveyed sufficiently to obtain a reliable prevalence estimate. In areas where estimated prevalence is <1 %, CWD will be regarded as a sporadic disease and populations will be monitored at ~5-yr intervals to detect increases in prevalence. Areas where estimated prevalence is >2% will be regarded as new endemic foci, and will be monitored annually to track changes in prevalence as described above. We also plan to continue conducting surveys of at least 5 other select deer populations where CWD has not been reported �116 previously. Target areas will include both high-risk (e.g., Middle Park, Piceance Basin) and low risk (e.g., San Luis Valley, Gunnison, southern Front Range) populations. Surveys will be designed such that the probability of failure to detect at least 1 case of CWD in an apparently-unaffected deer or elk population will be ,::::0.01even if herd prevalence is 1%. Finally, we will attempt to gather a sufficient number of random samples from DAUs statewide (n-l,OOO) to determine whether or not CWD should be generally regarded as endemic in Colorado's deer populations. A tentative schedule for the foregoing surveillance is outlined in Table 2. Harvest surveys will be conducted using techniques developed in Colorado since 1991. Because regulatory requirements for head submissions increase sample sizes >4-fold, submissions will be mandatory in most GMUs where surveys are being conducted. Limited licenses may also be used in select GMUs to aid in increasing compliance. Unstaffed barrel sites appear to be the most efficient method for sample collection, and will continue to be used as the primary method for conducting harvest surveys. Brainstem (obex) from ~ I-yr-old deer and elk will be.collected for CWD testing, and immunohistochemistry (IHC) using USDA monoclonal antibody F89/160.1.5 (O'Rourke et al., 1998) will be used as the primary screening tool. Reported prevalence estimates will continue to be based on numbers ofIHC:·positive cases; because IHC appears more sensitive than histopathology in detecting preclinical CWD cases, IHC-based prevalence estimates should provide a relatively unbiased measure of disease trends over time and allow more direct comparison between field data and simulation model predictions. EXPERIMENTAL MANAGEMENT There is no precedent for attempting to manage a TSE in free-ranging wildlife. Moreover, the biological need for such management is unclear at present. Programs for managing or eliminating TSEs of domestic livestock have proven only marginally successful, and the lack of obvious success in eradicating scrapie or BSE from endemic countries, compounded by epidemiological differences between CWD and other TSEs, make such programs rather poor models for prospective CWD management. Several fundainental features of CWD epidemiology are understood poorly, if at all; in particular, the influences of host (e.g., population density, intraspecific vs. interspecific transmission, genetics) and environmental factors (e.g., contamination, reservoirs, weather) on CWD dynamics remain undescribed. In light of the uncertainties associated with its epidemiology and ecology, we believe CWD management should be approached as an experiment, thereby allowing us to learn as we . take actions intended to reduce prevalence and limit distribution. The CDOW's initial approaches for attempting to control CWD in free-ranging deer and elk were outlined in an action plan for CWD drafted in 1995 (Miller et al., 1995; Appendix A). In addition to calling for a comprehensive public information campaign, this plan identified tactics for limiting distribution and reducing prevalence of CWD based on contemporary knowledge of the problem. Prompt removal of affected animals and enforcement of feeding prohibitions were recommended to help reduce CWD prevalence and transmission. Policies and regulations were also recommended, and subsequently instituted, to prevent wider distribution of CWD via human activities (e.g., trans locations, rehabilitation, game ranching). The plan also identified a need for more extensive surveillance to gather data on CWD distribution and prevalence. As more data on CWD distribution and prevalence have been gathered over the last 2.5 yrs, it has become apparent that CWD is more widely distributed and more prevalent in northeastern Colorado (and elsewhere) than initially believed. It follows that policy, goals, and strategies for CWD management in Colorado should be reevaluated in the context of new data, as well as changes in social and political attitudes toward animal TSEs. �117 Management policy: At present, the CDOW has disease in wild deer andelk. The disagreement over how to resolve increase recreational opportunities no stated lack ofa conflicts for deer policy related to the management of chronic wasting clear policy statement has led to internal confusion and with other policies and Long Range Plan objectives (e.g., and elk hunters). Recommendation: We recommend adoption of the following policy to guide decisions on deer and elk population management. in DAUs where CWD is endemic: Pl. The Colorado Division of Wildlife is committed to minimizing the threat that chronic wasting disease (CWD) poses to Colorado's native deer and elk resources. In DAUs where CWD is endemic, goals of reducing the occurrence and the further spread of CWD will serve as the primary basis for setting management objectives. Prospective management goals: We have. identified four overarching goals that could serve as a basis for decisions related to managing CWD: 1. 2. 3. 4. "natural regulation", containment, reduction, eradication. These goals represent a continuum of possible levels of intervention, ranging from benign. rs neglect to aggressive action. There are clear advantages and disadvantages associated with each of .. these prospective management goals. Although each progressive level offers more complete control of the problem, better control is accompanied by ever-increasing technical, fiscal, and political challenges, as outlined below: l."Naturai regulation": Directing no specific management intervention toward CWD beyond culling of reported clinical cases represents the status quo approach to both disease arid deer/elk population management. It could be argued that the ongoing activities described above constitute an attempt to manage CWD. Although the impacts of this approach on CWD prevalence in endemic foci remain undetermined, prevalence in endemic foci appears to have remained stable over at least the last J yrs. This approach offers the fewest technical challenges, may be regarded by some as "safest'unlight of the myriad of uncertainties in CWD epidemiology, and in the shortterm would be most sparing oflocal resources. Unfortunately, simulation models forecast that problems with CWD will likely get worse in affected populations if left largely unmanaged, the disease could become more widely distributed, and there are tangible potential impacts on wildlife resources, recreational opportunities, and revenues. In addition, perceived "inaction" may be unacceptable to some sportsmen's groups, agricultural interests, and perhaps the general public, and could lead to the loss of management authority by CDOW. Finally, opting for "natural regulation" seems biologically irresponsible. 2. Containment: Managing primarily to prevent spread of CWD from endemic foci would require additional knowledge of the mechanisms and probable routes of CWD dissemination, but many of the important strategic components needed to support this strategy are already in place. The likelihood of trans locating infected animals from endemic areas to other areas in Colorado or �118 elsewhere was significantly reduced by regulations adopted in 1996 (ch 14 ref). Between existing data from previous movement studies (e.g., Kufeld et al., 1989; Kufeld and Bowden, 1995) and data from studies initiated more recently, major natural emigration corridors from eastern Larimer County could probably be identified and subsequently might be interrupted via intensive culling operations. This approach, if successful, would limit the magnitude of the CWD problem while largely sparing local resources. This alternative may be more acceptable to sportsmen's groups and the general public than to agricultural interests, but is probably biologically justifiable. However, such an approach would require an essentially infinite long-term commitment and yet will not eliminate CWD entirely; prevalence could conceivably increase in endemic areas. There would likely be some local impacts on resources and recreational opportunities, perhaps accompanied by local public resistance. 3. Reduction: It may be possible to manage affected deer or elk populations to reduce CWD prevalence in endemic foci. A goal of prevalence reduction clearly demonstrates intent to limit the magnitude of the problem. It seems likely that this. approach would also provide for containment of CWD. Attempting to reduce CWD prevalence might improve "consumer confidence" among hunters in endemic GMUs. This alternative may be the most widely acceptable among sportsmen's groups, agricultural interests, and the general public in terms of responsible disease and resource management. It is probably biologically justifiable in view of projected long-term effects ofCWD on infected populations and the potential for CWD to spread from endemic foci if left unmanaged. As with the foregoing goal of disease containment, prevalence reduction will require a long-term commitment and may not eliminate CWD from endemic areas; depending on specific tactics employed, prevalence could even increase despite management efforts. This approach would likely cause more severe impacts on local resources and recreational opportunities, and consequently would be more likely to foster local public resistance to proposed management actions. 4. Eradication: Eradication is probably the most desirable goal of any attempt to manage CWD. Whether it would be feasible to completely eliminate CWD from Colorado remains highly questionable. Eradicating CWD would be the best means of restoring for "consumer confidence" among northeastern Colorado deer and elk hunters, and would presumably nullify concerns of traditional and alternative livestock interests about the potential for transmission to privately-owned animals. Perhaps most importantly, eliminating CWD would be the most effective way to preempt threats of its eventual spread to other native deer and elk populations in Colorado and elsewhere. Although desirable in concept, CWD eradication is probably infeasible given the complexities and uncertainties of its epidemiology. Eradication of CWD would require long-term commitment, and probably would exaet catastrophic impacts on resources and recreational opportunities in affected areas; broader ecological impacts (e.g., on predator-prey balances) could also be severe. Public resistance is likely to emerge, at least locally, as details of such a management program emerge. Finally, it is questionable whether the potential ecological costs of CWD eradication are biologically justifiable in light of the uncertainty about long-term impacts of the disease itself. Recommendations: Based on current understanding of CWD epidemiology, prevalence, and distribution in free-ranging deer in northeastern Colorado, we believe eradication is an extreme and unjustifiable management goal at this time. In light of the magnitude of prevalence and distribution CWD has reached in Larimer County deer populations under historical management regimes, however, continuing to rely on current management approaches seems equally unjustifiable. Consequently, we recommend an intermediate approach with two goals for managing CWD in deer and elk: �119 Gl. limit distribution ofCWD occurs. to no more than the 5 deer DAUs and 2 elk DAUs where it already G2. reduce average CWD prevalence among deer and elk to <1 % in each endemic DAU and <2% in each endemic GMU; and . '~ We believe achieving the foregoing management goals will serve to protect Colorado's deer and elk resources, restore confidence in the quality of deer and elk harvested in northeastern Colorado, and ameliorate political pressures from agricultural constituencies concerned about potential for transmission of CWD to domestic livestock or privately owned wildlife. Prospective management strategies: Effective strategies for managing CWD or any other TSE in free-ranging wildlife have never been identified or evaluated. Many conventional disease management options are precluded from consideration because vaccines, therapeutics, and live-animal tests for CWD are presently unavailable (Table 3). Contrary to experiences with domestic sheep (Goldmann et al., 1994), available data indicate relatively uniform genetic susceptibility to CWD among deer (K. I. O'Rourke and M. W. Miller, unpulb. data); consequently, genetic selection would probably have no impact on prevalence even if tools forits implementation were available. Similarly, controlling reproduction alone probably would be ineffective even if practical tools were available because maternal transmission alone is unlikely to be sustaining CWD prevalence at levels currently observed in some affected deer .populations. All remaining options we can identify to reduce CWD prevalence (Table 3) involve some form of aggressive population control directed at diminishing or preempting-disease transmission -- themagnitude and duration of such control will be driven by management objectives and population responses. Based on current understanding ofCWD epidemiology (Miller 1997, Miller et al., 1998, M.W. Miller, unpubl. data), we believe that strategies capable of lowering CWD transmission rates by reducing numbers of infected animals and point sources of environmental contamination (e.g., feeders) should be most effective in lowering prevalence; although the precise mechanisms of transmission have not been identified, those details will probably have little influence on the resolution of CWD management at the population level. General models predict that random culling should be less effective than selective culling in reducing disease prevalence (Barlow, 1996; McCarty and Miller, 1998). However, field data indicate that management strategies based on culling clinical suspects probably will miss a large proportion of the infected (and potentially infectious) individuals in a population; moreover, there is no practical way to apply "test and slaughter" regimes without an antemortem diagnostic-test for preclinical CWD. Accelerated culling of early clinical cases, either by humans or via predators, may help reduce CWD transmission and prevalence. Alternatively, if the probability of CWD transmission is density-dependent, then reducing deer or elk densities in endemic areas should lead to reduced CWD prevalence; to date, the hypothesis of density-dependent disease transmission remains untested. Obvious strategies for limiting CWD distribution remain equally elusive. The mechanisms driving the spread of CWD among deer and elk populations are even less apparent than those driving disease dynamics. It is possible that CWD distribution to date has been influenced by regular and/or random movements of deer and, less commonly, elk. Previous studies (Kufeld et al., 1989; Kufeld and Bowden, 1995; S. Steinert, pers. comm.; others?) have shown that proportions of both foothills and river bottom deer populations in northeastern Colorado are either migratory or mobile; the movement patterns documented in these studies provide at least a partial explanation for observed CWD distribution. It is also possible that CWD influences its own distribution by changing behaviors, range ..... �120 fidelity, and movement pattens of affected animals; observations of repetitive pacing in captive deer and elk affected by CWD could be manifested as extensive wandering movements of free-ranging individuals. Whether or not movement patterns or dispersal rates are density-dependent remains unclear; either way, reducing population size should also reduce the probability of affected individuals canying CWD to new areas. (However, we do recognize the possibility that at some lower threshold reduced density may actually promote deer and elk movements, particularly during the breeding season). Recommendations: Based on current uncertainty about how to effectively disrupt the processes that influence CWD epidemiology, prevalence, and distribution in free-ranging deer in northeastern Colorado, we recommend initiating controlled management experiments directed toward testing one (or both) of the following hypotheses: HI. CWD transmission and prevalence infoothills and river bottom deer populations can be diminished by a) selective culling; b) increasing natural mortality via mountain lion predation; OR c) reducing deer density in affected populations. H2. CWD distribution in foothills and river bottom habitats can be diminished by a) selective culling; . b) increasing natural mortality via mountain lion predation; OR c) reducing deer density in affected populations. We further recommend that the experiment(s) be designed and conducted in a way that facilitates achievement of two short-term management objectives: MI. to stem dispersal ofCWD via deer movements along the South Platte River corridor (GMUs 94, 951,96,91, and 92). M2. to reduce CWD prevalence among deer inhabiting the four most severely affected GMUs (9, 191, 19,20). Because CWD prevalence is low «1%) in elk, we recommend that populations in DAUs E4 and E9 should be maintained at or below current densities and monitored via harvest surveys at least every 5 yrs to detect potential increases in prevalence that.might warrant more aggressive management action. :. Design of Management Experiment We recommend using an adaptive resource management approach (Walters, 198_) to test candidate strategies for reducing CWD prevalence and distribution. Based on initial evaluations of these candidate strategies (Table 4), more detailed management experiments will be developed; as part of that process, decisions will be made on which, if any, approaches to pursue and where CWD management will be attempted. Tactics for accomplishing management approaches (e.g., harvest, selective culling, ground/aerial gunning, changes in predator harvest, etc.) will follow from internal consensus on which tactics are viable under the social and political constraints imposed by the area(s) selected for study. Prospective management tactics: Tactics supporting selected management strategies: �121 - harvest? (role of public hunting?) - shift emphasis from recreational opportunity to population management - modifications of GMU/ "management area" boundaries to focus mgt & distribute pressure - season structurellength & bag limits- reg changes - schedule - culling? - ground vs aerial vs capture & kill options - public vs agency c; - private land issues - fertility control? (probably not for reductions, but maybe to maintain) - agent, delivery, cost? - reduce predator quotas? (close seasons?) - predator introductions? - poisoning? - other? , '; . <" Issues surrounding CWD management - Should public hunting continue in endemic areas? What are public desires/expectations related to CWD management? Can we impose intensive population management on private lands? What are reasonable expectations for funding and long-term agency/political/public Others? commitment? Outstanding tasks (partial list) Human dimensions -- Public service -Funding (PBE)-- For both internal and external publics, seek consensus on: what direction to take, which strategies are most viablelleast objectionable, etc., impacts on recreational opportunity, trade-offs Seek/gain cooperation, access, etc. (follows from lID issues above) Estimate support required for evaluation of management activities, as well as fiscal impacts of candidate management approaches LITERATURE CITED Goldmann, W., N. Hunter, G. Smith, 1. Foster, and 1. Hope. 1994. PrP genotype and agent effects in scrapie: change in.allelic interaction with different isolates of agent in sheep, a natural host of scrapie. 1. Gen. Virol. 75:989-995. Miller, M. W. 1997. Monitoring and managing chronic wasting disease in Colorado. Pages _-_ in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-I0, Work Plan 2, Job 17. Colorado Division of Wildlife; Fort Collins, Colorado, USA. (in press) Miller, M. W. 1998. Monitoring and managing chronic wasting disease in deer. Pages _-_ in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-l1, Work Package 3001, Task 3. Colorado Division of Wildlife, Fort Collins, Colorado, USA. (in press) Miller, M. W. 1998. Monitoring and managing chronic wasting disease in elk. Pages _-_ in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-ll, Work Package 3002, Task 3. Colorado Division of Wildlife, Fort Collins, Colorado, USA. (in press) �122 Miller, M. W., M. A. Wild, and E. S. Williams. 1998. Epidemiology of chronic wasting disease in captive Rocky Mountain elk. J. Wildl. Dis. 34:532-536. Spraker, T. R., M. W. Miller, E. S. Williams, D. M. Getzy, W. J. Adrian, G. G. Schoonveld, R. A. Spowart, K. 1. O'Rourke, J. M. Miller, and P. A. Merz. 1997. Spongiform encephalopathy in free-ranging mule deer (Odocoileus hemionus), white-tailed deer (Odocoileus virginianus), and Rocky Mountain elk (Cervus elaphus nelsoni) in northcentral Colorado. J. Wildl. Dis. 33:1-6. Williams, E. S., and S. Young. 1980. Chronic Wasting disease of captive mule deer: A spongiform encephalopathy. Journal of Wildlife Diseases 16:89-98. and __ . 1982. Spongiform encephalopathy of Rocky Mountain elk. Journal of Wildlife Diseases 18: 465-471. and __ . 1992. Spongiform encephalopathies in Cervidae. Revue Scientifique et Technique Office International des Epizooties 11:551-567. ---' and __ . 1993. Neuropathology of chronic wasting disease in mule deer (Odocoileus hemionus) and elk (Cervus elaphus nelsoni). Veterinary Pathology 30:36-45. _-J _-J Table 1. Profiles used in chronic wasting disease targeted surveillance. • Species: • Age: mule deer white-tailed deer elk 2:. 18 months • Signs: emaciated and abnormal behavior &lor indifference to human activity &lor increased salivation &lor tremor, stumbling, incoordination &lor difficulty or inefficiency in chewing/swallowing increased drinking and urination &lor �123 Table 2. Tentative schedule for systematic statewide chronic wasting disease surveys of deer populations using harvest and/or road-kill samples. . CWD Status GMUs Years surveyed Endemic foci (:::::2%) 9, 19, 191,20 1996-2006 Endemic (~1%) .29 90,91,92,93 94 95,951,96 1997-2000 1997-1998;2004-2006 1997-2000 1996-1998,2004-2006 "High Risk" 87 :}8, 28, 37, 371 6, 16, 17, 161, 171 22 1997-2000 1998 1999-2001 1999 Other areas . 104 66,67 83 "Uncompahgre" "Southern Front Range" Random Statewide ..r- -. ~·i .. 1996-2000 1997-1998 1994-1993, 1998-1999 2000 2001 2002-2003 [Note: Survey targets and schedules subject to changes influenced by new data and availability of resources supporting CWD surveillance activities.] Table 3. Prospective strategies for chronic wasting disease management. Strategies for limiting distribution - preclude human-caused translocations (mostly done) - interrupt migration corridors (fencing?) - change habitat (improve/degrade to influence animal distribution?) - alter migratory behavior (how?) - reduce density (relationship between density & distribution untested) - other? Strategies for reducing prevalence - vaccination (no vaccine available) - treatment (no effective therapeutic drugs available) - genetic selection (evidence of uniform susceptibility) - fertility control (maternal transmission probably a minor contribution) - eliminate clinical suspects (misses carriers; ineffective to date) - test & slaughter (no reliable, practical live animal test) - selective culling (what criteria? mechanism?) - foster natural mortality (e.g., predation) to promote selective culling - reduce density (relationship between density & transmission untested) - other? �124 Table 4. Initial CWD management tactics to be evaluated via field studies and/or simulation modeling. Candidate tactic Schedule Selective culling via observation (± baiting) Modified "test and slaughter" (model/field) Selective culling via predation (modeling) Density reduction (modeling) Fertility control (modeling) 2/99-4/99 5/99-4/00 5/99-9/99 5/99-9/99 5/99-9/99 EXPERMIENTAL PLAN MANAGEMENT PROPOSAL -- NOT CURRENTLY INCLUDED IN DRAFT In all, eight management areas will be included in this before-and-after-controlled-intervention (BACI)experiment (Fig._); management areas will be paired, with four areas targeted for density reduction (50% reduction over a 2-yr period, then maintained at 50% of pretreatment density for 5 yr)' and four held under status quo management regimes (i.e., discourage feeding, eliminate suspects, etc.) as control areas (Table _). We will stagger the start of management treatments to minimize confounding effects of temporal variation on CWD prevalence and distribution (Table _). Deer density and CWD prevalence will be estimated annually in each management area, and mean CWD prevalences before and after management intervention will be compared to test density-dependent transmission hypotheses and assess the efficacy of treatment in reducing CWD prevalence. Similarly, CWD prevalence and distribution in GMUs adjacent to treatment and control areas will be compared to test hypotheses on density-dependent dispersal and assess the efficacy of treatment in stemming CWD dispersal. Table _. Treatment assignments and start dates for areas included in CWD management experiment. Location Treatment Control Start date northern Larimer County GMU9 GMU 191 1999-2000 southern Larimer County GMU 1ge/20ea GMU 19w120wb 2000-2001 South Platte River" bottom/plains GMU91 GMU92 1999-2000 South Platte River bottom/plains GMU96 GMU951 2000-2001 b GMU 19e120e will be the portions ofGMUs 19 and 20 east of the Pingree Park Road and ??? GMU 19w/20w will be the portions of GMUs 19 and 20 west of the Pingree Park Road and ??? 1 This preliminary treatment regime is subject to modification guided by simulation modeling exercises that will be completed later this year. �125 Colorado Division of Wildlife Wildlife Research Report July 1999 JOB PROGRESS REPORT Smteof __ ~ ~C~o~lo~r=ad~o~ _ Cost Center 3430 Mammals Program Project No. W..:...:..,_-1=5=3--=-R=-_",1=-2_,.....-_ Work Package No. ___,3::....:0=0~1 _ Deer Management Task No. -'4'-_ Regulation of Mule Deer and Elk Population Growth by F ertilitv Control Period Covered: July 1, 1998 - June 30, 1999 -Author: Dan L. Baker Personnel: T. M. Nett, M. A. Wild ABSTRACI' We conducted preliminary investigations to evaluate the effectiveness and side effects of two contraceptive agents in captive mule deer and elk. We treated female mule deer with a permanent fertility control agent (GnRH-PAP) and female elk with a temporary, reversible contraceptive. In mule deer, serum luteinizing hormone (LH) levels were reduced by 55-78% for at least 6 months following treatment. For elk, serum LH concentrations were reduced to baseline levels for 90 days posttreatment. Body weight dynamics, blood chemistry, hematology, and social behavior of female mule deer and elk appeared normal compared to untreated animals. �126 �127 REGULATION OF MULE DEER POPULATION GROWTH BY FERTILITY CONTROL . Dan L. Baker P. N. OBJECTIVES 1. To develop a practical and acceptable technology to inhibit reproduction in mammalian species which cause damage or constitute a significant public nuisance. 2. To demonstrate the feasibility of such technology in a field application. 3. To predict population impacts of alternative contraceptive regimens using simulation modeling. SEGMENT OBJECTIVES 1. To develop and test GnRH-toxin conjugate in captive mule deer and elk. 2. To develop a si~ulation populations. model to evaluate the ability of contraceptives to regulate large ungulate INTRODUCTION Segment Objective 1: GnRH-ToxinlAgonist Controlling the abundance of animals is fundamental to contemporary wildlife management. This is particularly true for wild ungulates. The most compelling motivation for regulating ungulate numbers is that overabundance causes problems that can be biological, economical, or political in scope (Jewel and Holt, 1981). Resolving these problems requires controlling population growth. Wild ungulate populations have traditionally been regulated by influencing death rates using controlled harvest or CUlling. However, there are an increasing number of circumstances where these traditional methods are not feasible. The use of fertility control to decrease birth rates is one of the most promising approaches to the long-term control of overabundant wild ungulates. During the past decade, research aimed at developing effective contraceptives for free-ranging wildlife populations has accelerated. These efforts have resulted in development and testing of a wide variety of potential contraceptive agents (Kirkpatrick and Turner 1985, Warren et aI. 1995). One of the most promising new non-steroidal, non-vaccine, approaches to contraception involves synthetic analogs of gonadotropin-releasing hormone (GnRH). GnRH is a molecule produced in the hypothalamus of the brain. It directs specific cells in the pituitary gland to synthesize and secrete two important reproductive hormones; follicle stimulating hormone (FSH) and luteinizing hormone (LH). These latter two hormones, known as gonadotrophs, control the proper functioning of the ovaries in the female and testes in the male. Analogs of GnRH have the potential to either permanently or temporarily inhibit reproduction. For most free-ranging wild ungulate applications, permanent sterilization or a combination of permanent sterilization and culling have been proposed as the most efficacious approaches to population management (Hone 1992, Garrot 1995, Hobbs et al. 1999, in press). For this application, �128 superactive analogs ofGnRH are coupled to a cytotoxin. The GnRH-toxin conjugate specifically targets the gonadotroph cells and permanently inhibits the ability of the cell to secrete FSH and LH. This approach has several potential advantages over other methods of contraception. These include: 1) a single treatment should permanently sterilize an animal 2) the same treatment should be effective in both males and females and in different mammalian species 3) GnRH-toxin conjugate will be metabolized from the body within a few days of treatment 4) the proteinaceous nature of GnRH-toxin conjugate eliminates the possibility of passage through the food chain. 5) the small volume required for effective contraception would facilitate microencapsulation and administration by syringe dart or biodegradable projectiles. In other situations where wildlife managers need to maintain flexibility in the use of fertility control, reversible contraception may be desirable. Examples of these situations include 1) wild ungulate populations exposed to periodic, severe, unanticipated winter mortality, 2) populations with low genetic variability, 3) populations that cannot be effectively monitored, 4) populations where public attitudes are opposed to permanent contraception, or 5) populations where non-lethal hunting recreation is a primary management objective. In these situations, superactive analogs of GnRH without the toxin subunit would be more appropriate. The inhibitory actions of long-term GnRH analog agonist on the ovulatory cycle of humans and other mammals is well-established (Casper and Yen 1979, Fraser 1983, Fraser et al. 1987, Concannon et al. 1991). Constant administration of high .doses of GnRH agonist results in down regulation of the pituitary GnRH receptors and suppression ofsecretionofLH andESH. Continued· treatment suppresses LH secretion, preventing the maintenance of normal.luteal function, and thus prevents viable pregnancy. Inhibition of ovulation caused by chronic administration of GnRH agonist has been successful in several species, including dogs (Vickery et al. 1989), cattle (Herschler and Vickery 1981), sheep (McNeilly and Fraser 1987), white-tailed deer (Becker and Katz 1995), and elk (Baker and Nett, 1999). Evidence from studies on pituitary receptors and gonadotropin content in experimental animals treated by long-term infusion ofGnRH agonist shows that sustained release is themost effective approach for temporarily suppressing pituitary-gonadal function (Clayton 1982, Sandow 1982). The practicality of this approach, however, is dependent upon development of a long-acting, slow-release preparation of agonist that can be remotely delivered. Recently, a practical mode of administration using subcutaneous implants has overcome the need for constant mechanical infusion of the analog. Slow release formulations of superactive GnRH agonist are now commercially available and have been shown to be effective in suppressing the pituitary ovarian axis for up to 6 months in a variety of mammalian species (Fraser et al. 1987, Asch et al. 1985). To our knowledge, only limited investigations have been conducted with either of these fertility control techniques on wild ungulates (Baker 1994, Baker et al. 1995, Becker and Katz 1995). Thus the specific objectives of these investigations were: 1) To evaluate the effectiveness and duration ofa single dose application of GnRH-toxin conjugate and GnRH - Agonist in preventing normal production of reproductive hormones in captive mule deer and elk. 2) To evaluate the effects of these contraceptives on the general health, blood chemistry, and hematology in mule deer and elk. �129 Segment Objective 2: SimulationModeling Using either culling or contraceptives to meet deer population objectives will require, wildlife managers to choose specific tactics of treatment Choices must be made on the number and age to treat, frequency of treatment, species, sex, etc, Decisions on the best tactics will depend on comparing the effects of alternative management actions on population behavior. To assist in these decisions, we developed an interactive simulation model of deer population dynamics to allow managers to evaluate alternative treatment regimes on simulated populations before applying them to real animals. Critical to model performance is knowledge of population demographics of the RMA deer herd and requires information on sex and age composition, recruitment, pregnancy rates, fetal sex ratio, and estimates of population density. This information together with knowledge of the habitat resources available to deer will provide a sound biological basis for management of deer populations at the RMA. METHODS AND MATERIALS GnRH- Toxin! Agonist Experiment 1: Evaluation of GnRH-PAP in mule. deer. Previously, we conducted controlled experiments with captive mule deer to determine the most effective dose of GnRH analog and the season of the year when treatments would be most effective in preventing fertility (Baker 1997). Here, we evaluate the effectiveness of GnRH..,P AP in preventing normal production of luteinizing hormone (LH) and the duration of effectiveness. We will evaluate contraceptive effectiveness in 4 ovariectomized mule deer. . Protocol for Ovariectomy in Mule Deer Tonic secretion of pituitary LH is the result of an interplay between a stimulatory input from the brain and an inhibitory feedback from the gonads. In the intact female, estradiol secreted from the gonads is a potent negative feedback hormone on LH secretion during anestrus (Goodman and Karsch 1980). Since measurement ofLH secretion is the primary indicator of GnRH-toxin conjugate effectiveness, it is imperative that female mule deer in this experiment be ovariectomized. Ovariectomies of mule deer were conducted at FWRF during the week of May 11, 1998. Deer were isolated and fasted for about 24 hr prior to surgery to alleviate regurgitation and aspiration of rumen contents. On the day of surgery, anesthesia was induced with intramuscular (IM) administration of 100 mg xylazine w/o 500 mg ketamine depending on response of the animal. Deer were then intubated and surgical anesthesia was maintained using a rebreathing circuit with isoflurane. Anesthetized deer were prepared for surgery by clipping hair in the abdominal area. They were then carried to a designated surgery area., placed in right lateral recumbency, and the surgical site prepared using standard surgical scrub. The surgical area was prepared with sterile towels and a surgical drape. Ovariectomy was performed via mid-ventral laparotomy and require 20 - 30 min per animal. Surgeons were attired in a sterile gown, mask, cap, disposable shoe covers and sterile gloves. Surgical assistants wore cap, mask, and disposable shoe covers. The ovarian artery was ligated prior to removal of the ovary. The incision was closed using a continuous suture and the skin closed with interrupted mattress stitches. We reversed anesthesia with yohimbine at a dose of 0.2 mg/kg (N). To minimize infection, deer were given ceftiofur sodium (1.1 mg/kg IV) administered perioperatively. Phenylbutazone (4 mglkg) was administered orally when deer were partially recovered from anesthesia �130 and every 48 hr for up to one week, if needed. Animals were placed in isolation pens for 24 hr and were observed every 5 min during recovery from anesthesia. This procedure followed ARBL Standard Operating Procedure #1 - Protocol for OvariectomylLuteectomy in Ewes and was modified for use in mule deer at FWRF. GnRH-toxin Conjugate Protocol Approximately 6 weeks following ovariectomy, GnRH-toxin was administered. Four ovariectomized deer were moved from 5 ha pastures to individual isolation pens, sedated with xylazine (100 mg 1M), and administered IV an optimum dose of GnRH-toxin (3 J-lgl50 kgBW). Deer were then placed in individual isolation pens and fitted nonsurgically with indwelling jugular catheters. Indwelling catheters remained in deer for up to 6 weeks and were checked and flushed daily. Deer remained in individual isolation pens for the first 6 weeks of the trial. This minimized tranquilization and general handling of deer and allowed daily evaluation of intake and general health of experimental animals. After 6 weeks, catheters were removed and deer were returned to 5 ha pastures. For subsequent trials, deer were removed from 5 ha pastures one day prior to GnRH trials and fitted with indwelling jugular catheters. Sampling was conducted the next day, catheters removed and deer returned to 5 ha pastures. One week after GnRH-toxin treatment, GnRH analog challenge trials were conducted. Experimental animals were moved to individual isolation pens and 3 J-lgl50 kg BW GnRI:i analog (Baker 1997) was administered through the indwelling cannula Blood samples (5ml) were collected at 0,30,60,90, 120, 180,240,300, and 360 min postinjection. After collections, blood was held at 4 C for 24 hours and serum obtained by centrifugation. Serum was stored at -20 C until analyzed. GnRH ' analog challenge trials were repeated each weekfor 6 weeks, then twice monthly for 3 months, then _' once monthly for 1 year. . Serum concentrations ofLH were quantified by means of ovine LH RIA (Niswender et al. 1969). The limit of sensitivity of the assay is 0.4 nglml. Response of the pituitary to GnRH-toxin conjugate was be assessed by 1) maximum LH response achieved postinjection, 2) total amount ofLH secreted (nglml/min); estimated by calculating the area under the LH curve, and 3) an LH response of < 2.5 ng/ml following GnRH challenge was be considered sufficiently low to prevent ovulation. Experiment 2: Evaluation of GnRH- Agonist (Lupron) in elk. Here, we conducted a pilot experiment to evaluate the effectiveness of GnRH agonist analog in suppressing ovarian function in elk. We determined the most effective dose and evaluated the duration of effectiveness. We also used these studies to evaluate the safety and side effects (if any) of treatment on a small number of animals before proceeding to a larger investigation. The information collected in the pilot study was used to assess individual variation in treatment responses and to conduct statistical power analysis to increase efficiency of future research efforts, We conducted controlled experiments with adult, tame female elk at the Colorado Division of Wildlife's Foothills Wildlife Research Facility, Fort Collins, Colorado during November 1998 to March 1999, Objective: To determine the amount of GnRH agonist analog that will induce half-maximal release of LH in female elk during estrus and evaluate duration of effectiveness. Design: We observed the LH response offemale elk to three levels ofGnRH agonist (0, 19,39, and 78 J-lglhour) in a completely randomized design with two animals per level. Based on results from previous studies, we chose a range of doses that should include the biologically effective range for elk. �131 We evaluated the effective duration of treatments by monitoring ovarian function throughout the remainder of the breeding season (Nov - Apr). Dosage Trial - This experiment was conducted during the breeding season to insure that the pituitary gland was at its most active state when stimulated by the GnRH agonist. On Day 1 of the experiment, 9 female elk were moved from 5 ha pastures to individual isolation pens, sedated with xylazine hydrochloride (50-100 mg/animal, IM), fitted nonsurgically with indwelling jugular catheters and administered subcutaneously one of three experimental doses of GnRH agonist analog. The sampling period began at 1000 h. Blood samples (5 ml) were collected at 0,60, 120, 180,240,300,360,480 min, 12,24,36,48,84, and 240 h postinjection. Animals remained in isolation pens during the 10 day trial. Elk were observed daily for side effects of treatments (if any) and catheters flushed with sterile saline solution. Following the last blood collection, catheters were removed and animals returned to 5 ha paddocks. After collection, blood was held at 4 -c for 24 h until serum was obtained by centrifugation. Serum was then stored at - 20°C until analyzed for LH. Serum concentrations ofLH was quantified by means of ovine LH RIA (Niswender et al. 1969). The limit of sensitivity of the assay was 0.4 ng/ml. Analysis: Responsiveness of the pituitary to GnRH agonist analog challenge was assessed in four ways: 1) maximum LH response achieved postinjection minus baseline 2) time required to reach maximum LH 3) total amount ofLH secreted (ng/mllmin) 4) the amount ofLH required to induce half maximal release ofLH (EDso). Response to treatments was analyzed with a one way analysis of variance for a completely random design with 3 levels of dose as treatments and 3 individual animals as replications. Simulation Model We developed an interactive simulation model to allow RMA biologist to evaluate alternative mule deer culling strategies. This model combines extant knowledge of mule deer population biology with measured population parameters at the RMA to predict the proportion of male and female deer that need to be removed each year to accomplish management objectives for this herd. Table 1. Principal parameters in culling simulation model for mule deer at RMA. Definition Value Adult/yearling female survival Adult/yearling male survival Fawn survival coefficient a Fawn survival coefficient 13 Fetal sex ratio ('Yo males) December recruitment 0.90 0.90 1.20 -0.053 0.47-0.62 0.59-0.72 Measured No No No No Yes Yes Reference Whittaker, 1992 Whittaker, 1992 Bartmann et aI., 1992 Bartmann et aI., 1992 �132 RESULTS AND DISCUSSION GnRH Toxin/Agonist Experiment 1: Evaluation of GnRH-PAP in mule deer. Mean serum LH concentrations were reduced in all experimental animals compared to pretreatment levels following treatment with GnRH-PAP (Fig 1). Mean levels ofLH (ng/ml) were initially reduced 72% of pretreatment levels after 15 days, then increased to about 51% (se ± 0.75) after 33 days and have remained relatively unchanged through 100 days posttreatment. Body weight dynamics, blood chemistry, hematology and social behavior of treated deer appear normal compared to non-treated female deer. 125 100 I - ...J c: II) E 75 C"d II) •... II) •... a. - 50 0 ~ 25 o Pre Toxin 33 15 48 100 Days Figure 1. Luteinizing hormone release in ovariectomized female mule deer treated with GnRH-PAP during the breeding season. Experiment 2: Evaluation of GnRH-Agonist in Rocky Mountain Elk. Administration of GnRH-Agonist implants resulted in a prompt and expected increase in serum LH levels in all female elk. We observed an average peak LH response of 15.52 ng/ml (se ± 0.94) which occurred approximately 3.5 h posttreatment. Peak response (ng/ml) and time to peak (h) were not different (p > 0.01) among treatment levels. Serum LH levels returned to baseline after 12 hours for all elk. Subsequent challenge with an intravenous injection of 1 f.,l GnRHI50 kg BW at 35,75,110, and 130 days posttreatment resulted in a significant (p < 0.02) increase in LH levels of control elk compared to elk treated with GnRH-Agonist implants (Fig. 2). We did not observe a significant (p < 0.001) treatment difference among doses of GnRH-Agonist. Lack of treatment effect was not due to high variation among animals but instead to small differences among treatments responses. Serum LH levels of treated elk remained at baseline (0.34 ng/ml) for the duration of the experiment (130 days). These findings are consistent with other evidence for desensitization of the pituitary gonadotrophs due �133 --f- Control - -is - Low --0-- Mad ...+... High 25 20 E 0; c 15 :r: ...J ~ w 10 a.. 5 o o 35 75 110 130 TIME (DAYS) Figure 2. Effects of subcutaneous administration otGnRH-Agonist on serum LH concentrations in elk. Doses were: control = 0, low = 32.5 mg, medium = 65 mg, and high = 130 mg. to prolonged and sustained stimulation. The results of this study demonstrate that one subcutaneous implant containing at least 32.5 mg of GnRH-Agonist can suppress pituitary-ovarian function for at least 90 days in elk. Simulation Model . Professional culling of adult mule deer at the RMA was implemented on a prescribed basis beginning in November 1994. Prior to 1994, the mule deer population was increasing at a rate of approximately 20 % per year. Since 1994, annual culling has reduced the population growth rate to 4% per year and allowed resource managers to meet deer management objectives for this herd. To meet these objectives has required that resource managers remove approximately 10- 15% of the adult females and 6-10% of the young males from the population each year. These estimates vary each year depending on productivity of the population, changes in sex and age ratios, and deer habitat resources. The mathematical model used to estimate the number of animals that must be culled annually to maintain a target, steady state population appears to provide a reasonably accurate representation of mule deer population dynamics at the RMA.· The efficacy of this model, however, is dependent on intensive monitoring of the population and adapting management actions to a changing environment. LITERATURE CITED Asch, R H, F. J. Rojas, T. R Tice, and A. V. Schally. 1985. Studies of a controlled - release microcapsule formulation of an LH-RH agonist in the rhesus monkey menstrual cycle. International J. Fertility 56:78-81. Baker, D. L. 1997. Regulation of mule deer population growth by fertility control: laboratory, field, and simulation experiments. Pages 81- 87 in Wildlife Research Report, Mammals Research, �134 Federal Aid Projects, Job Progress Report, Project W-153-R-4, SP1, Jl. Colorado Division of Wildlife, Fort Collins, Colorado, USA. Baker, D. L., and N. T. Hobbs. 1996. Regulation of mule deer population growth by fertility control: laboratory, field, and simulation experiments. Pages 113 -148 in Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-4, SPl, J1. Colorado Division of Wildlife, Fort Collins, Colorado, USA. Baker, D. L, M. W. Miller, and T. M. Nett. 1995. Gonadotropin-releasing hormone analog-induced patterns of luteinizing hormone secretion in female wapiti (Cervus elaphus nelson i) during the breeding season, anestrus, and pregnancy. Bio. Reprod. 52: 1193-1197. Becker, S. E., Katz, L. S. 1995. Effects of gonadotropin - releasing hormone agonist on serum LH concentrations in female white-tailed deer. Small Ruminant Research 18: 145-150. Casper, R. F., and S. S. C. Yen. 1979. Induction of luteolysis in the human with a long-acting analog of luteinizing hormone-releasing factor. Science 205 :408-41 O. Clayton, R. N. 1982. GnRH modulation of its own pituitary receptors: evidence ofbiphasic regulation. Endocrinology 111: 152-161. Concannon, P. W., and V. N. Meyers-Wallen, 1991. Current and proposed methods for contraception and termination of pregnancy in dogs and cats. 1. Am. Vet. Med. Assoc. 198:1214-1225. Fraser, H. M. 1983. Effect of treatment for one year with a luteinizing hormone-releasing hormone agonist on ovarian, thyroidal, and adrenal function and menstruation in the stump tailed monkey (Macaca arctoides). Endocrinology 112:245-253. __ , M., 1. Sandow, H. Seidel, and W. von Rechenberg. 1987. An implant of a gonadotropin releasing hormone agonist (burserelin) which suppresses ovarian function in the macaque for 3-5 months. Acta Endocrinological 15: 521-427. Garrot, R. A. 1995. Effective management of free-ranging ungulate populations using contraception. Wildlife Society Bulletin 23:445-452. Goodman, R. L., and F. 1. Karsch. 1980. Pulsatile secretion of luteinizing hormone: differential suppression by ovarian steroids. Endocrinology 107:1286-1290. Herschler, R. C., and B. H. Vickery. 1981. The effects ofLHRH ethylamide on the estrous cycle, weight gain, and feed efficiency in feedlot heifer. Amer. 1. of Vet. Res. 42: 1405-1408. Hobbs, N. T., D. L. Baker, and R. B. Gill. 1999. A general theory describing effects offertility control on populations of ungulates. Journal of Wildlife Management (in press). Hone, 1. 1992. Rate of increase and fertility control. 1. Applied Ecology 29:695-698. Jewell, P. A., and S. Holt. 1981. Problems in management oflocally abundant wild animals. Academic Press. 321pp. Kirkpatrick, 1. F., and 1. W. Turner, Jr. 1985. Chemical fertility control and wildlife management. Bioscience 35:485-491. McNeilly, A. S., and H. M. Fraser. 1987. Effect of gonadotropin-releasing hormone agonist-induced suppression ofLH and FSH on follicle growth and corpus luteum function in the ewe. 1. Endocrinology 115:273-282. Niswender, G. D., L. E. Reichert, Jr., A. R. Midgley, Jr., and A. V. Nalbandov. 1969. Radioimmunoassay for bovine and ovine luteinizing hormone. Endocrinology 84: 1166-1173. ___ . 1973. Influence of the site of conjugation on the specificity of antibodies to progesterone. Steroids 22:413-424. Sandow,1. 1982. Inhibition of pituitary and testicular function by LHRH analogs. Pages 19-39 in Jeffcoate S. L. and Sandlier (eds). Progress towards a male contraceptive. Wiley and Sons, Chichester. Warren, R. 1., L. M. White, W. R. Lance. 1995. Management of urban deer populations with contraceptives: practicality and agency concerns. Wildlife Society Bulletin 23: 441-444. �135 Colorado Division of Wildlife Wildlife Research Report July 1999 JOB PROGRESS REPORT Smteof ~C~o~lo~r~ad~o~ _ ,Project No. W.!.!--1~5~3~-R~-..!..12~ __ Work Package No. __ ~3~0:..:::0..!..1 _ Task No.' -=5:..__ _ Cost Center 3430 Mammals Program Deer Conservation Investigating Factors Contributing to Declining Mule Deer Numbers Period Covered: July 1 1998 - June 30, 1999 Authors: T. M. Pojar and W. F. Andelt '" Personnel.. R Arant, D. L. Baker, B. Banulis, D. Hartmann, G. Bock, D. C. Bowden, M. Caddy, D. Coven, B. Devries, B. Diamond, B. Dreher, 1. Ellenberger, V. Graham, D. Gustine, P. Hayden, G. Hust, C. Krumm, A Larsen, K. Larsen, S. Larsen, D. Masden, D. Matiatos, M W. Miller, C. Oliver. J. Olterman, M. Potter, E. Scott, L. Spicer, D. Steele, C. Wagner,:a. Watkins, M. Wild, M. Zeeman, S. Znamenacek. ABSTRACT Long-term data indicate that fawn:doe ratios have shown a significant and consistent decline over the past 20 years in most deer management units in Colorado as well as in most of the western states, This has resulted in a general decline in mule deer (Odocoileus hemionus) density and diminished recreational potential from this major resource. It is important for the Division of Wildlife to elucidate the factors contributing to the reduced doe:fawn radios and the declining deer populations. Reproduction and neonatal survival are 2 important components of population recruitment. Therefore, 2 studies were initiated during this segment to investigate pregnancy and fetal rates of adult does and the survival of neonatal fawns. Both of these studies were done on the Uncompahgre Plateau and, in addition, a supplemental neonate fawn survival study was done in Middle park. Thirty-seven of 40 (93%) adult does on the Uncompahgre Plateau were detected pregnant with transrectal ultrasound. The proportion of does detected pregnant with ultrasound did not differ (P = 0.817) from the proportion of does pregnant (91.4% of303 does) in previous Colorado studies. The average number of fetuses/doe, 1.70, (n=40) for adult does on the Uncompahgre Plateau was not less (Z = 0.340, P = 0.367) than the fetal rate of 1.74 (n=276) in previous Colorado studies. We therefore conclude that does are getting bred and producing fetuses at normal rates for the species and that this component of , the reproductive cycle is most likely not responsible for declining December fawn:doe ratios and, 'consequently, declining deer populations. The field operation was done during February 5-8, 1999; a draft of the manuscript to be submitted to a peer reviewed publication is in Appendix I. The objective of the fawn survival investigation was to put radios on 60 neonatal fawns on the Uncompahgre and 20 neonatal fawns in Middle Park. By June 30, 1999, we had put out 50 and 14 radios on fawns on the �136 Uncompahgre and in Middle Park, respectively for a total of 64 radioed fawns. First fawns were captured on June 9 in both areas and fawn capture peaked during June 13-24 with 76% of the total captured during this time; 19 fawns (30%) were captured during June 22-24. The first mortality was recorded on June 13 with the peak of mortality (50% of total) during June 28-July 12. Causes of mortality when divided between predation, sickness/starvation, and unknown was 47%, 47%, and 6%, respectively. As of this date (August 18, 1999) 30 of64 fawns have died for a mortality rate of 46.9%, Fawns were monitored every day (except weekends) from capture with the exception of the period of July 5-27 when they were monitored twice a day. After July 27 they were again monitored daily except on weekends. Eleven of the 14 fawns that died of causes other that predation (or unknown causes) have been necropsied thus far; all had in common severe thymic atrophy indicating a compromised immune system and chronic stress. It is difficult to discern if the stressor(s) predisposed them to the variety of other pathogens that appeared to be factors in their deaths. Some of the disease agents identified were: Cryptosporidia, Pasteurella, E. Coli, BVD, and a hemorrhagic disease (Blue Tongue or EHD). It should be noted that mortality attributed to predation is probably liberal because the fawn's sickness (including diarrhea in many cases) and weakened condition probably predisposed them to olfactory or visual location by a predator. Positively identifying scavenging vs. predation was difficult even when they were monitored once or twice a day because dense vegetation precluded tracking for evidence of pursuit and kill sites as can be done with snow cover. �137 MULE DEER PREGNANCY AND FETAL RATES William F. Andelt and Thomas M. Pojar P. N. OBJECTNES 1. Estimate pregnancy and fetal rates of adult mule deer does on the Uncompahgre Plateau during the 1998-1999 reproductive cycle. 2. Publish results in a peer-reviewed scientific journal (Appendix I). SEGMENT OBJECTNES 1. Estimate accuracy of an ultrasound system for detecting' pregnancy and number of fetuses in mule deer does maintained at the Colorado Division of Wildlife's Foothills Research Facility in Northwest Fort Collins and in mule deer does collected on the USFWS Rocky Mountain Arsenal. 2. Estimate pregnancy and fetal rates of adult mule deer in DAU D-19 (Uncompahgre). 3. Compare pregnancy and fetal rates of adult mule deer to historic rates in Colorado. 4. Analyze data and prepare an annual Federal Aid Job Progress report. RESULTS See Appendix I for the Research Program Narrative and a draft of the manuscript to be submitted to a peer reviewed publication. �138 �139 lMPACT OF PREDATION AND VEGETATIVE COVER ON MULE DEER FAWN SURVIVAL Thomas M. Pojar and William F. Andelt P. N. OBJECTIVES 1. Identify agents of neonatal mule deer fawn mortality from birth to 6 months of age. SEGMENT OBJECTIVES 1. Capture and radio collar 30-35 neonatal fawns each on the Uncompahgre Plateau (D-19), Middle Park (D-9), and Red Feathers (D-4). 2. Measure vegetative density and height at fawn bed sites. 3. Measure distance between successive bed sites for individual fawns. 4. Compare fawn survival and causes of mortality among 3 vegetatively and ecologically different study areas. 5. Correlate height and density of vegetation at fawn birth sites and bed sites with percent of fawns killed by predators. 6. Compare height and density of vegetation at fawn bed sites and random sites. INTRODUCTION The Research Program Narrative is in Appendix II. Because of budget redirection to accomplish the estimation of pregnancy and fetal rates via ultrasound the objectives outlined in the PN were modified to fit within budget, personnel, and logistic constraints. Major changes include: 1. Limit the fawn collaring effort to the Uncompahgre and Middle Park rather than attempt to do it on 3 study areas. Because of the size and diversity of vegetative communities on the Uncompahgre and because of intense political interest in this area, the target sampling intensity was increased to 60 fawns. The target sample in Middle park was reduced to 20 fawns due to the redirection of resources. 2. Abandon the idea of measuring vegetation at bed sites because of evidence that persistent disturbance of bed sites reduces survival of young ungulates (Philips 1998). 3. The logistics of placing an adequate crew in remote fawning areas and the unknown of whether or not fawns could be captured in sufficient numbers given deer density, terrain, and vegetative cover encountered negated many of the objectives in the PN. In essence, this investigation was reduced to placing emphasis on determining if fawns can be captured under these conditions and, if so, monitor their survival and causes of mortality. Summer fawn survival and causes of mortality are 2 important factors impacting deer population performance. STUDY AREAS There are 2 study areas involved in this investigation, the Uncompahgre Plateau and Middle Park. The Uncompahgre Plateau includes Game Management Units 61 and 62. It is a high elevation plateau, oriented northwest-southeast, formed by a structural uplift that is characterized by steep canyons in both directions from the divide ridge that runs the length of the plateau. The soils are primarily shallow with areas of rich deeper soils that support aspen (Populus tremuloides) communities at intermediate to higher elevations. High elevations have extensive stands of spruce-fir �140 (Picea-Abies) and fir-aspen (Abies-Populus tremuloides). The crest of the plateau peaks at about 9,500 ft (2,898 m) and declines in both directions through oakbrush (Quercus gambellii), pinyonjuniper (Pinus edulis-Juniperus osteosperma), and large sagebrush (Artemisia tridentata) parks as the elevation decreases. A thorough description of the physiography, geography, climate, and vegetation can be found in Anderson et. aI (1992) and Kufeld (1979). Middle Park is a high mountain park with similar plant communities as the Uncompahgre Plateau at similar elevations. A description of the area can be found in Tiedeman et. aI (1987). MElHODS Radio collared does were used to locate fawning areas although no special effort was made to capture the fawns of radioed does. The strategy for capturing fawns involved observing does for behavioral and physical signs, such as udder development, of having fawned. If it was determined that these indicators suggested that the doe had fawned, the area was searched for the fawn(s). Once located, the fawn was approached from behind (out of direct eyesight) and restrained for weighing, measuring, and putting on the radio collar. ·The collars used were expandable with elastic and loops sewed with cotton thread to accommodate growth and to drop off at about 6 months; they weighed about 65 g. RESULTS A total of 64 fawns were captured and radio collared; 50 on the Uncomphagre Plateau and 14 in Middle Park. The first fawns were captured on June 9 with 76% of the total captures made during June 13-24; the peak 3-day period of capture was June 22-24 when 30% of the captures were made (Figure 1). Total fawn mortality as of this writing (August 18) is 30 with 14 from a condition that is generally described as sickness/starvation (S/S), 14 from predation, and 2 from unknown causes where only the collar was found (Table 1). The 14 that died from SIS were all collected, frozen or stored on ice, and later necropsied at the Colorado State University Veterinary Teaching Hospital in Fort Collins. Results thus far are preliminary with some of the lab tests still pending and that is the reason for categorizing these deaths as general SIS. Some disease agents have been identified but it is not known if these were the cause of the fawns deaths. A common condition in all of the fawns is that they had atrophied thymus glands indicating chronic stress and probable immunodeficiency which could have predisposed them to succumbing to the pathogens that they encountered. Some of the pathogens 25 20 (/) c: i& u, -- 15 .c 10 0 •... Cl> E ::::J Z 5 6/1-6/3 6rT-6/9 6/13-6/15 6/19-6/21 6125-6/27 6/4--616 6/10-6/12 6/16-6/18 6/22-6124 6/28-6/30 Date Figure 1. Dates of fawn capture combined Uncompahgre Plateau and Middle Park. from 2 areas in Colorado. 1999- �141 identified are: the parasite Crhyptosporidia, Pasterrella, E. Coli, Bovine Virus Diarrhea (BVD), and a hemorrhagic disease (Blue Tongue or Epizootic Hemorrhagic Disease (EHD». In addition to the 30 dead radioed fawns, we found 2 stillborns, 2 other intact fawn carcasses, and one adult doe carcass. One of the stillborns was fresh enough to necropsy and was small {1.5 kg) and also had an atrophied thymus. The lab results on the adult doe are not complete but she showed some signs of a hemorrhagic condition; the other carcases were in stages of decomposition that precluded materials collection. Determining if a fawn died from some other cause or was killed by a predator is difficult. Because the fawns were monitored on a daily or twice daily schedule, all carcasses that were found partially eaten were attributed to predation; some errors were undoubtedly made by doing this which would inflate the deaths attributed to predation. Vegetation was too thick to discern tracks of a chase and capture as is possible during winter with snow cover, so unless the predation was actually witnessed, there will always be doubt as to the whether or not predation was the cause of death. In 2 instances, only the collar was found with no other tracks or supporting evidence of predation. These are categorized as "unknown" but are likely predator kills. It is unlikely that the collars were slipped because they fit snugly around the neck. One of these collars had a triangular rip that could have been caused by a predator tooth (or barbed wire), the other was without marks or blood so it's categorization is even less certain. With nearly half of the fawns dying of SIS it is probable that some portion of the fawns eaten by a predator were in some stage of "sic knessl starvation" resulting in them being more susceptible to discovery by a predator ~ Several of the fawns that died had diarrhea which would have broken their .shield of scentlessness that protects them from predators early in life. Other fawns that were in a diseased or weaken state may have lacked normal hiding or escape behavior that would have predisposed them to either visual or olfactory location by a predator in the vicinity. The classical evidence that an animal was alive when bitten (i.e. predated) by a predator is subcutaneous hemorrhage; this, however, does not establish that the prey was in a healthy state when preyed upon. The peak of mortalities from all causes was during June 28-July 11 with 16 of30 (53%) dying during this 14-day period (Figure 2); 5 from SIS and 11 from predation. One fawn died on the 2nd day after capture and was judged to be a day old at capture. It was found to have no milk in it's abomasum and had a small amount of forage in it's rumen both of which indicate that it had not nursed and was starving. It is not possible to determine if this fawn was abandoned because of handling or if the dam was not providing milk; a doe was in the area (and acting maternal) during capture and 2 days later when the fawn carcass was recovered. If the fawn was truly a day old when captured and had not nursed by that time it is possible that there was a bonding or lactation problem on the part of the doe. 5 CI) Q) E (ij 4 t 0 ~ c: 3 3: -.8 <IS u.. 2 0 E 1 ::s z 0 .•.... N 0 .•... CD 00 .•.... fb .•.... C{i CD ~ g ch C:! <0 ~ ,..._ N .•... 0 .•... 00 .•.... fb .•... -.. ,..._ j:::: Dates ..,. . ~ ~ ,..._ g, 00 ~ It) £2 00 .•... .•.... , ~ 00 Figure 2. Neonatal fawn mortality from all causes for the Uncompahgre and Middle Park, Colorado, 1999. ,..._ .•... &b .•.... -.. 00 Plateau �142 All of the other SIS occurred on the 3rd, or later, day after capture. Six fawns died between the 3ni and 5th day after capture and the rest of them (77%) died after the 6th day of capture (Figure 3). It is interesting to note that there were 2 somewhat long time spans where there were no mortalities from any cause - 6 days, July 14-19, and 17 days, July 26- August 13. It seems that if there were constant predator pressure on neonatal fawns as a food source and that sickness were not a factor in predator location of fawns, then there would not be these long spans of no mortality on the marked fawns. However, it is possible that just by chance alone, predators did not encounter a marked fawn during these time periods. One radio was verified to have failed and It is believed that 2 other radios ceased to work because extensive ground and aerial searches failed to make signal contact. A set of twins that were radioed recently disappeared after domestic sheep were herded into the area they were inhabiting; no signal was obtained with a ground search of the area so an aerial search will be made. 12,------------- -, 10 CI.I ~ tIS u.. '0 8 6 ~ g :z: 4 2 <0 r= 1 Days From Capture Figure 3. Days from capture that fawns died from any cause. Total for fawns captured on the Uncompahgre Plateau and Middle Par1<.Colorado. 1999. CONCLUSIONS The following conclusions can be made from this investigation. 1. It is possible to capture neonatal fawns under the conditions found on the Uncompahgre Plateau and in Middle Park. Success in capturing fawns is dependent mostly on doe density which dictates the number of does that the capture crew can encounter during a search. Road network is also important because of the greater mobility of the capture crew and, therefore, a greater probability of encountering does .. And finally, experience of the capture crew is a major factor in the success of finding fawns. Dense vegetative cover and roadless areas make sighting of does and capture of fawns unproductive for reasonable sample sizes. 2. There is evidence that there are stress factors impacting the 2 mule deer populations that were sampled. Atrophied thymus glands of all the intact carcasses recovered along with a variety of pathogenic agents identified suggests that the stressor(s) resulted in a suppressed immune response in the fawns making them susceptible to encountered pathogens. It has been demonstrated that thymus gland size in mule deer is seasonally cyclic with the peak during summer (good nutrition) and the trough during winter (limited resources) (Anderson et aI. 1974). Ozoga and Verme (1978:794) have demonstrated the relation between thymus size and dietary plane in white-tailed deer (0. Virginianus) fawns in controlled tests. They also noted that the thymus glands offawns dying of disease or malnutrition within a month of birth were "extremely small" compared to fawns dying of other causes (accidents). �143 LITERATORE CITED Anderson, A E., D. C. Bowden, and D. M. Kattner. 1992. The puma on Uncompahgre Plateau, Colorado. Technical Publication No. 40, Colorado Division of Wildlife. Anderson, A E., D. E. Medin, and D. C. Bowden. 1974. Growth and morphometry of the carcass, selected bones, organs, and glands of mule deer. Wildlife Monographs, No. 39. Kufeld, R C. 1979. History and current status of the mule deer population of the east side of the Uncompahgre Plateau. Division Report No. 11, Colorado Division of Wildlife. Ozoga, J. J. and L. J. Verme. 1978. The thymus gland as a nutritional status indicator in deer. Journal of Wildlife Management, 42:791-798. Philips, G. E. 1998. Effects of human-induced disturbance during calving season on reproductive success of elk in the upper Eagle River valley. Ph. D. Dissertation, Colorado State University, Fort Collins. .Tiedeman, J. A, R E. Francis, C. Terwilliger, Jr., and L. H. Carpenter. 1987. Shrub-steppe habitat types of Middle Park, Colorado. USDA Forest Service Research Paper RM-273, Rocky Mountain Forest and Range Experiment Station, Fort Collins. Table 1. Neonatal fawns captured and radio collared on the Uncompahgre Plateau and Middle Park, Colorado, 1999. Radio frequencies in the 149-150 KHz band are Uncompahgre fawns and 173 KHz radios are Middle Park fawns. ID Date Sex Weight (kg) Hindfoot Date Probable 149.410/99 06/1611999 M 3.5 24.4 07/0111999 Predation 150.010/99 0611511999 F 0.0 24.9 150.020/99 06/0911999 F 0.0 24.7 150.030/99 06114/1999 F 0.0 27.3 06/2911999 Sick/Starve 150.040/99 06114/1999 F 0.0 27.6 07/0611999 Sick/Starve 150.050/99 06/1511999 F 0.0 27.9 150.070/99 0611711999 M 3.8 26.0 08/1811999 Predation 150.080/99 06/2211999 F 4.4 27.0 06118/1999 Sick/Starve 06/29/1999 Predation 150.090/99 06/2111999 F 5.9 27.6 150.100/99 06/2111999 M 5.2 26.4 150.110/99 06114/1999 U 2.3 0.0 150.120/99 06/1711999 F 5.0 26.0 150.130/99 0611411999 F 2.5 24.8 150.140/99 06110/1999 F 3.3 24.4 150.150/99 06/1711999 M 5.3 26.7 07/2011999 Predation 150.160/99 06/2211999 F 0.0 26.0 06/3011999 Predation 150.170/99 0611511999 F 3.7 26.0 150.180/99 06/2111999 F 6.1 29.2 07/2511999 Sick/Starve 07/0711999 Sick/Starve 150.190/99 0611511999 M 4.0 26.0 150.200/99 06/2211999 M 3.3 24.1 150.210/99 06/2411999 F 4.4 26.4 150.220/99 06/1611999 M 3.8 24.4 150.230/99 06/2211999 F 5.3 28.6 150.270/99 06/23/1999 M 5.6 26.4 �144 Weight (kg) Hindfoot Date Probable M 3.1 25.0 06118/1999 Sick/Starve M 4.6 26.0 0611611999 M 0.0 26.7 150.310/99 06/2811999 F 5.6 28.6 150.320/99 06123/1999 M 6.0 27.9 150.340/99 06/2311999 F 5.6 27.6 06/30/1999 Sick/Starve 150.350/99 0611711999 M 0.0 28.7 06/22/1999 Sick/Starve 150.360/99 06/2411999 M 6.0 26.7 150.370/99 06/2311999 M 6.0 27.3 150.380/99 0612411999 M 2.4 23.2 07/1111999 Predation 150.390/99 06/2411999 F 3.8 26.7 07/0111999 Predation 150.400/99 06/2211999 M 5.0 27.3 150.420/99 06128/1999 F 4.2 26.2 150.430/99 06124/1999 M 5.3 26.7 0711111999 Unknown 0811611999 Predation ID Date Sex 150.280al99 06/15/1999 150.280b/99 06/28/1999 150.290/99 150.440/99 06/2511999 M 6.0 29.2 150.45Q/99 06115/1999 F 3.5 25.7 150.460/99 06124/1999 M 3.1 23.5 150.480/99 06/3011999 F 4.2 24.8 08/16/1999 Sick/Starve 150.510/99 0611611999 F 3.6 24.1 07/0111999 Predation 150.520/99 06116/1999 F 3.5 24.8 07/0811999 Predation 150.530/99 06114/1999 M 3.6 25.4 150.540/99 06/29/1999 M 4.5 26.0 150.570199 06/24/1999 F 2.5 21.9 0711311999 Sick/Starve 150.600/99 06/2911999 M 6.0 28.9 150.610/99 06/18/1999 M 3.6 25.4 07/23/1999 Sick/Starve 150.620/99 06/30/1999 M 4.5 24.4 07/0511999 Predation 173.510/99 06/0911999 M 4.5 26.7 0611311999 Sick/Starve 173.540/99 0611111999 F 3.8 26.0 07/2211999 Predation 173.550/99 0612411999 F 4.7 27.3 173.560/99 06/2111999 F 4.1 26.0 173.590/99 06/2211999 M 5.5 28.6 173.600/99 06/2111999 F 4.1 26.0 173.610/99 06/2111999 M 5.1 28.6 06/25/1999 Sick/Starve 07/0811999 Predation -. 173.630/99 06/2111999 F 3.8 25.4 173.640al99 06/22/1999 M 4.5 26.7 06/2511999 Sick/Starve 173.640b/99 06/30/1999 M 4.2 25.4 08/16/1999 Predation 173.660/99 06/29/1999 M 6.5 27.9 173.670/99 06/2511999 F 6.0 29.2 173.690/99 06/19/1999 U 0.0 0.0 07/0511999 Unknown 173.700/99 06/1811999 M 4.7 27.0 'Due to the preliminary nature of this report, the causes of death have been put into only 3 categories, sick/starve, predation, and unknown. The unknown category includes 2 radios that were found that had no other evidence present. It is suspected that these were actually predator kills because the collars fit snug enough that slipping the collar was unlikely, in addition, one of the collars had a small triangular rip in it that could be consistent with a predator tooth tear. �145 APPENDIX I Research Program Narrative and Draft Manuscript MULE DEER PREGNANCY AND FETAL RATES ON THE UNCOMPAHGRE PLATEAU January 12, 1999 Principal Investigators: WILLIAM F. ANDELT Department of Fishery and Wildlife Biology Colorado State University Fort Collins, CO 80523 970-491-7093 mOMAS M. POJAR Colorado Division of Wildlife 317 W. Prospect Fort Collins, CO 80526 970-472-4308 �146 �147 PROGRAM NARRATIVE State of. ___;C::::;o::::..:l~or:..::a:::::d~o_ Project No .. W..!.!._-~15~3::....-.:!:.::R,__ _ Work Package No. ---,3~0~0~1,-_ Task No. -=5'--_ Cost Center 3430 Mammals Program Deer Conservation Mule Deer Pregnancy and Fetal Rates on the Uncompahgre Plateau NEED Fawn:doe (f:d) ratios obtained in December on the Uncompahgre Plateau (Game Management Units 61 and 62, Data Analysis Unit (DAU) D-19) have declined (t = -3.39, P = 0.004) by an average of 1.8 fawns: 100 does per year from 1982-1998. December ratios ranged from a high of79f:l00d in 1982 to a low of 32f: 1OOdin 1996 and 34f: 100d in 1997; there were 51.9f: 1OOdin 1998. Besides low December f:d ratios, over-winter (December - May) fawn survival on the Uncompahgre Plateau was 49% during the winter of 1997-1998 (Bartmann and Pojar 1998a). The low December f:d ratios and poor over-winter survival of fawns for the Uncompahgre deer herd is of concern to the public and game managers. It is unknown if the low December ratios are due to a failure to breed, reduced fetal production, resorption/abortion offetuses, or low summer and fall fawn survival (Bartmann 1998). Pregnancy and Fetal Rates Pregnancy rates are key information to determine if does are breeding. Recent data show that the pregnancy rate during January 1998 of93.0% for 29 does ~ 1 year old in the Red Feather DAU (Bartmann and Pojar 1998b) was similar to 92.0% of 163 does ~ 2 years old in the same area during 1961-1964 (Medin and Anderson 1979). A 94.8% pregnancy rate for does ~ 2 years old (n = 114) and a 94.0% pregnancy rate for does> 1 year old (n = 134) in Middle Park (DAU D-9) was reported by Gill (1971) during 1969-197l. Other pregnancy rates and locations reported in the literature are as follows: 100% for does ~ 2 years old (n = 18) on the Forbes-Trinchera Ranch, Colorado (Freddy 1988), 89% (n = 47) during 1973 and 82% (n = 83) during 1978 in the Piceance Basin, Colorado (Bartmann 1998), and 94% of mule deer does> 1 year old in a California study (Salwasser et al. 1978). The 1998 data collected in Colorado appears to be normal and does not indicate that pregnancy rate is a contributing factor to the low f:d ratios, however these data were not collected in the Uncompahgre DAU where December f:d ratios were lowest. Fetal rates, the number of fetuses per pregnant doe, that are in a normal range for the species would indicate that nutritional and other needs of breeding does are being met. The fetal rate for does > 2 years old was 1.83 (n = 41) in the Cache la Poudre River drainage from 1961-1965 (Medin and Anderson 1979), 1.82 (n = 114) in Middle Park (Gill 1971), 1.89 (n = 18) on the Forbes-Trinchera Ranch (Freddy 1988), and l.46 (n = 61) and 1.65 (n = 37) in the Piceance Basin (G. C. White, Colorado State University, personal communication). Salwasser et al. (1978) reported a fetal rate of l.62 for 2 and 3 year old mule deer does (n = 47) and a rate of l.85 for does> 4 years old (n = 34) in California To help ascertain the cause of low fawn:doe ratios on the Uncompahgre Plateau of Colorado, an investigation to estimate: a) the proportion of ~ 2 year old does that become pregnant and b) the number of fetuses produced is needed. Estimates of pregnancy and fetal rates will indicate if these critical reproductive parameters are within the normal range for the species and will contribute to the understanding of the causes of the observed low f.d ratios and low recruitment into the Uncompahgre mule deer population. �148 OBJECTIVES We propose to estimate pregnancy and fetal rates of adult mule deer does on the Uncompahgre Plateau during the 1998-1999 reproductive cycle. HYPOTHESES H"l: H,,2: Pregnancy rates of adult mule deer does on the Uncompahgre Plateau during the 1998-1999 reproductive cycle do not differ from historic pregnancy rates (93%) in Colorado. Fetal rates of adult mule deer does on the Uncompahgre Plateau during the 1998-1999 reproductive cycle do not differ from historic fetal rates (1.74 fawns per adult doe) in Colorado. EXPECTED RESULTS OR BENEFITS This research will help ascertain if pregnancy and fetal rates are outside the normal range expected for mule deer in Colorado and if they are contributing factors to the observed low December fawn:doe ratios. Knowledge of fetal rates is essential for determining what factors might be causing the low fawn:doe ratios observed in Colorado. Recruitment into a mule deer population is dependent on adequate pregnancy and fetal rates, and young survival through breeding age. This investigation will provide information on the early stages of the recruitment process, i.e. whether or not does are becoming pregnant and producing normal litters. These data are imperative before research and management experiments are conducted to estimate the effect of various experimental manipulations on fawn:doe ratios. APPROACH Capture and Handling of Does Forty mature does C:::. 2 years old) will be captured using a helicopter and netgun on the Uncompahgre Plateau during February 1999. The deer usually will be pursued for < 2 minutes. We expect < 2% of the captured does to sustain injuries or mortalities (R. M. Bartmann, Colorado Division of Wildlife and A. W. Alldredge, Colorado State University, personal communications). Captured does will be hobbled, blindfolded, and transported via helicopterto a base station, usually < 5 miles from the capture site to.minimize stress, where they will be processed. Does captured < 2 miles from the processing station will be released at the station, whereas does captured> 2 miles from the station will be released at the capture site. We anticipate that does will not be held for more than 30 minutes. Pregnancy and Fetal Detection The does will be manually restrained without the use of a tranquilizer to allow quick release at the processing site, unless we find sedation is necessary. If necessary, does will be sedated with ketamine and xylazine and reversed with yohimbine. The does will be placed in lateral or sternal recumbency, depending on location of the fetuses. We will use trans rectal ultrasonography, pioneered by Lindahl (1971) and applied in field situations by Barrett (1981), Smith and Lindzey (1982). White et al. (1989), and Stephenson et al. (1995), to estimate pregnancy and fetal rate. A portable ultrasound unit available from the Colorado State University Veterinary Teaching Hospital will be used; the �149 equipment will be transported in a mobile processing station to within 5 miles of capture sites to minimize transport time and stress on the does. Blood will be collected by carotid venipuncture and serum pregnancy-specific protein-B (PSPB) (Sasser et al, 1986) will be used to provide a second estimate of pregnancy rates (Stephenson et al. 1995). The use of ultrasonography to detect pregnancy is a well tested method and provides accurate estimates of pregnancy rates in elk (Cervus elephus) (Bingham et al. 1990, Revol and Wilson 1991, Willard et al, 1994). Transrectal ultrasonography was found to accurately detect pregnancy in domestic sheep (Schrick and Inskeep 1993) and fallow deer (Dama dama; Lenz et al. 1993). Because the method is not as well tested to detect the number of fetuses carried by pregnant female mule deer, we will have controls in our study to document the accuracy of the method and make adjustment in the observed fetal rate if necessary. Ten potentially pregnant does that are held in captivity in the Colorado Division of Wildlife Big Game Research Facility on the Foothills Campus, Fort Collins will be untrasounded for pregnancy and fetal rates as a control on the methodology and to test the proposed handling and restraint methods to be used on does during the field operation. These does will be under close surveillance throughout pregnancy and during fawning to verify the ultrasound results. In addition, we will have access to ~ 10 deer to be collected in January or February on the Rocky Mountain Arsenal. These does will be ultrasounded and then necropsied for immediate verification of the effectiveness of detecting the number of fetuses. Ninety-five percent of Colorado mule deer conception dates, based on back-dating from fetal forehead-rump measurements, over a 3 year span (n=135) were reported to be between November 18 and December 12 (Gill 1972). Ultrasounding for fetal rate is most successful on small ungulates during 30-60 days into pregnancy (LaRue Johnson, CSU Veterinary Teaching Hospital, personal communication). We will target the time period ofJanuary 20 through February 10 to do the. ultrasounding. Dr. LaRue Johnson will perform the ultrasounding procedures. Animal Care and Use All capture and handling of animals will comply with the standards of the Colorado State University and Colorado Division of Wildlife Animal Care and Use committees. We will secure trespass permission from study site owners prior to the study. Sample Sizes and Power of Tests The following power analysis (Table 1) was developed to estimate sample sizes necessary to detect various mean departures of fetal rates from historic levels for adult does. For the historic data, we used the weighted mean of fetal rates for adult does from the Poudre River drainage (n = 41, = 1.83, SD = 0.44; Medin and Anderson 1979), Middle park (n = 119, x = 1.86, SD = 0.57; Gill 1971), the Forbes-Trinchera Ranch (n = 18, x = 1.89, SD = 0.47; Freddy 1988), and the Piceance Basin ([1973: n = 61, x= 1.46, SD = 0.77][1978: n = 37, x = 1.65, SD = 0.72]; G. C. White, Colorado State University, personal communication) to estimate the historic mean (1.74) and standard deviation (0.64). We used an expected 1999 standard deviation equal to the historic standard deviation (0.64; G. C. White, personal communication). An alpha of 0.05 was used in our estimates of power. Power of 0.8 or higher is desired. x �150 Table 1. One-tailed power to estimate if 1999 fetal rates differ from historic rate of 1.74 fawns per mature doe (;:: 2 years old). Sample size (n) Fetal rate Power Sample size (n) Fetal rate Power 20 20 20 20 30 30 30 30 40 40 40 40 1.3 1.4 1.5 1.6 1.3 1.4 1.5 1.6 1.3 1.4 1.5 1.6 0.91 0.74 0.49 0.24 0.97 0.87 0.62 0.31 0.99 0.93 0.72 0.36 50 50 50 50 60 60 60 60 80 80 80 80 1.3 1.4 1.5 1.6 1.3 1.4 1.5 1.6 1.3 1.4 1.5 1.6 1.00 0.97 0.79 0.41 1.00 0.98 0.84 0.46 1.00 0.99 0.91 0.53 Regarding sample sizes, the desired differential in fetal rate that we wish to detect is somewhat problematic. We are primarily interested in ascertaining if pregnancy rates and fetal rates are a major contributor to the observed low December fawn:doe ratio (0.36 fawns/doe predicted by the regression equation in 1998) on the Uncompahgre Plateau. Considering historic fetal rates of L 74 fetuses/doe, there apparently is a major failure to breed, inadequate fetal production, a major loss of fetuses, high neonatal fawn mortality, or a combination of these factors which result in only 0.36 fawns/doe (predicted by regression) remaining in December 1998. On average, fawn:doe ratios in Colorado have declined by about 0.015 fawns/doe/year during the last 20 years which results in a decline of about 0.3 fawns/doe over the 20 years. Fawn:doe ratios on the Uncompahgre Plateau have declined by 0.018 fawns/doe/year during the last 17 years (1982-1998), resulting in a decline of 0.30 fawns/doe over the 17 years. Thus, to determine if fetal rates contribute to the observed changes, we will attempt to detect a change of 0.30 fawns/doe. A change of 0.30 fawns/doe represents 21 % of the 1.4 (1.74 - 0.36 [predicted by the regression equation for 1998] = 1.4) reduction in the fawn:doe ratio from fetal counts to December age-ratio counts. Thus, we believe that detecting 21% (±) of the overall loss of fetuses/fawns from the fetal stage of reproduction to December will provide strong evidence that fetal production is or is not a major contributing factor to low fawn:doe ratios. To detect a change of 0.30 fawns/doe, about 40 does will be necessary for our evaluation. Data Analyses Fetal rates obtained during 1999 on the Uncompahgre Plateau will be compared to historic rates with a Z test. Significance level to reject the null hypothesis will be at P < 0.05. LOCATION This study will be conducted on the Uncompahgre 62 (D-19). Plateau, Big Game Management Units 61 and �151 WORK SCHEDULE Jan-Feb 1999 Mar-Jun 1999 Capture and ultrasound does Data analysis and write report PERSONNEL Thomas M. Pojar F. Andelt LaRue W. Johnson David C. Bowden Kenneth P. Burnham Co-Principal Investigator, Colorado Division of WildlifeWilliam Co-Principal Investigator, Department of Fishery and Wildlife Biology, Colorado State University Co-Principal Investigator, College of Veterinary Medicine and Biomedical Sciences, Colorado State University Statistical Consultant, Department of Statistics, Colorado State University Colorado Cooperative Fish and Wildlife Research Unit, Department of Fishery and Wildlife Biology, Colorado State University ESTIMATED COST Item Helicopter capture Veterinary consulting Ultrasound rental Travel expenses Vehicle Mileage TOTAL 10% contingency GRAND TOTAL Number 40 does 1 1 15 days 1 month 2,000 miles CostlItem $ 400 $1,000 $ 600 $ 95 $ 400 $ 0.15 Total $16,000 $ 1,000 $ 600 $ 1,425 $ 400 $ 300 $19,725 1,972 $21,697 s LITERA TURE CITED Barrett, R H. 1981. Pregnancy diagnosis with doppler ultrasonic fetal pulse detectors. Wildlife Society Bulletin 9:60-62. Bartmann, R M. 1998. A prospectus on mule deer in Colorado. Colorado Division of Wildlife, Fort Collins, Colorado, Unpublished report. Bartmann, R M., and T. M. Pojar 1998a Experimental deer inventory. Colorado Division of Wildlife, FederalAid in Wildlife Restoration Job Progress Report, Project W-153-R-ll, July. _, and _. 1998b. Deer reproduction assessment. Colorado Division of Wildlife, Federal Aid in Wildlife Restoration Job Progress Report, Project W-153-R-11, July. Bingham, C. M., P. R. Wilson, and A S. Davies. 1990. Real-time ultrasonography for pregnancy diagnosis and estimation of fetal age in farmed red deer. Veterinary Record 126:102-106. Freddy, D. 1. 1988. Effect of elk harvest systems on elk breeding biology. Colorado Division of Wildlife, Federal Aid in Wildlife Restoration Job Progress Report, Project W-153-R-2, July. Gill, R B. 1971. Middle Park deer study - productivity and mortality. Colorado Division of Wildlife, Federal Aid in Wildlife Restoration Job Progress Report, Project W-38-R-25. July:189-207. _. 1972. Productivity studies of mule deer in Middle Park, Colorado: Second Annual Mule Deer Workshop, Elko, Nevada, January 12. Xerox. Lenz, M. F., A W. English, and A Dradjat. 1993. Real-time ultrasonography for pregnancy diagnosis and foetal ageing in fallow deer. Australian Veterinary Journal 70:373-375. �152 Lindahl, I. L. 1971. Pregnancy diagnosis in the ewe by intrarectal doppler. Journal of Animal Science 32:922-925. Medin, D. E., and A. E. Anderson. 1979. Modeling the dynamics of a Colorado mule deer population. Wildlife Monographs 68:1-77. Revel, B., and P. R Wilson. 1991. Ultrasonography of the reproductive tract and early pregnancy in red .deer. Veterinary Record 128:229-233. Salwasser, H., S. A. Holl, G. A. Ashcraft. 1978. Fawn production and survival in the North Kings River deer herd. California Fish and Game 64:38-52. Sasser, R G., C. A. Ruder, and K. A. Ivani. 1986. Pregnancy detection in farm animals by radioimmunoassay of a pregnancy-specific protein in serum. in 1. Hau, ed., Pregnancy proteins in animals. Walter de Gruyer & Co. Berlin. Schrick, F. N., and E. K. Inskeep. 1993. Determination of early pregnancy in ewes utilizing transrectal ultrasonography. Theriogenology 40:295-306. Smith, R B., and F. G. Lindzey. 1982. Use of ultrasound for detecting pregnancy in mule deer. Journal of Wildlife Management 46:1089-1092. Stephenson, T. R, 1. W. Testa. G. P. Adams, R G. Sasser, C. C. Schwartz, and K. 1. Hundertmark. 1995. Diagnosis of pregnancy and twinning in moose by ultrasonography and serum assay. Alces 31:167-172. Welker, H. 1. 1986. Fawn mortality in the Lake Hollow deer herd, Tehama County, California California Fish and Game 72:99-102. White, I. R, W. A. C. KcKelvy, S. Busby, A. Sneddon, and W. 1. Hamilton. 1989. Diagnosis of pregnancy and prediction of fetal age in red deer by real-time untrasonic scanning. The Veterinary Record 124:395-397. Willard, S. T., R G. Sasser, J. C. Gillespie, J. T. Jaques, T. H. Welsh, Jr., and RD. Randel. 1994. Methods for pregnancy determination and the effects of body condition on pregnancy status in Rocky Mountain elk (Cervus elephus nelsonii. Theriogenology 42:1095-1102. �153 DRAFT MANUSCRIPT ESTIMATION OF MULE DEER PREGNANCY AND FETAL RATES ON THE UNCOMPAHGRE PLATEAU, COLORADO USING TRANSRECTAL ULTRASOUND William F. Andelt, Thomas M. Pojar, and LaRue W. Johnson Abstract Mule deer (Odocoileus hemionus) populations apparently have declined in several western states during the 1990's. In Colorado, mule deer fawn:doe ratios in December have declined by 0.15 fawns/doe/year from 1972 through 1995. Significant declines in the fawn:doe ratio have especially occurred on the Uncompahgre Plateau near Montrose, Colorado. Thus, we estimated adult mule deer pregnancy and fetal rates on the Uncompahgre Plateau during the 1998-1999 reproductive cycle to determine if lower pregnancy or fetal production was the cause of the deer decline. Thirty-seven of 40 (93%) adult does on the Uncompahgre Plateau were detected pregnant with ultrasound, and 36 of the 40 does were detected pregnant with pregnancy-specific protein-B (PSPB). The proportion of does detected pregnant with ultrasound does not differ (Xl) = 0.053, P = 0.817) from the proportion of does pregnant (91.4% of303 does) in previous studies in Colorado. The average number of fetuses/doe for all adult does (pregnant and non-pregnant) on the Uncompahgre Plateau (n = 40, '! = 1.70, SE = 0.109) was not less (Z = 0.340, P = 0.367) than in previous studies (n = 276, '! = 1.74, SE = 0.038). The average number of fetuses/doe for pregnant adult does on the Uncompahgre Plateau (n = 37, '!= 1.84, SE = 0.082) also was not less (Z = 0.260, P = 0.397) than in previous studies (n=258, '! = 1.86, SE = 0.028). INTRODUCTION Mule deer (Odocoileus hemionus) populations apparently have declined in several western states of the United States during the 1990's (Unsworth et al. 1999). In Colorado, mule deer fawn:doe ratios in December have declined by 0.15 fawns/doe/year from 1972 through 1995 (G. C. White, Colorado State University, unpublished data). In particular, fawn:doe ratios on the Uncompahgre Plateau of Colorado have declined (I = -3.39, P = 0.004) by an average of 0.18 fawns/doe/year from 1982-1998, and have ranged from a high of 79 fawns/100 does in 1982 to a low of 32 fawns/l00 does in 1996 and 34 fawns/100 does in 1997 (Colorado Division of Wildlife, unpublished data). In addition to low December fawn:doe ratios, over-winter (December - May) fawn survival on the Uncompahgre Plateau was 49% during the winter of 1997-1998 (Bartmann and Pojar 1998a). The low December fawn:doe ratios are a concern to the public and game managers. Pregnancy rates of collected adult (2:2years old) female mule deer have ranged from 97 to 100% during the 1960's, 1970's, and 1980's in Colorado (Gill 1971, 1972; Medin and Anderson 1979; Freddy 1987, 1988). The average numbers offetuses per adult doe (fetal rate) have ranged from an average of 1.83 to 1.91 (Medin and Anderson 1979; Gill 1971, 1972; Freddy 1987, 1988) during the same period in various areas of Colorado. Several hypotheses exist to explain the decline in December mule deer fawn:doe ratios including a failure to breed, reduced fetal production, resorption/abortion of fetuses, or low summer and fall fawn survival (R. M. Bartmann, Colorado Division of Wildlife, personal communication) mediated by low �154 buck doe ratios and a protracted fawning period, poor habitat quality and reduced nutrition or hiding cover, more predators and increased predation, competition with elk (Cervus elaphus), disease, poisonous plants; drought, or perhaps a combination of these factors. Our objectives in this study were to determine adult mule deer pregnancy and fetal rates on the Uncompahgre Plateau, Colorado during the 1998-1999 reproductive cycle. We hypothesized that pregnancy and fetal rates did not differ from rates obtained during the 1960's, 1970's, and 1980's in Colorado. Transrectal ultrasonography has been used to accurately detect pregnancy in domestic sheep (Schrick and Inskeep 1993), fallow deer (Dama dama; Lenz et aI. 1993), red deer (Cervus elaphus; Bingham et aI. 1990, Wilson and Bingham 1990, Revol and Wilson 1991), and elk (Willard et aI. 1994). Pregnancy-specific protein-B has been used to reliably detect pregnancy in mule deer and white-tailed deer (Odocoi/eus virginianus; Wood et aI. 1986), muskoxen (Ovibos moschatus; Rowell et aI. 1989), fallow deer (Wilker et aI. 1993), moose (Alces alces; Haigh et aI. 1993, Stephenson et aI. 1995), and elk (Willard et aI. 1994, Noyes et aI. 1997). Because neither technique has been used extensively in mule deer, with the exception of Smith and Lindzey (1982) and Wood et aI. (1986), we also verified the accuracy of both techniques. STIJDY AREAS To assess accuracy in determining pregnancy and number of fetuses per mule deer doe, we investigated captive does at the Colorado Division of Wildlife's Foothills Research Facility in Northwest Fort Collins,and wild does that were culled on the USFWS Rocky Mountain Arsenal, Commerce City, Colorado. In our main study, we ultrasounded does on the Uncompahgre Plateau (Game Management Units 61 and 62, Data Analysis Unit (DAU) D-19) in southwestern Colorado. The plateau is about 100 km long and 29-40 km wide, and is bounded by the San Miguel and Dolores rivers on the southwest and west, the Uncompahgre and Gunnison rivers on the east, the Colorado river on the North, and an escarpment of Dakota sandstone above Pleasant Valley and Dallas creeks on the southeast (Anderson et aI. 1992). Topography ranges from gently rolling to steep canyons. Deer were captured on winter range consisting of Pinyon pine (Pinus edulis)-Utahjuniper (Juniperus osteospenna)-sagebrush interspersed with open sagebrush parks. The climate is semi-arid, with warm summers and cold winter. METHODS We evaluated 10__ captive and 15 wild mule deer does to ascertain the accuracy of ultrasound and PSPB for detecting pregnancy and fetal rates. We restrained 6 (5 or 6 years old) of the 10 captive does in a capture pen and immobilized them with an intramuscular injection of 420-500 mg ofketamine and 80-100 mg ofxylazine (a 5:1 mixture) on 25 January 1999. We also immobilized 4 yearling (about 1.6 years old) captive does with a dart gun using 250 mg oftelazol and 125 mg ofxylazine. After processing, tranquilization was reversed with 10 mg of yohimbine. We evaluated the 15 wild does (2 yearling and 13 adults), which were culled for herd management purposes by the U.S. Fish and Wildlife Service, on 31 January 1999. We captured 40 adult mule deer does on the Uncompahgre Plateau with a netgun fired from a Hughes 500C helicopter (Helicopter Capture Services, Marysvale, Utah, USA; Barrett et aI. 1982). The does were captured on deer winter range in proportion to their density within 8 strata located around the edge of the plateau. The deer were manually restrained, hobbled, blindfolded, and transported by helicopter to a base station, usually <6 km from the capture site, where they were processed on 5-8 February 1999. We aged each doe as ajuvenile, yearling, or adult by tooth replacement and wear, weighed each doe, and injected the first 32 does subcutaneously �155 with 5 ml of vitamins A (500,000 iu's), D3 (50,000 iu's), and E (1,500,000 iu's). We attached a 148 MHz telemetry collar, and, after processing, released all except 1 doe at the processing sites; the 1 doe was transported by helicopter to the capture site and released there because a steep canyon separated the capture and processing sites. In addition, 1 juvenile female, 2 yearling females, and 2 males were captured. The juvenile was immediately released at the processing site, the 2 yearlings were processed similar to the adult does and released at the processing site, and the 2 males were immediately released at the capture site. We processed and released does within about 30 minutes of capture. We placed the 25 does used for accuracy evaluation and the 40 study does in dorsal recumbency and used a portable ultrasound unit (Aloka 500V, Aloka, Wallingford, Connecticut, USA) with a 5 MHZ linear-array probe, powered by a portable generator in the field, to ascertain pregnancy and number of fetuses/doe via trans rectal ultrasonography (Smith and Lindzey 1982, Bingham et al. 1990, Stephenson et al. 1995). We also collected about 10 ml of blood viajugular venipuncture to assess levels ofPSPB and ascertain pregnancy status. The blood was allowed to clot, centrifuged, and, the serum was decanted and stored on ice and then frozen. The serum was submitted to BioTracking (Moscow, Idaho, USA) to assess levels ofPSPB (Stephenson et al. 1995, Noyes et al. 1997). We ascertained accuracy of the ultrasound and PSPB procedures by performing necropsies and counting the number of fetuses in 1 captive doe that died from chronic wasting disease and 3 other does that were euthanized because of chronic wasting disease (M. A. Wild, Colorado Division of Wildlife, personal communication). We also ascertained accuracy of the ultrasound and PSPB by counting the number of fawns born during May-June 1999 for the other 6 captive does, whereas we necropsied the does collected on the Rocky Mountain Arsenal to determine the number of fetuses. One of us (L WJ) had extensive experience using ultrasound techniques and conducted all ultrasound procedures. Apriori, we conducted a power analysis to estimated the sample size necessary to detect a departure in fetal rates from previous studies of fetal rates in Colorado. For the previous studies, we used the weighted mean of fetal rates for adult does from .the Poudre River drainage (n = 41, x = 1.83, SD = 0.44; Medin and Anderson 1979), Middle Park (n = 119, x= 1.86, SD = 0.57; Gill 1971), the Forbes-Trinchera Ranch (n = 33, x = 1.91, SD = 0.46; Freddy 1987, 1988), and the Piceance Basin ([1973: n = 61, x= 1.46, SD = 0.77][1978: n = 37, x= 1.65, SD = 0.72]; G. C. White, Colorado State University, personal communication) to estimate a historic mean (1.74) and standard deviation (0.64). We used an expected 1999 standard deviation equal to the historic standard deviation (0.64; G. C. White, personal communication) in our analysis. We were primarily interested in ascertaining if pregnancy rates and fetal rates were a major contributor to the observed low December fawn:doe ratio (0.36 fawns/doe predicted by the regression equation for 1998 using data on fawn:doe ratios from 1982-1998) on the Uncompahgre Plateau. Considering historic fetal rates of 1.74 fetuses/doe, and a predicted fawn:doe ratio of 0.36 remaining on the Uncompahgre Plateau in December 1998, there is considerable loss of fetuses or fawns. On average, fawn:doe ratios in Colorado have declined by about 0.015 fawns/doe/year from 1972 to 1995 which results in a decline of about 0.3 fawns/doe over the 23 years. Fawn:doe ratios on the Uncompahgre Plateau have declined by 0.018 fawns/doe/year during the last 17 years (1982-1998), resulting in a decline of 0.30 fawns/doe over the 17 years. Thus, we attempted to determine if fetal rates contributed to the observed change of 0.30 fawns/doe which represents only 21% of the observed reduction in the fawn:doe ratio of 1.4 (1.74 - 0.36 [predicted by the regression equation for 1998] = 1.4). Thus, our sample size of 40 does was aprior estimated to have power of 80% for detecting a change of 0.3 fawns/doe at an alpha of 0.05. The proportion of adult does pregnant during 1999 was compared to historic data with a chisquare test (proc Freq, SAS Institute 1988). Fetal rates obtained during 1999 were compared to historic rates with a Z-test. An alpha level 0[0.05 was considered significant. �156 RESULTS Via necropsy or observing newborn fawns, we ascertained that 1 of the 10 captive does did not have a fetus, 3 does produced 1 fetus or fawn, and 6 does produced twin fetuses or fawns. Via ultrasound, we estimated that 3 captive does had single fetuses, 6 does had twins, and 1 doe had triplets. Thus, for each of 3 does, we estimated 1 more fetus via ultrasound compared to necropsies or observation of newborn fawns. All does were detected pregnant via PSPB levels. We necropsied the 15 culled mule deer does (2 yearlings and 13 adults) and found 5 does (2 yearlings and 3 adults) were barren, 3 does each contained 1 fetus, 6 does contained twins, and 1 doe contained triplets. Using ultrasound, we accurately detected all 15 does as pregnant or not pregnant. We also detected the correct number of fetuses per doe for 12 of the 15 does. We incorrectly detected 3 fetuses via ultrasound in 1 doe but she had only 2 fetuses, and we detected only 1 of 2 fetuses in 2 other does. PSPB levels indicated that 12 of the 15 does were pregnant; two of the 12 does were not detected-pregnant via both ultrasound and necropsy .. Thirty-seven of the 40 (93%) adult does on the Uncompahgre Plateau were detected pregnant with ultrasound. This proportion does not differ (XZJ = 0.053, P = 0.817) from the proportion of does pregnant (91.4% of303 does) in previous studies in Colorado (Gill 1971, 1972; Medin and Anderson 1979; Freddy 1987, 1988; R M. Bartmann, Colorado Division of Wildlife, unpublished data). The average number of fetuses/doe for all adult does (pregnant and non-pregnant) on the Uncompahgre Plateau (n = 40, ~= 1.70, SE = 0.109) was not less (Z= 0.340, P = 0.367) than in previous studies (n = 276, ~= 1.74, SE = 0.038; Gill 1971, 1972; Medin and Anderson 1979; Freddy 1987, 1988; R M. Bartmann, Colorado Division of Wildlife, unpublished data). The average number of fetuses/doe for pregnant adult does on the Uncompahgre Plateau (n = 37, ~ = 1.84, SE = 0.082) also was not less (Z = 0.260, P = 0.397) than in previous studies (n = 258, ~ = 1.86, SE = 0.028). PSPB levels concurred with ultrasound in ascertaining pregnancy in 39 of the 40 does; PSPB levels indicated 1 doe was not pregnant whereas 2 fetuses were visually detected via ultrasound. DISCUSSION We failed to detect a difference in pregnancy and fetal rates of adult female mule deer between our and previous studies in Colorado. Bartmann and Pojar (1998b) also found a relative high pregnancy rate of93.0% for 29 does ~1 year old near Red Feather, Colorado during January 1998. Thus, a failure to breed or maintain pregnancy thru at least early February can be discounted as the cause of the low fawn:doe ratios observed on the Uncompahgre Plateau. We restricted our. analyses of pregnancy and fetal rates to adult (~2 years old) mule deer does. We did not include yearlings in our sample because a lower proportion of yearlings become pregnant and they have lower fetal rates which would have biased our comparisons to historic rates unless the proportions of yearlings in both samples were equal. �157 LITERATIJRECITED Anderson, A. E., D. C. Bowden, and D. M. Kattner. 1992. The puma on Uncompahgre Plateau, Colorado. Colorado Division of Wildlife Technical Publication No. 40. Fort Collins, Colorado, USA. Barrett, M. W., J. W. Nolan, and L. D. Roy. 1982. Evaluation of a hand-held net-gun to capture large mammals. Wildlife Society Bulletin 10:108-114. Bartmann, R M. 1998. A prospectus on mule deer in Colorado. Colorado Division of Wildlife, Fort Collins, Colorado, USA. Unpublished report. _, and T. M. Pojar 1998a. Experimental deer inventory. Colorado Division of Wildlife, Federal Aid in Wildlife Restoration, Project W-153-R-11, Progress Report . ._, and _. 1998b. Deer reproduction assessment. Colorado Division of Wildlife, Federal Aid in Wildlife Restoration, Project W-153-R-11, Progress Report. Bingham, C. M., P. R Wilson, and A. S. Davies. 1990. Real-time ultrasonography for pregnancy diagnosis and estimation of fetal age in farmed red deer. The Veterinary Record 126:102-106. Freddy, D. J. 1987. Effect of elk harvest systems on elk breeding biology. Colorado Division of Wildlife, Federal Aid in Wildlife Restoration, Project 01-03-047, Progress Report. Freddy, D. J. 1988. Effect of elk harvest systems on elk breeding biology. Colorado Division of Wildlife, Federal Aid in Wildlife Restoration, Project W-153-R-2, Progress Report. Gill, R B. 1971. Middle Park deer study - productivity and mortality. Colorado Division of Wildlife, Federal Aid in Wildlife Restoration, Project W-38-R-25, Progress Report. _. 1972. Productivity studies of mule deer in Middle Park, Colorado. Second Annual Mule Deer Workshop, Elko, Nevada Haigh, J. C., W. J. Dalton, C. A. Ruder, and R G. Sasser. 1993. Diagnosis of pregnancy in moose using a bovine assay for pregnancy-specific protein B. Theriogenology 40:905-911. Lenz, M. F., A. W. English, and A. Dradjat. 1993. Real-time ultrasonography for pregnancy diagnosis and foetal ageing in fallow deer. Australian Veterinary Journal 70:373-375. Medin, D. E., and A. E. Anderson. 1979. Modeling the dynamics of a Colorado mule deer population. Wildlife Monographs 68: 1-77. Noyes, J. H., R G. Sasser, B. K. Johnson, L. D. Bryant, and B. Alexander. 1997. Accuracy of pregnancy detection by serum protein (PSPB) in elk. Wildlife Society Bulletin 25:695-698. Revol, B., and P. R Wilson. 1991. Ultrasonography of the reproductive tract and early pregnancy in red deer. The Veterinary Record 128:229-233. Rowell, J. E., P. F. Flood, C. A. Ruder, and R G. Sasser. 1989. Pregnancy-specific protein in the plasma of captive muskoxen. Journal of Wildlife Management 53:899-901. . Salwasser, H., S. A. Hell, G. A. Ashcraft. 1978. Fawn production and survival in the North Kings River deer herd. California Fish and Game 64:38-52. SAS Institute Inc. 1988. SAS/STAT User's guide, release 6.03 edition, SAS Institute Inc., Cary, North Carolina, USA. Schrick, F. N., and E. K. Inskeep. 1993. Determination of early pregnancy in ewes utilizing transrectal ultrasonography. Theriogenology 40:295-306. Smith, R B., and F. G. Lindzey. 1982. Use of ultrasound for detecting pregnancy in mule deer. Journal of Wildlife Management 46:1089-1092. Stephenson, T. R, J. W. Testa G. P. Adams, R. G. Sasser, C. C. Schwartz, and K. J. Hundertmark. 1995. Diagnosis of pregnancy and twinning in moose by ultrasonography and serum assay. Alces 31:167-172. Unsworth, J. W., D. F. Pac, G. C. White, R. M. Bartmann. 1999. Mule deer survival in Colorado, Idaho, and Montana. Journal of Wildlife Management 63:315-326. �158 Wilker, c., B. Ball, T. Reimers, G. Sasser, M. Brunner, B. Alexander, and M. Giaquinto. 1993. Use of pregnancy-specific protein-B and estrone sulfate for determination of pregnancy on day 49 in fallow deer (Dama dama). Willard, S. T., R G. Sasser, J. C. Gillespie, J. T. Jaques, T. H. Welsh, Jr., and RD. Randel. 1994. Methods for pregnancy determination and the effects of body condition on pregnancy status in Rocky Mountain elk (Cervus elephus nelsoni). Theriogenology 42:1095-1102. Wilson, P. R, and C. M. Bingham. 1990. Accuracy of pregnancy diagnosis and prediction of calving date in red deer using real-time ultrasound scanning. The Veterinary Record 126:133-135. Wood, A. K., R E. Short, A. Darling, G. L. Dusek, R G. Sasser, and C. A. Ruder. 1986. Serum assays for detecting pregnancy in mule and white-tailed deer. Journal of Wildlife Management 50:684--687. �159 APPENDIX II Research Program Narrative IMPACT OF PREDATION AND VEGETATIVE COVER ON NEONATAL MULE DEER FAWN SURVIVAL 21 October 1998 Principal Investigators: THOMAS M. POJAR Colorado Division of Wildlife 317 W. Prospect Fort Collins, CO 80526 970-472-4308 WILLIAM F. ANDELT Department of Fishery and Wildlife Biology Colorado State University Fort Collins, CO 80523 970-491- 7093 �160 �161 PROGRAM NARRATIVE State of --"C=o=lo=r=ad=o~ _ Project No. W!..!.--.:..:15::..::3:....-R~ _ Work Package No. _;3:::..;0:<...:0'-"1 _ Task No. ----:.4 _ Cost Center 3430 Mammals Program Deer Conservation Impact of Predation and Vegetative Cover on Mule Deer Fawn Survival NEED There is evidence that the mule deer (Odocoileus hemionus) population in Colorado has declined during recent years due mostly to low fawn survival and subsequent low population recruitment (Bartmann 1997). Colorado Division of Wildlife quadrat surveys to estimate population size suggest that some mule deer herds have declined by 31 % since 1992 while some have not shown much change during this time (Bartmann 1997). On 2 areas where recent over-winter (1997-98) survival was estimated, rates of fawn and adult doe survival was 74% (n = 38) and 100% (n = 30) for the Red Feather herd (D-4) and 49% (n = 39) and 84% (n =31) for the Uncompahgre herd (D-19), respectively (Bartmann and Pojar 1998a). This supports the contention that some of the purported population declines may be influenced by over-winter fawn survival. However, some herds have low December fawn:doe ratios indicating possible deficiencies in 2 major population growth mechanisms - initial fawn production (pregnancy rates) and neonatal fawn survival (Bartmann 1998). Pregnancy rates during January 1998 were 93% for 29 does >1 year old in the Red Feather DAU (Bartmann and Pojar 1998b) which was similar to 92.0% ofl63 does (>2 years old) pregnant in the same area during 1961-1964 (Medin and Anderson 1979), a 94.8% pregnancy rate for adult does (n = 114) and a 94.0% pregnancy rate for adult and yearling does (n = 134) in Middle Park (Gill 1971), and rates of 89% (n = 47) during 1973 and 82% (n = 83) during 1978 in the Piceance Basin (Bartmann 1998). The 1998 data indicate that the proportion of does becoming pregnant probably was not contributing to the low fawn:doe ratios, however these data were not collected in the Uncompahgre DAU where ratios were lowest. Pregnancy rates will be obtained in the Uncompahgre DAU during January 1999 to ascertain if these rates are contributing to the low fawn:doe ratios. Fetal rates for does older than yearlings were 1.83 (n = 41) in the Cache la Poudre River drainage from 1961-1965 (Medin and Anderson 1979) and 1.82 (n = 114) in Middle Park (Gill 1971). If fetal rates are currently as high as they were in the 1960's, then high rates of fawn mortality are likely responsible for the low. and declining fawn:doe ratios. Thus, determining the causes and magnitude of fawn mortality from birth through December will be extremely important for understanding mule deer population trends in Colorado. Low neonatal fawn survival is probably the most sensitive indicator that the environmental needs of populations are not being met (Gaillard et al. 1998). Fawn:doe ratios obtained in December have declined by an average of 1.5 fawns per 100 does per year from 1972-1995 (Bartmann 1997, G. C. White, Dept. Fishery and Wildlife Biology, Colorado State University, pers. commun.). December 1997 fawn:doe ratios were 34.2 in the Uncompahgre DAU and 58.8 in the Red Feather DAU (Bartmann and Pojar 1998a). The December fawn:doe ratios and over-winter survival rates indicate that the status of the Uncompahgre deer herd is of greatest concern. Gruell (1986) hypothesizes that historic (1880-1930) domestic livestock grazing and fire suppression caused successional changes in rangeland communities favoring shrubby species resulting in optimal mule deer habitat during the period 1930-1960. These conditions resulted in the irruption and overpopulation of deer herds and may have reduced present day carrying capacity which may be �162 mediated by lower nutrition andlor greater susceptibility to predation (see also Salwasser et al. 1978). Fire suppression and removal of herbaceous species by grazing can foster conditions that favor woody species invasion (Skovlin 1991) and decadent/senescent stands of shrubby species (Clements and Young 1996) thereby reducing the canying capacity for deer. Once the stable state of a particular vegetation type has been disrupted, especially in arid or semi-arid rangeland types, the vegetative community may stabilize at a lower successional state that is highly resistant to change without drastic perturbations (Laycock 1991). It is possible this situation is in process on the Uncompahgre Plateau (and other areas of the West) with the constant expansion of the pinon-juniper vegetative community into sagebrush and grassland communities. A low fawn:doe ratio on the Uncompahgre DAU could result from several causes including malnutrition and predation. Competition with elk (Cervus elaphus) for food or space on wintering grounds, or for fawning sites on summer range may cause poor neonate survival through decreased nutrition or increased susceptibility to predation. Livestock grazing can reduce forbs and fawn hiding cover (Loft et al. 1987) which can influence nutrition and predation (Smith and LeCount 1979, Bowyer and Bleich 1984), but there likely is less livestock grazing now thanwhen deer were more abundant indicating recent grazing may not be the cause of the decline. A short-term drought may have reduced forage for lactating does, may have protracted the fawning period making fawns more susceptible to predators as in white-tailed deer (Odocoi/eus virginianus, Andelt et al. 1987), or reduced alternate prey for predators which may be exacerbating the low fawn:doe ratios. Smith and LeCount (1979) reported that survival of mule deer fawns was associated with winter forb yield and October-April rainfall of the winter-spring period preceding the fawning period. Coyotes (Canis latrans) and other predators kill and eat mule deer fawns and adults, however the population impact of this predation is variable. Connolly (1978) cited 31 studies which reported predation as a limiting or regulating influence on ungulate populations in North America Connolly (1978) also cited 27 studies which indicated that predators did not limit or control the size of ungulate populations. Filonov (1980) provides evidence that the total loss to natural mortality for ungulates is relatively stable and that it is maintained by a complex system of compensating mechanisms, one of which is predation. Because of functional substitution of mortality factors, the various reasons for natural mortality are independent of the total mortality and the range of mortality rates remains somewhat constant. An example of compensatory mortality was demonstrated in the Piceance Basin of Colorado where predator control reduced overwinter mule deer mortality to predators, but the effect was compensated for by additional mortality due to starvation (Bartrnann et al. 1992). We need to determine the causes of neonatal fawn mortality in areas with low fawn:doe ratios, such as Uncompahgre Plateau, so that we know where to focus experimental research and management. Predator density is one factor that may have a significant impact on fawn survival independent of vegetational or nutritional components of the habitat (Smith et al. 1986). Coyotes, black bears (Ursus americanus), mountain lions (Felis concolor), and bobcats (Felis rufus) are found on the Uncompahgre Plateau and in other areas of Colorado, and may have a limiting effect on mule deer. Coyotes have been responsible for high neonatal white-tailed deer fawn (Cook et al. 1971, Garner et al. 1976, Litvaitis and Bartush 1980, Stout 1982), high neonatal mule deer fawn (Steigers and Flinders 1980, Steigers 1981), and high overwinter mule deer fawn (White et al. 1987) mortality. Beck (1995) reported a density of O. 93 black bears/mf on the Uncompahgre Plateau, and black bears can prey on mule deer fawns (Smith 1983), white-tailed deer fawns (Ozoga and Verme 1982, 1986; Mathews and Porter 1988), and elk calves (Schlegel 1976). Anderson et al. (1992) estimated that a minimum population of34 mountain lions on 3,120 km2 of the Uncompahgre Plateau may have killed 1,885 to 2,060 mule deer or about 8 to 12% of the estimated wintering deer population during 1987. Welker (1986) reported that mountain lions killed 2 or 3 of 16 radioed mule deer fawns; Bleich and Taylor (1998) found that mountain lion predation on adult does was the primary source of mortality in northeastem California herds. Bobcats also kill deer (Litvaitis et al. 1986) and pronghorn fawns �163 (Antilocapra americana) (Autenrieth 1982, 1984). Although these predators kill deer, their impact on mule deer fawn:doe ratios in Colorado is unknown. If predators are responsible for the low fawn:doe ratios, we need to determine which predator is the major contributor, and if some factors, such as low hiding cover, less alternate prey (Horejsi 1982, Andelt et al. 1987), a protracted fawning season, predator abundance, and possibly a high predator:deer ratio are making fawns more vulnerable to predation. Height of vegetation apparently plays an important role in the survival of neonate ungulates. Riley and Dood (1984) and Boyer et al. (1998) reported that mule deer fawns and (Decker 1991) reported that white-tailed deer fawns selected habitat types with dense vegetative cover. Horejsi (1982) reported that in years of drought, mule deer fawn survival was lower in areas oflivestock grazing and attributed this to decreased ground cover. Benzon (1996) reported that white-tailed deer fawns in the Black Hills, South Dakota, selected an understory of grasses for diurnal bed sites; height and density of vegetation was higher at bed sites than what was randomly available. Tucker and Garner (1983) reported that bed sites of pronghorn <4 weeks old were found in cover taller than that in surrounding areas. Autenrieth (1982, 1984) suggested that taller cover was important for reducing pronghorn fawn mortality to predators. Rothchild (1993) reported that vegetation was taller at the bed sites of pronghorn fawns than at random sites and the vegetation also was taller at bed sites of surviving pronghorn fawns compared to fawns that did not survive. Rothchild et al. (1984) suggested that pronghorn fawns with larger home ranges may sustain higher coyote predation in tall grass prairie of east central Kansas. Fawns have an infinite number of potential bed site locations within their home ranges. Measurement of random sites that are possible bed sites should be within a typical home range of a fawn. Home ranges of mule deer fawns in northern Colorado were variable and estimated at 130 ha from 10 June through August (Geduldig 1981). Average summer home range size offawns was 185 ha in the Missouri River Breaks, Montana and home range size decreased with increased population size (Riley and Dood 1984). Home range size of fawns increased with age in south-central Washington and averaged 257 ha for fawns 60 days or older (Steigers and Flinders 1980). Mule deer fawns traveled variable distances each day, and these distances more than doubled when fawns were 61-90 days old (640 m) compared to when 1-30 days old (319 m) (Steigers and Flinders 1980). OBJECTIVES We propose to identify the agents of mortality and quantify the extent of depredations on neonatal mule deer fawns. We also propose to correlate height, density, and composition of vegetation at fawn birth and bed sites with fawn survival. HYPOTHESES Our hypotheses are divided under two research areas, predation and vegetative cover. 1. What is the contribution of coyotes and other predators to neonatal mule deer fawn mortality? Hoi: The proportions of fawns killed by coyotes, black bears, mountain lions, and bobcats do not differ from one another. H.,2: Predation on fawns is not related to sex or activity of fawns; activity as measured by the distance between successive bed sites. Ho3: Predation on fawns to 8 weeks of age is not related to the average distance between a fawn and its mother. H04: Survival offawns and causes of mortality do not differ among 3 vegetatively and ecologically different study areas. Hos: Fawn birth weights and measurements do not differ among study areas. Roo: Distances between fawn bed sites do not differ among study areas. �164 2. Do height and density, and/or composition of vegetation at fawn birth and bed sites correlate with the percent of fawns killed by predators? H"I: Predation rates on fawns is not related to height and density (visual obstruction readings), and/or composition of vegetation at birth and bed sites. Ho2: Height and density (visual obstruction readings, VOR), and/or composition of vegetation do not differ between fawn bed sites and random sites. Ho3: Height and density (VOR), and/or composition of vegetation at fawn bed sites and random sites do not vary among study areas. EXPECTED RESULTS OR BENEFITS This research will quantify the amount and causes of neonatal fawn mortality and contribution of various predators to this mortality. The influence of vegetative characteristics at birth and bed sites on subsequent fawn survival will be estimated. Knowledge of these factors will assist in evaluating the causes of low December fawn:doe ratios and will guide future research to determine the effect of various experimental manipulations on neonatal fawn survival. For political, social, and economic reasons, we need to have knowledge about the causes of low fawn:doe ratios before corrective action is taken. APPROACH Animal Care and Use: All capture and handling of animals will comply with the standards of the Colorado Division of Wildlife and Colorado State University Animal Care and Use Committees. We will obtain a collectors permit from the Colorado Division of Wildlife and secure trespass permission from study site owners prior to the study. Fawn Capture: In a related study, 40 radioed mature does will be monitored on each study area to determine adult doe survival (Federal Aid Project W-153-R); fawns will be located primarily by monitoring these does and capturing their fawns. Our objective will be to capture and radio 25-35 fawns on each of 3 study areas. Three study areas will be selected based on : a) contrasting vegetative/ecological characteristics, b) a range in fawn:doe ratios indicating contrasting fawn survival rates, and c) availability of radio collared does to minimize fawn capture costs and ensure randomness in fawn capture. Three candidate areas which meet all 3 characteristics are: The Uncompahgre, Middle Park, and Red Feathers DAUs. Fawns <10 days old will be captured and fitted with expandable, drop-off radio collars. Fawns younger than 24 hr will not be captured to allow dam-newborn fawn bonding (White et al. 1972). Precautions will be taken to minimize injury or abandonment by the dam from the capture operation based on recommendations of Beale and Smith (1973) and Livezey (1990). The radio collars and all handling equipment will be stored in containers with native shrub/tree (sagebrush or juniper) material to help mask "unnatural" odors. Plastic gloves will be worn by the handlers. Fawn Processing: Captured fawns will be blindfolded, weighed, and the right hind foot will be measured as an index of skeletal size. If age is not known from observed parturition, it will be estimated by hoof condition and length, umbilical condition, and general body size, activity, and hair coat condition will be subjective measures of age. Each fawn will be individually marked with a plastic ear tag in the right ear; the ear tag will have engraved numbers which contrast to the color of the tag. An expandable radio transmitter (with mortality sensor) collar will be fitted; the collar will be designed to drop off after about 6 months. Before release, the fawn will be examined for injuries and it's general health and condition will be noted and recorded. Each fawn will be carefully observed after release and, . if possible, observed until it is rejoined by it's mother. �165 Fawn and Doe Monitoring: Each fawn and doe will be radio located daily for the first 6 weeks of the fawn's life, every 3rd day for the next month, and then once weekly through the end of November. Precautions will be taken to avoid disturbance of the fawn or dam during radio relocation except when prescribed bed site measurements are taken (see below). Radios on mortality signal will be located and the carcass (if available) will be examined for cause of death using criteria outlined in White (1973), Wade and Bowns (1984), Acorn and Dorrance (1990), and Andelt et al. (1998). If the carcass is missing, the general area will be surveyed for signs of predation and our best judgement of cause of death will be made or the cause will be recorded as "unknown". Birth and Bed Site Measurements: Vegetative cover will be measured at birth sites when they can be positively identified. Measurements will be taken at bed sites every 5th day to minimize disturbance of individual fawns. If a bedded fawn is found and does not flush, the point of observation will be flagged and GPS coordinates will be recorded. Distance to the bed sitewill be measured with a laser range finding scope (e.g. Bushnell "yardage Pro") and the direction to the site will be measured with a hand held compass. Marked bed sites will be measured the next day to minimize any changes in vegetation from time of use. The objective will be to obtain lObed site measurements for each fawn during their first 8 weeks of life. Micro habitat vegetational measurements of birth and bed sites will be a modification of those taken by Benzon (1996). Cover of shrubs by species, grass, and forbs will be estimated using 4 0.1 m2 (20 x 50 em) quadrants (Daubenmire 1959) 'placed in the 4 cardinal directions length-wise from the bed center; the 20 em end of the frame will be centered on and touching the bed site center stake. Visual obstruction from the birthlbed site will be estimated from the 4 cardinal directions using a robel pole viewed from a distance of 4 m and height of 0.5 m from the site (Robel et al. 1970). No adjustments will be made for relief in the terrain. For every birth or bed site measured, we will take identical measurements on IO.random sites within aBO ha square area centered on the bed site. The random sites will be located using random coordinates. Sample Sizes and Power of Tests: Estimates of the means and variances of2 or more treatments are necessary before conducting power analyses. Estimates of these parameters are not known for the proportions of fawns killed by various predators, and the effect of vegetation height on susceptibility of fawns to predation. However, some educated guesses about the magnitude of these parameters will help determine sample sizes. We calculated power for our hypotheses of primary management interest including: 1) proportions of fawns killed by various predators, and 2) the effect of vegetation height on susceptibility of fawns to predation. We calculated for vegetation height vs. susceptibility to predation by using 2 height categories, whereas the actual analysis' will consist of regression using a continuum of vegetation heights. Thus, the power of the actual experiment likely will be somewhat less than presented below. We did not calculate power for our hypothesis of differences in vegetation height at fawn bed sites and random sites because sample sizes will be inherently larger that number 2 above. An alpha of 0.05 was used in all estimates of power. Power of 0.8 or higher is desired. Data Analyses: The proportions of fawns killed and the proportions killed by various causes of mortality (predation, disease, malnutrition) will be compared among study areas with chi-square (proc Freq, SAS Institute, Inc. 1988). The proportion of fawns killed by various predators and the proportions of male and female fawns killed by predators will be compared with chi-square. The effect of distance between bed sites and the distance between fawns and their mothers on fawn survival will be estimated with logistic regression (Proc Genmod, SAS Institute Inc. 1993) by blocking on study areas. Fawn birth weights and measurements, and distances between consecutive bed sites will be compared among study areas with analysis of variance. The effect of vegetation height and density (VOR) on survival of fawns will be estimated with logistic regression (proc Genmod, SAS Institute Inc. 1993) by blocking on study �166 Power for Evaluating if the Expected Relative Proportions of Fawns Killed by Coyotes, Black Bears, and Mountain Lions Varies from Equality (i.e. 0.33 killed by each predator): Fawns killed en) 10 20 30 50 75 100 10 20 30 50 75 100 Coyotes 0.5 0.5 0.5 0.5 0.5 0.5 0.7 0.7 0.7 0.7 0.7 0.7 Proportion killed by various predators Black Bears Mountain Lions 0.25 0.25 0.25 0.25 0.25 0.25 0.25 0.25 0.25 0.25 0.25 0.25 0.15 0.15 0.15 0.15 0.15 0.15 0.15 0.15 0.15 0.15 0.15 0.15 Power 0.16 0.27 0.39 0.60 0.79 0.90 0.59 0.89 0.98 1.00 1.00 1.00 I-Sided Power for the Effect of Vegetation Height on Survival of Fawns to 2 Months: Sample sizes Short 20 40 40 80 40 40 40 Tall 20 40 40 80 40 40 40 ProPQrtion surviving Short Tall 0.7 0.8 0.7 0.8 0.7 0.9 0.7 0.9 0.5 0.64 0.5. 0.7 0.5 0.8 Power 0.11 0.18 0.61 0.88 0.27 0.73 0.80 areas (Appendix 1). The height and density of vegetation at fawn bed sites and random sites will be compared with analysis of variance by blocking on study areas. The proportion of vegetation that is composed of forbs, grass, and shrubs will be compared between bed sites and random sites and among study areas with analysis of variance. Significance level to reject the null hypothesis will be at P < 0.05. LOCATION This study will be done on 3 areas, Uncompahgre Plateau (D-19), Middle Park (D-9), and Red Feathers (D-4). WORK SCHEDULE Oct-Dec 1998 Jan-May 1999 Jun-Dec 1999 Jan-Jun 2000 Complete Program Narrative and confirm study area selection Procure and test field equipment and hire summer crew Capture, radio collar, and track fawns Data analysis and report writing PERSONNEL Thomas M. Pojar Co-Principal Investigator, CDOW William F. Andelt Co-Principal Investigator, Department of Fishery and Wildlife Biology, Colorado State University �167 David C. Bowden Statistical Consultant, Department of Statistics, Colorado State University Gary C. White Analytical Consultant, Department of Fishery and Wildlife Biology, Colorado State University ESTIMATED COST Personal Services PFTE costs Andelt Pojar TFTE costs Operating Vehicle (4x4) Radios Field equipment Travel Expenses Capitol TOTAL 5% contingency GRAND TOTAL FY 98-99· FY 99-00b Totalc $ 56,250 $ 45,000 $ 18,216 $ 75,000 $ 60,000 $ 26,717 $131,250 $105,000 $ 44,933 $ 18,306 $ 22,000 $ 13,500 $ 1,800 $ 33,900 na na $ 2,250 0 $197,867 $ 9,893 $207,760 $ 52,206 $ 22,000 $ 13,500 $ 4,050 0 $372,939 $ 18,647 $391,586 o $175,072 $ 8,754 $183,826 "Cost for FY 1998-99, representing field operations during May-June, 1999. "Cost for completion of the 1999 field operation during FY 1999-00, July-December 1999. "Total cost for 1 complete field season spanning 2 fiscal years. LITERATURE CITED Acorn, R. C., and M. 1. Dorrance. 1990. Methods of investigating predation of livestock. Alberta Agric. Prot. Branch. Edmonton. 36pp. Andelt, W. F., M. Bruscino, and C. Niemeyer. 1998. Interpreting the physical evidence of predation on big game animals and domestic livestock. Unpublished. Andelt, W. F., 1. G. Kie, F. F. Knowlton, and K. Cardwell. 1987. Variation in coyote diets associated with season and successional changes in vegetation. Journal of Wildlife Management 51:273277. Anderson, A. E., D. C. Bowden, and D. M. Kattner. 1992. The puma on Uncompahgre Plateau, Colorado. Colorado Division of Wildlife Technical Publication No. 40. Fort Collins, Colorado, USA. Anderson, A. E., and D. E. Medin. 1965a. Reproductive studies. Colorado Game, Fish, and Parks Department, Federal Aid in Wildlife Restoration Job Progress Report, Project W-38-R-4. January:165-193. Anderson, A. E., and D. E. Medin. 1965b. Reproductive studies. Colorado Game, Fish, and Parks Department, Federal Aid in Wildlife Restoration Job Progress Report, Project W-38-R-4. January Part 4:522-552. �168 Anderson, A. E., and D. E. Medin. 1966. Reproductive studies. Colorado Game, Fish, and Parks Department, Federal Aid in Wildlife Restoration Job Progress Report, Project W-38-R-4. January Part 2:275-290. Autenrieth, R. E. 1982. Pronghorn fawn habitat use and vulnerability to predation. Pronghorn Antelope Workshop, Dickinson, ND. 10:1l2-13l. Autenrieth, R. E. 1984. Little lost pronghorn fawn study - condition, habitat use and mortality. Pronghorn Antelope Workshop, Corpus Christi, Texas. 11:49-70. Bartmann, R. M. 1997. Mule deer population status in Colorado - update. Colorado Division of Wildlife, Fort Collins, Colorado, Unpublished report. Bartmann, R. M. 1998. A prospectus on mule deer in Colorado. Colorado Division of Wildlife, Fort Collins, Colorado, Unpublished report. Bartmann, R. M., and T. M. Pojar 1998b. Deer reproduction assessment. Colorado Division of Wildlife, Federal Aid in Wildlife Restoration Job Progress Report, Project W-153-R-l1, July. Bartmann, R. M., and T. M. Pojar 1998a Experimental deer inventory. Colorado Division of Wildlife, Federal Aid in Wildlife Restoration Job Progress Report, Project W-153-R-ll, July. Bartmann, R. M., G. C. White, and L. H. Carpenter. 1992. Compensatory mortality in a Colorado mule deer population. Wildlife Monographs 121:1-39. Beale, D. M., and A. D. Smith. 1973. Mortality of pronghorn antelope fawns in western Utah. Journal of Wildlife Management. 37:343-352. Beck, T. 1995. Development of black bear inventory techniques. Colorado Division of Wildlife, Federal Aid in Wildlife Restoration Job Progress Report, Project W-153-R-8, WP 5, Job 2, July. Benzon, T. A. 1996. Mortality and habitat use of white-tailed deer fawns in the Northern Black Hills, South Dakota, 1991-1994. Completion Report. Pittman-Robertson Project W-75-R-34. Bowyer, R. T., and V. C. Bleich. 1984. Effects of cattle grazing on selected habitats of southern mule deer. California Fish and Game 70:240-247. Boyer, R. T., 1. G. Kie, and V. Van Ballenberghe. 1998. Habitat selection by neonatal black-tailed deer: climate, forage, or risk of predation. Journal of Mammalogy 79:415-425. Clements, C. D., and 1. A. Young. 1997. A viewpoint: Rangeland health and mule deer habitat. Journal of Range Management 50:129-138. Connolly, G. E. 1978. Predators and predator control in big game management. Pages 369-394 in 1. L. Schmidt and D. L. Gilbert, eds. Big game of North America Stackpole Books, Harrisburg, Pa Daubenmire, R. 1959. A canopy-coverage method of vegetation analysis. Northwest Science. 33:4364. Filonov, C. 1980. Predator-prey problems in nature reserves of the European part of the RSFSR. Journal of Wildlife Management. 44:389-396. Gaillard, J. M., M. Festa-Bianchet, and N. G. Yoccoz. 1998. Population dynamics oflarge herbivores: variable recruitment with constant adult survival. Trends in Ecology and Evolution 13:58-63. Garner, G. W., J. A. Morrison, and J. C. Lewis. 1976. Mortality of white-tailed deer fawns in the Wichita Mountains, Oklahoma. Proceedings of the Annual Conference of Southeastern Fish and Wildlife Agencies 30:493-506. Geduldig. H. L. 1981. Summer home range of mule deer fawns. Journal of Wildlife Management 45:726-728. Gill, R. B. 1971. Middle Park deer study - productivity and mortality. Colorado Division of Wildlife, Federal Aid in Wildlife Restoration Job Progress Report, Project W-38~R-25. July:189-207. Gruell, G. E. 1986. Post-1900 mule deer irruptions in the Intermountain West. United States Department of agriculture, Forest Service. Intermountain Research Station, Ogden, Utah. General Technical Report INT-206. �169 Horejsi, R G. 1982. Mule deer fawn survival on cattle-grazed and ungrazed desert ranges. Arizona Game and Fish Department Federal Aid Project Report, Phoenix. 43pp. Laycock, W. A. 1991. Stable states and thresholds of range condition on North American rangelands: A viewpoint. Journal of Range Management 44:427-433. Litvaitis, J. A and W. S. Bartush. 1980. Coyote-deer interactions during the fawning season in Oklahoma Southwestern Naturalist 25: 117-118. Litvaitis, J. A, A G. Clark, and J. H. Hunt. 1986. Prey selection and fat deposits of bobcats (Felis rufus) during autunm and winter in Maine. Journal ofMammalogy 67:389-392. Livezey, K. B. 1990. Toward the reduction of marking-induced abandonment of newborn ungulates. Wildlife Society Bulletin 18:193-203. . Loft, E. R, J. W. Menke, J. G. Kie, and R. C. Bertram. 1987. Influence of cattle stocking rate on the structural profile of deer hiding cover. Journal of Wildlife Management 51:655-664. Mathews, N. E., and W. F. Porter. 1988. Black bear predation of white-tailed deer neonates in the central Adirondacks. Canadian Journal of Zoology 66: 1241-1242. Medin, D. E., and A. E. Anderson. 1979. Modeling the dynamics of a Colorado mule deer population. Wildlife Monographs 68: 1-77. Ozoga, J. J., and L. J. Verme. 1982. Predation by black bears on new-born white-tailed deer. Journal ofMammalogy 63:695-696. Ozoga, J. J., and L. J. Verme. 1986. Relation of maternal age to fawn-rearing success in white-tailed deer. Journal of Wildlife Management 50:480-486. Riley, S. J. and A. R Dood. 1984. Summer movements, home range, habitat use, and behavior of rriule deer fawns. Journal of Wildlife Management 48: 1302-1310. _ Robel, R J., J. N. Briggs, A. D. Dayton, and L. C. Hulbert. 1970. Relationships between visual obstruction measurements and weight of grassland vegetation. Journal of Range Management. 23:295-297. Rothchild, S. L. 1993. Mortality, home range, and habitat use of pronghorn fawns within tallgrass prairie of eastern Kansas .. M.S. thesis, Emporia State University. 85pp. Rothchild, S. L., E. J. Finck, and K. E. Sexson. 1994. Mortality and bedding site selection of pronghorn fawns in tall grass prairie. Proceedings of the Pronghorn Antelope Workshop 16:104-116. Salwasser, H., S. A Holl, G. A Ashcraft. 1978. Fawn production and survival in the North Kings River deer herd. California Fish and Game 64:38-52. SAS Institute Inc. 1988. SAS/STAT user's guide. Version 6.03. SAS Institute Inc., Cary, North Carolina. SAS Institute Inc. 1993. SASR Technical Report P-243, SAS/STATR Software: The GENMOD Procedure, Release 6.09, Cary, North Carolina Schlegel, M. 1976. Factors affecting calf elk survival in Northcentral Idaho. A progress report. Proceedings of the Western Association of State Game and Fish Commissioners 56:342-355. Skovlin, J. M. 1991. Fifty years of research progress: a historical document on the Starky Experimental Forest and Range. General Technical Report PNW-GTR-266. Portland, Oregon: U. S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. Smith, R 1983. Mule deer reproduction and survival in the LaSal Mountains of Utah, 1983. M.S. thesis, Utah State University, Logan. Smith, R H., and A LeCount. 1979. Some factors affecting survival of desert mule deer fawns. Journal of Wildlife Management 43:657-665 Smith, R H., D. J. Neff, and N. G. Woolsey. 1986. Pronghorn response to coyote control - a benefit cost analysis. Wildlife Society Bulletin 14:226-231. . Steigers, W. D., Jr. 1981. Habitat use and mortality of mule deer fawns in western South Dakota Ph.D. Dissertation, Brigham Young University. 206pp. �170 Steigers, W. D., Jr., and 1. T. Flinders. 1980. Mortality arid movements of mule deer fawns in Washington. Journal of Wildlife Management 44:381-388. Stout, G. C. 1982. Effects of coyote reduction on white-tailed deer productivity on Fort Sill, Oklahoma. Wildlife Society Bulletin 10:329-332. Tucker, R. D. and G. W. Gamer. 1983. Habitat selection and vegetational characteristics of antelope fawn bedsites in West Texas. Wade, D. A, and J. E. Bowns. 1984. Procedures for evaluating predation on livestock and wildlife. Texas Agricultural Experiment Station Bulletin B-1429, College Station. Welker, H. J. 1986. Fawn mortality in the Lake Hollow deer herd, Tehama County, California California Fish and Game 72:99-102. White, G. C., R. A Garrott, R. M. Bartmann, L. H. Carpenter, and A W. Alldredge. 1987. Survival of mule deer in northwest Colorado. Journal of Wildlife Management 51:852-859. White, M. 1973. Description of remains of deer fawns killed by coyotes. Journal of Marnmalogy 54:291-293. . White, M., F. F. Knowlton, and W. C. Glazener. 1972. Effects of dam-newborn fawn behavior on capture and mortality. Journal of Wildlife Management 36:897-906. Prepared by _ Thomas M. Pojar Wildlife Researcher Colorado Division of Wildlife William F. Andelt Associate Professor Department of Fishery and Wildlife Biology Colorado State University Appendix 1. Logistic regression table for the effect of vegetation height on survival offawns. The analyses will be conducted with SAS using Proc Genmod. Numerator Source Height Areas DF 1 2 M.S. height area Denominator DF M.S. Fawns -5 Error (Model) Fawns -5 Error (Model) � https://cpw.cvlcollections.org/files/original/961ea51651a9932493969854b32f3877.pdf ba9d86624255db5db3928285ec10db65 PDF Text Text 171 Colorado Division of Wildlife Wildlife Research Report July 1999 JOB PROGRESS REPORT Smteof ~C~o~lo~r~ad~o~ _ Mammals Research Project No. ---'W.:.,.--=1...:,5;::;..3-..:.R::....-=12=--_ Elk Investigations Work Package __ --=3::...:0:;,,::0:,:2'-_ Task No. ---=-1 _ Estimating Survival Rates of Elk and Developing Techniques to Estimate Population Size Period Covered: July 1, 1998 - June 30, 1999 Author: D. 1. Freddy Personnel: R. Adams; T. Beck J. Broderick, G. Byrne, D. Crane, R. Del Piccolo, J. Ellenberger, V. Graham, D. Homan, R. Kahn, K. Madariaga, D. Masden, C. Mehaffy, P. Neil, J. Olterman.J, Thompson, P. Will, K. Wright, S. Yamashita, D. Younkin, CD OW; D. Bowden, G. White.T, Gross, M. Kneeland, W. Andelt, CSU; D. Ouren, D. Schneider, T. Fancher USGS-BRD, BLM Glenwood Springs, CO, USFS Rifle, CO, Rocky Mountain Elk Foundation, cooperating. ABSTRACT We monitored survival and movements of 169 previously radio-collared elk composed of 136 adult females and 33 adult males from 1 July 1998 through 30 June 1999. Survival of adult females during summer-fall was 0.77 ± 0.07 (n = 136) inclusive of hunting deaths and 1.00 (n = 105) with hunting deaths censored while survival during winter-spring was 0.98 0.03 (n = 103). The 2 deaths of adult females during winter-spring were attributed to accident and malnutrition. Survival of adult males during summer-fall was 0.34 0.17 (n = 32) inclusive of hunting deaths and 1.00 (n = 11) with hunting deaths censored while survival during winter-spring was 1.00 (n = 11). A combination of unlimited either-sex elk hunting licenses and high numbers of antlerless elk hunting permits during rifle hunting seasons in 1998 contributed to removing 23% of the adult females which was sufficient to temper population growth. Progress continues towards developing habitat use models to predict areas important to managing elk. Draft manuscripts on elk survival rates and techniques to estimate population size are progressing. ± ± �172 �173 ESTIMATING SURVIVAL RATES OF ELK AND DEVELOPING ESTIMATE POPULATION SIZE TECHNIQUES TO David 1. Freddy P. N. OBJECTIVE Estimate survival rates of adult female, adult male, and calf elk and develop techniques to estimate population size. SEGMENT OBJECTIVES 1. Estimate winter, summer, and annual survival rates of adult female and male elk from fates of previously radio-collared elk. 2. Develop an initial habitat use prediction model for elk inhabiting oakbrush dominated habitats based upon locations of radio-collared elk obtained from 1993 -98. 3. Analyze data, summarize annually in Federal Aid Job Progress Report, and prepare draft manuscripts regarding elk survival rates and techniques to estimate population size. INTRODUCTION Our objectives are to provide reliable estimates of survival rates for calves during winter and for adult females and males throughout the year and to develop and test a system to estimate elk densities on winter ranges. Estimates of calf survival were obtained during winters 1993-94 through 1996-97 and summarized in Freddy (1997). We are continuing to obtain estimates of survival for adult males and females. For adults, we are interested in survival rates inclusive of hunting mortalities to document human-induced rates of survival and exclusive of hunting mortalities to estimate natural rates of survival. We are evaluating a system for estimating population size or density that incorporates estimates of sighting bias in conjunction with random sampling of search quadrats as sample units. Our winter study area encompasses about 324 rnf (839 knr') in the eastern half of Game Management Unit 42 south and east of Rifle, Colorado. This area is part of the Grand Mesa Elk Data Analysis Unit E-14. Elk winter habitats include juniper-pinyon woodland (Juniperus osteosperma-Pinus edulis), oakbrush-mountain shrub (Quercus gambelii-Amelanchier alnifolia), aspen (Populus tremuloides), sagebrush (Artemisia tridentata), and agricultural fields below elevations of9,800 ft. In summer, elk use oakbrush-mountain shrub, aspen, and subalpine fir-Engelmann spruce (Abies lasiocarpa-Picea engelmannii) meadow systems throughout the Grand Mesa region up to elevations of 12,000 ft. (Freddy 1993). METHODS We placed radio collars (172-176MHz) having mortality sensors on elk calves age 6 months and adults age 2:18 months during December 1993-1996 (Freddy 1997). Elk were captured using helicopter net-gunning and portable corral traps (Freddy 1997). During 1998-99, we monitored 169 adult elk, 136 females and 33 males, having functioning radios as of July 1998. Collars were white and had black identification symbols on dorsal surfaces of collars to allow visual identification of individual elk. �174 Estimating Elk Survival Rates We monitored survival of radioed elk with aerial surveys using a Cessna 185 at 2-4 week intervals from December 1993 to June 1999 and with daily ground surveys from December-April each year from 1993-94 to 1996-97. Survival rates (S) of radioed elk were calculated using the binomial estimator with a variance, V AR(S) = S(I-S)/n collars (White and Garrott 1990). Survival rates are expressed as the mean estimate the 95% confidence interval. We used x2-contingency tests to compare survival rates between sexes and among time intervals (pROC FREQ, SAS 1988). Sample sizes refer to numbers of individual radioed elk. We defined 3 major time intervals for survival analyses: winter-spring was 1 December to 14 June, summer-fall was 15 June to 30 November, and annual was 1 December to 30 November to coincide with timing of capture and collaring. For adult elk during these time intervals, we calculated survival rates inclusive of natural and hunting-related mortalities, exclusive of hunting mortalities, and exclusive of natural mortalities. Excluding, or censoring hunting mortalities, provided estimates of natural survival rates. Excluding natural mortalities but including hunting mortalities provided estimates of hunting removal rates. Censoring elk associated with categories of mortalities reduced sample sizes used to estimate survival rates. Elk were also censored ifradios failed or if their life/death status was unknown after an extended period of time. Archery, muzzleloading, and rifle hunting seasons occurred from about 1 September to 15 November each year during the summer-fall time interval. Late-rifle seasons occurred in some years . and restricted hunters to taking only antlerless elk from about 25 November-31 January which included portions of the summer-fall and winter-spring time intervals. Yearling spike-antlered males were generally not legal quarry in most areas frequented by radioed elk. Male elk were legal quarry when branch-antlered usually at age 2 years. Cause of death of radioed elk recovered in the field was estimated from presence or absence of gunshot wounds or bite wounds on carcass, predator tracks or scat at carcass site, physical positioning of carcass remains whether buried, covered, scattered, or consolidated (Wade and Browns 1982, Halfpenny and Biesiot 1986), relative amount of internal fat and marrow fat if present with carcass, and results of histopathology and marrow fat analyses. Fat content (percent dry matter) of bone marrow and estimates of age based on dental cementum were obtained for dead elk by the Colorado Division of Wildlife Laboratory while histopathology analyses were provided by the Colorado State University Veterinary Diagnostic Laboratory. Photographs were taken of nearly all mortalities so that physical evidence could be reviewed and judged by outside experts (pers. comm. T. Beck, W. Andelt). ± Elk Movements and Habitat Use Models We continued to precisely locate selected radioed elk at least once per month since their capture to document seasonal movements using a Cessna 185. These elk were originally selected at random from within trap zones and equalized by age class in January 1994. Elk from the original sample that died were replaced each subsequent January with randomly selected elk that were usually of the same sex, age class, and trap zone as elk that died. All replacement elk in January 1999 were ;:::2-years old. As of 1 January 1999, these 43 elk were classified as 32 adult females and 11 adult males males. During June 1999, we again located an additional 28 adult females that were originally selected at random in 1995 to document their locations during the calving period. Females from the original 1995 sample that died were replaced each subsequent June with adult females randomly selected from the same trap zones of elk that died. As needed, we located other elk to document unusual movements. We continue to cooperate with the USGS-BRD, USFS, and BLM with the support of the Rocky Mountain Elk Foundation to create a GIS database allowing analyses relating elk locations, movements, and home ranges to habitat components with the intent of developing a predictive habitat use model. During 1998, initial efforts were undertaken to develop a model with Dr. 1. Gross at the Natural Resource Ecology Laboratory at Colorado State University. �175 Elk habitat models are being developed by using a staged process. Major stages included initial data analysis and reduction, development of predictive variables (i.e., GIS coverages), and . development and statistical analysis of alternative habitat models. Initial data analysis included an extensive process to locate and correct errors in the data set, create new variables (e.g., age, class, season), and to subset the data so that analyses included only the animals of interest. This first stage of this process was accomplished by developing a database application in MicroSoft Access© that validated data records and produced new tables with the data needed for further analysis. The entire data set consisted of 5,321 elk locations. We reduced the data set to include only data from elk that were randomly selected and systematically located during years 1993-1998. This reduced total elk locations to 2,888. Locations were subset into groups by sex, age (calf, yearling, adult), and season. Seasons were biologically defined as spring (1 April- 30 May), summer (1 June - 31 August), fall (1 September - 30 November), and winter (1 December - 31 March). Using ArcInfo© and Ar~VieW©, we linked each location to GIS coverages for elevation, slope, aspect (EW and NS), distance to a road, and to tasselcap indices for brightness, greenness, and wetness. To evaluate the ability of these variables to predict elk observations, we used ordinary least squares (Upton and Fingleton 1985) to identify significant relationships arid coefficients that were used for logistic regression. RESULTS AND DISCUSSION Between 1 December 1993 and 14 June 1999,235 radioed elk died of which 31 were calves 611 months old and 204 were adults ~12 months old (Table 1, Appendix I). Calves died of natural causes (Freddy 1997) while for adults hunting accounted for 93% of 204 deaths. Hunting accounted for 98% of 102 deaths for adult males and 88% of 102 deaths for adult females. . Survival Estimates Adult Females Summer-Fall Survival during summer-fall 1998 for adult females all age 2:.24 months was 0.77 ± 0.07 (n = 136) inclusive of hunting deaths and 1.00 (n = 105) with hunting deaths censored (Table 6). Net survival in 1998 was the lowest measured during 5 summer-fall periods due to increased hunting deaths that resulted in a hunting removal rate of 23%. Reduced survival due to hunting in 1998 was evident in all cohorts offemales (Tables 2-5,7,8). Natural survival rates remained >0.99 during all 5 years (Table 6). Increased hunting removal rates were caused by a combination of increased numbers of limitedentry rifle antlerless permits and first-time use of unlimited over-the-counter rifle either-sex permits. Unlimited either-sex permits were valid in Game Management Units (GMUs) 43,52, and 521 during the second and third rifle seasons while limited antIerless permits were available for GMUs 42 and 421 for the first, second, and third rifle seasons. These GMUs collectively encompassed the area inhabited by radioed female elk. Approximate locations of each radioed female were obtained prior to each season, each elk was assigned to a GMU, and life/death status of each elk was known prior to and following each season. During the second rifle season, adult female removal rates were greater in areas having unlimited either-sex permits (26%) than in areas having limited antIerless permits (6%) (P=O.OOI, X2 =10.7, dJ=I). During third rifle season, removal rates were not different between types of permits (P=0.53, X2 =0.39, dJ=I)(Table 10). Removal rates with antlerless permits were consistent between seasons while removal rates with either-sex permits declined drastically during the third season (Table 10). In this geographic area, unlimited either-sex permits provided a burst of female removal only during the second season. �178 Prepared by _ David J. Freddy Life/Science Researcher Table l. Causes of deaths in radio-collared elk between 1 December 1993 and 14 June 1999. Calves (M=male, F=female) were age 6-11 months and collared at age 6 months. Yearling males and females .were age 12-17 months and juvenile males and females were age 18-23 months and all were collared at age 6 months. Adult males and females were age >24 months at time of death. Elk Age/Sex Class at Death Adult Total Cause of Death Calves Yearling Juvenile M F M All and -Code M F M F M F F Natural Causes 0 3 Malnutrition-6 2 0 0 0 2 5 2 0 Unknown-Suspect Malnutrition-31 0 0 3 1 3 1 0 0 0 0 4 Predation-Lion-3 0 0 1 8 5 13 8 4 0 0 0 0 Predation-Bear- 35 0 0 0 0 0 0 0 0 0 0 0 0 0 Unknown Predator-5 0 0 0 0 0 1 1 1 Unknown-Suspect 0 Predation-30 2 1 1 2 3 8 11 5 0 0 Accident-Birthing-32 0 0 0 2 0 2 0 0 0 0 2 Accident-Fell-IO 0 1 0 0 2 2 0 0 0 1 3 Unknown Cause-l l 2 0 0 2 4 1 0 0 1 2 6 Subtotals 10 26 45 14 Q Q Q £ £ 12 11 Le~1 Hunting Arc ery-33 0 0 0 0 9 3 9 5 0 2 14 ·4 Muzzleloading-34 0 0 0 0 8 2 8 12 0 2 0 0 0 0 0 1 0 1 0 0 1 Arch~lMuzzle-27 28 28 Rifle- irst-46 0 0 0 0 0 0 11 11 39 15 15 17 Rifle-Second-47 0 0 0 0 0 17 32 0 0 0 9 12 9 12 Rifle- Third-48 0 0 0 0 21 0 0 0 0 0 6 6 6 Rifle-Late-29 0 0 0 69 52 56 69 125 Subtotals Q Q Q Q Q 1. Wound~Loss 0 2 2 2 Archery uzzle-24 0 0 0 0 0 2 4 0 0 0 1 1 2 3 Archery-52 0 0 1 1 0 0 1 3 1 Rifle-First-43 0 0 0 0 3 4 Rifle-Second-a-t 0 0 0 0 0 4 10 4 10 14 0 2 2 Rifle- Third-45 0 0 0 0 0 1 1 3 0 0 r 1 Rifle-Unk.Reg.-25 0 0 0 0 0 0 0 1 0 0 0 Rifle-Late-26 0 0 0 0 0 3 3 3 10 20 Subtotals 32 Q Q Q Q 11 £1 1 1 Illegal Hunting 0 5 Rifle-First-49 0 5 0 0 0 0 5 0 0 6 Rifle-Second-50 0 5 0 0 I 0 0 6 0 0 -. 0 0 Rifle-Third-51 1 0 0 0 0 1 0 1 0 0 0 1 0 0 Rifle-Unk.Reg.-9 0 0 0 0 1 1 0 I 0 0 0 0 1 0 1 Archery Season-8 0 0 Out-of-Season-7 0 0 0 1 0 2 0 3 3 0 0 0 12" Subtotals }. Q Q Q 11 1 £ £ 11 Presumed Hunting" Missing-ArchMuzz-21 0 0 1 0 0 0 0 0 0 1 1 3 3 Missing-Rifle Ist-40 0 0 0 0 0 0 3 3 6 Missing-Rifle 2nd-41 0 0 0 2 6 2 7 0 1 0 9 Missing-Rifle 3rd-42 0 0 0 0 0 0 0 0 0 0 0 Subtotals 10 16 Q Q Q Q .2 Q .2 .2 1 Totals 17 14 13 6 2 3 87 93 119 116 235 • These elk were illegally shot as spike-antlered yearling males: (173.190/93) (173.232/93) (173.309/93) (173.320/93) (173.919194) (174.059/94) (174.140/95) (174.200/95) (174.679/95) (174.729195) (174.861195) (173.210/96). (173.309/93) disappeared in 1994 during first rifle season when spike-antlered yearling males were not legal and was assumed to have been taken illegally. All other illegal deaths were confirmed. b These elk disappeared during hunting seasons and remained missing for several months and are assumed dead and legally harvested: (172.207193) (172.269193) (172.308193) (172.498193) (172.649/93) (172.800/93) (172.961/93) (172.991193) (173.390/93) (173.439193) (173.760/94) (174.001/94) (174.181/94) (174.770195) (174.929/96) (175.199/96). �Table 2. Survival rates, inclusive ofnonhunting and hunting mortalities, for winter-spring (WS, 1 December-14 June) and summer-fall (SF, 15 June-30 November) time periods from 1 December 1993-14 June 1999 for the cohort of calves age 6 months radio-collared in December 1993. Survival rate for male and female calves pooled when age 6-11 months was 0.92 (95% CI 0.86-0.98, n=73). Survival rates (S) calculated as a mean estimate of (alive)/(alive + dead) and variance S(1-S)/n collars. Elk Elk Age (months) and Time Period (WS or SF dates) 6-11 (mos) 12-17 18-23 24-29 30-35 36-41 42-47 48-53 54-59 60-65 66-71 6-71 SF WS SF WS SF WS SF WS SF WS ALL 1993-94 1994 .1994-95 1995 1995-96 1996 1996-97 1997 1997-98 1998 1998-99 1993-99 0.88 0.78 0.99 32 0 4b 0 4 1.00 0.76 0.99 28 0 0 0 0 0.21 1.00 0.06 0.37 4 0 0 0 0 0.50 0.00 0.00 4 22 0 2 1.00 0.00 1.00 2r 0 0 0 0 0.00 0.97 0.87 1.00 35 0 1.00 0.92 1.00 34 0 0 0 0 0.97 0.91 1.00 33 0 1 0 1 0.84 0.91 1.00 32 0 5 0 5 1.00 0.71 0.97 27 0 0 0 0 0.89 WS MALES Survival 0.89 L 95%CI U 95%CI n collars 36 Censored" Died 4 Nonhunt 4 Hoot 0 28< 0 224 0 22 1 0 1 0 1 0.00 33 1& 3-'& 33 4 29 FEMALES Survival 0.95 L95%CI U95%CI n collars 37 Censored" Died 2 Nonhunt 2 Hoot 0 Ih 0 1 0.97 34 0 1 0 1 27 0 31 0 3 0.96 0.76 1.00 24 0 1 1 0 0.87 0.87 1.00 23 0 31 0 3 1.00 0.72 1.00 18 0 0 0 0 0.51 0.34 0.69 35 21t 17 '3 14 2 • Censored denotes collar failure and/or animallifeldeath status not known. b Collar (173.309/93) disappeared 1994 hunting seasons and assumed dead. e Includes (173.241193) collar failure August 1995 but seen January 1996. d Collars (173.390/93),(173.439/93) disappeared 1995 hunting seasons and assumed dead. • Censored elk; (173.241193) failed August 1995, (173.381193) slipped collar December 1995. r Includes (173.340/93) alive May 1997. I Censored elk (173.249/93) slipped collar September 1997. k Collar (172.800/93) disappeared 1994 hunting seasons and assumed dead. i Collar (172.961193) disappeared 1997 hunting seasons and assumed dead. j Collar (172.991193) disappeared 1998 hunting seasons and assumed dead. k Censored elk (172.849/93) failed May 1999, (173.151193) failed February 1999. ..... .....) \0 �Table 3. Survival rates, inclusive ofnonhunting and hunting mortalities, for winter-spring (WS, 1 December-14 June) and summer-fall (SF, 15 June-30 November) time periods from 1 December 1994-14 June 1999 for the cohort of calves age 6 months radio-collared in December 1994. Survival rate for male and female calves pooled when age 6-11 months was 0.90 (95% CI 0.83-0.97, n=69). Survival rates (S) calculated as a mean estimate of (alive)/(alive + dead) and variance S(1-S)/n collars. Elk Age (months) and Time Period (WS or SF dates) 6-11 (mos) 12-17 WS SF 1994-95 1995 0.91 0.81 1.00 33b 0 3 3 0 0.90 0.79 1.00 30b 0 .0.97 0.78 0.99 32 0 1 0 1 0.97 0.91 1.00 30 0 1 1 0 18-23 ./ 24-29 30-35 36-41 42-47 48-53 54-59 6-59 WS SF WS SF WS SF WS ALL 1995-96 1996 1996-97 1997 1997-98 1998 1997-98 1994-98 MALES Survival L 95%CI U 95%CI n collars Censored" Died Nonhunt Hunt 3 0 3 0.96 0.88 1.00 26 1c 1 1 0 0.08 0.00 0.19 25 0 23 0 23 1.00 2d 0 0 0 0 0.50 0.00 1.00 2 0 1 0 1 0.93 0.90 1.00 29 0.96 0.83 1.00 27 0 1 0 1 0.85 0.89 1.00 26 0 4 0 ·4 1.00 0.70 0.99 22 0 0 0 0 1.00 0.00 0.00 1 0 0 0 0 0 18 0 0 0 31 2 31 4 27 0.64 1.00 0.42 0.85 14 0 0 0 0 FEMALES Survival 0.89 L 95%CI U 95%CI n collars 36 Censored" Died 4 Nonhunt 4 Hunt 0 I" 2 0 2 • Censored denotes collar failure and/or animallifeldeath status not known. Includes (173.949194) collar failure December 1994 but seen January 1996 • Censored elk:(173.949194) collar failure, which was killed in fJISt rifle season 1996. d Includes (174.030/94) alive June 1997. • Censored elk: (173.719194) slipped collar May 1996. f Collar (173.760/94) disappeared 1998 hunting seasons and assumed dead. • Censored elk: (174.030/94) slipped collar September 1998. b 22 0 gf 0 8 0.40 0.23 0.57 35 0 21 5 16 •...• 00 0 �Table 4. Survival rates, inclusive ofnonhunting and hunting mortalities, for winter-spring (WS, 1 December-14 June) and summer-fall (SF, 15 June-30 November) time periods from 1 December 1995-14 June 1999 for the cohort of calves age 6 months radio-collared in December 1995. Survival rate for male and female calves pooled when age 6-11 months was 0.88 in 1995 (95% CI 0.81-0.96, n=69). Survival rates (S) calculated as a mean estimate of (alive)/(alive+dead) and variance of S(I-S)/n collars. 6-11(mos} WS 1995-96 12-17 SF 19.96 Elk Age (months) and Time Period (WS or SF dates) 24-29 36-41 18-23 30-35 SF WS SF WS 1996-97 1997 1997-98 1998 MALES Survival L 95%CI U 95%CI n collars Censored" Died Nonhunt Hunt 0.86 0.74 0.98 35 2b 5 5 0 0.83 0.68 0.97 29 1c 5 0 5 0.96 0.87 1.00 24 0 1 1 0 0.23 0.04 0.41 22 4 Id I' 17 0 17 0 0 0 FEMALES Survival L 95%CI U 95%CI n collars Censored' Died Nonhunt Hunt 0.91 0.81 1.00 34 0 3 3 0 0.87 0.75 0.99 31 0 4 0 0.93 0.82 1.00 27 0 2 1 1 0.83 0.66 0.99 23 2f 4 0 4 1.00 4 1.00 18 I' 0 0 0 0.25 0.00 0.86 4 0 3 0 3 0.89 0.73 1.00 18 0 2 0 ·2 42-47 WS 1998-99 1.00 1 0 0 0 0 1.00 16 0 0 0 0 6-47 ALL 1995-99 0.03 0.00 0.10 32 5 31 6 25 0.52 0.33 0.70 .31 3 15 4 11 • Censored denotes collar failure and/or animal life/death status not known. Censored elk (174.619/95) slipped collar May 1996, (174.800/95) capture mortality. • Censored elk (174.660/95) slipped collar July 1996. d Censored elk (174.671195) likely collar failure July 1997. • Censored elk (174.190/95) slipped collar May 1998. r Censored elk (174.441195) slipped collar August 1997 and (174.580/95) collar failure July 1997. I Censored elk (174.470195) collar failure December 1997. . b .... 00 .... �Table 5. Survival rates, inclusive ofnonhunting and hunting mortalities, for winter-spring (WS, 1 December-14 June) and summer-fall (SF, 15 June-30 November) time periods from 1 December 1993-14 June 1999 for the cohort of calves age 6 months radio-collared in December 1996. Survival rate for male and female calves pooled when age 6-11 months was 0.86 (95% CI 0.81-0.96, n=69). Survival rates (S) calculated as a mean estimate of (alive)/(alive+dead) and variance ofS(I-S)/n collars. Elk Age (months) and Time Period (WS or SF dates) 6-11 (mos) 12-17 18-23 24-29 WS SF WS SF 1996-97 1997 1997-98 1998 1998-99 1996-98 0.85 0.73 0.98 34 0.97 0.90 1.00 29 0 1 0 1 1.00 1.00 28 0 0 0 0 0.36 0.17 0.54 28 0 184 0 18 0.29 0.14 0.45 34 1 24 1.00 0.80 30-35 6-35 WS ALL MALES Survival L 95%CI U 95%CI n collars Censored' Died Nonhunt HWlt Ib 5 5 0 10 0 0 0 0 5 19 FEMALES Survival 0.86 L 95%CI U 95%CI n collars 35 Censored' Died 5 Nonhunt 5 Hunt 0 1.00 0.74 0.98 30 0 0 0 0 30 0 0 0 0 30 0 6< 0 6 • Censored denotes collar failure and/or animallifeldeath status not known. Censored elk. (174.619/96) capture mortality. C Collar (174.929/96) disappeared 1998 hunting seasons and assumed dead. d Collar (175.199/96) disappeared 1998 hunting seasons and assumed dead. b 1.00 0.65 0.95 24 0 0 0 0 0.69 0.53 0.85 35 0 11 5 6 o •... 00 N �Table 6. Survival rates, inclusive ofnonhunting and hunting mortalities, for winter-spring (WS 1 December-14 June) and summer-fall (SF, 15 June30 November) time periods from 1 December 1993 - 14 June 1999 for all radio-collared adult female elk age 2:,12months. Survival rates (S) calculated as a mean estimate of (alive)/(alive + dead) and variance S(1-S)/n collars. Adult WS Adult SF Adult WS 1993-94 1994 1994-95 Elk Age and Time Period (WS or SF dates) Adult Adult Adult Adult Adult SF WS SF WS SF 1995 1995-96 1996 1996-97 .1 Survival L 95%CI U 9S%CI n collars Censored" Died Nonhunt Hunt 0.96 0.91 1.00 68b• 0 0.87 0.80 0.94 100bf 0 3 13< 1 2 I 12 0.96 0.92 1.00 9Sbl 0 4 1 3 • Censored denotes collar failure or life/death status unknown. Includes (172.011/93) collar failure April 1994, seen January 1996 , Collars (172.800/93,172.649/93) disappeared 1994 hunt seasons. d Collar (172.207/93) disappeared 1995 hunt seasons, assumed dead. • Composed of 6-18+ and 62-30+ months old females. f Composed of35-12+, 6-24+, and 59-36+ months old females. • Composed of34-18+ and 61-30+ months old females. b Composed of32-12+, 34-24+, and 63-36+ months old females. I Composed of30-18+ and 89-30+ months old females. J Censored elk (172.011/93) failure and (173.719/94) slipped collar. k Composed of31-12+, 29-24+, and 83-36+ months old females. b 0.94 0.90 0.98 129bh 0 8d 0 8 0.94 0.90 0.98 119i 2) 7 4 3 0.89 0.84 0.94 1431: 0 16 1 15 0.98 0.95 1.00 1271 0 3 1 2 1997 0.91· 0.87 0.96 152m 2D 13 0 13 Adult WS Adult SF Adult WS 1997-98 1998 1998-99 0.99 0.97 1.00 138· IP 2 1 1 0.77 0.70 0.84 136<1 0 31' 0 31 0.98 0.95 1.00 103' 21 2 2 0 I Composed of27-18+ and 100-30+ month old females . ••Composed of30-12+, 23-24+, and 99-36+ months old females. • Censored elk (174.441/95) slipped collar August 1997 and (174.S80/95) likely failure June 1997. • Composed of30-18+ and 108-30+ months old females . P Censored elk (174.470/95) likely collar failure August 1997. q Composed of30-24+ and 106-36+ months old females. r Collars (172.269/93,172.308/93,172.498/93,172.991/93, 173,760/94, 174.929/96) disappeared 1998 hunt seasons and assumed dead. • Composed of24-30+ and 81-42+ months old females. I Censored elk (172.849/93, 173.151/93) failed collars. ,_. OQ W �186 APPENDIX I. Mortalities of235 radio-collared elk from 1 December 1993 through 14 June 1999. Age is aQQfoximate age in ~ears of elk at death: C=calf age 6-11 monthsz Y=yearling age 12-23 months. Frequency ID! Death TraQ Year caEtured Site Zone Sex Age Date Cause of death & code number 172.030/93 172.050/93 172.039/93 172.070/93 172.080/93 172.090/93 172.090/94 172.101193 172.128/93 172.139/93 172.160/93 172.181193 172.201193 172.207/93 172.229/93 172.258/93 172.269/93 172.277/93 172.290/93 172.290/94 172.300193 172.308/93 172.369/93 172.369/94 172.409/93 172.448/93 172.459/93 172.498/93 172.509/93 172.519/93 172.530/93 172.542/93 172.542/94 172.549/93 172.570/93 172.581193 172.581194 172.590/93 172.610/93 172.620/93 172.639/93 172.649/93 172.658/93 172.670/93 172.678/93 172.690/93 172.699/93 172.730/95 172.749/95 172.758/95 172.800/93 172.821/93 172.890/93 172.899/93 172.950/93 172.961193 172.991193 173.000/93 BR EG GR BR GM GR SM SR GC BC CC MC GS SG MC HY MC GS SG SM AC GH AC FM AC ·MH MH FS FS FS FS FS FM PG PG PG FM PG PG PG PG PG PG PG PG FM FM LS LS LS SR EG BC BC GH MG SG WM A B B A A B D B B C C C C D C C C C D D E D E B E E E H H H H H B H H H B H H H H H H H H B B H H H B B C C D D D G F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F 5 12 4 11 6 11 11 8 8 17 3 3 3 4 8 3 12 3 6 4 8 8 3 4 8 12 8 12 3 6 8 6 15 7 16 3 3 5 3 5 16 2 5 4 9 8 7 19 11 15 Y 3 3 C 2 4 5 C 27-Oct-95 31-Oct-98 14-Jun-95 12-Feb-96 05-Nov-94 16-Jan-94 12-Oct-98 14-Oct-96 31-Oct-96 19-Oct-95 23-Oct-94 03-Nov-94 15-Nov-94 30-Nov-95 29-Oct-98 04-0ct-94 29-Oct-98 O4-Oct-94 23-Oct-94 16-Sep-96 30-0ct-97 29-Oct-98 18~94 14-Jan-96 29-Dec-94 . 01-Nov-98 24-Oct-96 15-Oct-98 21-Oct-95 12-Oct-98 29-Oct-98 01-Feb-94 06-Jan-99 28-Nov-95 03-Nov-94 17-Oct-94 08-Dec-95 i4-Apr-96 O4-Oct-94 04-Nov-98 14-Jun-96 30-Nov-94 02-Nov-97 16-Jan-95 22-Dec-94 24-Jan-94 05-Nov-95 05-May-99 07-Aug-96 28-Oct-98 30-Nov-94 08-Sep-96 06-Nov-96 18-Mar-94 18-Oct-95 30-Oct-97 29-Oct-98 25-Apr-94 Legal kill second rifle season-47 Legal kill third rifle season-48 Unknown-suspect predation-30 Unknown-l l Legal kill third rifle ~n-48 Wounding loss late rifle season-26 Legal kill first rifle season-46 Legal kill first rifle season-46 Wounding loss second rifle season-44 Wounding loss first rifle season-43 Legal kill second rifle season-47 Wounding loss second rifle season-44 Wounding loss second rifle season-44 Disappear first rifle season-40 Legal kill second rifle season-47 Legal kill archerylmuzzle season-27 Disappear second rifle season-41 Wounding loss archery/muzzle-24 Legal kill second rifle season-47 Legal kill muzzleloading season-34 Wounding loss second rifle season-44 Disappear second rifle season-41 Legal kill archery season-33 Legal kill late rifle season-29 Wounding loss late rifle season-26 Legal kill third rifle season-48 Legal kill second rifle season-47 Disappear [JISt rifle season-40 Legal kill second rifle season-47 Legal kill first rifle season-46 Wounding loss second rifle season-44 Mountain lion predation-3 Accident-irrigation ditch-If Legal kill late rifle season-29 Wounding loss second rifle season-44 Legal kill first rifle season-46 Legal kill late rifle season-29 Unknown-11 Accident. birthinglca1ving-32 Legal kill third rifle season-48 Accident. birthinglca1ving-32 Disappear first rifle season-40 Legal kill third rifle season-48 Legal kill late rifle season-29 Wounding loss late rifle season-26 megal kill-7 Legal kill third rifle season-48 Malnutrition-6 Unknown-suspect predation-30 Wounding loss second rifle season-44 Disappear second rifle season-41 Legal kill archery season-33 Legal kill third rifle season-48 Mountain lion predation-3 Legal kill first rifle season-46 Disappear second rifle season-41 Disappear second rifle season-41 . MaInutrition-6 --------------------------------------------------------------------------------------------- �187 Appendix I. (continued) Frequency IDI Trap Year captured Site Zone 173.000/94 173.010/93 173.041193 173.050/93 173.060/93 173.070/93 173.081193 173.090/93 173.100/93 173.120/93 173.140/93 173.160/93 173.171193 173.190/93 173.190/96 173.201193 173.210/93 173.210/96 173.219/93 173.219/96 . 173.232/93 173.232/96 173.262193 173.262/94 173.269/93 . .173.279/93 173.289/93 173.289/94 173.300/93 173.300/96 173.309/93 173.320/93 173.320/94 173.332/93 173.332/96 173.340/93 173.351193 173.351196 173.359/93 173.359/96 173.370/93 173.370/96 173.381196 173.390/93 173.402/93 173.410/93 173.410/96 173.420/93 173.429/93 173.439/93 173.450/93 173.461/93 173.461194 173.461196 173.469/93 173.469/94 173.479/93 173.479/96 173.492/93 HM WM GM MH MH MH FS FS FS GM PG FS FM BC BC GR GC LM BC CG BR BC BC HM MC MC MC PR GS MC HY HY HM GH BC SG SG BG AC MC MH RG RG MG MG WM EW WM MH PG PG PG CM GM FS PR FS RG FS B G A E E E H H H A H H B C C B B D C D AMY C C B C C C A C C C C B D C D D E E C E E E D D G G G E H H H AMY A H A H E H Death Sex· Age F F M F F F F F F M F F F M M M M M M M 4 4 2 5 2 5 3 4 4 2 3 3 5 Y 2 2 2 2 2 2 M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M M 2 C 2 2 3 C 2 2 2 Y Y 2 2 2 4 2 2 2 2 2 C C 2 2 2 1 2 2 2 2 C M M M M M M 2 C 2 2 2 2 Date 31-Oct-98 07-Jan-98 19-Oct-95 31-Oct-98 27-Dec-95 03-Mar-98 22-Oct-96 24-Oct-97 30-0ct-97 14-Oct-95 31-Oct-96 13-Nov-96 10-Oct-98 29-Oct-94 1O-Oct-98 30-Nov-95 19-Oct-95 30-Oct-97 06-Nov-95 21-Oct-98 15-Nov-94 31-Oct-98 18-Mar-94 12-Oct-96 06-Nov-95 12-Oct-96 22-Mar-94 18-Sep-96 24-Oct-95 10-Oct-98 30-Nov-94 20-Oct-94 14-Sep-96 12-Sep-95 10-Oct-98 20-Oct-97 05-Nov-95 27-Oct-98 06-Sep-95 09-Nov-98 08-Nov-95 08-Jan-97 19-Feb-97 30-Nov-95 18-Oct-95 02-Nov-95 29-Sep-98 24-Oct-95 14-Sep-95 30-Nov-95 15-Oct-95 07-Feb-94 19-5ep-95 22-Oct-98 25-Apr-94 20-Oct-96 1O-Sep-95 l7-Oct-98 03-Sep-95 Cause of death & code number Legalkill third rifle season-48 Accident-fell-l0 Legalkill first rifle season-46 Legalkill third rifle season-48 Legalkill late rifle season-29 Illegalkill-7 Legalkill secondrifle season-47 Legalkill secondrifle season-47 Legalkill secondrifle season-47 Legalkill first rifle season-46 WOWldinglosssecondrifleseason-44 Legalkill third rifle season-48 Legalkill first rifle season-46 megal kill secondrifle season-50 Legalkill first rifle season-46 Woundingloss third rifle season-45 Legalkill first rifle season-46 Illegalkill archery/muzz.season-8 Legalkill third rifle season-48 Legalkill secondrifle season-47 illegal kill secondrifle season-50 Legalkill third rifle season-48 Unknown-ll Legalkill first rifle season-46 Legalkill third rifle season-48 Legalkill first rifle season-46 Unknown-ll Legalkill muzzleloadingseason-34 Legalkill secondrifle season-47 Legalkill first rifle season-46 Illegalkill first rifle season-49 megal kill first rifle season-49 Legalkill archeryseason-33 Legalkill muzzleloadingseason-34 Legalkill first rifle season-46 Legalkill secondrifle season-47 Legalkill third rifle season-48 Legalkill secondrifle season-47 Legalkill archeryseason-33 Woundingloss third rifle season-45 Legalkill first rifle season-46 Mountainlion predation-3 Unknown-suspectpredation-30 Disappearfirst rifle season-40 Legalkill first rifle season-46 Woundingloss secondrifle season-44 Woundingloss archery/muzzleseasons-24 Legalkill secondrifle season-47 Legalkill archeryseason-33 Disappearfirst rifle season-40 Legalkill first rifle season-46 Malnutrition-6 Woundingloss archeryseason-50 Legalkill secondrifle season-47 Malnutrition-6 Legalkill secondrifle season-47 Legalkill archeryseason-33 Legalkill secondrifle season-47 Legalkill archeryseason-33 !~~~~~~]--------~~-----~-------~----]----7--]]~~~!:?~-~g~!~~E:~~~~:_~~~~ _ �188 Appendix I. (continued) Frequency ID! Year Captured Trap. Site Zone 173.510/93 FM FM FM 173.521193 173.530/94 173.540/94 173.549/94 173.569/94 173.580/94 173.589/94 173.610/94 173.640/94 173.640/96 173.649/94 173.659/94 173.679/94 173.689/94 173.719/96 173.739/94 173.750/94 173.760194 173.789/94 173.819/94 173.829/94 173.840/94 173.849194 173.859/94 173.870/94 173.919/94 173.929/94 173.939/94 173.959/94 173.970/94 173.981/94 173.990/94 174.001194 174.009194 174.019/94 174.039/94 174.049194 174.059/94 174.069/94 174.080/94 174.090/94 174.101/94 174.109/94 174.119/94 174.119/95 174.129/94 174.140194 174.140/95 174.150/94 174.160/94 174.170/94 174.170/96 174.181194 174.200/95 174.210/95 174.220/95 174.230/95 174.230/96 00 HM PR PR 00 BC MC GM BC MC MP BG MC BG BG MR MR MM MM MM SM SM SM BC BC BC BC MC MP BC BG BG BG MR MM MR BG MR MR MM MM MM GB MM MM GB MM MM FM MC FM MS MS MS MS EW ~~~~~~2~ ~ Death Sex Age B B B B B A A B C C A C CF4 D E C E E E E F F F D D D C C C C C D C E E E E F E E E E F F FMC B F FMC B F F B C B F F FMC FMC G M ~ ~ Date Cause of Death & Code Number 2 28-Aug-95 17-Oct-95 31-Oct-98 13-Oct-% 07-Sep-95 l6-Oct-97 21-Dec-95 01-Jun-95 14-Nov-97 30-Mar-95 16-Mar-97 1l-Oct-97 25-Oct-98 29-Oct-98 31-Oct-% 03-Nov-98 29-Oct-98 18-Oct-98 29-Oct-98 20-Mar-95 30-Oct-97 IO-Jan-97 29-Sep-98 20-Oct-98 23-Oct-96 21-Feb-95 02-Nov-95 l6-Oct-96 18-Sep-96 02-Nov-96 20-Oct-96 02-Nov-96 20-Oct-96 30-Nov-% 12-Oct-96 13-Nov-96 05-Sep-96 14-Sep-96 17-Oct-95 26-Sep-96 ll-Oct-97 20-Oct-96 24-May-96 19-Oct-96 01-Mar-95 30-Nov-97 14-Oct-96 14-Mar-95 30-Nov-96 14-Oct-96 15-Sep-96 25-Apr-95 10-Oct-98 30-Nov-96 18-Oct-96 ll-Oct-97 08-Apr-96 24-Apr-96 l3-Oct-98 Legal kill archeryseason-33 Legal kill first rifle season-46 Legal kill third rifle season-48 Legal kill first rifle season-46 Legal kill archeryseason-33 Woundingloss first rifle season-43 Unknown-ll Unknown-suspectpredation-30 Woundingloss third rifle season-45 Unknown-suspectpredation-30 Malnutrition-6 Legal kill first rifle season-46 Legal kill secondrifle season-47 Woundingloss first rifle season-43 Legalkillfrrstrifleseason-46 Legal kill third rifle season-48 Woundingloss secondrifle season-44 . Legal kill secondrifle season-47 Disappear secondrifle season-41 Unknown-suspectmalnutrition-31 Legal kill secondrifle season-47 Legal kill late rifle season-29 Woundingloss archery/muzzleseasons-24 Legal kill secondrifle season-47 Legal kill secondrifle season-47 Mountain lion predation-3 illegal kill secondrifle season-50 Legal kill first rifle season-46 Legal kill archeryseason-33 Legal kill third rifle season-48 Legal kill secondrifle season-47 Legal kill third rifle season-48 Legal kill secondrifle season-47 Disappeararchery/muzzleseason-21 Legal kill first rifle season-47 illegaI kill rifle season-9 Legal kill archeryseason-33 Legal kill muzzleloadingseason-34 illegal kill first rifle season-49 Woundingloss archery/muzzle-24 Legal kill first rifle season-46 Legal kill secondrifle season-47 Unknown-suspectpredation-30 Legal kill secondrifle season-47 Mountain lion predation-3 Woundingloss rifle season-25 Legal kill first rifle season-46 Mountainlionpredation-3 llIegalkillthirdrifleseason-51 Legal kill first rifle season-46 Legal kill muzzleloadingseason-34 Mountain lion predation-3 Legal kill first rifle season-46 Disappear secondrifle season-41 llIegaIkill first rifle season-49 Legal kill first rifle season-46 Unknown-suspectmalnutrition-31 Mountain lion predation-3 Legal kill first rifle season-46 ~ ~I:~~~~~ ~~~~~~~~P!~~~:~ M M F M F F F F F F F F 2 2 7 2 Y 3 Y C 3 C C 3 F F F F F F F F F F F F F M M M M M M M M M M M M M M M M M M 4 2 2 4 4 4 C 3 2 4 4 2 C Y 2 2 2 2 2 2 2 2 2 2 2 Y 2 3 2 Y 2 M M 2 2 M M M M M M M M Y 2 2 C 2 2 Y 2 _ �189 AEEendix I. {continued} Frequency ID! Tral! Site Zone Year caEtured 174.240/96 SR B MD 174.301195 E MS 174.319/95 F MD 174.329/95 E MD 174.339/95 E 174.349/95 MD E 174.360/95 MD E 174.401/95 AP D 174.420/95 AP D 174.478/95 WD e 174.491195 LB e 174.500/95 LB e 174.520/95 KR B 174.520/96 GM A 174.551/95 we G 174.560/95 BL A 174.609/95 GB B MD 174.629/95 E 174.639/95 MD E 174.650/95 HG E 174.679/95 HG E 174.689/95 AP D 174.689/96 EW G 174.700/95 AP D 174.710/95 AP D 174.729/95 VM D 174.740/95 VM D 174.750/95 VM D 174.760/95 VM D 174.770/95 WD e 174.780/95 WD e 174.789/95 WD e 174.789/96 EW G 174.800/96 BG E WD 174.809/95 e 174.820/95 WD e 174.830/95 LB e 174.841195 LB e 174.851195 LB e 174.861195 KR B 174.870/95 LB e 174.880/95 KR B 174.889/95 we G 174.899/95 BL A 174.910/96 LM D 174.929/96 LM D 174.959/96 LM D 174.998/96 BG E 175.059/96 Be e 175.130196 GM A 175.139/96 GM A 175.149/96 SR B 175.160/96 Be e 175.170196 LM D 175.189/96 RG E EW 175.199/96 G 175.209/96 MS F END Death Sex Age M 2 F 3 F Y F Y F e 2 F F Y F 2 F 2 2 F 2 F e F F Y F e 3 F F Y e F 2 M M 2 M 3 M Y e M e M M 2 M 2 M Y M 2 M 2 M 3 M 2 2 M e M 2 M M e y M 2 M M 2 3 M M 2 Y M M 2 2 M 2 M 2 M F e 2 F F 2 F 2 F e F e 2 F 2 F 2 M e M 2 M M 2 2 M Date 12-Sep-98 20-Sep-98 02-Oct-% 03-Sep-96 27-Mar-96 l3-Oct-97 22-Apr-97 22-Sep-% 1l-Oct-97 13-Sep-97 10-Sep-97 16-Apr-96 21-Sep-96 25-Apr-97 14-Oct-98 14-May-97 15-Apr-96 11-Oct-97 21-Oct-97 14-Sep-98 13-Nov-96 05-Feb-96 14-May-97 16-Oct-97 15-Oct-97 17-Oct-% 08-Nov-97 05-Nov-97 15-Sep-98 16-Oct-97 II-Oct-97 22-May-96 28-Oct-98 09-Apr-97 22-Apr-97 14-Oct-97 l3-Oct-97 12-Sep-98 22-Oct-97 31-Oct-96 30-Oct-97 11-Oct-97 20-Oct-97 30-Oct-97 27-Feb-97 29-Oct-98 29-Oct-98 21-Oct-98 13-Feb-97 24-Mar-97 28-Oct-98 19-Oct-98 17-Sep-98 26-Mar-97 l3-Oct-98 29-Oct-98 24-Oct-98 Cause of death & code number Legal kill muzzleloading season-34 Legal kill archery season-33 Wounding loss archery season-52 Legal kill archery season-33 Unknown predator-S Legal kill first rifle Season-46 Unknown-suspect predation-30 Legal kill muzzleloading season-34 Legal kill first rifle season-46 Legal kill muzzleloading season-34 Wounding loss archery season-52 Unknown-suspect predation-30 Legal kill muzzleloading season-34 Mountain lion predation-3 Legal kill first rifle season-46 Illegal kill-7 Unknown-suspect predation-30 Legal kill first rifle season-46 Legal kill second rifle season-47 Legal kill muzzleloading season-34 Illegal kill second rifle season-50 .Mountain lion predation-3 Unknown-suspect maInutrition-31 Wounding loss first rifle season-43 Legal kill first rifle season-46 Illegal kill first rifle season-49 Legal kill third rifle season-48 Legal kill third rifle season-48 Legal kill archery season-33 Disappear first rifle season-40 Legal kill first rifle season-46 Unknown-suspect predation-30 Wounding loss second rifle season-44 Mountain lion predation-S Accident, fell-lO Legal kill first rifle season-46 Legal kill first rifle season-46 Legal kill muzzleloading season-34 Illegal kill second rifle season-50 Illegal kill second rifle season-50 Wounding loss second rifle season-44 Legal kill first rifle season-46 Legal kill second rifle season-47 Wounding loss second rifle season-44 Unknown-suspect predation-30 Disappear second rifle season-41 Wounding loss second rifle season-44 Legal kill second rifle season-47 Unknown-l l Mountain lion predation-3 Legal kill second rifle season-47 Legal kill second rifle season-47 Legal kill muzzleloading season-34 Unknown-suspect maInutrition-31 Legal kill first rifle season-46 Disappear second rifle season-41 Legal kill second rifle season-47 �190 Appendix Il, Abstract from paper presented at Western States and Provinces Elk and Mule Deer Workshop, Salt Lake City, Utah, March 1999. ESTIMATING ELK DENSITIES IN COLORADO: PROGRESS WITH PERPLEXITIES DAVID J. FREDDY, Colorado Division of Wildlife, Research Center, 317 West Prospect Road, Fort Collins, CO, USA 80526 DAVID C. BOWDEN, Department of Statistics, Colorado State University, Fort Collins, CO 80523 USA GARY C. WIllTE, Department of Fishery and Wildlife Biology, Colorado State Univeristy, Fort Collins, CO 80523 USA Abstract: During winters 1994-1998, we evaluated helicopter survey methodologies to estimate elk densities injuniper-pinyon (Juniperus osteosperma-Pinus edulis) and oakbrush-mountain shrub (Quercus gambelii-Amelanchier alnifolia) habitats. We developed sighting bias correction models to correct for groups of elk not observed when counting elk on 2.59-km2 (l-mf) quadrats. Within a 350-km2 area, we compared elk densities obtained from a stratified random quadrat sampling system corrected for sighting bias with independent estimates of densities obtained from mark-resight models using known numbers of radio-collared elk (120-137 elk) within the quadrat survey area. Although observers detected about 80% of the elk groups on quadrats during sighting bias trials and resulting sighting bias models increased estimated elk densities on quadrats by 10-12%, quadrat densities were 2:,25%lower than mark-resight densities. Estimated densities of elk ranged from 6-10 elk/krrr', We propose that errors in counting numbers of elk in a group may contribute more to underestimating elk densities than errors in detecting groups of elk. �191 Colorado Division of Wildlife Wildlife Research Report July 1999 JOB FINAL REPORT Smteof ~C~o~lo~r~ad~o~ _ Cost Center 3430 Mammals Research Project No. ~W.;_-..::.1=53=_-=R:....-.:..::12::.__ _ Elk Conservation Work Package __ ----'3~0:;..:0=2'_____'_ _ T~kNo. 2=_ _ Elk Movements in Response to Early-season Hunting in the White River Area .. Period Covered: July 1, 1998 - June 30, 1999 Author: M. M. Conner Personnel: G. White, D. Freddy, R Kahn, 1.Maddison, J. Ellenberger ABSTRACT Understanding causes of elk movement to private land is important to wildlife managers who use hunting on public land as a population management tool. In the White River area of northwest Colorado, late-summer elk movements onto private lands made it difficult to harvest enough elk to maintain specified population levels. I conducted a 2-year field experiment to determine if early-season hunting (archery and muzzleloading) caused elk movement to private land during late summer. The study area was split into north and south treatment areas: each area received both an early- and lateopening treatment. Early-opening treatment was archery season that opened 1 week earlier than the historical opening, and late-opening treatment was an archery season that opened 2 weeks later, . yielding a 23-week difference in opening dates. The 80-88 adult female elk used in this study were captured from random locations throughout the study area I relocated radiocollared elk approximately 2 times per week for a 3-month period surrounding early- and late-opening dates each year. Using a Geographical Information System (GIS) generated map, I classified each elk location as being on public or private land. Elk receiving the late-opening treatment moved 10 days later than elk receiving the early treatment (P = 0.013, one sided ANOVA). Because there appeared to be an interaction between area and treatment, I performed a post hoc analysis separately by area In the north area, elk receiving the late-opening treatment moved 14 days later than elk receiving the early treatment (P + 0.006), one sided r-test), compared to a 6-day difference for elk in the south area (P = 0.218). Approximately twice as many radiocollared elk moved to. private land on the north area (44%) compared to the south area (23%). The experimental effect on the number of elk directly influenced by early-season hunting can be estimated by the abrupt increase in the proportion of elk moving to private land when hunting opened. Proportion of elk on private land increased 8-18% at the opening of early season, The greater response of elk to hunter activity on the north area may be due to topographical and migration differences between areas. The south area had densely forested canyons and cliffs, inaccessible to motor vehicles, which provided elk good refuge from hunters on public land. In �192 contrast, the north area offered less shelter on public land, but had refuge in large private-land tracts that bordered the area During the study period, elk moving to private land in the north were just beginning their fall migration, whereas elk in the south area were completing their migration by moving to private land. Elk on the north area that move onto private land do not risk depleting their winter forage, whileelk on the south area may degrade their winter range by grazing an extra couple of months on their wintering grounds. Colorado Division of Wildlife surveyed 34% and 37% of hunters using the study area during a post-hunt telephone survey in 1996 and 1997 to estimate the number of hunters afield per day with a maximum 95% Clof ±150 hunters. Daily use varied between 115-1,130 hunters per treatment area, with corresponding hunter densities of 0.006-0.51 hunters/knr', The proportion of elk on private land was not strongly affected by hunter density. Domestic sheep were accused of causing elk movements to private land during early-season hunting. I collected a location for at least one band of sheep per flight. From this data, I used nonparametric statistics and bootstrapping techniques to calculate the probability that elk avoided sheep at 3 spatial scales. Locations of elk were random with respect to sheep when all elk were used in analyses (P> 0.342), and when only elk within 1km of a sheep band were used (P> 0.912). Although elk may avoid sheep at <1 km, it is unlikely that they avoid sheepat large enough distances to cause widespread elk movements to private land. Although early-season opening may not be the sole cause of elk movements to private land, the experimental results show that early-season hunting caused some elk to move to private land, especially on the north area The increase of elk on private land directly attributable. to opening of hunting was 8-18%. Hence, the CDOW can reduce, at most, approximately 20% of elk movement to private land by manipulating early-season hunting. However, if elk movements are concentrated in problem areas, then manipulation of opening date may result in a proportionally higher reduction of elk movements in those areas. Managers concerned with reducing elk movement to private land should consider the public and private land refuges available to elk, as well as historical elk migration patterns. �193 ELK MOVEMENT IN RESPONSE TO EARLY-SEASON HUNTING IN THE WHITE RIVER AREA Mary M. Conner P. N. OBJECTIVES 1. Test whether early-season hunting causes movement of elk from areas of heavy hunting pressure to areas of lower hunting pressure. SEGMENT NARRATIVES 1. Estimate the mean date of movement of elk from non-refuge areas, and determine whether the timing is equivalent to the opening of archery season. 2. Evaluate alternative hypotheses about causes of elk movement such as livestock grazing, woodcutting, recreationalists, weather or forage quality. .. ..3. Publish analyses of elk movement in response to archery hunting in peer-reviewed scientific journals. INTRODUCTION Following is the most recent available draft of the Ph.D. Thesis summarizing results of research activities directed toward accomplishment of Program Narrative and Segment Narrative Objectives. ��195 DISSERTATION ELK MOVEMENT IN RESPONSE TO EARLY-SEASON HUNTING IN THE WHITE RIVER AREA, COLORADO Submitted by Mary M. Conner Department of Fishery and Wildlife Biology In partial fulfillment of the requirements for the Degree of Doctor of Philosophy Colorado State University Fort Collins, Colorado Fall 1999 �196 COLORADO STATE UNIVERSITY June 28, 1999 WE HEREBY RECOMMEND THAT THE DISSERTATION BY MARY M. CONNER ENTITLED ELK MOVEMENTS PREPARED UNDER OUR SUPERVISION IN RESPONSE TO EARLY-SEASON THE WHITE RIVER AREA BE ACCEPTED AS FULFILLING IN PART REQUIREMENTS DEGREE OF DOCTOR OF PIIll.OSOPHY. Committee on Graduate Work Advisor Department Head HUNTING IN FOR THE �197 ABSTRACT OF DISSERTATION ELK MOVEMENT IN RESPONSE TO EARLY-SEASON HUNTING IN THE WHITE RIVER AREA, COLORADO Understanding causes of elk movement to private land is important to wildlife managers who use hunting on public land as a population management tool. During the 1980s, late-summer elk movements onto private land in the White River area of northwest Colorado made it difficult to harvest enough elk to maintain specified population levels. I conducted a 2-year field experiment to determine if early-season hunting (archery and muzzleloading) caused elk movement to private land during late summer. The study area was split into north and south treatment areas; each area received both an early- and late-opening treatment. Early-opening treatment was an archery season that opened 1 week earlier than the historical opening, and late-opening treatment was an archery season that opened 2 weeks later, yielding a 3-week difference in opening dates. The 80-88 adult female elk used in this study were captured from random locations throughout the study area. I relocated radiocollared elk approximately 2 times per week for a 3-month period surrounding early- and late-opening dates each year. Using a Geographical Information System (GIS) generated map, I classified each elk location as being on public or private land. Elk receiving the late-opening treatment moved 10 days later than elk receiving the early treatment (P = 0.013, one-sided ANOVA). Because elk on the north area appeared to respond differently to treatment than elk on the south area, I performed a posl hoc analysis separately by area. In the north area, elk receiving the lateopening treatment moved 14 days later than elk receiving the early treatment (P = 0.006, one-sided r-test), compared to a 6-day difference for elk in the south area (P = 0.218). Approximately twice as many radiocollared elk moved to private land on the north area (44%) compared to the south area (23%). The experimental effect on the number of elk directly influenced by early-season hunting can be estimated by the abrupt increase in the proportion of elk moving to private land when hunting opened. Proportion of elk on private land increased 8-18% at the opening of early season. The greater response of elk to hunter activity on the north area may be due to topographical and migration differences between areas. The south area had densely forested canyons and cliffs, inaccessible to motor vehicles, which provided elk good refuge from hunters on public land. In contrast, the north area offered less shelter on public land, but had refuge in large private-land tracts that bordered the area. During the study period, elk moving to private land in the north area were just beginning their fall migration, whereas elk in the south area were completing their migration by moving to private land. Elk on the north area that move onto private land do not risk depleting their winter forage, while elk on the south area may degrade their winter range by grazing an extra couple months on their wintering grounds. Colorado Division of Wildlife surveyed 34% and 37% of early-season hunters using the study area during a post-hunt telephone survey in 1996 and 1997. The goal was to survey enough hunters to estimate, during the study period, the number of hunters afield per day with a maximum width 95% CI of±l50 hunters. Daily use varied between 115-1,130 hunters per treatment area, with corresponding hunter densities of 0.06-0.51 hunters/km', The proportion of elk on private land was not affected by hunter density. Elk in the White River area responded more to the opening of hunting season than to hunter density. It may be that hunter density must surpass or drop below a threshold before elk begin to respond directly to hunters, and hunter densities did not cross any threshold during the study. If hunter density continues to be a suspected cause of increased elk movement to private land, then an experiment manipulating hunter density should be conducted. Domestic sheep also were considered to cause elk movements to prisate land during early-season hunting. I collected a location for at least one band of sheep per flight. From these data, I used non-parametric statistics and randomization techniques to calculate the probability that elk avoided sheep at 3 spatial scales. Locations of elk were random with respect to sheep when all elk were used in analyses (P = 0.737 in 1996, P = 0.199 in 1997), when only elk within 5 km ofa sheep band were used (P = 0.342 in 1996, P = 0.990 in 1997), and when only elk within 1 km ofa sheep band were used (P> 0.969 in 1996, P = 0.912 in 1997). Although elk may avoid sheep at �198 <1 Ian, it is unlikely that they avoid sheep at large enough distances to cause widespread elk movements to private land. Although early-season opening may not be the sole cause of elk movements to private land, the experimental results show that early-season hunting caused some elk to move to private land, especially on the north area. The direct effect of hunting season is reflected in the 8-18% increase in proportion of elk on private land that occurred on opening day. Hence, in the White River area, the CDOW can reduce, at most, approximately 20% of elk movement to private land by manipulating early-season hunting. However, if elk movements are concentrated in problem areas, then manipulation of opening date may result in a proportionally higher reduction of elk movements in those areas. The elk not affected by opening of hunting season may be elk for which private land is enroute to their wintering area. Late summer movement onto private land may have become part of a learned migration behavior, changing hunt season opening and/or hunter density may have little effect on these elk. Future experiments and management actions should consider an early hunting season on private land, where problems occur, to retain elk on public land during late summer. Mary M. Conner Department of Fishery and Wildlife Biology Colorado State University Fort Collins, CO 80523 Fa1l1999 �199 TABLE OF CONTENTS ABSTRACT OF DISSERTATION ACKNOWLEDGMENTS INTRODUCTION . . . ELK MOVEMENTS IN RESPONSE TO DISTURBANCE Elk Movement in Response to Human Activities Elk Movement in Response to Hunting . , . . PROJECT OVERVIEW . LITERATURE CITED ....•.....................................................•.......... FIGURE .........................................................................•.... CHAPTER 1: ELK MOVEMENT COLORADO INTRODUCTION IN RESPONSE TO EARLY-SEASON HUNTING IN NORTHWEST . ...........•.......................................................•.... STUDY AREA ••.•..........................................................•..•........ METHODS ................•.......••...••........••.•.•.......................•....... Experimental design Data collection Data analysis REsULTS ........................ . ~.. . . . . . . . . . . . . . . . . . . . ••••....•.•••••.••••...•••.•..........••••••.••..•............•...•.••.••..•. DISCUSSION .••.•.•.••.••.••..•........••..........••.......................•...•...... MANAGEMENT IMPLICATIONS .............••..........................••...••........•.•.. LITERATURE CITED ....................••..•.•........•........•........................ TABLES ...•....................................................•..........•.......... FIGURES ...............•...•......................................................•.. CHAPTER2: EFFECT OF HUNTER DENSITY ON ELK MOVEMENT TO PRIVATE LAND IN NORTHWEST COLORADO . INTRODUCTION . STUDY AREA •.....••.............................•.....••...........••.........•.•.... METHODS ••••...••...••.•..........•................•............••........•......... Radio-telemetry Data Collection Hunter Survey Data Collection Estimating Number of Hunters Estimating Number of Hunters Afield per Day Estimating Mean Number of Days per Hunter and Hunter-days Elk Movement versus Hunter Density RESULTS . . . . . . ...•.........••..•...•....•............................................•..... Hunter Survey Data Elk Movement versus Hunter Density DISCUSSION .....................................•.•...•............•.................. MANAGEMENT IMPLICATIONS ......•..........•••......•.....••.....•..•.................• LITERATURE CITED TABLES FIGURES ......•......••.•............................................................ .:.'- . . . . ~-., :,-'".! ELK LOCATIONS IN RELATION TO DOMESTIC SHEEP BANDS AND A METHOD TO TEST FOR AVOIDANCE OR ATTRACTION BETWEEN ANIMALS . CHAPTER3: INTRODUCTION STUDY AREA METHODS Data Collection Data Analysis . . . . . �200 RESULTS DISCUSSION . Experimental Results Methodology for Determining Attraction and Avoidance MANAGEMENT IMPLICATIONS LITERATURE CITED TABLES ...........................................................•.................. FIGURES . . . . . APPENDIX 1: SAMPLE SIZE CALCULATION FOR HUNTER SURVEY •••••••••••••••••••••••• APPENDIX 2: STANDARD ERRORS FOR DAILY GMU ESTIMATES OF HUNTERS AND ATVS AFIELD APPENDIX 3: MAPS OF ELK AND DOMESTIC SHEEP LOCATIONS ON SELECTED FLIGHTS ••• ACKNOWLEDGMENTS i"l m: Funding was provided by Colorado Division of Wildlife (CDOW). Colorado Habitat Partners Program in the Northwest Region provided funding for flights to collect elk locations. I greatly appreciate the financial support. From the CDOW, I would like to extend a special thanks to Dave Freddy for his insightful elk discussions, reviews, and helping to procure financial support. A second special thanks to Dick Bartmann; without his help, organization, funding procurement, and common sense I would have struggled with all the logistics of collaring and tracking over 80 elk. Also from the CDOW, thanks to Jeff Madison for all his help with obtaining funding from Colorado Habitat Partners Program. My major professor, Dr. Gary White, has been quite an inspiration to me. What can I say, he is one of the smartest people I know and I have been lucky to study with such an adept thinker. My committee member, Dr. Ken Burnham, has always been a great help with statistical and mathematical issues that were difficult for me, and never missed a detail in my dissertation work. His help greatly improved my dissertation. Dr. Bill Alldredge always had the common elk-sense aspect and practical questions. Without him, I might have forgotten the most important issues about elk in their natural environment Finally, Dr. Bruce Wunder provided me with the interesting big picture questions that I would not have considered with out him. Thank you to all my committee; I enjoyed and benefitted from your input and help tremendously. The proximate reasons that I was able to complete this academic endeavor were excellent academic advising and a good funding source, but the ultimate reason is the relationships that have nourished me since . childhood. Primarily, my parents are the reason I have been able to start and complete my doctorate. My mother, who importuned me to "finish what you begin", and my dear father, with his natural Buddhist nature, both allowed me and encouraged me to do whatever it is to which I set out. Additionally, their financial support during graduate school kept me out of the poorhouse and kept my car out of the junkyard. I must thank my sister, who rolled her eyes and told me to "get a grip" many times during my graduate studies. I then thank my extended family, the cast of characters to whom I am related. They make me laugh and somehow think that I could do anything if that's the stock from which I originate. To my long time friends who work hard, but certainly expect to vacation well, and have little tolerance of any stupid, wimpy PhD excuses, thank you. My friends M. Kiuchi, P. Kurz, M. Zimmer, E. Soderstrom, A. Peet, D. Yonenaka, N. Larsen, M. Riker, T. Howe, A. Bruno, and T. Weller deserve special acknowledgements. Of the mentors from my academic career, I would like to thank and acknowledge Sister Rita Basta, Dr. Mike Jaeger, Dr. Dale McCullough, and Dr. Wayne Getz. Wayne, a special thanks to you, although you will never know how much you inspired me. Iwould also like to thank and acknowledge the myriad of graduate students who have made my life bright and graduate school stimulating. Most importantly, infinite thanks to my dear husband, John Shivik. He has been a stalwart support, fellow adventurer, paper reviewer, research partner, coffeemaker, cocktail alchemist, bad joke teller, and all round husband extrodinaire. Thank you, thank you, thank you all. I hope you like what you've been so instrumental in producing - this one's for you. �201 INTRODUCTION ELK MOVEMENTS IN RESPONSE TO DISTURBANCE Migration between high-elevation summer range and low-elevation winter range is common among Rocky Mountain elk (Cervus elaphus nelsoni) in mountainous regions (Adams 1982). High-elevation ranges provide high levels of nutrition during the summer, but snow makes forage inaccessible during the winter. Fall migrations typically take place before the snow becomes too deep for animals to forage. In the White River area of Colorado, elk migrate between summer range, located in high-elevation national forest and wilderness areas, and lowelevation wintering grounds, located as near as the base of the summer range or as far as 80 Ian away. Historically, fall elk migration took place in December (Boyd 1970), but since the 1980s, fall movements have apparently advanced into August and September (Gray et al. 1994). During the development of the 1994 elk management plan for the White River herd, early elk movement from summer ranges onto public land was the most common complaint (Gray et al. 1994). In response to complaints, Colorado Division ofWildIife (CDOW) conducted a pilot study on 20 radiocollared cow elk from 1992-1995 to determine if and when elk were moving onto private land. Results from the pilot study indicated a correlation between early-season hunting activity and elk movement to private lands. However, it did not establish a causal relationship. Before instituting any management changes, CDOW needed an experiment to determine whether early-season hunting activity was causing latesummer elk movements. I designed and conducted such an experiment in 1996-1997, and that experiment is the topic of my dissertation. In this introduction, I present background material on elk responses to an array of human activities in general and to hunting in particular. I intend this background material to provide a framework for understanding my experiment to determine the cause of late-summer elk movements in the White River area . -..r Elk Movement in Response to Human Activities Elk responses to recreational activities such as hiking, cross-country skiing, and automobile driving depend on habituation to humans and on the presence of hunting. In Rocky Mountain National Park, where elk were accustomed to people and were not hunted, elk showed little response to approaches of people in automobiles or on foot, day or night (Schultz and Bailey 1978). In a study of another unhunted elk population in Yellowstone National Park, elk response to cross-country skiers depended on how accustomed elk were to people. In areas of low human activity during the winter, elk responded with flight distances of 125-1,700 m, while in an area of high human activity, the flight distance was considerably shorter, ranging from 0 to 300 m (Cassirer et al. 1992). Interestingly, elk in the area of high human activity showed a three-fold increase in flight distance when they were disturbed outside the area where people were present 24 hours a day (Cassirer et al. 1992). Thus, even habituated elk appear to flee human activity in areas where they do not expect such activity. Besides habituating to humans, elk may learn to differentiate between dangerous and benign situations with respect to road avoidance. In Rocky Mountain National Park, where elk were not hunted and traffic volume was high, Schultz and Bailey (1978) found that none of their 14 delineated elk behaviors changed with traffic volume, and there was little or no avoidance of the roads in winter. In contrast, in Roosevelt National Forest, which is adjacent to Rocky Mountain National Park, Rost (1975) found that a population of hunted elk avoided roads in winter. Wright (1983) found that mean distance ofradiocollared elk to jeep trails more than doubled, increasing from 800 m to 2,100 m when hunting season opened. In addition to danger level, the level of road activity also affected elk movements. After roads were closed in an experimental treatment, Cole et al. (1997) found a reduction in daily movement, home range size, and core area, all measures of elk movement Czech (1991) found a significant increase in mean elk distances to a road after it was opened to the general public with a corresponding increase in traffic. Elk also cross heavily traveled roads less often than lightly traveled roads (Hershey and Legee 1982). A third factor, cover, may also determine the degree.of elk response to disturbance. Logging disturbances changed normal elk movements (Edge and Marcum 1985): elk moved significantly longer distances away from logging areas than toward them. But both distance from disturbances, and areas of high elk use, were correlated to the amount of cover in the area (Edge and Marcum 1985). Similarly, Czech (1991) found that elk tolerance of logging operations was correlated positively with proximity to hiding cover. In one of the few manipulative experiments on elk movements, elk calves subjected to simulated surface-mining activities sought coniferous forest significantly more often than undisturbed calves (Kuck et al. 1985). Oil drilling has the same effect as mining: elk increased their use of forested habitats during drilling and post-drilling periods, compared with pre-drilling periods (Van Dyke and Klein 1996). Collectively, these disturbance experiments build a strong case for a relationship between elk movement and availability of cover. �202 All studies that evaluated elk movements after the disturbance ended found displacement to be a temporary condition in most situations. The Yellowstone elk displaced by cross-country skiers typically returned close to their original locations after people left the area (Cassirer et al. 1992). Road proximity resumed pre-hunting distances after hunting season closed (Wright 1983), and mean distance to roads decreased soon after the roads were closed for the season (Hershey and Legee 1982). During weekends, and immediately following the end oflogging activities, elk moved back into logged areas (Ward 1976, Edge and Marcum 1985). ~! Elk Movement in Response to Hunting Documented responses of elk to hunting include movement away from hunters or heavily hunted areas, increased movement, movement to thick cover, shifts in circadian patterns, and elevation shifts. Most responses seem to occur before winter migration, but some studies attribute changes in migration patterns to hunting pressures. For example, Knight (1970) and Morgantini and Hudson (1979) found a reversal of the normal downward autumn migration of elk, coinciding with the opening of the hunting season. Responses of elk to hunting activity may depend on the amount, location and quality of hiding cover, as well as the topography of the area. Altmann (1956) described an evasive "migration" of elk hunted adjacent to Yellowstone National Park. With the opening of hunting season, movements of these elk became relatively long (513 km), and the elk ceased long movements only when they reached the sanctuary of the Park. Similarly, Martinka (1969) found that 12 marked elk, which were in the National Elk Refuge just prior to hunting season, moved between 8 and 22 km, to areas closed to hunting in Grand Teton National Park. Other studies have found lessdramatic movement to refuge areas. With opening of hunting season, elk moved to densely forested (Irwin and Peek 1979) or shrubby (Morgantini and Hudson 1979) areas adjacent to their typical areas of activity, which provided refuge from hunting. Unpublished data for 1992-1995 from a pilot study on elk in the White River area (M. M. Conner, Colorado State University, unpublished data) found that distance moved around opening date (± 1 week from opening date) differed depending on habitat cover in the vicinity. Elk located in rugged and relatively inaccessible areas with access to thick timber and rough terrain (n = 11) moved an average of 858 mlday (95% CI = 602, 1113). In contrast, elk located in road-accessible areas (n = 15), with little access to vegetative and topographic cover, moved an average of 1258 mlday (95% CI = 1011, 1505) to refuge on private land. Thus, elk without access to nearby habitat refuges moved significantly farther (401 mlday: 95% CI = 45, 756) than elk with access (P = 0.014, l-sided), Areas of rough topography and dense timber may serve as refuge; if such refuge is available, elk may not move far even when subject to high hunting pressure. Elk may respond to hunting method or density of hunters. Wright (1983) examined elk response to different hunting seasons: archery and muzzleloading, rifle-special elk, rifle-special deer, and deer and elk rifle. The greatest density of elk hunters was during the rifle-special elk season, and distances traveled by elk were also greatest during that season. Elk traveled three to four times farther during this season than during archery and muzzleloading season. Wright (1983) concluded that archery and muzzleloading hunters did not change the behavior of radiocollared cow elk. However, this result might be explained by low hunter densities (0.13 archery and muzzleloading hunters/km', compared to 1.42 rifle hunters/km') rather than by the hunting method. In a 1985 study in the White River area, Consolidation Coal Company radiocollared 23 elk to evaluate their movement onto the company's protected land during early-season hunting. Eighty-seven percent of the elk were still found on National Forest lands midway through archery and muzzleloading seasons (Camp Dresser and McKee, Inc. 1986). But the lack of movement may be explained by a 63% reduction in archery hunters from 1984 to 1985, due to a change in hunting regulations (Grayet al. 1994). Zahn (1974), Lemke (1975), and Hershey and Legee (1982) noticed an increase in movements during the first 10-12 days of hunting season, followed by normal movement during the remainder of the season. These authors noted that hunting pressure was heavy during the first 10-12 days of hunting and relatively light for the remainder of the season. These studies of hunting suggest that elk behavior may change only when the density of hunters reaches a critical threshold. While some elk appear to have learned to move to refuge areas, elk already in refuges may learn to remain there in response to hunting pressure. In Jackson Hole, both resident and migratory elk use the National Elk Refuge for their winter range (Martinka 1969). Of the 183 marked elk in the study, resident elk were defined as elk found at any time from July through August within 15 km of the winter range, while migratory elk were defined as all other elk. In the fall, both resident and migratory elk had to cross hunting areas to move to their winter range in the National Elk Refuge. Resident elk were sensitive to hunter presence and tended to hang back in areas closed to hunting, while migratory elk were less hesitant to move through hunting areas to the winter range. These studies of elk moving to, or staying in, refuge areas support the theory that elk are responding, possibly with learned behavior, to hunting pressure. �203 Other studies found evidence to indicate that elk are not responding to hunting pressure, but are being differentially removed from heavily hunted areas. In Wyoming, hunting pressure focused on a resident population of elk just east of Yellowstone National Park, drastically reducing their numbers (Rudd et al. 1983). Like the Jackson Hole herd, migratory elk that summered in protected areas of Yellowstone National Park wintered with resident elk in unprotected range just outside of the park. Hunting in the unprotected range occurred before the migratory elk arrived from the park, drastically reducing the herd of resident elk (Rudd et al. 1983). The proportion of migratory to resident elk increased in the unprotected range, apparently due to hunting itself and not to huntinginduced shifts in elk behavior. Boyce (1991) noted that migration of elk wintering in the National Elk Preserve in Jackson Hole changed dramatically between the 1950s and 1980s. The primary migration routes shifted from openhunting national forest lands to routes through Grand Teton National Park, where hunting was more restricted. Between 1950 and 1954,22% of the migrating elk were estimated to have moved through Grand Teton National Park; by 1980-1984 the figure was 51%. Boyce (1991) hypothesized that this change may be due to differential harvest of the migrating animals and not due to shifts in elk behavior. However, there has been no experiment to test whether the cause of migration changes is differential removal or learned response. Similar to results from non-lethal distmbance studies, Zahn (1974), Lemke (1975), and Hershey and Legee (1982) found that elk response to hunter distmbance was short-lived. These studies noted that elk movements return to normal after the initial 10-12 days of opening day, and that hunting is most intense during those initial days. Elk not only discontinued their erratic and increased movements, but also returned to their pre-season activity areas, indicating that hunting activity was a temporary distwbance. In conclusion, elk show highly plastic responses to human activity, habituating to people in areas where ..there is no hunting, but actively avoiding hunters. Elk response to human activity depends on the amount of. and .the danger of, contact with humans. For example, avoidance of roads varies with the amount of human use and the :danger level. The presence of dense cover for hiding seems to attenuate responses to human activity. Finally, elk responses to human activity do not seem to persist after the activity subsides. All studies of elk responses to hunting have been observational and any causal relationship between hunting and elk movements remains untested. PROJECf OVERVIEW Game Management Units (GMUs) 12, 23, 24, and 33 in northwestern Colorado compose the majority of the area of the White River elk herd. The White River herd has been one of the most heavily hunted, managed, and documented elk herds in Colorado (Boyd 1970, Freddy 1987, Gray et al. 1994). Like all elk herds in Colorado, the White River herd was decimated during the late 1800s, eventually being reduced to a few hundred elk (Bryant and Maser 1982). Since regulated hunting began in 1929, the population has been growing and was estimated to be 31,000 in 1993, the upper bound for CDOW management objectives (Gray et al. 1994). The number of hunters using the area has grown along with the elk herd, with especially significant increases in the number of early-season hunters (archery and muzzleloading) between 1984 and 1992 (Fig. I.1). In the White River area, summer range is high-elevation public forest, and winter range is lower-elevation private land. Historically, the main White River elk migration from summer to winter ranges took place between late November and early January (Boyd 1970). However, Freddy (1987) noted that elk were beginning to be found in lower elevations, characteristically winter range, during the September and October hunting seasons, which was uncommon in the 1960s. Furthermore, there is now evidence that elk are moving in August and early September, during early-season archery hunting (Grayet al. 1994). Typically, early-season hunters did not hunt private lands; thus, private lands provided elk a refuge during early-season hunts. Late-summer elk movements to private land have led to complaints by local private landowners and resource managers (Gray et al. 1994). For private landowners, complaints focused on crop damage. For CDOW, the primacy problem is herd size: because hunters have limited access to private lands, it is difficult for CDOW to maintain or reduce herd size via harvest (1. Madison and C. Reichert, CDOW, personal communication). Hunter equity is a third issue: if a large proportion of elk are on private land, then the best hunting will be monopolized by those with enough money to buy trespass permits (1. Madison and C. Reichert, CDOW;'personal communication), Complaints about late-summer elk movement during the mid- to late-1980s prompted the CDOW to conduct a preliminary study on late-summer elk movement to private land in the White River area. In 1992 the CDOW trapped and radiocollared 20 adult female elk. A 4-year pilot study from 1992 to 1995 was conducted on 14 to 20 of these radiocollared elk from GMUs 12,23,24, and 33. During the study, 70-100% ofradiocollared elk located on public land areas moved to private or wilderness areas, and the mean day of movement was not different from opening day of early-season hunting. For elk moving from public to private land (n = 29) during the 4- year pilot study, the mean date of movement was 3 days (95% CI = -4,11) from opening date (M. Conner, Colorado �204 State University, unpublished data). Although it did not establish a causal relationship, this study provided strong evidence of a correlation between elk movements and the opening of early-season hunting. To determine if archery and muzzleloading hunting caused elk movements in the White River area in the White River area, I conducted a 2-year experiment. In the White River area, other factors that may contribute to elk movement include historical movement patterns, long-term, learned movement responses, recreational activity, wood cutting, and livestock activity. Any or all of these factors may contribute to elk movement from July through October. However, because the pilot study identified archery hunting as the prime suspect for late-summer movements, and because the CDOW was interested in manageable rather than academic issues, this study focused on the effects of archery hunting on elk movement. In Chapter 1, I report on the field experiment designed to determine whether archery hunting causes elk movement: this experiment compares the timing of elk movements, and the proportion of elk on or moving to private land, with the opening date of archery season. In Chapter 2, I estimate daily hunter densities and compare elk movement to private lands with hunter density. In Chapter 3, I discuss the impact of domestic sheep grazing on elk movement. Each chapter is written to stand alone for publication and follows manuscript guidelines for The Journal of Wildlife Management. LITERATURE CITED Adams, A. W. 1982. Migration. Pages 301-321 in 1. W. Thomas and D.E. Toweill, editors. Elk of North America: ecology and management. Stackpole Books, Harrisburg, Pennsylvania. USA. Altmann, M 1956. Patterns of herd behavior in free-ranging elk of Wyoming, Cervuscanadensis nelsoni. Zoologica 41:65-71. . Boyce, M. S. 1991. Migratory behavior and management of elk (Cervus elaphus). Applied Animal Behaviour Science 29:239-250. Boyd, R J. 1970. Elk of the White River Plateau, Colorado. Colorado Division of Game, Fish and Parks Technical Publication 25, Fort Collins, Colorado, USA. . Bryant, L. D., and C. Maser. 1982. Classification and Distribution. Pages 1-59 in 1. W. Thomas and D.E. Toweill, editors. Elk of North America: ecology and management. Stackpole Books, Harrisburg, Pennsylvania. USA. Camp Dresser and McKee, Inc. 1986. Meeker PRLA elk migration study: monitoring report, volume 2. Prepared for Consolidation Coal Company. Camp Dresser and McKee, Inc., Denver, Colorado, USA. Cassirer, E. F., D. 1. Freddy, and E. D. Ables. 1992. Elk responses to distwbances by cross-country skiers in Yellowstone National Park. Wildlife Society Bulletin 20:375-381. Cole, E. K., M. D. Pope; and R G. Anthony. 1997. Effects of road management on movement and survival of Roosevelt elk. Journal of Wildlife Management 61: 1115-1126. Craighead, 1. 1., G. Atwell, and B. W. O'Gara. 1972. Elk migrations in and near Yellowstone National Park. Wildlife Monographs 29. Czech, B. 1991. Elk behavior in response to human distwbance at Mount St. Helens National Volcanic Monument. Applied Animal Behaviour Science 29:269-277. Edge, W. D., and C. L. Marcum. 1985. Movements of elk in reaction to logging disturbances. Journal of Wildlife Management 49:926-930. Freddy, D. J. 1987. The White River elk herd: A perspective, 1960-85. Colorado Division of Wildlife Technical Publication 37 Fort Collins, Colorado, USA. Gray, J. P., G. Byrne, and J. Madison. 1994. White River elk data analysis unit plan: game management units: 11,211,12,13,131,231,23,24,25,26,33. Colorado Division of Wildlife Internal Report, Grand Junction, Colorado, USA. Hershey, T. J., and T. A. Leege. 1982. Elk movements and habitat use on a managed forest in north-central Idaho. Idaho Department ofFish and Game Wildlife Bulletin 10, Boise, Idaho, USA. Irwin, L. L., and J. M. Peek. 1979. Relationships between road closures and elk behavior in northern Idaho. Pages 199-204 in M. S. Boyce and L. D. Hayden-Wing, editors. North American elk: ecology, behavior, and 'inanagement.· University of Wyoming, Laramie, Wyoming, USA. ott Knight, R R. 1970. The Sun River elk herd. Wildlife Monographs 23. Kuck, L., G. L. Hompland, and E. H. Merrill. 1985. Elk calf response to simulated distwbance in southeast Idaho. Journal of Wildlife Management 49:751-757. Lemke, T. O. 1975. Movement and seasonal ranges of the Burdette Creek elk herd, and an investigation of sport hunting. Montana Fish and Game Department Job Final Report 32.01, Job No. BG-3.15, Missoula. Montana. USA. �205 Martinka, C. 1. 1969. Population ecology ofswnmer resident elk in Jackson Hole, Wyoming. Journal of Wildlife Management 33:465-481. Morgantini, L. E., and R. J. Hudson. 1979. Human distribution and habitat selection by elk. Pages 132-139 in M. S. Boyce and L. D. Hayden-Wing, editors. North American elk: ecology, behavior, and management. University of Wyoming, Laramie, Wyoming, USA. Rost, G. R. 1975. Responses of deer and elk to roads. Thesis, Colorado State University, Fort Collins, Colorado, USA. Rudd, W. J., A. L. Ward, and L. L. Irwin. 1983. Do split hunting seasons influence elk migrations from Yellowstone National Park? Wildlife Society Bulletin 11:328-331. Schultz, R. D., and J. A. Bailey. 1978. Responses of national park elk to human activity. Journal of Wildlife Management 42:91-100. Van Dyke, F., and W. C. Klein. 1996. Response of elk to installation of oil wells. Journal ofMammalogy 77:1028-1041. Ward, A. L. 1976. Elk behavior in relation to timber harvest operations and traffic on Medicine Bow Range in south-central Wyoming. Pages 32-43 in S. Hieb, editor. Proceedings of the Elk-Logging-Roads Symposium. University ofIdaho, Moscow, Idaho, USA. Wright, K. L. 1983. Elk movements, habitat use, and the effects of hunting activity on elk behavior near Gunnison, Colorado. Thesis, Colorado State University, Fort Collins, Colorado, USA. Zahn, H. M. 1974. Seasonal movements of the Burdette Creek elk herd. Montana Fish and Game Department Job Final Report 32.01, Job BG-3.13, Missoula, Montana, USA. 7000 (/J •.. 6000 ~ Q) c :::l .J::. c: ,. 95%CI --- -" 5000 0 (/J <U 4000 Q) (/J I >. "i:: <U 3000 Q) 0 ~ 2000 Q) .0 § 1000 z 0 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 Year * Variance estimates not available 1984-1987. Figure 11. Number of archery and muzzieloading hunters using GMUs 12, 23, 24, and 33 in northwest Colorado, 1984-1994. Data from Deer, Elk, and Antelope Management (DEAMAN) database and software, Colorado Division of Wildlife, unpublished data. �206 �207 CHAPTER 1: ELK MOVEMENT IN RESPONSE TO EARLY-SEASON COLORADO HUNTING IN NORTHWEST INTRODUCTION Since the early 1900s, Rocky Mountain elk (Cervus elaphus nelsoni) populations have expanded their range from remote areas to much of the Rocky Mountain States (Bryant and Maser 1982). As elk range expanded, so too did elk contact with humans and human disturbances. Elk responses to hooting, road traffic, hikers, cross country skiers, snowmobiles, logging, mining, and oil drilling have been studied in order to reduce elk-human conflicts. Problems with elk responding to hooting center around elk moving onto hooting refuges, which are areas with no or low levels of hooting such as private land, national parks, or densely forested areas. Documented responses of elk to hooting include movement away from hooters or heavily hooted areas (Altmann 1956, Martinka 1969, Wright 1983), increased movement (Zahn 1974, Lemke 1975, Hershey and Legee 1982, Wright 1983), movement to dense vegetative cover (Irwin and Peek 1979, Morgantini and Hudson 1979), movement to private land (Wright 1983) or national parks (Altmann 1956, Martinka 1969), shifts in circadian patterns (Strohmeyer and Peek 1996), and elevation shifts in migration patterns (Knight 1970, Morgantini and Hudson 1979). Most responses to hooting occur before migration to winter range, but some studies have noted changes in migration patterns that were attributed to hooting pressures (Martinka 1969, Boyce 1991). All previous studies of elk responses to hooting have been observational, there have been "noexperiments designed to test whether hooting activity causes elk movements. Elk inhabiting the White River area of northwest Colorado are no exception to the pattern of increased range expansion and increased conflict with humans. Prior to 1960, the fall migration of White River elk from high elevation public land, summer range, to lower elevation private land, winter range, (Fig 1.1) occurred during the month of December (Boyd 1970). However, Freddy (1987) noted that elk were beginning to be found in lower elevations during the September and October hooting seasons, which was an uncommon occurrence in the 1960s. Since the 1980s, fall movements have apparently advanced into August and September (Gray et al. 1994). Archery and muzzleloading hooting (early-season hooting) opens on the third Saturday of August in the White River area; around the time of the alleged elk movements onto private land. Since the 1960s, the number of earlyseason hooters using the White River area has grown, with especially significant increases between 1984 and 1992 (Fig. 1.2). Typically, archery hooters did not hoot private lands; thus, private lands provided elk a refuge during early-season hoots. As archery and muzzleloading hooter numbers increased, so too did complaints about early elk movements onto private land. For the private landowner, complaints about early elk movements focused on crop damage. For the Colorado Division of Wildlife (CDOW), the primary problem was herd size: it was difficult for CDOW to maintain or reduce herd size via harvest because hooters have limited access to private lands, (J. Madison and C. Reichert, Colorado Division of Wildlife, personal communication). Because of complaints during the mid- to late 1980s, CDOW conducted a preliminary study on late summer elk movement to private land in the White River area. From 1992 to 1995,20 radiocollared elk were intensively monitored during August and September, approximately one month before and one month after opening day of archery hooting. Although the proportion and date of elk moving indicated a correlation between elk movement and opening of early-season hooting, a causal relationship was not established. Before instituting any management changes, the CDOW required an experiment to determine if early-season hooting activity was causing late summer elk movements. My objective was to determine if archery hooting caused elk movement to private land during late summer. Specifically, I conducted a 2-year field experiment in.whichopening date of archery hooting was manipulated in a crossover design. The study area was splitinto 2 treatment areas; each area received an earlyand late-opening treatment. I tested the null hypotheses that; (1) the mean date of movement to private land was not different between early and late treatments, and (2) the proportion of elk on private land was not affected by the opening of archery hooting. �208 STUDY AREA The White River area of northwestern Colorado covered approximately 4,540 km2 and was composed of Game Management Units (GMUs) 12,23,24, and 33. In Colorado, GMUs were delineated to distribute hunters through allocation of hunting licenses. White River area land ownership was 34% private land and 66% public land (Fig 1.1), with 82% of public land being United States Forest Service (USFS). USFS land was mostly at high elevation and toward the center of the study area, with some Bureau of Land Management (BLM) areas at lower elevations in the southern sections of the study area. Private land was lower in elevation and comprised mainly ranches and coal mines. Topography, climate, and vegetation varied Widelythroughout the study area. Elevation ranged from 1,629 to 3,700 m. The central part of the study area was high elevation public land, split by the White River valley. The high elevation areas dropped off to lower elevations to the north, south, and western edges of the study area. Higher elevations had severe winters with heavy snowfall, while lower elevations had comparatively mild winters. Mean annual precipitation at 3,000 m in the National Forest areas was about 100 cm, compared with about 30 cm at lower elevations of <2,000 m. Vegetation types at elevations >2,600 m were in the montane/subalpine zone and included groves of aspen (Populus tremuloides), Engelmann spruce (Picea engelmanni), and alpine fir (Abies lasiocarpa) interspersed with grassy meadows. Middle elevations (1,9802,600 m) were a transitional zone that consisted primarily of pinion pine (Pinus edulis), jumper (Juniperus scopulorum), and big sagebrush (Artemisia tridentata). Lower elevations «2,000 m) in the southern and northern parts of the study area were in the Great Basin zone with sage grasslands. Oakbrush patches were found at the middle and lower elevations with major shrubs that included Gambel's oak (Quercus gambelii), serviceberry (Amelanchier alnifolia), mountain mahogany (Cercocarpus montanus), chokecherry (Prunus virginiana), snowberry (Symphoricarpos utahensis), bitterbrush (Purshia tridentata), and rabbitbrush (Chrysothamnus nauseosus). Higher and middle elevation areas provided summer and fall forage for elk, while lower elevations were typically used by elk during winter. Boyd (1970) provides a detailed description of the study area. Hunting for elk was economically important to local residents in the study area (Gray et al. 1994). Guides, outfitters, local service sectors, and private landowners (who sell trespass permits) made a large proportion of their annual income during the elk and deer hunting seasons, with most hunting revenues accrued during rifle hunting seasons. METHODS Experimental design The White River and North Fork of the White River divided the study area roughly east-west into 2 halves (Fig. 1.3). GMU 12 and parts of GMUs 23 and 24 composed the north treatment area, while GMU 33 and parts of GMUs 23 and 24 composed the south treatment area. Because only 2 elk moved from one side of the White River to the other within a year during the 4-year pilot study (M. Conner, Colorado State University, unpublished data), I felt there would be little interchange between treatment areas. Two treatments were applied: an archery season that opened 1 week earlier, and another that opened 2 weeks later than normal, yielding a 3week difference in opening dates. During the first year of the study, the early-opening treatment was randomly assigned to the south treatment area. Treatments were reversed the second year of the study. Thus, in 1996, archery hunting opened early, 24 August, in the south area, and late, 14 September, in the north area. When treatments were reversed in 1997, archery hunting opened early, 23 August, in the north area, and late, 13 September, in the south area. The number of archery and muzzleloading licenses issued for elk and deer during ."", the stuey:'were basedon the mean number per year calculated for 1990-1994 (4,985; Colorado Division of .•....•.." Wildlife unpublished data), with licenses allocated approximately 50% per treatment area. Restricting licenses kept hunter density consistent during the study and consistent with years of high elk movement complaints. Sample size for number of collared elk was based on the primary response variable, mean date of movement, for the following parameters; a. = 0.05, ~ = 90%, effect size = 7-day difference in mean date of movement between treatments, and estimated variance = 22.7 days. The variance was based on pilot study data (M. Conner, Colorado State University, unpublished data). For these parameters, 36 cows per treatment area , -':. �209 were required. To allow for some telemetry failures or erratic elk movements off the study area, extra cows were collared. Thus, 40 cows per treatment area were collared for the study. Elk capture locations were randomly picked from a uniform distribution by a computer. Random Universal Transverse Mercator (UTM) coordinates were generated until 80 points lay within the study area's national forest boundaries. Elk were captured by helicopter netgunning (Barrett et al. 1982) during mid-July in 1996 and 1997. Elk were captured on their summer range to assure a random sample for the July-October study period. The helicopter pilot flew to each randomly selected location and captured the first adult female elk found near that location. Although it was not practical to capture exactly as dictated by a random number algorithm, due to constraints on private land access and time, a reasonably representative sample was obtained with 2 elk being captured at some locations (Fig. l.3). In addition to the 80 elk captured in 1996,8 radiocollared elk from the pilot study were used in 1996 analyses. In 1997, 10 additional elk were captured to replace animals that died, bringing the total number of collared animals back to 80, with 40 per treatment area. Colorado State University Institutional Animal Care and Use Committee approved the capture and handling methods (protocol 95-180A-Ol). Each elk was fitted with a 148-152 MHz radio collar with a mortality sensor. A high percentage of bulls are lost to hunting, which could reduce sample size below levels required to maintain the power of the hypothesis tests. Because this study was to take place during a hunting season, when bulls are desirable game, I wished to collar only adult females to minimize loss of sample size. During fall rutting season, August-October, there is a high degree of association (97%) between male and female elk (Franklin and Lieb 1979). I collared only adult female elk to represent the movement patterns of both sexes and maintain the power of the hypothesis tests. Data collection The relevant time frame for determining elk movement in response to archery hunting was defined as ±1 month of opening day. Because early treatment opening was 23-24 August and late treatment opening was 13-14 September, I collected locations 20 July-IO October. The study period ended slightly less than a month after late treatment opening date to avoid confounding elk movements due to archery hunting with effects due to rifle hunting, which opened 13-14 October. All analyses were done on data collected over the entire 3-month study period. I relocated radiocollared elk between 0700-1500 hr using a Cessna 182 or 185 fixed-wing aircraft with a 2-element Yagi antenna mounted to each strut of the airplane. For each elk relocation, UTM coordinates were recorded with a Global Positioning System that was not differentially corrected. I collected elk locations 2 times a week with 2-4 days between collections. A Geographical Information System (GIS) map was used to record elk locations, land ownership, and treatment boundaries. The GIS map was digitized from a United States Geological Survey map at a scale of 1:100,000. For each location ofa telemetered cow, a 0 was assigned if the location was on public land (e.g. Flat Tops Wilderness Area, USFS, State Wildlife, and BLM lands) and a 1 was assigned if the location was on private land. Because I was interested in gross classifications of elk locations (public land versus private land), I did not do a formal check of the telemetry error. However, I did a cursory check of telemetry error when picking up collars from a different study in the area. The mean error on the collars I collected was 333 m (n = 24,95% CI = 265,401), with a range of 117-683 m. Data analysis The primary response variables were mean date of movement for elk that changed classification (occupying public and moving to private land) and proportion of elk on private land. I calculated the.date of movement for each elk using logistic regression oflocation on public (y = 0) and private land (y = 1) versus date. The date of movement was defined as the date at which the logistic curve crossed 0.5 on the y-axis, indicating an equal probability of being on public or private land. Only dates of movement from public to private land during the study period were included in analyses of mean date of movement. If hunting had no effect on elk movements, then there should be no difference between the mean date of movement to private land between early- and late-opening date treatments. To test for a treatment effect on mean date of movement, I accounted for treatment, area, and year effects. The corresponding ANOVA model was: �210 where: Yljkl = J.l = (1.1 = date of movement for the fh radiocollared elk, receiving treatment overall mean date of movement (for all elk, both years), Ok = fixed effects due to treatment i (early or late opening), fixed effects due to year j (year 1 or year 2), fixed effects due to area k (either north or south), and sljk/ = random error for the ~j = I'" radiocollared i, in year j and in area k, elk. Specific hypotheses tested were: Hoi: Mean date of elk movement for early-opening treatment treatment; (1.1 ;;:: 0; = mean date of elk movement for late-opening HAl: Mean date of elk movement for early-opening treatment < mean date of elk movement for late-opening treatment; (1.1 < 0; lIm: Mean date of elk movement for year 1 of experiment HAl: Mean date of elk movement for year 1 of experiment = mean :F- mean date of elk movement for year 2; ~j == 0; date of elk movement for year 2; ~j:F- 0; Ho3: Mean date of movement for north-area elk = mean date of movement for south-area elk, Ok== 0; HAl: Mean date of movement for north-area elk :F- mean date of movement for south-area elk, Ok:F- O. Hypothesis 1 was the experimental hypothesis; hypotheses 2 and 3 accounted for other sources of variation. I used PROe GLM (SAS Institute 1990) to estimate and test differences in mean date of movement by area, treatment, and year. Type m sum of squares were used for hypothesis tests. To determine whether elk receiving both early and late treatments on public land, regardless of area or year, were affected by opening date, I used a paired I-test blocked on individual elk. This individual blocking reduced confounding effects and better accounted for repeated measures taken on animals receiving both treatments; thus, the analysis was limited to those individuals that received both early- and late-opening date treatments. Fewer elk received both treatments than received a single treatment because some elk died after the first study period, some elk were on public land only one year (they stayed on private land the other year), and some changed treatment areas between years. For the elk getting both treatments, I subtracted their early date of movement from their late date of movement to account for the fact that this was a repeated measurement on the elk. I tested the null hypothesis: Ho: Difference in date of movement for late treatment - early treatment <== 0; HA: Difference in date of movement for late treatment - early treatment> O. I used logistic regression models to predict the proportion of elk on private land on day i (p i) from elk location data (public or private land). Models included date as a covariate, and hunting season, area and year effects as categorical variables. JUlian date was used to model date effects so that year effects could be considered separately from study period date. Hunting season referred to the period before opening of archery season (hunting season = 0) and after opening (hunting season = 1). I began with a global model, which included date, hunting season, area, year, plus all 2-, 3- and 4-way interactions of date, hunting season, area, and year (fable 1.1, Model 1). I then developed 9 additional a priori hypothetical models for proportion of elk on private land (fable 1.1, Models 2-10). After analyzing the a priori models, I built 3 additional models to see if a better model could be built (fable 1.1, Models 11-13). �211 I followed the methodology of Burnham and Anderson (1998) to select an appropriate model. I used Akaike's Information Criterion (Akaike 1973), adjusted for overdispersion and corrected for small sample bias (QAlCc), as the basis for objectively ranking models and selecting an appropriate "best approximating" model (Burnham et al. 1995). QAlCc was defined as: =_ QAlCc 2(lnl) +2K c + 2K(K +1) n-K -I ' where In £ is the natural logarithm of the likelihood function evaluated at the maximum likelihood estimates for a c given model, K is the number of estimable parameters from that model, n is sample size, and is the estimated overdispersion factor. QAlCc was used instead of AlCc because repeated measurements were taken on radiocollared elk. Lack of independence in the data, in this case repeated measures on elk, may lead to overdispersion or "extra-binomial variation" (Burnham and Anderson 1988). I estimated the overdispersion c) parameter ( from the global model, and then used this estimate to adjust (inflate) the variance estimates of model parameters and predicted values, and to calculate QAlCc for all models (Burnham and Anderson 1998). Parameter c, estimates, and QAlCc values were calculated using PROC GENMOD (SAS Institute 1993). The sample size, n, was the number of elk locations collected during the study period. The best approximating model was selected based on minimum QAlCc. Models were ranked and compared using DQSAlCc (Leberton et al. 1992, Burnham and Anderson 1998) and normalized QAlCc weights (Buckland et al. 1997, Burnham and Anderson 1998). For the suite of models being compared, AQAlCc was computed for each model as: AQAlCc", = QAlCc", - QAlCc"'ln , where QAlCc ••was the QAlCc value for the mth model and QAlCcm1nwas the minimum QAlCc among the suite models being compared. Essentially, AQAlCc", is an estimate of the distance between the best approximating model and model m. Normalized AlCc weights (w",) were estimated for each mth model: -.!..~QAICc e 2 w m-- III 1 ~::e ' --~QAICc R 2 r r=l where R refers to the R models chosen for evaluation. AQAlCc and normalized QAlCc weights were used to address model selection uncertainty. Generally, models within 1-2 QAlCc units of the selected model were considered competing models for explaining elk movements to private land. Another element of uncertainty in estimates of model precision is selection of the appropriate model (Buckland et al. 1997). If several sets of data, generated from the same underlying process, were evaluated using AlCc, there would be some variation in the selected best model (Burnham and Anderson 1998). Estimates of precision from anyone best model do not include model selection variance and may be biased low. Therefore, after model selection, I used model averaging to better estimate the precision of the estimates of daily proportion of elk on private land from all the models I developed. Average estimates of the daily proportion of elk on private land were calculated for each day i across all models considered. Because all models were logit models, modelaveraged daily proportion of elk on public land, Pi' and var(Pi) were calculated in the logit scale and backtransformed to calculate the appropriate confidence interval, which was slightly asymmetric. The transfomation used was: logit(p;) = _ logit(pi) In[ I !fo; J. R = LWm m=l 10git(Pim), and �212 -;;- e logit( PI ) . Pi = 1+ e1ogit(PI) , where Wm is the normalized AlCc weight. The variance of the model-averaged conditional sampling variance and model uncertainty was estimated: estimates, which included where M", is the mth model. The 95% confidence interval for the daily estimated proportion of elk on private land was calculated as: e +95%CI~ogit(PI)] -------::-, 1+ e +95%CI~ogit(PI)] and To estimate the effect of hunting season on the proportion of elk on private land, I estimated the difference between the proportion of elk on private land immediately before and after opening date: e1ogit(Pb) I+e1ogit(Pb) P P , where a and b are the model-averaged proportion of elk on private land after and before opening date. The variance in estimated proportion of elk on private land before and after opening of archery hunting was calculated: �213 Procedures for estimating covariance of model-averaged values are in the development stage and untested (K. Burnham, Colorado Cooperative Fish and Wildlife Research Unit, personal communication). Thus, Iassumed the covariance to be zero. This omission may result in a slightly inflated variance, and a possibility of not detecting a true effect of hunting season. The variance of the model-averaged estimated proportion of elk on private land was calculated using the delta method (Seber 1982): Let U = logit(p a), then v8r{pa) = v;:I ~] , and "ll+ell :;...r~) •--=-v du ••••.. :;...r) du _ ( dU)2 v••••.. ;:...r)_ [ U -=-- --=U - v"'\Ya dPa dPa dPa e" 2 ]2;:...r) vat•.. U. (l+e") Substituting logit(p a) back in for u: The var(Pb) was calculated similarly. The 95% CI was calculated: - - - 95%CI(Pa - h)::: (Pa - - ,,- h) ± 1.96se(Pa - h)· -i-:",: Because previous studies used changes in daily distances moved and elevation shifts to evaluate elk responses to disturbance, Icalculated these metrics for comparison. Mean daily distances moved and mean daily elevations during the study period were calculated from the 80-radiocollared elk. Distances were calculated from consecutive UTM locations for each elk as straight-line distances. Because elk were not located at fixed time �214 intervals, distances between locations were expressed per day by dividing the distances moved by the number of days between locations. Daily distances were not absolute measures of distance moved, rather they were indices of movement. Elevations for each elk location came from a statewide 1:100,000-scale elevation map. RESULTS Results from the ANOVA on mean date of elk movement indicated a treatment effect (P = 0.013, one sided), but no area or year effects (P > 0.100; Table 1.2). However, north- and south-area elk showed different responses to opening of archery hunting. To evaluate this apparent difference, Ipreformed a post hoc analysis separately by area. This analysis revealed a 14-day difference (P = 0.006, one sided) in mean date of movement between treatments for elk in the north area, versus a 6-day difference (P = 0.218, one sided) between treatments for elk in the south area (Table l.3). Area and year were confounded within sequence of treatments this study. That is, elk in the north area received the late-early treatment sequence, while elk in the south area received the early-late treatment sequence. Thus, it was impossible to determine whether a sequence effect was really a difference resulting from (1) a difference between treatment areas because of location of private-land refuges, topography, etc., or (2) elk receiving a late-opening treatment the first year followed by an early-opening treatment the second year behave differently than elk receiving the early-late sequence. Ireference this confounded effect as an area effect, which I will discuss later. Sixteen of the 80-radiocollared elk received both early and late treatments and moved from public to private land during the study period. That is, there were 32 paired movements (16 elk receiving 2 treatments and moving form public to private both years) compared to 60 unpaired movements (elk receiving treatments and moving from public to private land at least 1 year). Some elk moved to private land immediately after capture in 1996, and some elk never returned to public land in 1997, thus fewer than 80 elk were available for treatment each year. The difference between the paired and unpaired movements occurred because 4 elk moved the first year but died before the second year, 3 elk were collared in 1997 and moved only the second year, 11 elk only moved one year because they were on public land one year and private land the other year, 1 elk changed treatment areas and received late-late treatments (counting as 2 unpaired movements), and 8 elk moved from public to private land one year only. The difference in mean date of movement for the 16 elk receiving both early and late treatments was 5 days (95% CI = -3,13; P = 0.097 one sided). ' Using model-averaged values, Igenerated predicted curves of proportion of elk on private land versus Julian day, hunt season, area, and year (Fig. l.4 and l.5). All logistic models to estimate the daily proportion of elk on private land had a significant positive date effect indicating that proportion of elk on private land increased from 19 July-lO October both years of the study (Fig. 1.4 and 1.5). The logistic regression model with the lowest AlCc (Table 1.1, model 13) was: logit(p;) = -4.715 -1.294(area)+ 4.987(hunt season) + O.OI5(Julianday) + O.009(areax Julian day) - 0.0 I8(hunt season x Julian day) , Pi where is the predicted proportion of elk on private land on Julian day i. The top 2 models, both post hoc models, (Table 1.1, models 13 and 12) were 1.85 AlCc units from each other, while the remaining 11 models were over 3 AlCc units greater than model 13 (Table 1.4). Hence, Iconsidered models 13 and 12 as competing models. Model 12, the second model, was identical to model 13 except for an additional area x hunt season interaction. Thus, area, hunt season, Julian dateaarea x Julian day, hunt season x Julian day, and area x hunt season may be important predictors of the proportion of elk on private land. The area effect arose from a higher proportion of elk on private land, in general, on the north area versus the south area, the hunt season effect arose from a higher proportion of elk on private land after hunting opened versus before, and the Julian date effect arose from the increase in elk on private land through time (the study period). The area x Julian day interaction arose from the steeper slope for the north area versus the south area, and the hunt season x Julian day interaction arose from the steeper slope before hunting season opened versus after it opened. �215 The experimental effect on the number of elk directly influenced by early-season hunting can be estimated by the abrupt increase in proportion of elk moving to private land when hunting opened. I used the difference in the model-averaged predicted proportion of elk on private land immediately before and immediately after opening day to estimate the direct effect of opening of hunting. Elk on private land increased 8-18% at the opening of early-season hunting (Table 1.5). During the study period, a significantly higher proportion of radiocollared elk moved to private land in the north treatment area compared to south area (P = 0.035; Table 1.6). On average, elk moved 613 m/day (95% CI = 594,632). The mean daily distance moved by elk before hunting season opened was 651 m/day (95% CI = 619, 683) and 575 m/day (95% CI = 540, 610) after opening (Fig. 1.6 and 1.7). During the study, the mean elevation of elk changed by -2.1 m/day (95% CI = -2.3, -1.9). Elk moved down in elevation an average of -2.4 m/day (95% CI = -2.9, -1.9) before hunting season opened and down -0.7 m/day (95% CI = -1.2, -0.2) after opening (Fig. 1.8 and 1.9). DISCUSSION Experimental Results Although elk movements in the White River area may not be solely caused by archery hunting activity, the experimental results show that archery hunting activity has an effect on elk movements despite other factors, especially in the northern treatment area. The strength of a 2 x 2 crossover design is that it blocks or controls for two sources of external variation, allowing for much stronger inference of cause and effect than the same design without a strict crossover (Ratti and Garton 1994). The crossover is also more efficient than a randomized design (Ott 1993); that is, a randomized design requires a larger sample size. In this experiment, area and year were the external sources of variation controlled for by the crossover design, and opening date of archery hunting was the treatment. Any factor that occurred on both halves of the study area, such as recreational activity, grazing, weather, forage phenology etc., that also caused elk to move, would lessen detection of an effect due to hunting activity. For example, if recreational activity caused some elk movement during the study period, then these recreationally induced movements would be unaffected by opening date of archery hunting and would serve to attenuate detection of archery hunting effects. If no elk were responding to archery hunting activity, either because they moved in response to another factor or they did not move at all, then I would fail to reject the null hypothesis and would conclude that archery hunting was not affecting elk movement. In the White River elk population, some elk may move regardless of what happens with archery hunting, because their movement was a response to some other factor; some elk may not move; and some elk may move due to changes in archery hunting activity. In the north treatment area, elk receiving the early treatment moved 14 days earlier than elk receiving the late treatment, indicating that some elk moved in response to archery season. Similarly, the effects of archery hunting on the daily proportion of elk on private land can also be distinguished from other effects. The positive slope for proportion of elk on private land resulted from steady movement of elk to private land during the study period and suggested that factors other than archery hunting may have affected elk movements. However, some of the movement to private land was directly attributable to hunting. That is, 23-44% of the radiocollared elk moved to private land during the study period, and 8-18% of this movement occurred at the opening of archery season. The significant jump in proportion of elk on private land at archery opening, beyond the steady increase, indicates that archery hunting activity did affect movement beyond that caused by other factors on both the north and south treatment areas. Sequence or carry-over effects can dilute detection of treatment effects in this type of study. A carryover effect occurs when the sequence of treatments affects responses to treatments (Ott 1993). In the case of White River elk, a carry-over effect would exist if south-area elk, receiving the early-opening treatment the first year, moved early the second year, before the late-opening date. A carry-over effect would be manifest on the north area if elk receiving the late-opening treatment the first year, moved late the second year, after the earlyopening date. In this experiment, elk on the north area moved slightly before opening date both years, while elk on the south area moved in accordance with opening date. Thus, I concluded that there was no carry-over effect due to the sequence of the treatments. I attributed differences in movements to area rather than sequence effects. A post hoc analyses on mean date of movement and daily proportion of elk on private land revealed that treatment effects were greater in the north area. When designing the study, I assumed that the north and south �216 areas were similar and would serve as controls for each other. Alleged elk movements to private land were perceived to be similar problem on both areas. The differences may have occurred because habitat, cover, topography, or any other area factors important to elk movement were different on the two treatment areas. Fortunately, each area received both treatments, so I could look at each area as its own control and evaluate each area separately. In the north treatment area there was a significant 14-day difference in mean date of movement between early and late treatments, while in the south treatment area there was a non-significant 6-day difference. In the north treatment area, about twice as many elk moved to private land (44%) compared to elk in the south area (23%). The difference in treatment response in the north and south areas may be due to cover and refuge configuration. In general, animals must balance the tradeoff between feeding and danger (Krebs and Davies 1993); elk may reduce use of open areas and use cover more frequently in times of perceived or real danger. Elk calves subjected to simulated surface-mining activities used coniferous forest significantly more often compared to undisturbed calves (Kuck et al. 1985). Similarly, elk subjected to oil drilling significantly increased use of forested habitats during drilling and post drilling periods, compared with the pre-drilling period (Van Dyke and Klein 1996). Where logging activities disturbed elk, areas of high elk use were negatively correlated to the amount of cover in the area (Edge and Marcum 1985, Edge et al. 1985, Czech 1991). In the White River area, the north area was rolling, with open parks interspersed with a few high peaks, while the higher elevation south area was essentially a plateau cut by steep narrow canyons and with steep, densely forested cliffs defining the plateau. These densely forested canyons and cliffs, inaccessible to motor vehicles, offered good refuge from hunters on public land. In contrast, the north area offered less shelter on public land, but had refuge in large private-land tracts that bordered the area. The south treatment area had less private land adjacent to public land, as BLM land was abundant at lower elevations. It may be that elk in the south area moved into forest refuges on public land after archery hunting opened, while elk in the north area moved to private-land refuges. More generally, elk movements during hunting seem to depend on location and quality of hiding cover. Elk decreased their daily movements after attaining sanctuary from hunting in national parks (Altmann 1956, Martinka 1969) or thick vegetation inaccessible to most hunters (Lemke 1975, Irwin and Peek 1979, Morgantini and Hudson 1979). I found a similar general pattern; elk on the north and south areas reduced their daily movements an average of 76 m/day after hunting season opened and they attained refuge from hunting on private land or in thick vegetation. a Scale Issues Three main issues became evident during the course of the study that fall under the rubric of large-scale field experiments: (1) a difference between movement responses for elk in the north and south areas, (2) a difference in the experimental results compared to pilot study results, and (3) failure of many elk to receive both treatments. As discussed, I think the difference in treatment responses between north and south elk were due to proximity of cover and private land hunting sanctuary. Additionally, I speculate that differences in wintering grounds and migration routes between the two treatment areas also affected movement responses. In the south treatment area, wintering grounds were at the south end of the study area. Southern elk were impaired from migrating beyond the study area by barriers created by Interstate 70 and the Colorado River. In the north treatment area, although private-land refuge was adjacent to the study area, wintering grounds were 30-80 km west of the study area. During the study period, elk moving to private land in the north area were just beginning their fall migration, whereas elk in the south area were completing their migration by moving to private land. It may be that there was no energetic cost to northern elk to move to private land in late summer, while southern elk would risk depleting their winter forage by moving to private land in late summer. There was less movement during.the experiment than I expected based on pilot study data. During the experiment, 21-48% of elk moved to private land compared to 70-100% during the pilot study. I think this disparity arose because the pilot study lacked a random sample from the entire area. Pilot study animals were collared primarily in areas with perceived movement problems. When the spatial scope of sampling expanded such that the entire study area was randomly sampled, many elk were collared in areas with no previous record of movement. Corresponding to the increased spatial variability in elk capture location was an increased variability in elk movement responses: elk in some areas did not move. The pilot study animals were not representatives of the entire study area, rather they represented areas with movement problems. �217 I designed this experiment with the intent that it be a crossover experiment, which used a crossover ANOVA model to test whether the mean date of elk movement from public land to private land was different between treatments. Exposure by each animal to both treatments was a requirement of the crossover analysis, and this requirement was impossible to implement in this large-scale field experiment. I did not have the control to ensure that each animal received both treatments; therefore, I analyzed the data as paired and unpaired data. Interpretation of mean date of movement results would have been more robust if all elk received both treatments. Mean date of movement between treatments was less dramatic for the paired analysis (5 days) compared to the unpaired analysis (10 days), though the difference was not significant. This difference may represent sampling variance; the small size of the paired sample makes this possibility likely. MANAGEMENT IMPLICATIONS Although there was a statistically significant effect of archery hunting on timing and number of elk moving to private land in the north treatment area, the question remains as to whether this effect was significant to elk management. The experimental effect on number of elk moving can be estimated by the abrupt increase in proportion of elk moving to private land when hunting opened. This jump was 8-18%; hence, the CDOW can reduce, at most, approximately 20% of the elk movement to private lands by manipulating archery hunting activity. However, 20% of a herd may represent a large enough number of animals to make management changes worthwhile. Additionally, if elk movements are concentrated in problem areas, then manipulation of opening date may result in a proportionally higher reduction of elk movements in those areas. Managers must decide whether the contribution of archery hunting activity to elk movement is large enough to warrant management action. Elk responded to archery hunting activity differently in north and south treatment areas; I concluded that archery hunting activity had little or no effect in the south area, but had a statistically significant effect on timing and proportion moving in the north area. Thus, managers concerned with elk movement need to be cautious if citing White River results as a reason for management actions; elk movement was dependent on the habitat and topography of an area, and these factors must be considered in any plan to reduce elk movements. Furthermore, because this study manipulated opening date of hunting activity but kept hunter numbers constant, the question remains of what percentage of the elk would still move ifhunting were reduced or eliminated. Managers may also want to consider the time needed to make permanent changes in elk movement patterns. Elk responses to disturbance may be long- or short-lived depending on the type of disturbance and the type of elk response. For example, studies that evaluated elk movements after disturbances ceased found that displacement appears to be temporary in most situations. Elk displaced by cross-country skiers returned to their original locations after people left the area (Cassirer et al. 1992). During weekends, and immediately following the end oflogging activities, elk moved back into logged areas (Ward 1976, Edge and Marcum 1985). Even for more potentially lethal hunting disturbances, mean distances to roads returned to pre-hunting distances after hunting season closed (Hershey and Legee 1982, Wright 1983). Zahn (1974), Lemke (1975), and Hershey and Legee (1982) also found that elk response to hunter disturbance was short-lived. In the White River area, this short-lived response seems to characterize south-area elk. In 1996, hunting season opened early, 24 August, in the south area, and closed 14 September. A drop in the proportion of elk on private land occurred after closing date (Fig. 1.4a), indicating that some elk moved back to public land after hunting season closed. This was also seen in elevation shifts of south-area elk: after hunting season closed in 1996, mean daily elevations increased (Fig 1.9a), indicating that elk moved back up to public land after the disturbance ended. Elk in the south treatment area seem to exhibit a short-term response to hunting disturbances. On the other hand, if the elk response to disturbance was a migration shift, then the movement response may last longer. Elk that have learned to move to unhunted refuges and choose migration routes through lightly hunted areas (Altmann 1956, Martinka 1969) may continue these patterns if no cost was incurred by the change. In the north area, the steady movement may be only partially a direct effect of hunting season. Some elk may learn to move to private land as a conditioned response to avoid hunting danger, and this movement may have become paired with some unconditioned stimuli, such as shortening day length. Because movement in the north area is the beginning of migration, and because timing of migration is thought to be stimulated by daylight cues (Krebs and Davies 1993), north-area elk may now move when the days begin to shorten in addition to the stimulus of hunter activity. As older cow elk have been found to assume leadership roles within a group (Franklin and Lieb 1979), �218 social learning may pass on this pattern to future generations. Thus, elk on the north area may move to private land in response to direct cues of hunter activity and indirect cues of day length. When considering elk movements in response to hunting activity, wildlife managers should consider the type of movements being exhibited by the elk as well as the refuge configuration in the area. Although reducing hunter numbers may reduce movement where elk responses to disturbance are short lived (probably short movements to habitat refuges), different management practices may be needed where migration patterns have become a ftxed behavior. If elk have established a migration pattern onto private land, private land hunts, on lands to which elk move, may be a good solution to break the migration pattern and deter early elk movements onto private land. LITERATURE CITED Akaike, H. 1973. Information theory and an extension of the maximum likelihood principle. Pages 267-281 in B. N. Petran and F. Csaki, editors. International symposium on information theory. Second edition. Akademiai Kiadi, Budapest, Hungary. Altmann, M. 1956. Patterns of herd behavior in free-ranging elk of Wyoming, Cervus canadensis nelsoni. Zoologica 41 :65-71. Barrett, M. W., J. W. Nolan, and L. D. Roy. 1982. Evaluation of a hand-held net gun to capture large mammals. Wildlife Society Bulletin 10:108-114. .. Boyce, M S. 1991. Migratory behavior and management of elk (Cervus elaphus). Applied Animal Behavioral Science 29:239-250. Boyd, R J. 1970. Elk of the White River Plateau, Colorado. Colorado Division of Game, Fish and Parks Technical Publication 25, Fort Collins, Colorado, USA Buckland, S. T., K. P. Burnham, and N. H. Augustin. 1997. Model selection: an integral part of inference. Biometrics 53:603-618. Burnham, K. P., D. R Anderson, and G. C. White. 1995. Model selection strategy in the analysis of capturerecapture data. Biometrics 51:888-898. Burnham, K. P., and D. R Anderson. 1998. Model selection and inference: a practical information-theoretic approach. Springer-Verlag, New York, New York, USA Bryant, L. D., and C. Maser. 1982. Classification and Distribution. Pages 1-59 in J. W. Thomas and D. E. Toweill, editors. Elk of North America: ecology and management Stackpole Books, Harrisburg, Pennsylvania, USA Cassirer, E. F., D. J. Freddy, and E. D. Ables. 1992. Elk responses to distmbances by cross-country skiers in Yellowstone National Park. Wildlife Society Bulletin 20:375-381. Czech, B. 1991. Elk behavior in response to human distmbance at Mount St Helens National Volcanic Monument. Applied Animal Behavioral Science 29:269-277. Edge, W. D., and C. L. Marcum. 1985. Movements of elk in reaction to logging disturbances. Journal of Wildlife Management 49:976-930. Edge, W. D., C. L. Marcum, and S. L. Olson. 1985. Effects oflogging activities on home-range fidelity of elk. Journal of Wildlife Management 49:741-744. Franklin, W. L., and 1. W. Lieb. 1979. The social organization of a sedentary population of North American elk: a model for understanding other populations. Pages 185-198 in M. S. Boyce and L. D. Hayden-Wing editors. North American elk: ecology, behavior, and management. University of Wyoming, Laramie, Wyoming, USA Freddy, D. J. 1987. The White River elk herd: A perspective, 1960-85. Colorado Division of Wildlife Technical Publication 37, Fort Collins, Colorado, USA Gray, J. P., G. Byrne, and 1. Madison. 1994. White River elk data analysis unit plan: game management units: 11,211,12,13, i31 ,231 ,23 ,24 ,25,26,33. Colorado Division of Wildlife Internal Report, Grand Junction, Colorado, USA. Hershey, T. 1., and T. A. Leege. 1982. Elk movements and habitat use on a managed forest in north-central Idaho. Idaho Department ofFish and Game Wildlife Bulletin 10, Boise, Idaho, USA. Irwin, L. L., and 1. M. Peek. 1979. Relationships between road closures and elk behavior in northern Idaho. Pages 199-204 in M. S. Boyce and L. D. Hayden-Wing editors. North American elk: ecology, behavior, and management. University of Wyoming, Laramie, Wyoming, USA Knight, R R 1970. The Sun River elk herd. Wildlife Monographs 23. Krebs, 1. R, and N. B. Davies. 1993. An introduction to behavioural ecology. Third edition. Blackwell Science Ltd., Oxford, England. �219 Kuck, L., G. L. Hompland, and E. H. Merrill. 1985. Elk calf response to simulated disturbance in southeast Idaho. Journal of Wildlife Management 49:751-757. Leberton, J-D., K. P. Burnham. 1. Clobert, and D. R Anderson. 1992. Modeling survival and testing biological hypotheses using marked animals: a unified approach with case studies. Ecological Monographs 62:67-118. Lemke, T. O. 1975. Movement and seasonal ranges of the Burdette Creek elk herd, and an investigation of sport hunting. Montana Fish and Game Department Final Report 32.01, Job Number BG-3.15, Missoula, Montana, USA. Martinka, C. 1. 1969. Population ecology of summer resident elk in Jackson Hole, Wyoming. Journal of Wildlife Management 33:465-48l. Morgantini, L. E., and R 1. Hudson. 1979. Human distribution and habitat selection by elk. Pages 132-139 in M. S. Boyce and L. D. Hayden-Wing editors. North American elk: ecology, behavior, and management. University of Wyoming, Laramie, Wyoming, USA. Ott, R L. 1993. An introduction to statistical methods and data analysis. Fourth edition. Wadsworth, Inc., Belmont, California, USA. Ratti, J. T., and E. O. Garton. 1994. Research and experimental design. Pages 1-23 in S. Hieb, editor. Research and management techniques for wildlife and habitats. Fifth edition. The Wildlife Society, Bethesda, Maryland, USA. SAS Institute. 1990. SAS/STAT® user's guide, version 6.0. Fourth edition. SAS Institute, Inc., Cary, North Carolina, USA. .SAS Institute. 1993. SAS/STAT® software: The GENMOD procedure, release 6.09. SAS® Technical Report P243, SAS Institute, Inc., Cary, North Carolina, USA. .. Seber, G. A. F. 1982. The estimation of animal abundance and related parameters. Second edition. Edward Arnold, London, England. 'Strohmeyer, D. C., and J. M Peek. 1996. Wapiti home range and movement patterns in a sagebrush desert. Northwest Science 70:79-87. Van Dyke, F., and W. C. Klein. 19%. Response of elk to installation of oil wells. Journal ofMamrna1ogy 77: 1028-1041. Ward, A. L. 1976. Elk behavior in relation to timber harvest operations and traffic on Medicine Bow Range in south-central Wyoming. Pages 32-43 in S. Hieb, editor. Proceedings of the Elk-Logging-Roads Symposium. University of Idaho, Moscow, Idaho, USA. Wright, K. L. 1983. Elk movements, habitat use, and the effects of hunting activity on elk behavior near Gunnison, Colorado. Thesis, Colorado State University, Fort Collins, Colorado, USA. Zahn, H. M. 1974. Seasonal movements of the Burdette Creek elk herd. Montana Fish and Game Department Final Report 32.01, Job Number BG-3.13, Missoula, Montana, USA. �220 Table 1.1. Description and representation of a priori (models 1-12) and post hoc (models 11-13) models relating effects of year, area, opening of archery hunting season, and Julian date to daily proportion of radiocollared elk found on private land (PI) from 20 July to 10 October in the White River area, Colorado, 1996-1997. Hypothesis Model structure 1. Global model: year, area, opening of archery hunting, Julian day, and all possible interactions f30+f31 (y)+f32(A)+f33(S)+f34(D)+f3s(yxA)+f36(YXS) +f37CYxD )+f3s(Ax S)+f39(AxD)+f310(SxD) +f311(yxAxS)+f312(yxAxD)+f313(yxSxD) +f314(AxSxD)+f31S(yxAxSxD) 2. No year x date interactions f30+f31 (y)+f32(A)+f33(S)+f34(D)+f3s(yxA)+f36(YXS) +f3~AxS)+f3s(AxD)+f39(SxD)+f310(YxAxS) +f3u(AxSxD) 3. No year x main effects (A and S) interactions f30+f31(y)+f32(A)+f33(S)+f34(D)+f3s(yxD) +f36(AxS)+f3~AxD)+f3s(SxD)+f39(AxSxD) 4. No year effects f3o+f31(A)+f32(S)+f33(D)+f34(AxS)+f3s(AxD) +f36(SxD)+f3~AxSxD) 5. No area x date interactions f30+f31 (y)+f32(A)+f33(S)+f34(D)+f3s(yxA)+f36(YXS) +f37CYxD)+f3s(AxS)+f39(SxD) +f31o(yxAxS)+f3ll(yXSxD) .. 6. No area x main effects (Yand S) interactions f30+f31(y)+f32(A)+f33(S)+f34(D)+f3S(yXS)+f36(yxD) +f3~AxD)+f3s(SxD)+f39(Y xSxD) 7. No area effects f30+f31(Y)+f32(S)+f33(D)+f34(yXS)+f3s(yxD) +f36(SxD)+f37CYxHxD) 8. No hunt season x date interactions f30+f31 (y)+f32(A)+f33(S)+f34(D)+f3s(yxA)+f36(YXS) +f37CYxD)+f3s(AxS)+f39(AxD)+f310(yxAxS) +f3ll(yxAxD) 9. No hunt season x main effects (A and Y) interactions f30+f31(y)+f32(A)+f33(S)+f34(D)+f3s(yxA)+f36(yxD) +f3~AxD)+f3s(SxD)+f39(Y xAxD) a 10. No hunt season effects f3o+f3.(A)+f32(Y)+f33(D)+f3iAxY)+f3s(AxD) +f36(YxD)+f3~Ax Y xD) 11. Only hunt season effects f3o+f3.(S)+f32(D)+f33(SxD) 12. Only hunt season and area effects f3o+f31(A)+f32(S)+f33(D)+f3iAxS)+f3s(AxD)+f36(SxD) 13. Only hunt season and area effects with no S x A interaction f3o+f3.(A)+f32(S)+f33(D)+f34(AxD)+f3s(SxD) • Y represents year with 1996 = 0 and 1997 = 1, A represents treatment area with south area = 0 and north area 1, S represents archery hunting season with 0 = before opening and 1 = after opening, and D represents the covariate date, which is Julian day. The dependent variable (PI) was in logit scale. = �221 Table 1.2. Days between opening dates, least-square mean differences with ±95% CI, and P-value for the null hypothesis of no difference between mean date of movement by treatment, year, and area for radiocollared elk in the White River study area, Colorado 1996-1997. Days between opening dates Treatment effect: (late treatment - early treatment) Days between mean dates of movement P 20 10±9 0.013* Year effect: (1997 - 1996) 1 4±9 0.346 Area effect: (south area - north area) 0 7±8 0.100 * One-sided test Table 1.3. Differences between opening dates, least square mean differences with ±95% CI, and one-sided P-value for the null hypothesis of no difference between mean date of movement for radiocollared elk in the White River study area, Colorado 1996-1997. Treatment area Days between Opening dates Days between mean dates of movement (late treatment - early treatment) P North 20 14± 10 0.006 South 20 6± 15 0.218 �222 Table l.4. Ranking of a priori (models 1-12) and post hoc (models 11-13) hypothesized models relating effects of year, area, opening of archery hunting season, and Julian day to daily proportion of radiocollared elk found on private land (PI) from 20 July to 10 October in the White River area, Colorado, 1996-1997. Models were ranked by QAICc values and normalized QAICc weights ( 11) m ). Model structure Model number a ~o+~1(A)+~2(S)+~3(D)+~4(AxD)+~5(SxD) ~o+~1(A)+~2(S)+~3(D)+~4(AxS)+~5(AxD) +~6(SxD) ~o+~1(Y)+~2( A)+f33(S)+~4(D)+~5(Y xA) +~6(Y xD)+~~AxD)+~8(SxD)+~9(Y xAxD) 13 12 9 b K QAICc AQAICc Wm 6 7 10 4718.68 4720.53 4721.92 0.00 l.85 3.24 0.495 0.196 0.098 C ~o+~1(A)+~2(S)+~3(D)+~4(AxS)+~5(AxD)+~6(S;D) +~~AxSxD) 4 8 4722.42 3.74 0.076 ~O+~1(y)+f32(A)+f33(S)+~4(D)+~5(YXS) +~6(Y xD)+~~AxD)+~s(SxD)+~9(Y xSxD) 6 10 4723.37 4.69 0.048 ~O+f31 (Y)+~2(A)+~3(S)+~4(D)+~5(Y xA) +~6(YxS)+~~AxS)+~8(AxD)+~9(SxD) +~lo(YxAxS)+f311(AxSxD) 2 12 4723.57 4.90 0.043 ~O+~1(y)+f32(A)+~3(S)+f34(D)+~5(YxD) +~6(AxS)+~~AxD)+~8(SxD)+~9(AxSxD) 3 10 4724.11 5.43 0.033 ~O+f31(y)+f32(A)+f33(S)+f34(D)+f35(YxA)+f36(YXS) +f3.,(YxD)+f3s(AxS)+f39(SxD)+f310(YxAxS) +f311(yxSxD) 5 12 4726.6 7.92 0.009 1 ~O+f31(y)+f32(A)+f33(S)+f34(D)+f35(YxA)+f36(YXS) +f3.,(YXD)+f38(AxS)+f39(AxD)+f310(SxD)+f311(yxAxS) +f312(yxAxD)+f313(yXSxD)+f314(AxSxD) +f315(yxAxSxD) 16 4730.41 11.74 0.001 f30+f31(Y)+~2(A)+f33(S)+f34(D)+f35(YxA)+f36(YXS) +f3.,(YXD)+f38(AxS)+~9(AxD)+f310(YxAxS) +f311(yxAxD) 8 12 4733.36 14.68 0.000 f3o+f31(A)+f32(y)+f33(D)+f34(Axy)+f35(AxD) +f36(YxD)+f3~Ax Y xD) 10 8 4745.57 26.89 0.000 ~O+f31(y)+f32(S)+f33(D)+f3 4(YxS)+f35(YxD)+f3iSxD) +f3~SxYxD) 7 8 4845.74 127.07 0.000 11 4 4855.07 136.39 0.000 f30+f31(S)+f32(D)+f33(SxD) • Y represents year with 1996 = 0 and 1997 = 1, A represents treatment area with south area = 0 and north area = 1, S represents archery hunting season with 0 = before opening and 1 = after opening, and D represents the covariate date, which is Julian day. The dependent variable (PI) was in logit scale. b Numbers correspond to those in Table l.1 C Number of estimable parameters. �223 Table 1.5. Model-averaged estimates of hunt season effect represented by the difference of proportion of radiocollared elk on private land immediately before and after opening day and 95% confidence intervals; White River area, Colorado, 1996-1997. trreatment Area 1997: 1996: 1996: 1997: North -early North -late South - early South - late Estimated proportion of elk on private land Immediately Immediately Difference 95%CI of difference before opening after opening 0.095-0.257 0.408 0.584 0.176 0.542 0.621 0.078 -0.001-0.158 0.089-0.231 0.160 0.256 0.416 0.099 0.028-0.170 0.317 0.416 Table 1.6. Mean proportion of radiocollared elk on private land during 1996-1997 for each treatment area in the White River area, Colorado, at the beginning (19 July) and end (10 October) of the study period. Treatment area North South Mean proportion of elk on private land 1996-1997 July 19 October 10 Change 95% CI of change 0.200 0.161 0.636 0.393 0.436 0.232 0.296-0.577 0.105-0.360 �224 t N km o 10 20 .~ BLM Land . Flat Tops Wilderness . fill -, <- e. GMU Boundaries ~ state Wildlife Land •• Forest Service Land White River :. Towns o study Area Boundary Colorado Figure 1.1. Alleged elk movements from high elevation public land to lower elevation private land, and land ownership in the White River study area, Colorado. 7000 ~ Q) .•... 6000 C ",. - •... -, , :::J ..c c 0 5000 (/J a:J Q) (/J 4000 I >. "C a:J 3000 0 2000 Q) L- a> .0 5 z 1000 o +---~---.---.---.r---~--.---~---.---.--~ 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 Year * Variance estimates not available 1984-1987. Figure 1.2. Number of archery and muzzleloading hunters using GMUs 12, 23, 24, and 33 in northwest Colorado, 1984-1994. Data from Deer, Elk, and Antelope Management (DEAMAN) database and software, Colorado Division of Wildlife, unpublished data. �225 t N .~ BlM Land :[ill Flat Tops Wlldemess ~2 State Wildlife Land km o 10 o Study Area Boundary •. Forest Service Land 20 .• Elk Capture locations •. Towns Figure 1.3. North and south treatment areas and July 1996 elk capture locations in the White River study area, Colorado. �226 Early Opening Late Opening a c 0.9 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 "'C c: ca Q) .•... ca > -I:: 0.7 .. 0.5 .•..--.. -!.~ ... ------.-_"!' - _. _. - _ .. - - - - .• -- -. ......,. 1'.•... _.- .. .... ~-- - ~. 0.4 - .• -.- -----.-.-~- .~.,..... -. 0.6 0.3 0.2 South 96 eo .•.. r:::: C. Opening date 0.8 Opening date North 97 0.1 0.0 .•.. ~ <0 co .•.. t:: IS) c: 0 - ~ Q) b d 0 0.9 c: 0.8 0 t 0 0.9 Opening date • -_ ....•...•..... 0.7 ... 0.6 0.8 0.6 c. 0.5 0.5 L- 0.4 0.4 0.3 0.3 0 a.. Opening date 0.7 ... - .. _.-- .•... .. ------ -- ...---.- 0.2 0.2 North 96 0.1 South 97 0.1 0.0 0.0 ex) ex) t:: t:: .•.. .•.. + Raw data ___ Predicted ___ 95%CI Figure 1.4. Model averaged predicted values, 95% CI, and raw data of proportion ofradiocollared elk on private land versus date for (a) early treatment on the south area 1996, (b) early treatment on the north area 1997, (c) late treatment on the north area 1996, and (d) late treatment on the south area 1997, July-October in the White River study area, Colorado. .. �227 0.9 ,---------------------------------------------, 0.8 a Early opening fitted curve\ 0.7 0.6 0.5 0.4 ~ • ••••• • ....... _. .~~-.. 0.3 ~ ~ __ .. • • d :.. .!!!! •••• • • - 0.2 •• Late opening • Late opening fitted curve 0.1 • Early opening 0.0 +---~--r__.~~--~--_r--~--,_--~--.__.--~ ----..- 1.0 f'- f'- co N ..-co co co ..-co --- <» 1.0 <» ~ co co Date 1.0 N ~ ..-<» <»..-en N <0 ~ <» - ..- ..-..- C") 0 0 0 0.9 0.8 b 0.7 0.6 0.5 0.4 • Late opening • Early opening Early opening fitted curve \ ••••• •....... ' .. 0.3 0.2 0.1 ..\ . - .. -.__.-; •• • , .• ••• • • I • ••• Late opening fitted curve 0.0 +-----.-.----.---r-..,...---.------.--.--------.-..------.---.' <» ~ ~ ~ co co <» Date ..- - -..-..o o Figure 1.5. Model averaged predicted values and raw data of proportion ofradiocollared elk on private land versus date for (a) north-treatment area and (b) south-treatment area during July-October 1996-1997 in the White River study area, Colorado. >. -c_tV E 1600~------------------------------------_. 1400 '-' 1200 -South early 1996 -North early 1997 South late 1997 Early opening Late opening .• " , - - -North late 1 -c Q) ~ 1000 Q) g 800 tV .•... :6 ~ ·iii -c 600 400 c: tV Q) 200 :iE 0 IX) .•.... -- t-- - - It) .•.... IX) N IX) IX) t-- - It) .•... N N IX) IX) -- - - - - 0) It) N 0) IX) Date N .•.... 0) .•.... <0 N 0) 0) 0) -- ('") a a .•.... a Figure 1.6. Mean distance moved per day by radiocollared elk during July-October 1996-1997 in the White River study area, Colorado. �230 �231 CHAPTER 2: EFFECT OF HUNTER DENSITY ON ELK MOVEMENT TO PRNATE LAND IN NORTHWEST COLORADO INTRODUCTION Animals may modulate their response to stimuli depending on the quality, quantity, context, and intensity of the stimuli. Prey species, such as elk (Cervus elaphus), may show responses to varying densities of predators, such as hunters. Elk may increase their use of a refuge to avoid hunters; that is, as the density of predators increases, so too does the probability that elk are found in a refuge area. Or, elk may ignore hunters at low densities, and move to refuge areas only when hunter densities increase above some tolerance and danger threshold. To avoid hunters, elk have moved to refuges in national parks (Altmann 1956, Martinka 1969), private land (Wright 1983, Chapter 1), or densely vegetated areas (Irwin and Peek 1979, Morgantini and Hudson 1979) adjacent to their typical areas of activity. However, little research as been done on general elk responses to differing levels of hunter pressure, and no research has directly evaluated the effects of hunter density on elk use of refuge. Since the 1960s, the number of early-season (archery and muzzleloading) hunters using the White River area of northwest Colorado has grown, with especially significant increases between 1984 and 1992 (Fig. 2.1, DEAMAN, Colorado Division of Wildlife, unpublished data). Typically, archery hooters did not hunt private lands; thus, private lands provided elk a refuge during early-season hunts. As archery and muzzleloading hunter numbers increased, so too did complaints about early elk movements onto private land. For the Colorado Division of Wildlife (CDOW), the primary problem was herd size: it was difficult for CDOW to maintain or reduce herd size via harvest because hunters have limited access to private lands, (1. Madison and C. Reichert, Colorado Division of Wildlife, personal communication). Managers hypothesized that reduction of early-season hunter pressure, achievable through limiting licenses, would reduce movement onto private land. As part of an experiment about elk movement in response to early-season hunting activity, CDOW conducted an intensive hunter survey in the fall of 1996 and 1997. The study area was split into 2 treatment areas. A telephone survey of early-season hunters was conducted to estimate the daily number of hunters using each treatment area with a 95% confidence interval of ±l50 hunters (total confidence interval of300 hunters). I used these data, along with elk location data, to determine how the density of early-season hunters affected elk movement to private land. My objectives were to: (I) summarize hunter survey data and provide estimates of hunter numbers, hunter-days, and mean days hunted per hunter by treatment area, Game Management Unit (GMU), hunt code, and day, (2) compute these estimates for hunters using ATVs, and (3) evaluate the relationship between hunter density and elk movement to private land. I used logistic regression and Akaike's Information Criteria (AlC) model selection (Burnham and Anderson 1998) to evaluate the relationship between hunter density and elk movement to private land. STUDY AREA The White River area of northwestern Colorado covered approximately 4,540 km2 and was composed of Game Management Units (GMUs) 12,23,24, and 33. In Colorado, GMUs were delineated to distribute hunters through allocation of hunting licenses. White River area land ownership was 34% private land and 66% public land (Fig 2.2), with 82% of public land being United States Forest Service (USFS). USFS land was mostly at high elevation and toward the center of the study area, with some Bureau of Land Management (BLM) areas at lower elevations in the southern sections of the study area. Private land was lower in elevation and comprised mainly ranches and coal mines. Topography, climate, and vegetation varied widely throughout the study area. Elevation ranged from 1,629 to 3,700 m. The central part of the study area was high elevation public land, split by the White River valley. The high elevation areas dropped off to lower elevations to the north, south, and western edges of the study area. Higher elevations had severe winters with heavy snowfall, while lower elevations had comparatively mild winters. Mean annual precipitation at 3,000 m in the National Forest areas was about 100 em, compared �232 with about 30 em at lower elevations of <2,000 m. Vegetation types at elevations >2,600 m were in the montane/subalpine zone and included groves of aspen (Populus tremuloides), Engelmann spruce (Picea engelmanni), and alpine fir (Abies lasiocarpa) interspersed with grassy meadows. Middle elevations (1,9802,600 m) were a transitional zone that consisted primarily of pinion pine (Pinus edulis), juniper (Juniperus scopulorum), and big sagebrush (Artemisia lridentata). Lower elevations «2,000 m) in the southern and northern parts of the study area were in the Great Basin zone with sage grasslands. Oakbrush patches were found at the middle and lower elevations with major shrubs that included Gambel's oak (Quercus gambelii), servicebeny (Amelanchier alnifolia), mountain mahogany (Cercocarpus montanus), chokecheny (Prunus virginiana), snowberry (Symphoricarpos ulahensis), bitterbrush (Purshia lridenlala), and rabbitbrush (Chrysothamnus nauseosus). Higher and middle elevation areas provided summer and fall forage for elk, while lower elevations were typically used by elk during winter. Boyd (1970) provides a detailed description of the study area. Hunting for large game was economically important to local residents in the study area (Grayet al. 1994). Guides, outfitters, local service sectors, and private landowners (who sell trespass permits) made a large proportion of their annual income during the elk and deer hunting seasons, with most hunting revenues accrued during rifle hunting season. METHODS Radio-telemetry Data CoUection As part of a study on effects of early-season opening date on elk movements to private land, the study area was divided roughly east-west into 2 halves along the White River and North Fork of the White River (Fig. 2.2). GMU 12 and parts ofGMUs 23 and 24 composed the north treatment area, while GMU 33 and parts of GMUs 23 and 24 composed the south treatment area. Two treatments were applied: an archery season that opened 1 week earlier, and another that opened 2 weeks later than normal, yielding a 3-week difference in opening dates. Each area received both an early- and late-opening treatment. During the first year of the study, the early-opening treatment was randomly assigned to the south treatment area. Treatments were reversed the second year of the study. Thus, in 1996, archery hunting opened early, 24 August, in the south area, and late, 14 September, in the north area. When treatments were reversed in 1997, archery hunting opened early, 23 August, in the north area, and late, 13 September, in the south area. Archery season was 4 weeks long during the earlyopening treatment and 3 weeks long during the late-opening treatment. Muzzleloading season was 1 week long and nested within archery season. Opening of muzzleloading season coincided with opening of archery season for the late-opening treatment, and occurred on the third weekend during the early-opening treatment. The 80 adult female elk used in this study were captured from random locations throughout the study area. Chapter 1 describes capture and collaring procedures in detail. Elk locations were collected 20 July-l0 October 1996-1997. I relocated radiocollared elk between 0700-1500 hr using a Cessna 182 or 185 fixed-wing aircraft with a 2-element Yagi antenna mounted to each strut of the airplane. For each elk relocation, Universal Transverse Mercator (UTM) coordinates were recorded with a Global Positioning System that was not differentially corrected. During the early-season hunt, I collected elk locations 2 times a week with 2-4 days between collections. A Geographical Information System (GIS) map was used to record elk locations, land ownership, and treatment boundaries. The GIS map was digitized from a United States Geological Survey map at a scale of 1:100,000. For each location of a telemetered cow, a 0 was assigned if the location was on public land (e.g. Flat Tops Wilderness Area, USFS, State Wildlife, and BLM lands) and a 1 was assigned if the location was on private land. Hunter Survey Data Collection Number of archery and muzzleloading hunting licenses for elk and deer in 1996 were based on the mean number calculated from 1990 - 1994 (4,985; Colorado Division of Wildlife unpublished data). Restricting licenses kept hunter density consistent during the study and consistent with years of high elk movement complaints. In the fall of 1996 and 1997, after archery and muzzleloading season ended, CDOW conducted a �233 telephone survey with a goal of sampling enough hunters to estimate the number of hunters afield on any day with a 95% confidence interval of±150 hunters (total confidence interval width = 300 hunters; Appendix I). Sample size was based on the binomial distribution and was conservative. That is, the maximum variance occurs when 50% of the license holders hunt, and enough license holders were surveyed to meet the ±150 hunters confidence interval under this condition. CDOW conducted the telephone survey following protocols typically used to collect hunter data. The sample was a simple random sample. The initial sample size was larger than required, with an expectation that 70-90% of hunters called will respond with a complete survey. If this requirement was not met, then additional hunters were again randomly chosen from those not sampled the first time. The survey began as soon as the hunting season closed, and continued for approximately 4 months. Telephone surveys have been found to provide valid estimates of harvest and days hunted (White 1993, Steinert et aI. 1994). Hunt codes: D-E-012-01-A E-E-012-Ol-A E-F-012-01-M E-M-OI2-01-M D-M-OI2-01-M D-E-033-01-A E-E-033-01-A E-F-033-01-M E-M-033-01-M D-M-033-01-M - archery deer, either sex, on north area, archery elk, either sex, on north area, muzzleloading elk, cow, on north area, muzzleloading elk, bull, on north area, muzzleloading deer, buck, on north area, archery deer, either sex, on south area, archery elk, either sex, on south area, muzzleloading elk, cow, on south area, muzzleloading elk, bull, on south area, and muzzleloading deer, buck, on south area, were surveyed. Hunters that purchased two licenses on the same treatment half (e.g. D-E-OI2-01-A and E-EOI2-01-A) were counted as one hunter and were proportionally assigned to hunt-code groups. The number of individuals purchasing licenses was the size of the statistical population sampled for estimates, and was used in all estimates of hunter number, hunter-days, and average days/hunter. Hunters were asked: 1. 2. 3. 4. 5. 6. 7. License identification number, Hunt code; If they hunted or did not hunt; In which GMUs they hunted; How many days they hunted per unit; If they hunted on day 1, on day 2, on day 3, etc. of the 7season; and If they used an ATV. From these data, 3 categories of information were estimated for hunt-code group, treatment area, and GMU: Number of hunters that hunted and number of hunters that used ATVs; Number of hunters afield per day, and number of hunters with ATVs afield per day; and Number of hunter-days and average number of days in the field per hunter and per hunter using an ATV. The following notation is used in estimator formulas, with i, j, k, l, and m indexing the group for which the estimator is being used: ",," i .. j k l m - day (day 1 is opening day, day 2 is 2ndday of the season etc.), 'c'" hunt code group (north treatment area contained hunt code groups: DE01201A, DM01201M, EE01201A, EF01201M, EMOI201M; south treatment area contained hunt code groups: DE03301A, DM03301M, EE03301A, EF03301M, EM03301M), GMU (12, 23, 24, or 33), individual hunter, and treatment area (north or south half of study area). �234 For example, Him is the estimated number of hunters afield on day i in treatment m, Hi/an is the estimated number of hunters afield on day i in GMU k, treatment area m, Hlan is the estimated number of hunters that hunted in GMU k, treatment area m, etc. Additional notation is: N Number of individuals purchasing licenses, N'· - Number of licenses sold, n Number of license holders surveyed who responded with at least partial information, Number of license holders surveyed who reported hunting, n , - a H- Number of license holders surveyed who reported hunting with an ATV, Estimated number of hunters, A Estimated number of hunters using ATVs, Number of hunter-days reported for surveyed hunters, Number of hunter-days reported for surveyed hunters using ATV s, - d 1 u D - Number of hunter-days reported for surveyed hunters in a particular GMU, Estimated number of hunter-days, ._ . F - Estimated number of hunter-days for hunters using ATVs, d - Mean number of days afield per hunter, - Estimated probability that a license holder hunted, and 1 p r Mean number of days afield per hunter using an A TV, Estimated probability that a license holder hunted with an A TV. Day i, hunter I, and A TV use were entered as 0 or I. For example,. if a surveyed license holder hunted, then a 1 was entered, or, if they did not hunt, then a 0 was entered. Similarly, if a hunter hunted on-day i, then a I was entered, or, if the hunter did not hunt, then a 0 was entered for that day. If a hunter used an ATV . ~ then a 1 was entered, and the hunter was assumed to use the A TV every day hunting. The finite correction factor, (N';;, n) , was used for all estimates. N~te that N' was used in the finite correction because each license, and not individual, was randomly sampled. Thus, a license holder purchasing 2 licenses could be interviewed for 0, I, or 2 of the licenses. The number of licenses sold varied between 150- I ,219 by hunt code and the finite correction factor varied between 0.45-0.72 (Table 2.4 and 2.5). The 95% confidence intervals were constructed similarly for all variables, e.g.: or, where sample sizes were <100, the appropriate t-statistic was used in lieu of 1.96: Estimating Number 01Hunters Estimates of number of hunters ( H ) and their associated variances were calculated for each hunt code group (J) separately, then summed over huntcode groups{i) for treatment (m) or GMU (k). The number of hunters from hunt code group (j) that hunted in treatment area (m) was estimated: �235 5 H m =LHj and , j=1 5 Var(Hm) = L vM(H j)' j=1 Similarly, the number of hunters using ATVs (A) from hunt code group (j) that hunted treatment area (m) was estimated: ;.•.r)_ -, vCU\rj (N'· -n·) ;·(1-;·) J J J Nj J , nj 5 Am = LAj ,and j=1 5 vfu-{Am) L vM(A j) = . j=1 The number of hunters (H) from hunt code (J) that hunted in GMU (k) and treatment area(m) was estimated: A Pjk n"k J =-, nj 5 if Ian = L if jk j=1 , and �236 5 var(H Ian) = "I v3.r(H jk) . j=i Similarly, the number of hunters with ATVs (A) from hunt code (j) that hunted in GMU (k) and treatment area (m) was estimated: (N'· -n·) i·k(l-i·k) :;',.{A) vCU\rjk = J , J J Nj 5 Alan = LAjk J nj , and j=i 5 viJ.r(Akm) = L v3.r(Ajk)· j=i Estimating Number of Hunters Afield per Day Estimates of total number of hunters (iI) afield per day (l) and their associated daily variances were calculated for each hunt code group (j) separately, then summed over hunt code groups (j) for treatment (m) .. or GMU (k). The number of hunters afield on day (I) within treatment (m) was estimated: A Pij , =-, n..lj nj A Him 5 LHij, j=i A = and ,.," 5 vilr{ iI im ) = "I yare iI ij ) . j=i Similarly, the number of hunters afield using ATVs (A) estimated: on day (I) within treatment area (m) was �237 a.,lJ r... A =- lJ nj ' yar(; ..) = lJ (N'· -n·) ; ..(1-;..) J J lJ lJ N' , nj j :;'..tAA) :;'..tN v••.• \ ij = v••.• \ jrijA) 5 A Aim = A A = N j2 yar(rij)' L Aij , and A j=1 5 vfu-(A;m)= Lvfu-(Aij)' j=1 The number of hunters (I) afield (H) on day (I) in GMU (k), treatment area (m) was estimated: nj A :;'..t-) V••.•\Pijk A = 5 A ~ 2 (N'. _ n.) L (Pijkl - Pijk) J J 1=1 N'. =----J nj(nj-I) A H;km = L Hijk' and j=1 5 vfu-(Hilan) = L var(H ijk)' j=1 Similarly, the number of hunters (I) using ATVs (A ) afield on day (i) in GMU (k), treatment area (m) was estimated: A Ujkl rijkl :::--fiji, UjI �238 5 A A L Aijk A;km = , and j=l 5 vfu-(A;km) = L vfu-(Aijk ). j=l Estimating Mean Number of Days per Hunter and Hunter-days _ A Estimates of mean days afield per hunter (d ) and total hunter-days ( D ), and their associated variances were calculated for each hunt code group (J) separately. For each hunt code group (J), mean days afield per hunter (J ) was assumed to be independent of the proportion of license holders that hunted (p). Estimates of mean days afield per hunter by treatment and GMU were calculated directly, and associated variances were summed over hunt code groups (J) for treatment (m) or GMU (k). The mean days afield per hunter and total number of hunter -days were estimated for hunters (I) from hunt code group (J) that hunted treatment area (m): var(d-)= or.-v» J n'~ nj(nj Nj J nm Ldml -d m---.-, - 1=1 _ var(d m) 5 = _ L var(d j), P:I - L.(djl-dj) J .:..:/=::!.l 2 _ -1) �239 " 5" Dm=IDj,and j=1 5 var(D j)' var(Dm) = I j=1 Similarly, for each hunt code group (j), mean days afield per hunter using an ATV (f) independent of the proportion of license holders that hunted cr). was assumed to be The mean days afield per hunter using an ATV (1) and total number of hunter-days with A TVs (F) were estimated for hunters (l) from hunt code group (j) that hunted treatment area (m): fr·1 =A·-f· '}1 =N·;'·-f· 111' _ 5 varCf m) = _ I var(f) , j=1 " 5" Fm = IFj ,and j=1 5 vMCFm) = L var(Fj). j=1 For each hunt code group (j), mean days afield per hunter (d ) was assumed to be independent of the proportion of license holders that hunted ( p). Mean days afield per hunter ( d ) and total number of hunter-days (D) were estimated for hunters (l) from hunt code group (j) that hunted GMU (k) and treatment area (m): �240 ... .... Djk ~ .••. = Hjkdjk = NJ}jkdjk> _ 5 ~ vID-{dkm) = L v3r(djk), j=l 5 Dkm = "IDjk ,and j=1 5 van.Dkm) = Lvan.Djd. j=l Similarly, for each hunt code group (J), mean days afield per hunter using an A TV independent of the proportion of license holders that hunted (;.). ~ . (J ) was assumed to be Mean days afield per hunter (l) using an A TV (J) and number of hunter-days using A TVs (F) were estimated for hunters that hunted GMU (k) and treatment area (m): ijkJ = fjl hI, �241 5 A Iljkajk ~ j=l han = ~-:-s-- Iajk j=l 5 A A van]/an) = I Van]jk j=l 5 F/an = IFjk ,and j=l 5 vM(FIcm) = I vM(Fjk). j=l ). �242 Elk Movement versus Hunter Density I used logistic regression models to evaluate the effects of hunter density on elk movement to private land. The response variable representing elk movement to private land, proportion of elk on private land on day i (vi), was calculated from elk location data (each elk location was classified as public or private land). Hunter density was the estimated density of hunters afield on day i, in hunters/knr'. Hunter density was not an exact measurement, rather it was an estimate with an error term. Typically, using a predictor variable with an error term results in the regression coefficient biased toward zero (Fuller 1987). That is, I would be less likely to detect an effect of hunter density than actually existed. However, if the variance of the predictor variable (hunter density) is large compared to the variance of the estimates, then the effect of the error in the predictor variable can be effectively ignored during regression analysis (Draper and Smith 1966). The variance of hunter density (0.0098) was large compared to the variance of the estimates (maximum variance = 0.0002); hence I ignored the error in hunter density. Because this experiment was not designed to manipulate hunter density, I used model selection to evaluate the effect of hunter density beyond other possible confounding factors. I previously built an extensive suite of models to evaluate the effects of hunt season opening on elk movement to private land (Chapter 1, Table 1.1). Instead of generating another long list of models, I began with the 2 best models from the existing set (Chapter 1, Table 1.4, Models 12 and 13); these 2 models were <2 QAlCc units from each other and were competing models (fable 2.1, Models 1 and 2). My main goal was to evaluate the effect of hunter density on elk movement to private land, and compare this effect to hunt season opening. I replaced Julian day with hunter density to evaluate whether hunter density was a better predictor of elk on private land on day i than Julian day (fable 2.1, Models 3 and 4). To evaluate the importance of hunt season opening compared to hunter density, I first removed hunt season from Model 3, and then removed hunter density (fable 2.1, Models 5 and 6). Although treatment models did not work well predicting the proportion of elk on private land in the study manipulating opening date, I hypothesized that treatment may provide more information when hunter density was included in a model because hunter density patterns varied between earlyand late-opening treatments. To evaluate this idea, I added treatment to the best model that included hunter density (fable 2.1, Model 7), and replaced area with treatment (fable 2.1, Model 8). Finally, evaluate whether the cumulative interaction with hunters caused elk to move to private land, I replaced hunter density with cumulative hunter density in the best model that included hunter density (fable 2.1, ModeI9). Note that none of these models were strictly a priori because I began with previously run best models and built from these best models. I followed the methodology of Burnham and Anderson (1998) to select an appropriate model. I used Akaike's Information Criterion (Akaike 1973), adjusted for overdispersion and corrected for small sample bias (QAICc), as the basis for objectively ranking models and selecting an appropriate.t'best approximating" model (Burnham and Anderson 1998). QAICc was defmed as: QAICc =_ 2~l) c +2K + 2K(K +1) , n-K-1 where In f. is the natural logarithm of the likelihood function evaluated at the maximum likelihood estimates for c a given model, K is the number of estimable parameters from that model, n is sample size, and is the estimated overdispersion factor. QAICc was used instead of AlCc because repeated measurements were taken on radiocollared elk. Lack of independence in the data, in this case repeated measures on elk, may lead to TA" overdispersion 01' "extra-binomial variation" (Burnham and Anderson 1988). I estimated the overdispersion parameter (c) from the global model, and then used this estimate to adjust (inflate) the variance estimates of model parameters and predicted values, and to calculate QAlCc for all the models (Burnham and Anderson 1998). Parameter estimates, c, and QAlCc values were calculated using PROC GENMOD (SAS Institute 1993). The sample size, n, was the number of elk locations collected during the study period. l1'. ,,~. �243 The best approximating model was selected based on minimum QAICc. Models were ranked and compared using ~QAICc (Leberton et al. 1992, Burnham and Anderson 1998) and normalized QAICc weights (Buckland et al. 1997, Burnham and Anderson 1998). For the suite of models being compared, ~QAICc was computed for each model as: ~QAICc", = QAICc", - QAICc"'ln • where QAICC",was the QAICc value for the mth model and QAICcMln was the minimum QAICc among the suite models being compared. Essentially, ~QAICc", is an estimate of the distance between the best approximating model and model m. Normalized QAICc weights (w",) were estimated for each mth model: ~ e _.!.AAAlCc 2~ •• wm =---1 ---, R --ll.QAlCc ~:e 2 r r=1 where R refers to the R models chosen for evaluation. aQAICc and normalized QAICc weights were used to address model selection uncertainty. Generally, models within 1-2 QAICc units of the selected model were considered competing models for explaining elk movements to private land. RESULTS Hunter Survey Data CDOW surveyed 35-36% of early-season hunters using the study area in 1996 and 1997. Early-season hunters were 77% archers and 23% muzzleloaders. Estimates of hunter use with and without ATVs are provided by treatment (Table 2.2) and GMU (Table 2.3). Estimates are provided by hunt code for 1996 (Table 2.4) and 1997 (Table 2.5). The estimated number of hunters afield per day and number of hunters using ATVs are shown with their 95% confidence intervals by treatment (Fig. 2.3) and GMU (Fig. 2.4). In 1996, daily information was collected for 23 days for each hunter, although hunting season extended 30 days during the early-opening treatment. Therefore, 1996 graphs of the south area, where hunting opened early, show only the first 23 days of the season. In 1997, the survey was changed to collect information on hunter use for the entire 30-day season. Appendix 2 contains confidence intervals for daily GMU data. Elk Movement versus Hunter Density The best logistic regression model with the lowest QAICc value (Table 2.6, Modell) was: This was the best model from the manipulative experiment and did not have a hunter density covariate. In fact, the first model with a hunter density covariate was 35.2 QAICc units from the top model (Table 2.6, Model 9), and the hunter density regression coefficient was not significantly different than zero in any of the models in which it was included. Model 9, with cumulative hunter density preformed similarly (aQAICc < 2) to its counterpart, Model 6 with straight hunter density (Table 2.6). Model S, with hunter density, preformed poorly (aQAICc » 2) compared to its counterpart, Model 6, with hunt season replacing hunter density (Table 2.6). �244 DISCUSSION By design, hunter use of the study area was constant from year to year and between treatment areas. However, the daily pattern of use was different for early- and late-opening treatments. When hunting opened early, hunter use was relatively constant. There were several small peaks coinciding with each weekend during the season, and one larger peak coinciding with the opening of muzzleloading season. When hunting opened late, there was a large peak on opening weekend, partially because muzzleloading season opened with archery season. The late treatment peaked at higher densities of hunters (0.44-0.50 hunters/knr') compared to the early treatment (0.28-0.36 hunters/knr'). Several hunting studies indirectly provide insight about elk responses to different levels of hunting pressure. Wright (1983) examined elk response to different hunting seasons: archery and muzzleloading, versus 3 different types of rifle hunting seasons. Distances traveled by elk increased with hunter density, and were lowest during archery and muzzleloading season when hunter densities were the lowest. Although hunting method and hunter density were confounded, elk response to hunter density remains a possible explanation for increased elk movements. An earlier study in the White River area, designed to evaluate elk movement onto a coal company's protected land during early-season hunting, found that 13% of the radiocollared elk were on private lands midway through archery and muzzleloading seasons (Camp Dresser and McKee, Inc. 1986). During this study, 50% and 53% of the radiocollared elk were on private land on the same day during 1996 and 1997 respectively. The lack of movement onto private land in the earlier study may be explained by a 63% reduction in archery hunters for that year because of a change in hunting regulations (Gray et al. 1994). ThuS, elk movement to private land may decrease when hunting pressure drops. Finally, Zahn (1974), Lemke (1975), and Hershey and Legee (1982) noticed an increase in distances moved by elk during the first 10-12 days of hunting season, followed by normal movement during the remainder of the season. These authors noted that hunting pressure was heavy during the initial 10-12 day of hunting, and relatively light for the remainder of the season. Although all these hunting studies are observational, taken together a pattern emerges; at high hunter densities elk increase their movements to refuge, and at low densities they decrease their movements to refuge. However, there is a confounding between the effects of hunter density and the opening of hunting season opening. That is, the increase in distances moved by elk during the initial 10-12 days of hunting season (Zahn 1974, Lemke 1975, Legee 1982) could be due to high densities of hunters or to the opening of hunting season itself. Elk in the White River area seemed to respond more to the opening of hunting season than to hunter density. That is, models that included a hunt season effect did significantly better than models without hunt season effect (Chapter 1). In contrast, models with a Julian day covariate (fable 2.6, Models 1 and 2) preformed far better than models with a hunter density covariate (fable 2.6, Models 3-5 and Models 7-9). In fact, hunter density can not be considered as a predictor of proportion of elk on private land because models with hunter density were a minimum of35.2 AQAICc units from the top model. Additionally, at the opening of earlyseason hunting, 8-18% of the study elk moved to private land (Chapter I), while after opening, elk movement to private land was not related to hunter density (Fig. 2.5). Thus, hunter density had no effect on elk movements to private land in the White River area. It may be that hunter density must exceed a threshold before elk begin to respond directly to hunters, and hunter densities did not surpass this threshold during the study. MANAGEMENT IMPLICATIONS Ifhunter density continues to be a suspected cause of increased elk movement to private land, then an experiment manipulating hunter density should be conducted. Managers concerned with reducing elk movements to private land have an opportunity to evaluate effects of hunting pressure when setting license numbers for a particular area. Additionally, increasing hunter density on private land where problems are occurring may be the most direct strategy to retain elk on public land during late summer. Whatever the management strategy, manipulation of hunter density offers managers a good opportunity to conduct an adaptive management experiment to increase information regarding the management of elk movements to private land. �245 LITERATURE CITED Akaike, H. 1973. Information theory and an extension of the maximum likelihood principle. Pages 267-281 in B. N. Petran and F. Csaki, editors. International symposium on information theory. Second edition. Akademiai Kiadi, Budapest, Hungary. Altmann, M. 1956. Patterns of herd behavior in free-ranging elk of Wyoming, Cervus canadensis nelsoni. Zoologica 41:65-71. Boyd, R J. 1970. Elk of the White River Plateau, Colorado. Colorado Division of Game, Fish and Parks Technical Publication 25, Fort Collins, Colorado, USA. Buckland, S. T., K. P. Burnham, and N. H. Augustin. 1997. Model selection: an integral part of inference. Biometrics 53:603-618. Burnham, K. P., and D. R Anderson. 1998. Model selection and inference: a practical information-theoretic approach. Springer-Verlag, New York, New York, USA. Camp Dresser and McKee, Inc. 1986. Meeker PRLA elk migration study: monitoring report, volume 2. Prepared for Consolidation Coal Company. Camp Dresser and McKee, Inc., Denver, Colorado, USA. Draper, N. R and H. Smith Jr. 1966. Applied regression analysis. Second edition. John Wiley and Sons, New York, New York, USA. Fuller, W. A. 1987. Measurement error models. John Wiley and Sons, New York, New York, USA. Gray, J. P.; G: Byrne, and J. Madison. 1994. White River elk data analysis unit plan: game management units: 11,211,12,13,131,231,23,24,25,26,33. Colorado Division of Wildlife Internal Report, Grand Junction, Colorado, USA. Hershey, T. J., arid T. A. Leege. 1982. Elk movements and habitat use on a managed forest in north-central Idaho. Idaho Department ofFish and Game Wildlife Bulletin 10, Boise; Idaho, USA Irwin, L. L., and J. M. Peek. 1979. Relationships between road closures and elk behavior in northern Idaho. Pages 199-204 in M. S. Boyce and L. D. Hayden-Wing, editors. North American elk: ecology, behavior, and management. University of Wyoming, Laramie, Wyoming, USA Leberton, J-D., K. P. Burnham, J. Clobert, and D. R Anderson. 1992. Modeling survival and testing biological hypotheses using marked animals: a unified approach with case studies. Ecological Monographs 62:671I8. Lemke, T. O. 1975. Movement and seasonal ranges of the Burdette Creek elk herd, and an investigation of sport hunting. Montana Fish and Game Department Job Final Report 32.01, Job Number BG-3.15, Missoula, Montana, USA Martinka, C. J. 1969. Population ecology of summer resident elk in Jackson Hole, Wyoming. Journal of Wildlife Management 33 :465-481. Morgantini, L. E., and R J. Hudson. 1979. Human distribution and habitat selection by elk. Pages 132-139 in M. S. Boyce and L. D. Hayden-Wing, editors. North American elk: ecology, behavior, and management. University of Wyoming, Laramie, Wyoming, USA. SAS Institute. 1993. SAS/STAT® software: The GENMOD procedure, release 6.09. SAS® Technical Report P-243, SAS Institute, Inc., Cary, North Carolina, USA. Steinert, S. F., H. D. Riffel, and G. C. White. 1994. Comparisons of big game harvest estimates from check stations and telephone surveys. Journal of Wildlife Management 58:335-340. White, G. C. 1993. Precision of harvest estimates obtained from incomplete responses. Journal of Wildlife Management 57:129-134. Wright, K. L. 1983. Elk movements, habitat use, and the effects of hunting activity on elk behavior near Gunnison, Colorado. Thesis, Colorado State University, Fort Collins, Colorado, USA. Zahn, H. M. 1974. Seasonal movements of the Burdette Creek elk herd. Montana Fish.and Game Department Job Final Report 32.01, Job BG-3.13, Missoula, Montana, USA. �246 Table 2.1. Description and representation of models relating effects of area, treatment, hunt season opening, hunter density, and Julian date to daily proportion of radiocollared elk found on private land (VI) from 20 July-l 0 .October in the White River area, Colorado, 1996-1997. Hypothesis Model structure • 1. Lowest AlCc model from the experimental early-season opening date b 2. Second lowest AlCc model from the experimental manipulation of early-season opening date b 130+13. (A)+f32(S)+f33(D)+f3.(AxD)+f3s(SxD) +f36(AxS) 3. Hunter density is a better predictor than Julian day - Model 1with hunter density replacing the Julian day covariate C f3o+f3.(A)+(32(S)+f33(H)+f3.(AxH) 4. Hunter density is a better predictor than Julian day - Model 2 with hunter density replacing the Julian day covariate C 5. Hunter density is a better predictor than hunt season hunter density and area effects only 6. Hunt season is a better predictor than hunter density - hunt season and area effects only 7. Treatment is an important effect when hunter density is a covariate - add treatment to best model with hunter density 8. Treatment is an important effect when hunter density is a covariate - replace area with treatment manipulation of f3o+f3.(A)+f32(S)+f33(D)+f3.(AxD)+f3s(SxD) f3o+f3.(A)+(32(S)+f33(AxS) 9. Cumulative hunter density better predicts elk movement to private land - use best model that includes hunter density and replace the hunter density covariate with a cumulative hunter density covariate • A represents treatment area with south area = 0 and north area = 1, S represents archery hunting season with 0 before opening and 1 = after opening, D represents the covariate date, which is Julian day, and H represents the covariate hunter density on day i in hunters/knr', The dependent variable (VI) was in logit scale. b See Chapter I, Table 1.4. C SxH cannot be estimated because hunter density = 0 before hunt season opening. = �247 Table 2.2. Number of licenses sold (N), percent surveyed (s), estimates of hunter numbers (Ii ), hunter-days (D ) mean days huntedlhunter (d), hunters using ATVs (A ), hunter-days with ATVs (ft ), and mean number of ' days/hunter using an ATV ( 1), by treatment area during archery and muzzleloading season in the White River area 1996-1997. The half-width of the 95% confidence interval is shown for all estimates. Area North Year 1996 1997 Ii N' s 1,711 37.2 36.9 2,170 D b 1,362 ± 25 1,786 ± 20 C 8,999 ±390 10,716 ±426 d 6.4 ± 1.1 6.0±0.8 A 1 F 209 ±34 329 ±45 1,555 ± 314 2,134 ±392 7.2±4.1 6.4 ± 2.1 1996 2,253 35.1 6.2 ± 1.0 300 ±40 1,715 ± 30 10,920 ±455 2,205 ±380 7.1 ±2.8 1997 2,090 33.2 258 ±40 1,679 ±26 11,128 ± 419 6.4±0.9 2,083 ±408 7.6 ±2.5 • Because some individualspurchased 2 licenses,the numberof individualspurchasing licenses,which was used in all estimates,was slightlyless than the number of licensessold. b The estimatednumber of hunters is lower than the number of licenses sold because some hunters purchasing licenses did not hunt and because some hunters purchased 2 licenses. e Estimatednumber of hunter-days and hunter-dayswith ATVs are sums of estimated hunter-daysby hunt code (fables 2.4 and 2.5), and not the direct product of estimatednumber of hooters x mean number of days/hunter. South Table 2.3. Estimates of hunter numbers (fl ),hunter-days (D ), mean days huntedlhunter (d), hunters using ATVs (A ), hunter-days with ATVs (ft ), and mean number of dayslhunter using an ATV (j), by GMU during archery and muzzleloading season in the White River area 1996-1997. The half-width of the 95% confidence interval is shown for all estimates. Area North GMU Year 12 1996 1997 1996 1997 19% 1997 22 23 fl' 773 1,014 437 533 291 426 ±48 ± 58 ±44 ±52 ±38 ±49 Db d 4,842 ±441 6,295 ±484 2,653 ±355 3,274 ±483 1,504 ±277 2,425 ±401 6.0± 1.8 6.0± 1.2 5.8±2.0 5.9±2.2 5.0±2.5 5.3± 1.8 ft A 74 117 104 162 75 102 ±22 ±30 ±25 ±33 ±21 ±26 480 571 676 1,092 399 643 ± 181 ± 173 ±212 ±333 ± 154 ±296 jc 6.3 5.0 6.3 6.3 5.3 5.8 ±7.1 ±3.1 ±4.7 ±3.5 ±7.5 ±3.9 890 ±235 1996 3,893 ±409 151 ± 30 5.7 ± 3.6 715 ±53 5.3± 1.8 1997 111 ±28 700 ±201 6.0 ±2.6 494 ±51 2,770 ±358 5.5± 1.9 4.5 ±4.5 23 1996 600 ±51 3,272 ±375 75 ±21 342 ± 170 5.4±2.0 1997 8.5 ± 7.4 577 ±53 3,514 ±417 5.9±2.4 21 ± 12 195 ±308 33 1996 132 ±28 974 ±287 6.9 ±4.8 638 ±52 3,735 ±456 5.7±2.0 1997 143 ± 31 1,188 ±302 746 ±56 4,843 ±452 6.3± 1.4 8.0 ±3.1 • All GMU estimates are slightly greater than treatmentor hoot code estimates because some hunters hunted in more than oneGMU. b Estimated numberof hunter-days and hooter-dayswith ATVs are sums of estimatedhunter-daysby hunt code, and not the direct product of estimated number of hunters x mean numberof days/hunter. c Because of the small sample sizes, a log-basedconfidenceintervalshould be used in some cases so that values for the 95% CI of 1are >0. South 22 �Table 2.4. Estimates of hunter numbers (if ), hunter-days (D), mean days hunted/hunter (d), mean number of day !¥hunter using an ATV (i), by hunt code during archery and muzzleloading 95% confidence interval is shown for all estimates. Percent of Number of NORTH AREA . D if licenses sold • hunters surveyed 32.0 241 D-E-012-01-A 94±7 663 ±S3 973 32.5 E-E-012-01-A 874 ±22 6,368 ±367 165 51.5 E-F-012-01-M 647 ±69 126 ±8 hunters using ATVs (A), hunter-days with ATVs (ft), and N season in the White River area 1996. The half-width of the d A 7.1 ± 0.7 1S±8 7.3 ± 004 149 ± 31 ft 139 ± 67 1,198 ± 302 i 7.6 ± 2.0 8.1 ± 1.1 E-M-O 12-0 I-M 178 48.9 167 ±3 808 ±61 5.1 ±OA 4.8 ±OA 21 ±9 103 ± 48 D-M-012-01-M 154 46.8 101 ±7 513 ± 59 5.1 ± 0.4 8±5 50±38 4.8 ± 1.3 6.0 ± 3.2 292 ± 13 1,849 ± 187 6.3 ±0.6 59 ± 16 380 ± 128 6.4 ± 1.3 7,184 ±400 6.8 ±OA 4.8 ±OA 194 ± 35 1,579 ± 352 12 ±6 66 ±43 5.3 ±2.7 SOUTH AREA D-E-033-01-A E-E-33-01-A E-F-033-01-M 526 30.6 1,219 32.0 1,056 ±23 150 49.3 122±6 581 ± 55 16 ±7 86 ±41 8.1 ± 1.1 5.3 ± 1.0 47.5 160 140±6 817 ±70 5.8 ±0.4 16±8 93 ±50 5.9 ± 1.6 45.5 198 4.7 ±0.5 14±7 68 ±41 4.9 ± 1.9 105 ± 12 490 ±77 D-M-033-01-~ • Because some individuals purchases 2 licenses, the number of individual purchasing licenses was slightly less the number of licenses sold. The number of individuals purchasing licenses was used for all estimates. E-M-033-O I-M, Table 2.5. Estimates of hunter numbers (if ), hunter-days (D), mean days hunted/hunter (d), hunters using ATVs (A), hunter-days with ATVs (ft), and mean number of dayslhunter using an ATV (i), by hunt code during archery and muzzleloading season in the White River area 1997. The half-width of the 95% confidence interval is shown for all estimates. D-E-O 12-0 I-A Number of licenses sold • 402 NORTH AREA Percent of hunters surveyed if D d A 54.7 205 ±5 1,243 ±63 6.1 ± 0.3 41 ±8 253 ± 57 1,193 ± 16 6.3 ± 0.3 233 ±42 1,540 ± 377 E-E-O 12-0 I-A 1,277 27.9 E-F-O 12-0 1-M 166 43.4 134±8 E-M-012-01-M 180 48.3 160±5 D-M-O 12-0 I-M 145 45.5 SOUTH AREA D-E-033-01-A 94 ±6 7,456 ±407 ft j 6.3 ±0.8 6.6 ± 1.1 686 ±70 5.1 ±0.4 31 ± 11 193 ± 73 6.1±1.0 850 ±63 5i3 ±OA 21 ±9 131 ± 60 6.2 ± 1.4 480 ±53 5.1 ± 0.5 3±3 16 ± 17 5.0 ±O.O 416 30.0 223 ±7 1,503 ± 119 6.8 ±0.5 43 ± 13 330 ± 118 7.7 ± 1.4 1,185 28.4 1,058 ±23 7,640 ±388 7.2 ±0.3 166 ± 34 1,516 ± 386 7.1 ± 1.2 E-F-033-01-M 147 48.3 121 ±6 584 ±56 4.8 ±0.4 17 ±7 73 ±34 4.3 ±0.6 E-M-033-01-M 155 49.0 139±4 711 ± 57 5.1 ± 004 11 ±6 55 ±34 4.8 ± 1.6 E-E-33-01-A a 46.0 187 D-M-033-O I-M 5.1±1.2 ~IQ;1;Q,~ lQ2;1;42 21;1; lJ~ ;1;:Z ~82;1; ~J • Because some individuals purchases 2 licenses, the number of individual purchasing licenses was slightly less the number oflicenses sold. The number of individuals purchasing licenses was used for all estimates. .t:- oo �249 Table 2.6. Ranking of models relating effects of area, treatment, hunt season opening, hunter density, cumulative hunter density, and Julian day to daily proportion ofradiocollared elk found on private land (VI) from 20 July-IO October in the White River area, Colorado, 1996-1997. Models were ranked by QAICc values and normalized QAICc weights ( W m ). Model number Model structure • f30+f3I(A)+f32(S)+f33(D)+f3iAxD)+f3s(SxD) f30+f3I(A)+f32(S)+f33(D)+f34(AxD)+f3s(SxD) +f36(AxS) f3o+f3I(A)+f32(S)+f33(c)+f34(AxC) +f3s(AxS) f3o+f3I(A)+f32(S)+f33(AxS) f3o+f3I(A)+f32(S)+f33(H)+f34(AxH)+f3s(AxS) f3o+f3I(A)+f32(S)+f33(H)+f3iAxH) f3o+f3I(A)+f32(S)+f33(H)+f34(T)+f3s(AxH)+f36(AxS) +f3-ifxH) f3o+f3I(A)+f32(D)+f33(AxD) f30+f3I(T)+f32(S)+f33(H)+f34(1'XH)+f3s(l'xS) 1 b KC QAICc AQAICc 6 3760.28 0.00 0.729 wm 2 7 3762.26 1.97 0.271 9 6 3795.45 35.17 0.000 6 4 3796.29 36.01 0.000 4 6 3798.59 38.31 0.000 3 5 3800.41 40.13 0.000 7 5 8 8 3802.34 42.06 0.000 4 3867.39 107.11 0.000 6 3897.96 137.68 0.000 • A represents treatment area with south area = 0 and north area = I, S represents archery hooting season opening with before opening = 0 and after opening = I, T represents treatment with early opening = 0 and late opening = I, D represents the covariate date, which is Julian day, H represents the covariate hooter density on day i in hunters/km', and C represents the covariate cumulative hooter density on day i in hunters/knr', The dependent variable (vJ was in logit scale. b Numbers correspond to those in Table 2.1 C Number of estimable parameters. 7000 e -- 6000 (I) "". c: :J s: c: 5000 0 en - .... , CO en 4000 (I) I >. 1ij 3000 -._ (I) 0 2000 (I) ..c ~ 1000 z 0 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 Year * Variance estimates not available 1984-1987. Figure 2.1. Number of archery and muzzleloading hunters using GMUs 12,23,24, and 33 in Colorado, 19841994. Data from Deer, Elk, and Antelope Management (DEAMAN) database and software, Colorado Division of Wildlife unpublished data. �250 ~. BLMLand g. Flat Tops Wilderness B. State Wildlife Area o Km o 10 20 Treatment Area Boundary IIForest Service Land Figure 2.2. North and south treatment areas and land ownership in the White River study area, Colorado. �251 a c Early Opening 1,200 Late Opening 1,200 Opening of muzzleloading End of muzzleloading 1,000 1,000 800 - "'C Ci) 1 ~ 600 ii= CO 800 South 1996 600 400 L. North 1996 400 (J) .c 200 E :::s '"'_-" 0 e "'C .e ...,. ~ (X) .•.. s ...,. .•.. t: a; 0> ~ 0> ...,. .•.. 10 a a; d 1,200 1,200 E 0 .•.. b CO ~ 200 - .. -_ -- ..- -- ....... End of muzzJeloading Opening of muzzleloading 1,000 1,000 W 800 800 South 1997 North 1997 600 600 400 400 200 200 - __ .- .. _- - ..... -- •. - .... --.------- ... 0 M N to 0 M to ~ .•.. M a; 0> -- _- --- .• _-- ---- -_ --_-----:----------- 0 .•.. M 0 s a; Date •••••••••••. LEGEND Hunters Hunters using A TVs 95%CI Figure 2.3. Estimated daily number of hunters and hunters using A TVs in the (a) south treatment area 1996, (b) north treatment area 1997 , (c) north treatment area 1996, and (d) south treatment area 1997, during archery and muzzleloading season in the White River area, Colorado. The first date is opening date. Each date shown represents a Saturday. 1 There is a break in estimates for the south area in 1996 because survey data were only collected for 23 out 6fthe 3'0 days of the early treatment. The survey was extended to 30 days in 1997. j","; .. �252 a c 1996 1,000 1997 1,000 B 1 A 800 800 .A C B C B C "C CD 'i=ns 600 600 400 400 •••• CI) .c 200 E :::J 0 s::: '. .•..•.-._ ..•. s £:! <0 "C "' •..........•. 200 ...,..." .~. ..•.•.•. ..•. ~ t: OJ (i; a: (i; Sb ns .-E 0 s <0 N M 0 s £:! <0 d 1,000 1,000 'oiJ tn B 1 A 800 W 800 C A I I 600 600 400 400 200 200 0 ..• .~.-.~~: .......••• ~~:~ ~ ;;S t! co t: '" ..•. '" 0 s 0:> £:! en on a .., s -- -..._0 t! co Date LEGEND -GMU12 .•.•••••.GMU23 --GMU24 _.-•..._ GMU33 A B C Opening date of archeryseason on early-treatment. Opening date of muzzleloadingseasonand archeryseason on late treatment. Closing date of muzzleloadingseason. Figure 2.4. Estimated daily number of (a) hunters 1996, (b) hunters using ATVs in 1996, (c) hunters 1997, and (d) hunters using ATV s 1997, during archery and muzzleloading area, Colorado. I Each date shown represents a ~aturdaY'';J c' season in the White River + There is a break in estimates for GMU 23, 24, and 33 in 1996 because survey data were only collected for 23 out of the 30 days of the early treatment. The survey was extended to 30 days in 1997. �253 CHAPTER 3: ELK LOCATIONS IN RELATION TO DOMESTIC SHEEP BANDS AND A METHOD TO TEST FOR AVOIDANCE OR ATTRACTION BETWEEN ANIMALS INTRODUCTION Hearsay, accusation, tall tales, and rumor dominate the discussion about Rocky Mountain elk (Cervus elaphus nelsoni) movements in response to domestic sheep (Ovis aries). Elk movement responses to sheep are a concern of hunters, who feel that livestock grazing on public land is a disturbance that elk avoid. Hunters claim that domestic sheep force elk away from public hunting areas and onto private land, but there is little documentation of elk responses to domestic sheep. In one study of elk and livestock grazing, Clegg (1994) found that elk densities decreased rapidly after introduction of sheep, herders, and dogs. However, the distance of impact was not recorded making it difficult to assess whether sheep could move elk away from public hunting areas. Most studies of elk and sheep have originated from livestock concerns and concentrated on dietary overlaps and forage competition (pickford and Reid 1943, Nichols 1957, Stevens 1966, Olsen and Hansen 1977, MacCracken and Hansen 1981, Beck et al. 1996). In the White River area, elk and domestic sheep graze high mountain meadows. Approximately 26 bands of domestic sheep used the study area between May and October. Early-season hunting (archery and muzzleloading), which occurs between late August and mid-October, overlaps with sheep grazing. During . meetings about elk movements in response to early-season hunting, archery hunters expressed concern that domestic sheep cause elk to move off public land to private land during the early season. As part of the stakeholder process, CDOW agreed to collect locations of sheep bands in the area when collecting locations of 80 radiocollared elk for an archery hunting study. My objectives were to (1) determine if elk avoided sheep at several spatial scales and (2) develop a methodology applicable to radio-telemetry data to evaluate attraction or avoidance between animals. I used location data from radiocollared elk and visual locations of sheep bands from the summer and fall of 1996 and 1997. The null hypothesis was that elk locations were random with respect to sheep locations. I used nonparametric statistics and randomization techniques to calculate, under the null hypothesis, the probability that elk avoided sheep. STUDY AREA The White River area of northwestern Colorado covered approximately 4,540 km2 and was composed of Game Management Units (GMUs) 12,23,24, and 33. In Colorado, GMUs were delineated to distribute hunters through allocation of hunting licenses. White River area land ownership was 34% private land and 66% public land (Fig. 3.1), with 82% of public land being United States Forest Service (USFS). USFS land was mostly at high elevation and toward the center of the study area, with some Bureau of Land Management (BLM) areas at lower elevations in the southern sections of the study area. Private land was lower in elevation and comprised mainly ranches and coal mines. Topography, climate, and vegetation varied widely throughout the study area. Elevation ranged from 1,629 to 3,700 m. The central part of the study area was high elevation public land, split by the White River valley. The high elevation areas dropped off to lower elevations to the north, south, and western edges of the study area. Higher elevations had severe winters with heavy snowfall, while lower elevations had comparatively mild winters. Mean annual precipitation at 3,000 m in the National Forest areas was about 100 em, compared with about 30 cm at lower elevations of <2,000 m. Vegetation types at elevations >2,600 m were in the montane/subalpine zone and included groves of aspen (Populus tremuloidesy, Engelmann spruce (Picea engelmanni), and alpine fIT (Abies lasiocarpa) interspersed with grassy meadows. Middle elevations (1,9802,600 m) were a transitional zone that consisted primarily of pinion pine (Pinus edulis),juniper (Juniperus scopulorum), and big sagebrush (Artemisia tridentata). Lower elevations «2,000 m) in the southern and northern parts of the study area were in the Great Basin zone with sage grasslands. Oakbrush patches were found at the middle and lower elevations with major shrubs that included Gambel's oak (Quercus gambelii), serviceberry (Amelanchier alnifolia), mountain mahogany (Cercocarpus montanus), chokecherry (Prunus �254 virginianay; snowberry (Symphoricarpos utahensis), bitterbrush (Purshia tridentata), and rabbitbrush (Chrysothamnus nauseosus). Higher and middle elevation areas provided summer and fall forage for elk, while lower elevations were typically used by elk during winter. Boyd (1970) provides a detailed description of the study area. USFS issued permits for sheep grazing on national forest lands on the study area. During the study period, approximately 26 bands of domestic sheep used the study area. Each band was assigned to a particular area called an allobnent. Bands ranged in size from 750-1,350 sheep, and averaged approximately 1,043 sheep (95% CI = 981,1,106; USFS unpublished data from 1992-1995). Band sizes are approximate because actual numbers were often less than permitted numbers (Mary Massey, United States Forest Service, Meeker Colorado, personal communication). Sheep arrived on the study area 16 June-l l July, and left 10 September-IS October. Herders and dogs stayed with the sheep bands and moved them through their allobnent area. METHODS Data Collection The adult female elk used in this study were captured from randomly selected locations throughout the study area. Capture and collaring procedures are descried in detail in Chapter 1. Elk and sheep locations were collected 20 July-l0 October 1996-1997. I relocated radiocollared elk between 0700-1~OO hr using a fixed-wing aircraft with a 2-element Yagi antenna mounted to each strut of the airplane. For each elk relocation, Universal Transverse Mercator (UTM) coordinates were recorded with a Global Positioning System. I collected elk: locations 2 times a week with 2-4 days between collections. During each flight, the position of at least 1 sheep band was recorded. Because elk-sheep interactions were not the focus of the study, locations of sheep bands were opportunistically chosen. That is, I typically recorded the location of only one band of sheep seen and did not record any other bands, although up to 3 bands per day were sometimes seen. Thus, each band location was a sample of the population of 26 bands. Because I did not collect locations in a specific route each flight, the daily sample of sheep bands was fairly random (Fig. 3.1). Once a sheep band was seen, I flew over the band and recorded UfM coordinates at the approximate center of the band. Data Analysis + c All analyses were computed separately for each year. Distances between elk and sheep bands were calculated from UTM coordinates as straight-line distances. For each day a sheep band was located, I calculated the distance between the band and every radiocollared elk located that day. I called these paired distances because they were paired in time (same day). I then calculated the distance between the location of the sheep band, located that day, and the locations of each radiocollared elk located every other day. I called these unpaired distances because they were temporally unpaired. During the study period, 20-23 locations were collected on each elk per year. Thus, for each elk, there was I paired distance and 19-22 unpaired distances for each day locations were collected. Note that there was no way to compute the variance on 1 measurement; hence paired and unpaired distances could not be compared using parametric methods. However, a rank representing the paired distance compared to the unpaired distances, between 1 elk and I sheep band, represents 1 sample. Thus, I used a ranking method to evaluate avoidance between elk and sheep bands. For each elk and each day locations were collected, I ordered distances, paired and unpaired, from smallest to largest, and assigned a ranking to each distance. That is, the smallest distance between a sheep band and an elk was assigned a 1, the next smallest distance a 2 and so on. Thus, for each elk and each day data were collected, there was a ranking of 1 to number of distances collected on that elk (Fig. 3.2). If elk were located randomly with respect to sheep bands, then paired distances' would be equal to unpaired distances, and the mean rank of paired distances would be equal to the mean rank unpaired rankings. If elk avoided sheep, then paired distances would be farther from sheep, on average, than unpaired distances, and the mean rank of paired distances would be significantly greater than the mean rank of unpaired rankings. Conversely, if elk were attracted to the sheep, then paired distances would be closer to sheep, on average, than unpaired distances, and the mean rank of paired distances would be significantly less than the mean rank of unpaired rankings (Fig. 3.3). This assumes that elk do not avoid an area where sheep were located after the sheep leave the area. �255 For each year, I extracted the rank of the paired distances for each elk and day that both the elk and sheep band were located, and calculated the mean ranking of the paired distances. The statistical expressions of the null and alternative hypothesis were: Ho: The mean rank of paired distances = the expected mean rank of unpaired ranks, and HA: The mean rank of paired distances> the expected mean rank of unpaired ranks, which tested the study hypothesis: Ho: Locations of elk were random with respect to locations of sheep; elk did not avoid sheep, and HA: Locations of elk were father to sheep than expected at random; elk avoided sheep. Note that it was possible that elk were attracted to sheep because sheep are grazed in areas preferred by elk for grazing, but this was not the question of interest. In order to evaluate the probability of the observed paired rank, I generated the distribution of paired ranks under the null hypothesis that elk and sheep were located randomly with respect to each other. To develop the null distribution, I randomized on sheep bands. That is, I randomly assigned a day to each sheep band. For example, a sheep band located on day 2, may be randomly assigned to day 6. After randomly assigning a day to each sheep band, paired and unpaired distances were calculated as described for the observed data. From each sample, I extracted the ranks of the randomly "paired" rank. From the collection of samples via the randomization procedure, I generated the distribution of mean ranks for paired elk and sheep locations that were random, and created a null distribution representing no avoidance (or attraction) between elk and sheep. For each randomized sample, the mean rank was saved and the randomization was re-run until there were 999 mean rankings. The mean of the 999 randomized mean ranks was the expected mean rank. The observed mean rank was merged with the randomized mean ranks, and the mean ranks were ordered from largest to smallest. The percentile of the observed sample was the P-value of the hypothesis test; that is, the probability of observing the rank of the sample or a larger rank under the null hypothesis that elk are located randomly with respect to sheep (Mooney and Duval 1993). Spatial scale may influence the ability to detect avoidance or attraction between animals (Doncaster 1990, Turchin 1998). That is, 'if elk were> 10 km from a particular sheep band, then it was not likely that there was any interaction between those elk and the sheep band. To include only elk close enough to interact with the sheep bands, I repeated this analysis at 2 other spatial scales. First, I re-analyzed the data using elk and sheep bands that were <5 km from each other on any day. Finally, I re-analyzed the data using elk and sheep bands that were <I km of each other. I did not examine closer spatial scales because sample sizes became too small. RESULTS The observed ranks were distributed without any apparent attraction or avoidance for 1996 (Fig. 3.4) and 1997 (Fig. 3.5). For all elk, elk <5 km from a sheep band, and elk <1 km of a sheep band, the observed mean rank was not significantly greater than the expected mean rank under the null hypothesis that locations of elk were random with respect to sheep (fable 3.1). Thus, elk did not avoid sheep at the 3 spatial scales examined. For all spatial scales, the randomized mean rank remained constant (Table 3.1). In 1996, when approximately 22 locations were collected per elk, the randomized mean rank was 11.48-1l. 49 for the 3 spatial scales. In 1997, when approximately 23 locations were collected per elk, the randomized mean rank was 11.931l.97 for the 3 spatial scales. Maps of elk and domestic sheep locations on several flights are presented in Appendix 3. DISUCSSION Experimental Results Elk did not appear to avoid sheep at any of the spatial scales I examined. Elk may avoid sheep at <1 km, but I lacked sample sizes needed for fmer scale detection of avoidance. However, because elk did not avoid �256 sheep at > 1 km, it is extremely unlikely that domestic sheep caused any large scale movements of elk from public to private lands. Additionally, it may be that elk avoided the area used by sheep following sheep occupation. If this effect lasted for a long period of time, then the rank method used would be biased because would not detect this avoidance. If there is concern about elk avoidance of sheep at <1 km, or elk avoidance of an area after sheep occupation, then a study designed to answer these questions is needed. Although there is little documentation of elk response to domestic sheep disturbances, studies have been conducted on other, similar, non-lethal disturbances. Elk responses to recreationalists, cross country skiers, cattle, and roads may illuminate possible elk responses to domestic sheep. In areas of continuous human presence, such as Rocky Mountain Park, elk showed little response to approaches of people in automobiles or on foot, day or night (Schultz and Bailey 1978). However, in the less intensely used Medicine Bow National Forest, Ward et al. (1973) found that elk avoided recreationalists (campers, fishermen, and picnickers) at <800 m. Elk Response to cross country skiers depended on the habituation of elk; in areas of high human activity elk moved short distances away from skiers, while in areas of low human activity, elk movement away from skiers was over 3 times greater than that of habituated elk (Cassirer et al. 1992). With respect to cattle, elk showed no movement responses to cattle grazing at> 100 m (Ward et al. 1973, Clegg 1994). In addition to habituating to non-lethal disturbances, elk may learn to distinguish between dangerous and harmless disturbances. In Rocky Mountain National Park, where elk were not hunted, Schultz and Bailey (1978) found no avoidance of roads in winter. In contrast, in Roosevelt National Forest, which is adjacent to Rocky Mountain National Park, Rost . (1975) found that a population of hunted elk avoided roads in winter. Wright (1983)found that mean distance to dirt roads more than doubled when hunting season opened. In the White River area, where sheep have been grazed for the past 100 years, elk may have learned that sheep pose no threat, and like elk exposed to other nonlethal human activities, the White River elk may have habituated to sheep activities and ceased to avoid sheep. Methodology for Determining Attraction and Avoidance One of the difficulties in studying behavioral interactions between animals with radio telemetry data is lack of quantitative methods to determine if one animal's movements or locations are related to another animal's (White and Garrott 1990). Historically, radio-telemetry location data has been used to describe the size and shape of an animal's home range. Most studies collecting these type of data do not take into account the temporal information available in the telemetry data (White and Garrott 1990). This temporal information can be used to describe social interactions, such as avoidance or attraction, between animals. Two general approaches have been used to describe animal interaction. The first, static territorial overlap, describes the overlap of home ranges (Webster and Brooks 1981, Minta 1992) as an indication of attraction or avoidance. The second method, temporal or dynamic interactions, uses distance expectations based on expected home range use to test for attraction or avoidance (Dunn 1979, Macdonald et al. 1980, Doncaster 1990, Minta 1992). Both static and parametric temporal methods of testing for avoidance between animals require knowledge of home range size or use. There are difficulties in estimating the mean area and variance of a home range, even if estimation procedures are accurate (Worton 1987). Additionally, home range statistics vary depending on the method of estimation (White and Garrott 1990). Parametric methods that test for temporal interaction assume that home range use follows a bivariate normal distribution (Jennrich and Turner 1969, Dunn 1979). It is unlikely that animals use space in a bivariate normal manner because resources, mates, and cover are all likely to be clumped and non-normal in distribution. To combat unlikely representations of home range size of use, Doncaster (1990) suggests a nonparametric approach based on differences between observed and theoretical distributions of separation distances between 2 animals. All temporally paired distances and unpaired distances between 2 animals are calculated, and a critical distance is chosen for analysis of temporal interaction. Using this distance, data can be placed in a 2 x 2 contingency table showing frequencies of paired and unpaired distances closer and farther than this critical distance. If distances are closer than expected, then the animals are attracted to each other; if distances are farther than expected, then the animals are avoiding each other. The rank method presented here is similar to Doncaster's (1990) nonparametric test for attractions and avoidance. Both methods have the advantage of not requiring assumptions about home range size or use compared to the static and parametric dynamic methods. However, there are 2 main advantages of the rank method over Doncaster's (1990) nonparametric approach. First, the rank method requires no arbitrary critical distance to evaluate attraction or avoidance. Although it may appear that the various spatial scales I used are similar to a critical distance, they are not. This study was not originally designed to test for attraction and �257 avoidance between elk and sheep, and the spatial scale over which I collected data (4,540 knr') was too large for accurate testing of attraction and avoidance between animals. Because of the large scale, I examined various spatial scales to subset out animals close enough to sense each other. Studies designed to test for avoidance or attraction interactions would not be at such a large scale and would not need to subset out animals close to each other. Additionally, at any scale examined, I did not set any critical distance at which attraction or avoidance was to occur. Second, Doncaster's (1990) method uses a chi-square test statistic, which works best when expected cell counts are 2::5. The chi-square would not have worked well in this study because there was only 1 paired distance, which results in a count of 1 and 0 in cells containing the number of paired distances closer and farther than the critical distance. The chi-square test may not work well for any study with low sample sizes or unbalanced data (more locations on one animal than another). However, this problem is attenuated if a Fisher's exact test is used to evaluate the 2-way contingency table of paired and unpaired distances. MANAGEMENT IMPLICATIONS This study suggests that elk do not avoid domestic sheep at distances great enough to cause elk movements off public hunting grounds to private land. If concern about elk avoidance of sheep at distances of <1 km becomes serious, a more complete study, with radiocollared sheep from each band and a design tailored to the avoidance question, should be performed. The rank method used to test for elk avoidance of sheep could be used in many radio-telemetry data sets to evaluate attraction or avoidance between animals. The rank method, using a randomization procedure to generate the null distribution, is a flexible, powerful, and assumption reduced approach to test for attraction or avoidance behaviors between animals. The main requirement of the method is that simultaneous locations are taken on all animals in the study. LITERATURE CITED Beck, 1. L., J. T. Flinders, D. R Nelson, and C. L. Clyde. 1996. Dietary overlap and preference of elk and domestic sheep in aspen-dominated habitats in north-central Utah. Pages 81-85 in K. E. Evans, compiler. Sharing common ground on western rangeland: proceedings of a livestock/big game symposium. U.S. Forest Service, Intermountain Research Station, Ogden, Utah, USA. Boyd, R J. 1970. Elk of the White River Plateau, Colorado. Colorado Division of Game, Fish and Parks Technical Publication 25, Fort Collins, Colorado, USA. Cassirer, E. F., D. J. Freddy, and E. D. Ables. 1992. Elk responses to disturbances by cross-country skiers in Yellowstone National Park. Wildlife Society Bulletin 20:375-381. Clegg, K. 1994. Density and feeding habits of elk and deer in relation to livestock disturbance. Thesis, Utah State University, Logan, Utah, USA. Doncaster, C. P. 1990. Nonparametric estimates of interaction from radio-tracking data. Journal of Theoretical Biology 143:431-443. Dunn, J. E. 1979. A complete test for dynamic territorial interaction. Pages 159-169 in F. M. Long, editor. Proceedings for the second international conference on wildlife biotelemetry. University of Wyoming, Laramie, Wyoming, USA. Jennrich, R I., and F. B. Turner. 1969. Measurement of non-circular home range. Journal of Theoretical Biology 22:227-237. MacCracken, 1. G., and R M. Hansen. 1981. Diets of domestic sheep and other large herbivores in south central Colorado. Journal of Range Management 34:242-243. Macdonald, D. W., F. G. Ball, and N. G. Hough. 1980. The evaluation of home range size and configuration . using radio tracking data. Pages 405-424 in C. J. Amlaner, Jr. and D. W. Macdonald, editors. A handbook on biotelemetry and radio tracking. Pergamon Press, Oxford, England. Minta, S. C. 1992. Tests of spatial and temporal interaction among animals. Ecological Applications 2: 178204. Mooney, C. Z., and RD. Duval. 1993. Bootstrapping: a nonparametric approach to statistical inference. Sage Publishing Inc., Newbury Park, California, USA. �258 Nichols, L. Jr. 1957. Forage utilization by elk and domestic sheep in the White River National Forest. Thesis, Colorado State University, Fort Collins, Colorado, USA. Olsen, F. W., and R M. Hansen. 1977. Food relationships of wild free-roaming horses to livestock and big game, Red Desert, Wyoming. Journal of Range Management 30:17-20. Pickford, G. D. and E. H. Reid. 1943. Competition of elk and domestic livestock for summer range forage. Journal of Range Management 7:328-332. Rost, G. R 1975. Responses of deer and elk to roads. Thesis, Colorado State University, Fort Collins, Colorado, USA. Schultz, R D., and 1. A. Bailey. 1978. Responses of national park elk to human activity. Journal of Wildlife Management 42:91-100. Stevens, D. R 1966. Range relationships of elk and livestock, Crow Creek drainage, Montana. Journal of Wildlife Management 30:349-363. Turchin, P. 1998. Quantitative analysis of movement: measuring and modeling population redistribution in animals and plants. Sinauer, Sunderland, Massachusetts, USA. Ward, A. L., J. J. Cupel, A. L. Lea, C. Z. Oakeley, and R W. Weeks. 1973. Elk behavior in relation to cattle grazing, forest recreation, and traffic. Transcripts from the 38th American Wildlife Natural Resource Conference 38:327-337. Webster, B. A., and R J. Brooks. 1981. Social behavior of Microtus pennsylvanicus in relation to seasonal changes in demography. Journal of Mammalogy 64:738-751, White, G. C., and R A. Garrott. 1990. Analysis of radio-tracking data. Academic Press, San Diego, California, USA. Wright, K. L. 1983. Elk movements, habitat use, and the effects of hunting activity on elk behavior near Gunnison, Colorado. Thesis, Colorado State University, Fort Collins, Colorado, USA. Worton, B. J. 1987. A review of models of home range for animal movement. Ecological Modeling 38:277298. Table 3.1. Paired mean rank, randomized mean rank, 95% confidence interval of the randomized mean rank, and the probability of observing the sample rank or greater under the null hypothesis that elk locations are random with respect to sheep locations. Paired mean rank is the observed rank of temporally paired observations, and randomized mean rank is the expected rank of the temporally paired observations if elk are located randomly with respect to sheep. Number of Paired Randomized Randomized observed rankings mean rank mean rank 95%CI 1996 1,844 11.34 11.48 [11.47,11.49] 0.737 1997 1,790 12.14 11.97 [11.96, 11.98] 0.199 AU elk <5 km of a sheep hand on any day 150 1996 1997 124 11.71 10.59 11.48 11.93 [11.45, 11.52] [11.89,11.96] AU elk <1 km of a sheep hand on any day 34 1996 1997 19 9.09 9.84 11.49 12.00 [11.41,11.57] [11.90,12.10] P'?:.Ro AU elk :;" 0.342 0.990 _: 0.969 0.912 �259 t N Meeker Glenwood Rifle km 0 Springs !II 10 20 a ~ .A *• BLM Land Flat Tops Wilderness State Wildlife Land Study Area Boundary Forest Service Land Sheep Locations 1996 Sheep Locations 1997 Towns Figure 3.1. Sheep bands located during flights 20 July-l0 October 1996-1997, and land ownership in the White River Area, Colorado. Note that all white area (blank area) is private land. �260 Day Elk Temporally paired and unpaired distances Sheep Band PAIRED 1 2 unpaired unpaired 3 22 2 3 2 21 4 11 1 4 17 1 3 22 unpaired I Ullpalled unpaired unpaired 21 PAIRED 20 2 3 1 unpaired Ullpalled PAIRED unpaired Rank 22 22 15 7 2 i PAIRED 2 2 2 3 unpaired ~paired 21 1 unpaired Ulipalied PAIRED unpaired 9 15 7 21 . I . . 2 2 2 2 22 ·1 2 3 2 22 . unpaired 2 2 1 2 2 2 3 unpaired unpaired unpaired 22 4 2 22 PAIRED 11 88 88 88 1 2 3 PAIRED 18 1 7 2 22 unpaired unpaired . 88 88 88 88 22 i 2 3 88 22 2 11 unpaired 2 8 21 unpaired 1 unpaired 3 ':tr~ Figure 3.2. Representation of temporally paired and unpaired distances between elk and sheep. Only 1 sheep band was located per day, therefore sheep band 1 was located on day 1, sheep band 2 was located on day 2 §Jp. Ranks were randomly assigned as an example. Ranks of the temporally paired distances were extracted to . calculate the paired mean rank for a given year and spatial scale. "i} -c- ' �261 a =-7.1------- 200 f-----p-a-lre-d-~-;~~-ra-n-k Expected mean rank 150 >u = 5.5 c CD ::J 0" 100 e U- 50 I I 0 2 1 3 '~ 5 6 Rank 4 7 8 10 9 b 200 = 3.9 Paired mean rank Expected mean rank = 5.5 >- 150 o c CD g. e 100 U- 50 o I I' 1 2 3 4 5 6 Rank 7 8 11 9. 10 9 10 c = Paired mean rank 5.5 Expected mean rank 5.5 200 = ~15O_ c CD ::J 0"100 e U- 50 0 1 2 3 4 5 6 Rank 7 8 Figure 33. Theoretical distribution of paired ranks and paired mean rank under the expectation that (a) elk avoid domestic sheep, (b) elk are attracted to domestic sheep, or (c) elk are located randomly with respect to domestic sheep. Paired mean rank is the observed rank of temporally paired observations, and expected mean rank is the mean rank of all observations (temporally paired and unpaired) if elk are located randomly with respect to sheep. �262 a 120 ~--------------~--~~------~ = Paired mean rank 11.34 Expected mean rank = 11.48 100 i;'80 C CD g.60 CD .t40 20 o ~~~~~~~ 1 3 5 7 9 11 13 15 17 19 21 23 Rank b 15 Paired mean rank c 11.71 Expected mean rank c 11.48 >40 ·0 c :s CD C" e u, 5 I 0 1 3 5 7 I 9 11 13 15 17 19 21 23 Rank c 6,.--------------------------------, Paired mean rank 5 = 9.09 Expected mean rank = 11.49 1 o ~~~~~~~~~~~~~~~~ 1 3 5 7 9 11 13 15 17 19 21 23 Rank Figure 3.4. Observed distribution of paired ranks and paired mean rank for (a) all elk, (b) elk <5 km ofa sheep band, and (c) elk <I km ofa sheep band, 20 July-If October 1996 in the White River area, Colorado. Paired mean rank was the observed rank of temporally paired observations. Expected mean rank was the mean rank of all observations (temporally paired andunpaired) if elk were located randomly with respect to sheep, and was calculated from 999 samples that randomized on locations of sheep bands. �263 a 120 .-------~~~--~~~~----__. Paired mean rank c 12.14 Expected mean rank = 11.97 100 ()' 80 c ~ 60 c:r Q) U: 40 20 o ~~~~~~~~~~~~~~~~~ 1 3 5 7 9 11 13 15 17 19 21 23 Rank b 15 ~----------------------------, Paired mean rank = 10.69 Expected mean rank z:: 11.93 ~10 c Q) ::I 0" e u, 5 1 3 5 7 9 11 13 15 17 19 21 23 Rank c 6 ~-------------------------------, Paired mean rank 5' « 9.84 Expected mean rank « 12.00 1 o ~~~,,'-~'-~~~-'~-',,~04 1 3 5 7 9 11 13 15 17 19 21 23 Rank Figure 3.5. Observed distribution of paired ranks and paired mean rank for (a) all elk, (b) elk <5 Ian of a sheep band, and (c) elk <1 km ofa sheep band, 20 July-IO October 1997 in the White River area, Colorado. Paired mean rank was the observed rank of temporally paired observations .. Expected mean rank was the mean rank of all observations (temporally paired and unpaired) if elk were located randomly with respect to sheep, and was calculated from 999 samples that randomized on locations of sheep bands. �264 �265 APPENDIX 1: SAMPLE SIZE CALCULATION FOR HUNTER SURVEY Note, G. White (Colorado State University, personal communication) derived this sample size calculation. Hunt codes D-E-012-OI-A, E-E-012-01-A, E-F-012-01-M, E-M-012-01-M, D-M-012-01-M, D-E-033-01-A, E-E033-01-A, E-F-033-01-M, E-M-033-01-M, and D-M-033-OI-M used the study area during the study. To estimate total number of hunters on any day with a 95% confidence interval within ±I50 hunters, the binomial distribution and proportion of licenses holders was used to derive the number of license holder to be surveyed, where: N Number of licenses holders (number of individuals purchasing licenses), p Estimated proportion of hunters hunting, n Number of licenses holders sampled/surveyed, H Estimated number of hunters hunting, N-n N Finite population correction factor, if = pN. and ~r(H~)- N -n N2 va --- N :;rf~) _ N -n N2 p{l- p) Vcu.\p --- N n • The maximum variance will occur if 50% of the license holders hunt (i.e., p = 0.5), with the variance decreasing for larger or smaller percentages. For N license holders, the sample size (n) required to obtain a 95% confidence interval of ±I50 with a. = 0.05 and P = 0.5 is: On most days, the confidence interval will be less than ±I50 because not exactly 50% of the hunters will be hunting. For example, if 80% oflicense holders are hunting, then the variance of p is reduced from 0.25/n to 0.16/n (without the finite population correction factor). Thus, the confidence interval will be considerably smaller than ±150 hunters. �266 �267 APPENDIX 2: STANDARD ERRORS FOR DAILY GMU ESTIMATES OF HUNTERS AND ATVS AFIELD Estimated nwnber of hunters afield per day by GMU and standard errors of the estimates for 1996. DATE S724796 8125196 8126/96 8/27/96 8/28/96 8129196 8/30/96 8/31196 911196 912196 9/3/96 9/4/96 9/5196 9/6/96 9/7/96 9/8/96 9/9/96 9/10/96 9/11196 9/12196 9/13/96 9/14/96 9/15196 9/16/96 9/17/96 9/18/96 9/19/96 9120196 9121196 9122196 9123/96 9124/96 9125196 9126/96 9127/96 9128/96 9129/96 9130/96 10/1196 10/2196 10/3/96 10/4/96 10/5/96 10/6/96 RI2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 568 551 498 460 410 362 314 282 209 155 124 III 91 80 80 74 73 70 64 61 63 66 51 Estlmatea nunters aflelo H23 H24 210 127 211 135 176 129 157 119 151 106 122 85 124 84 154 121 150 132 118 126 109 105 112 105 106 94 116 88 161 119 147 118 120 116 114 115 113 113 112 109 103 113 526 421 531 400 381 265 343 243 296 197 241 163 211 133 117 186 113 47 77 30 72 32 62 25 50 22 43 28 75 36 64 33 37 26 33 29 :3"1 33 38 31 39 25 46 19 42 21 H33 21S 226 185 176 152 129 130 177 175 151 133 119 112 118 163 146 74 69 65 71 74 211 213 94 78 52 43 39 34 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Stanoaro error of estimates H23SE H24SE RI2SE 17 14 0 14 17 0 14 16 0 14 15 0 '15 13 0 11 14 0 11 14 0 15 14 0 15 14 0 14 14 0 13 13 0 13 13 0 13 13 0 14 12 0 16 14 0 15 14 0 14 14 0 14 14 0 14 13 0 14 13 0 13 14 0 25 22 23 22 25 23 20 16 23 16 20 22 15 19 21 14 17 21 13 16 20 12 15 19 13 9 18 11 7 16 7 II 14 6 14 10 6 12 9 7 12 9 8 12 II 7 11 11 II 7 8 7 II 8 7 II 8 10 7 8 II 7 8 11 6 9 10 6 9 H33SE IS 18 17 16 15 14 14 16 16 15 15 14 13 14 16 15 II II 10 11 II 16 16 8 8 6 6 6 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Note: H12 represents estimated number of hunter afield in GMU 12, H23 represents estimated number of hunter afield in GMU 23, etc. Hl2SE represents the standard error of the estimated number of hunters in GMUI2, H12SE represents the standard error of the estimated number of hunters in GMU23, etc. �268 Estimated number of hunters using ATVs afield per day by GMU and standard errors of the estimates for 1996. DATE 8/24/96 8125196 8126/96 8127/96 8128/96 8129/96 8/30/96 8/31196 911196 912196 9/3/96 9/4/96 915196 9/6/96 917196 918/96 9/9/96 9/10/96 9/11196 9/12196 9/13/96 9/14/96 9/15196 9/16/96 9/17/96 9/18/96 9/19/96 9120196 9121/96 9122196 9123196 9124/96 9125196 9126/96 9127/96 9/28/96 9129196 9/30/96 1011196 1012196 10/3/96 10/4/96 10/5/96 10/6/96 Standard error of estimates Estlmatea nunters using A I 'Vs afield HAI2 HA23 HA23SE HA24SE HA24 HA33 HAI2SE HAJ3SE 48 0 20 51 0 9 6 9 60 47 0 20 52 9 9 41 44 0 0 17 8 5 9 41 43 0 8 5 0 16 9 4 0 40 13 41 0 8 8 0 36 12 41 0 8 4 8 4 0 36 12 36 0 8 8 4 0 29 55 0 7 13 10 26 7 4 0 0 13 51 9 5 0 17 11 0 4 50 10 0 17 14 47 0 5 5 9 0 17 5 14 5 36 0 8 0 16 5 8 35 0 3 8 0 32 7 8 33 0 3 8 0 49 7 57 0 9 3 10 46 0 48 9 6 0 3 9 0 40 8 6 26 0 3 7 0 39 7 23 0 8 3 7 0 8 39 7 23 0 3 7 0 33 5 18 0 8 2 6 0 28 7 4 11 15 0 5 50 14 139 77 45 10 9 8 50 136 77 14 10 42 9 8 47 11 90 13 59 9 9 3 46 85 11 8 55 12 9 3 37 77 10 8 46 12 8 3 36 56 7 39 8 8 9 2 35 50 34 6 9 8 7 2 40 53 7 27 6 8 6 2 31 20 6 4 9 0 7 0 22 16 2 0 6 5 1 0 17 17 4 0 6 6 2 0 15 12 4 0 5 4 2 0 12 12 4 0 4 4 2 0 9 15 7 0 4 5 3 0 9 24 9 4 7 0 4 0 6 19 9 0 3 6 4 0 2 10 8 0 4 4 0 2 4 8 0 3 4 0 2 4 8 0 0 3 4 2 4 8 0 0 3 4 0 7 6 0 0 0 3 3 3 10 0 2 0 0 4 0 3 7 0 2 0 0 3 0 Note: H12 represents estimated number of hunter afield in GMU 12, H23 represents estimated number of hunter afield in GMU 23, etc. H12SE represents the standard error of the estimated number of hunters in GMU12, H12SE represents the standard error of the estimated number of hunters in GMU23, etc. �269 Estimated number of hunters afield per day by GMU and standard errors of the estimates for 1997. Standard error of estimates Eshmatea hunters afIeld DATE 8/23/97 8/24/97 8/25/97 8/26/97 8127197 8128/97 8129/97 8/30/97 8/31197 9/1197 912197 9/3/97 Hl2 328 H23 178 H24 124 314 282 252 168 138 123 115 119 117 226 206 191 220 211 191 151 139 9/4/97 115 915/97 9/6/97 107 130 9/1/97 141 918/97 9/9/97 9/10/97 9/11/97 9/12197 9/13/97 9114197 9/15197 9116197 9117/97 9/18/97 9/19/97 9120197 9121197 9122197 9123/97 9124/97 9125197 9/26/97 9127197 9128/97 9/29/97 9/30/97 10/1/97 10/2/97 10/3/97 1014197 1015/97 132 120 112 122 142 416 391 364 345 300 109 100 H33 o 17 15 o 20 20 20 14 I3 13 14 14 II 1I II 19 17 14 13 12 12 17 15 15 13 13 I3 II 10 10 10 11 11 12 16 15 12 12 12 16 17 23 il 12 11 82 o 66 119 97 81 77 98 96 87 56 49 51 62 61 o o o o o o o o o o o o o 101 87 65 67 68 68 71 484 483 63 66 67 60 61 477 470 462 409 399 .0 o 483 477 401 403 379 317 19 19 16 17 17 22 21 21 20 18 17 205 449 442 404 309 172 275 . 160 112 223 o o o o o o o 57 50 53 52 50 46 50 o o 44 50 75 72 79 o 77 o 39 87 o 37 37 40 45 71 67 91 51 98 o o o o o o o 149 307 242 224 207 135 123 115 114 105 81 73 H24SE 15 23 22 21 92 ll5 110 64 H23sE 17 o o o 109 67 H12SE 23 249 275 240 161 148 127 126 130 142 137 16 14 o o o o o 14 12 14 II 26 26 25 25 24 22 21 19 16 10 10 10 10 10 o 9 o 10 10 10 34 III o 24 105 o 8 8 8 9 9 H33SE 0 1-5 15 14 0 13 o 12 o o o o o o o o o o o o o o o o 12 26 26 26 24 24 22 20 20 o o 25 25 23 23 23 22 20 21 19 20 16 15 15 15 14 13 12 12 12 12 12 18 17 10 8 7 15 15 16 16 16 13 12 I3 14 14 15 14 Note: H12 represents estimated number of hunter afield in GMU 12, H23 represents estimated number of hunter afield in GMU 23, etc. H12SE represents the standard error of the estimated number of hunters in GMU12, H12SE represents the standard error of the estimated number of hunters in GMU23, etc. �270 Estimated number of hunters using ATVs afield per day by GMU and standard errors of the estimates for 1997. DATE 8723797 8124/97 8/25197 8/26/97 8127197 8/28/97 8129197 8/30/97 8/31197 9/1197 912197 9/3/97 9/4/97 915197 9/6/97 9nl97 918/97 9/9/97 9/10/97 9/11197 9/12197 9/13/97 9/14/97 9/15197 9/16/97 9/17/97 9/18/97 9/19/97 9120197 9121/97 9122197 9123/97 9124/97 9125/97 9126/97 9127/97 9128/97 9129/97 9/30/97 10/1/97 1012197 10/3/97 10/4/97 1015197 Estlmatea flunters arlelO HAI2 RA23 HA24 43 70 34 36 70 39 32. 39 56 48 30 35 48 40 30 45 33 30 26 25 30 21 27 30 27 28 18 24 31 16 18 18 36 18 36 12 15 33 12 8 29 9 7 34 10 5 30 8 12 30 8 5 30 8 2 30 8 3 28 5 21 7 5 1I0 29 50 32 no 50 112 30 51 20 113 48 22 106 48 81 II 39 10 77 33 7 52 30 7 35 19 0 14 7 0 14 7 0 8 7 0 8 7 0 8 7 0 15 3 0 19 3 0 19 3 0 19 3 0 19 3 0 19 3 0 19 3 0 12 3 0 12 3 HA33 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 99 101 94 85 78 63 56 64 58 54 40 37 35 42 48 39 29 22 24 22 34 34 34 Stanaara error or ~stlmates RA23SE HA24SE HAI2SE HA33SE 12 8 9 0 12 9 8 0 10 9 8 0 8 9 8 0 9 9 0 8 8 8 9 0 8 7 0 7 7 8 0 8 8 6 0 8 8 5 7 0 8 6 0 6 6 8 0 5 8 5 5 0 7 4 3 0 8 4 3 0 8 4 3 0 8 4 5 0 4 3 8 0 8 4 0 3 8 3 0 4 7 3 0 14 8 13 7 8 14 8 13 14 8 13 8 14 8 12 6 14 8 12 6 12 8 11 5 4 12 8 10 10 8 II 3 6 II 3 8 4 11 0 6 6 4 9 0 4 4 0 9 0 4 4 9 0 4 4 9 0 6 3 10 9 0 6 3 8 0 6 3 7 0 6 3 7 6 0 3 7 0 6 3 9 0 6 3 9 0 5 3 9 0 5 3 Note: H12 represents estimated number of hunter afield in GMU 12, H23 represents estimated number of hunter afield in GMU 23, etc. H12SE represents the standard error of the estimated number of hunters in GMUI2, H12SE represents the standard error of the estimated number of hunters in GMU23, etc . . '. �-~!a White River Elk Movement Study Elk and Sheep .,:":,: Locations on July White River Elk Movement 19, 1996 . Elk and Hamilton Sheep Locations Hamilton ~ Study on July 24, "I:l l"'l 1996 Z =~ (,H Km o 40 20 ~ 00 0 ~ l"'l ~ ~ ~ == 0 ~ l"'l >-l >-( 00 ~ 00 = l"'l l"'l "I:l ~ 0 ~ Rio Blanco Rio Blanco > >-l >-( 0 z 00 0 Z LEGEND ;';), «;. * A • 'OJ. BlM land Flat Tops Wilderness State Wildlife Land Study Area Boundary White River National Forest White River Elk location Sheep location Towns 00 l"'l ~ l"'l ~ >-l l"'l =~ e- >-( ~ = >-l 00 N -.l •.... �~, , ~ t-J White River Elk Movement White River Elk Movement Study Elk and Sheep Locations on July 31. Elk and 1996 Sheep Locations Hamilton Hamilton Krn o 20 40 LEGEND 'BLM Land Flat Tops Wlldemess State Wildlife Land Study Area Boundary White River National Forest - White River .•. Elk Location Sheep Location • Towns !* Study on August 4. 1996 �White River Elk Movement Study Elk and Sheep Locations on August White River Elk Movement Study 14, 1996 Elk and Hamilton Sheep Locations on August 28, 1996 Hamilton f<m o Rio Blanco 20 40 Rio LEGEND BlM land Flat Tops Wilderness State Wildlife Land Study Area Boundary White River National Forest White River A Elk location Sheep location • Towns * tv -.I V..l �White River Elk Movement Study Elk: and Sheep Locations on August 20, 1997 Elk and Hamilton White River Elk Movement Study Sheep Locations on September 6, 1997 Hamilton Krn o .... 20 LEGEND BLM Land Flat Tops Wilderness State Wildlife Land Study Area Boundary White River National Forest !* - White River A. Elk Location Sheep Location Towns • 40 N -..l 00 �':.,,' White River Elk Movement Study Elk and Sheep Locations on September White River Elk Movement Study 17, 1997 Elk and Hamilton Sheep Locations on Septermber 24, 1997 Hamilton Km o 20 40 LEGEND BLM Land !* - A • Flat Tops Wilderness State Wildlife Land Study Area Boundary White River National Forest White River Elk Location Sheep Location, Towns N -...l \0 �tv gg White River Elk Movement Study ~:~ c r.c Sh e e c ~occ~:o;:s 0:1 Cc iob e r 1, ~997 White River Elk Movement Study Elk and r:"he::,,~) L()(/J;:r)n~: on October 5, 1997 Hamilton Hamiiton Km o . 20 40 LEGEND BlM Land Flat Tops Wildemess Area State Wildlife Area Study Area Boundary White River National Forest - White River .•. Elk Location ~ Sheep Location • Towns ! �281 Colorado Division of Wildlife .Wildlife Research Report July 1999 JOB PROGRESS REPORT Smreof ~C~o~lo~rnd==o~ _ Cost Center 3430 Mammals Program Project No. ....:.W.:....-....::l.::..5=-3-..:.R:::....-=12=-_ Work Package No. _--=3:..::0~0.:::.2 _ Elk Management Task No. --=-3 _ Monitoring and Managing Chronic Wasting Disease in Elk Period Covered: July 1, 1998 - June 30, 1999 Authors: M. W. Miller and C. T. Larsen Personnel: K. I. O'Rourke, T. R. Spraker, E. Wheeler, M. A. Wild, and E. S. Williams ABSTRACT Elk from throughout Colorado were examined for occurrence of chronic wasting disease using targeted surveillance. Between June 1998 and May 1999, 7 chronic wasting disease (CWD) cases were diagnosed . from among 12 "suspect" elk submitted from endemic game management units (GMUs) in northeastern Colorado. CWD was not diagnosed in any of 4 additional "suspect" elk submitted from elsewhere in Colorado, No cases ofCWD occurred among 23 adult elk held at CDOW's Foothills Wildlife Research Facility. �282 �283 MONITORING AND MANAGING CHRONIC WASTING DISEASE IN ELK M. W. Miller and C. T. Larsen P. N. OBJECTIVES (1) Design, conduct, and report results of: (a) targeted surveillance to estimate and detect changes in distribution of chronic wasting disease (CWO) in free-ranging elk populations; and (b) harvest or road-kill surveys to estimate and detect changes in prevalence of CWO in enzootic elk populations. (2) Design, conduct, and report results of experimental studies using captive elk naturally or experimentally infected with CWO. AGREEMENT OBJECTIVES (1) ( 2) Conduct and report results of targeted surveillance to estimate and detect changes in distribution of CWD in free-ranging elk populations statewide. Observe epizootiological features of naturally-occurring CWO in captive elk. MATERIALS AND METHODS Surveillance We monitored elk populations throughout Colorado for occurrence of CWO using a combination of targeted surveillance and harvest or road-kill surveys. These were organized and conducted as follows: Targeted (= clinical disease) surveillance: Elk showing clinical signs consistent with those seen in chronic wasting disease were collected by field personnel statewide and brain tissues examined for evidence of spongiform encephalopathy. The "suspect case" profile was defined as follows: • Species: elk • Age: ~ 18 months • Signs: emaciated and abnormal behavior &lor indifference to human activity &lor increased salivation &lor tremor, stumbling, incoordination &lor difficulty or inefficiency in chewing/swallowing &lor increased drinking and urination Where possible, submissions were subjected to complete necropsy; in some situations, only heads were available for examination and sampling. In all cases, histopathology of brain tissue (Williams �284 and Young 1993) was used to diagnose CWD; in some cases, immunohistochemistry (O'Rourke et aI., 1998) or other ancillary tests were used to confirm or support diagnoses. Harvest surveys: No elk harvest surveys were planned for this segment. Epizootiological Studies Epidemiology of naturally-occurring CWD in captive elk (Miller and Wild): We observed captive adult elk (n = 23) held at CDOW's Foothills Wildlife Research Facility for clinical signs ofCWD and submitted all mortalities for complete necropsy and histopathological examination. RESULTS AND DISCUSSION Surveillance Targeted (= clinical disease) surveillance: Between June 1998 and May 1999, 7 chronic wasting disease (CWD) cases were diagnosed among 12 "suspect" elk submitted from endemic game management units (GMUs) in northeastern Colorado; CWD was not diagnosed in any of 4 additional "suspect" elk submitted from elsewhere in Colorado. Since 1981,23 clinical CWD cases have been diagnosed in elk from Larimer County GMUs (Fig. 1). Harvest surveys: No elk harvest surveys were conducted in this segment, and none are planned for 1999-2000. Epizootiological Studies Epidemiology of naturally-occurring CWD in captive elk (Miller and Wild): None of the 23 captive adult elk held at CDOW's Foothills Wildlife Research Facility developed clinical CWD during June 1998-May 1999; 3 female elk that died during that period were not infected. We will continue to monitor this naturally-infected captive herd and examine all mortalities for evidence of CWD. A manuscript describing CWD epidemiology in captive elk during 1986-1997 was published in July (Miller et al., 1998). ACKNOWLEDGMENTS The statewide CWD monitoring and surveillance program described here relies heavily on efforts of dedicated field personnel throughout the Colorado Division of Wildlife, and truly represents a division-wide effort to improve our understanding and management of this important disease problems. In addition to those specifically listed, we collectively thank all of those regional and area biologists, district and area wildlife managers, volunteers, elk hunters, and lOthers'who assisted by submitting suspect cases, harvested animals, or road-killed animals throughout the year. �285 LITERA TURE CITED Miller, M. W., M. A. Wild, and E. S. Williams. 1998. Epizootiology of chronic wasting disease in captive Rocky Montain elk. 1. Wildl. Dis. 34: 532-538. O'Rourke, K. I., T. V. Baszler, J. M. Miller, T. R. Spraker, I. Sadler-Riggleman, and D. P. Knowles. 1998. Monoclonal antibody F89/160.1.5 defines a conserved epitope on the ruminant prion protein. J. Clin. Microbiol. 36: 1750-1755. Williams, E. S., and S. Young. 1993. Neuropathology of chronic wasting disease in mule deer (Odocoi/eus hemionus) and elk (Cervus e/aphus nelsoni). Veterinary Pathology 30: 36-45. Prepared by _ Michael W. Miller Wildlife Research Veterinarian River River 50 100 Figure 1. Between March 1981 and June 1999, 23 clinical CWD cases have been diagnosed in elk from GMUs in Larimer County, Colorado. �286 �287 Colorado Division of Wildlife Wildlife Research Report July 1999 JOB PROGRESS REPORT Smreof Project No. Work Plan No. Task No. Period Covered: Authors: Personnel: ~C~o~lo~r~ad~o~ _ _,W_,_-....:1"""5.::.3..••. R"--~12=__ _ -=3;..:::.0.::...04..:...,_ _ ___:3::.._ _ Cost Center 3430 Mammals Program Management of Other Ungulates Strategies for Managing Pasteurellosis in Mountain Sheep Populations July 1, 1998 - June 30, 1999 M. W. Miller, H. J. McNeil, and J. L. George K. Larsen, S. Berry, J. Vayhinger ABSTRACT A pasteurellosis epidemic that began in the Sugarloaf Mountain subpopulation of the Tarryall-Kenosha bighorn herd complex extended to two adjacent subpopulations (Black Canyon, Twin Eagles) during winter 1998-1999. Sick and dead bighorns were found in the Twin Eagles subpopulation by December 1998, and in the Black Canyon subpopulation by January 1999. Based on preliminary data, vaccination with PhSV 911 mo earlier did not appear to improve bighorn survival during subsequent epidemics. A group of bighorns translocated from Dome Rock to the Holy Cross Wilderness received PhSV during winter 1998-1999. No adverse effects have been observed in free-ranging bighorns that received PhSV last winter, or during 1997-1998. We began developing and evaluating a modified-live P. haemolytica vaccine. Using polymerase chain reaction (PCR) techniques, we screened candidate carrier strains and further characterized bacterial genomes linked to leukotoxin production. Leukotoxin A (lleta) gene of P. haemolytica TI0 ECO-loowas successfully amplified, and subsequent sequencing revealed that the lleta gene of this bighom-derived strain was approximately homologous to the published lkta gene of P. haemolytica TI0. We successfully ligated 123 and 1767 bp gene fragments. Initial attempts to use PCR to increase yields for cloning failed, but such efforts continue. Plasmid transfers of lleta are planned for next segment. �288 �289 EXPERIMENTS TO IDENTIFY AND MANAGE STRESS IN MOUNTAIN SHEEP POPULATIONS M. W. Miller, H. J. McNeil, and J. L. George P. N. OBJECTIVES 1. Design, conduct, and report on experiments evaluating Pasteurella haemolytica vaccines and vaccine delivery systems and identify those with potential application in managing free-ranging bighorn populations. 2. Design, conduct, and report on field experiments evaluating management strategies for preventing pasteurellosis epizootics in bighorn populations SEGMENT OBJECTIVES 1. Report results of an experiment evaluating options for delivering a multivalent Pasteurella haemolytica supernatant vaccine to bighorn sheep. 2. Provide multivalent Pasteurella haemolytica supernatant vaccine for use in select bighorn sheep management activities statewide. 3. Begin development and evaluation of a modified-live Pasteurella haemolytica vaccine for use in bighorn sheep. STRATEGIES FOR MANAGING PASTEURELLOSIS IN MOUNTAIN SHEEP POPULATIONS Inability to control infectious disease outbreaks and subsequent mortality in mountain sheep populations represents a significant obstacle to long-term success in their management. Although the "bighorn pneumonia complex" has been studied intensively for over 3 decades, little is known about many aspects of its etiology and epizootiology. Moreover, management interventions recommended for preventing or controlling this problem remain untested. Although viral, bacterial, and parasitic agents have all been incriminated in these outbreaks, Pasteurella spp. are perhaps the most common pathogens associated with bronchopneumonia in bighorns (Miller, 1999). Two species, P. haemolytica and P. multocida, and several biotypes and/or serotypes within those species, have been isolated from bighorns during epizootics. Unfortunately, despite extensive diagnostic and experimental investigation, the epizootiology of pasteurellosis in wild bighorn populations is poorly understood. In the absence of knowledge about the epizootiology of pasteurellosis, effective strategies for managing pneumonia llr bighorn populations have not emerged. Here, we report on a series of ongoing research studies designed to improve knowledge about various aspects of pasteurellosis epizootiology and management in bighorn sheep. �290 METHODS AND MATERIALS Delivery of Pasteurella haemolytica supernatant vaccine to bighorn sheep (McNeil and Miller): Last segment we conducted an experiment evaluating options for delivering a multivalent Pasteurella haemolytica supernatant vaccine (PhSV) to bighorn sheep. A draft manuscript describing that research was accepted for publication in the Journal of Wildlife Diseases (see abstract, Appendix A). Use of Pasteurella haemolytica supernatant vaccine in free-ranging bighorn sheep (Miller, George, and Vayhinger): We continued evaluating survival of free-ranging bighorn sheep vaccinated with PhSV in the face ofa pneumonia epidemic during winter 1997-1998 by comparing survival rates of vaccinated bighorns to those of unvaccinated individuals. Our evaluation was conducted in conjunction with other ongoing studies of the Tarryall-Kenosha bighorn herd complex. In addition, PhSV was used in select field operations during the winter of 1998-1999 (Table 2). We also continued exploring ways of adding Pasteurella multocida antigens to our vaccine formulation. Development of a modified-live Pasteurella haemolytica vaccine for bighorn sheep (McNeil, Lo, and Miller): We prepared a study plan and began developing and evaluating a modified-live Pasteurella haemolytica vaccine. Highlights of preliminary laboratory technique development and evaluations are reported. Bacteria: Pasteurella haemolytica TI0 <Eco-lOO) was obtained from a bighorn sheep that died during a pasteurellosis epidemic in southcentral Colorado in 1990 (Kraabel et al., 1997). Bacterial stocks were kept frozen at -70° C in Brain Heart Infusion Broth (BlllB) with 15 % glycerol. Thawed stock was used to inoculate blood agar plates incubated at 37° C overnight (O/N). A small volume (50 rnl in a 175 rnl Erlenmeyer flask) of BlllB was inoculated with a single colony from the OIN agar plate and incubated at 37° C, 120 rpm OIN. The resulting culture was used to extract DNA for use in polymerase chain reactions (PCR). Preparation of genomic DNA: Two commercial kits were used to extract maximum amounts of genomic DNA fromP. haemolytica TI0 (ECO-lOO). A QIAGEN Genomic-tip" (QIAGEN Inc., Mississauga, ON, Canada) was used and DNA was extracted according to the Bacterial DNA Isolation Protocol in the QIAGEN Genomic DNA Handbook. A PUREGENE® genomic DNA isolation kit (Gentra Systems, MN, USA) was also used to improve yields. Genomic DNA was used as template for subsequent PCR. peR: PCR was utilized to amplify the leukotoxin A gene (lkta) of P. haemolytica TI0 <Eco-lOO).Many different primers and conditions were assessed. The gene was successfully amplified using primers based on the published P. haemolytica TI0 leukotoxin A gene sequence (Genbank Accession Z26247). 5'-ATGGGAACT AAACTAACCCT-3' 5'-TAAGCTGCTCTAGCAAATTG-3 , Conditions were optimized at 3.5 mM [Mg"'] using the genomic DNA prepared by the PUREGENE® method as template. The thermal cycler (Perkin Elmer-Cetus, Norwalk, Connecticut, USA) was programmed for 94 C for 30 seconds, 51 C for 30 seconds, 72 C for 3 minutes and 30 seconds. This was repeated for 30 cycles. Two sections of the lkta gene also were amplified via PCR for use in subsequent ligation. Similarities between the sequence of the lkta gene for EcO-lOO and P. haemolytica Al (Genbank Accession M20730) were considered. Previous work in our laboratory (R. Lo, unpubl. data) used two existing NaeI sites on the �291 lkta gene to delete a section of the gene and ligate the smaller fragments together to form a truncated gene. Although the Nael sites on the lkta Ec0-1oo were not a perfect match, using primer design, two sections were amplified. These sections incorporated exact NaeI restriction sites. The 123 base pair segment from the 5' end of the gene was amplified using the following primers: 5'-GGATCCCTIATGGGAACTAAAC-3 5'-GTCTGGCCGGCTIGGGTIAA-3' , The 1767 base pair segment from the 3' end of the gene was amplified using the following primers: 5'-TGCCGCCGGCTCTGTGGTI -3' 5'-CGAAACAATCTAGAGTIGCCAATC-3 , Conditions were optimized at 3.5 mM [Mg"'] using the genomic DNA prepared by the PUREGENE® method as template. The thermal cycler was programmed for 94 C for 30 seconds, 59 C for 30 seconds, 72 C for 3 minutes and 30 seconds. This was repeated for 30 cycles. PCR was also used to try to increase the amount of ligated product. 5'-GGATCCCTIATGGGAACTAAAC-3 , 5'-CGAAACAATCTAGAGTIGCCAATC-3 The following primers were used: , Conditions were optimized at 3.5 mM [Mg'"] using gel purified ligated product as the template. The thermal cycler was programmed for 94 C for 30 seconds, 64 C for 30 seconds, 72 C for 3 minutes and 30 seconds. This was repeated for 30 cycles. All PCRproducts were run on 0.75 % agarose gels (85 V for 23-45 minutes) and stained with ethidium bromide. Bands were visualized using ultra-violet (UV) light and the image captured by a GelDoc 1000 machine (Bio-Rad Laboratories, Mississauga, ON, Canada). peR Product PUrification: PCR products to be sequenced or used for further manipulation were purified using a QIAGEN QIAquick PCR purification kit". Purified product was run on gels as before purification to ensure that the band in question was still visible. In instances where extraneous bands were visible, gel extraction was used to isolate the desired product. A low melting point 1.0% agarose gel was loaded with product and run at low voltage overnight (8-9 V, 1216 hours). The gel was then stained with ethidium bromide and visualized using UV light on a transilluminator. The desired band was cut out using a sterile scalpel blade and put into a tube. The tube was heated to melt the agar. This material was then purified in the same manner as the non-gel purified PCR products. Restriction Enzyme Digestion: Purified 123 bp and 1767 bp fragments were digested using NaeI (promega 'Corporation, Madison, WI, USA) with the recommendedbuffer for 1 hour at 37 C. The enzyme was then inactivated at 60 C for 15 minutes. A lul aliquot was run on a 0.75% agarose gel to ensure product integrity. These products were to be ligated together. Restriction enzyme digestion was also used for mapping the ligated product. Digestions were performed using NaeI, ApaI (promega Corporation, Madison, WI, USA) and SacI (New England Biolabs Ltd., �294 APPENDIX A EFFECTS OF DELIVERY METHOD ON SEROLOGICAL RESPONSES OF BIGHORN SHEEP TO A MULTIVALENT PASTEURELLA HAEMOLYTICA SUPERNATANT VACCINE Heather 1. McNeil, I Michael W. Miller/ Jennifer A. Conlon," Ian K. Barker, I and Patrica E. Shewen' I 2 3 Department ofPathobiology, Ontario Veterinary College, University of Guelph, Guelph, Ontario, NIG 2Wl, Canada Colorado Division of Wildlife, Wildlife Research Center, 317 West Prospect Road, Fort Collins, Colorado 80526-2097, USA Merial Incorporated, 115 Transtech Drive, Athens, Georgia, 30601-1649, USA ABSTRACT: The efficacy and safety of a multivalent Pasteurella haemolytica supernatant vaccine (serotypes A2 and TI0) using different delivery systems were examined in captive Rocky Mountain bighorn sheep (Ovis canadensis canadensisi. Twenty bighorn sheep were grouped according to baseline leukotoxin neutralizing antibody titers (~2 or >210g2-1) and vaccination history (previously vaccinated or unvaccinated). Within these groups, animals were randomly assigned to one of two delivery treatments: hand injection (control) or biobullet implantation. All bighorns received a single dose from the same lot of vaccine (n = lO/treatment); four additional animals were injected intramuscularly with 0.9% saline as unvaccinated sentinels. Mild, transient lameness one day after hand injection or biobullet implantation was the only adverse effect. Serum neutralizing antibody titers to P. haemolytica leukotoxin differed among delivery treatments (P = 0.009) and among baseline titer/vaccination history groups (P = 0.013). Neutralizing titers were highest among hand-injected bighorns. Although neutralizing titers were lower among implanted bighorns than hand-injected controls at 1 wk (P = 0.002) and 2 wk (P = 0.021) after vaccination, seroconversion rates in response to implantation (6/10) and hand injection (9/10) did not differ (P = 0.303). Agglutinating antibody titers to TI 0 were high and did not vary over time or between delivery treatments. Agglutinating antibody titers to A2 in the hand-injected controls were not different (P ~ 0.07) than those in bighorns vaccinated with biobullet implantation. These data demonstrate that although hand injection elicits higher absolute titers, biobullet implantation may also stimulate effective antibody responses to P. haemolytica supernatant vaccine. Further evaluation ofbiobullet vaccination against pneumonic pasteurellosis in free-ranging populations of wild bighorn sheep is warranted. Key words: Bighorn sheep, Ovis canadensis, Pasteurella haemolytica, pasteurellosis, vaccine delivery, vaccination. �295 Colorado Division of Wildlife Wildlife Research Report July 1999 JOB FINAL REPORT Smreof Project No. Work Plan No. Job No. C~o~l~o~rad~o _ _,W:..!......• 1""'"5.::..3-...:.R.::..-~12=__ _ .::..30"-'0"-'4'-_ 4...!-.. _ Mammals Research Mountain Goat Investigations Mountain goat numbers, distribution, and dispersal in the northern Collegiate range. Period Covered: July 1, 1998 - June 30, 1999 Author: D. F. Reed ABSTRACT A manuscript titled "Mark-resight population estimates of mountain goats in Colorado" (Reed and Vayhinger [in preparation]) was prepared. Additionally, a manuscript titled "A conceptual interference competition model for introduced mountain goats" was prepared and submitted to the Journal of Wildlife Management (Reed [in review]) and collaboration with Natural Resources Ecology Laboratory (NREL) occurred to extent and refine some of the Mt Evans dam. �296 �297 MOUNTAIN GOAT NUMBERS, DISTRlBUTION, AND DISPERSAL IN THE NORTHERN COLLEGIATE RANGE Dale F. Reed P.N. OBJECTIVE To improve estimates of mountain goat populations by mark-resight methodology, to determine distribution, and to estimate dispersal rates in an increasing mountain goat population. SEGMENT OBJECTIVE 1. Analyze data and prepare manuscripts. STUDY AREA The study area is described in the Program Narrative (Reed 1995). METHODS AND MATERIALS The methods are outlined in the Program Narrative and 1995 and 1996 progress reports (Appendix A in Reed 1995, Reed 1996). RESULTS Results have been reported in Reed (1996), Reed (1997), and in a manuscript titled "Mark-resight population estimates of mountain goats in Colorado" (Reed and Vayhinger [in preparation]. Additionally, a manuscript titled "A conceptual interference competition model for introduced mountain goats" was prepared and submitted to the Journal of Wildlife Management (Reed [in review]) and collaboration with NREL (ref Rocky Mountain National. Park contract) occurred to extent and refine some of the Mt Evans data by evaluating spatial distribution, habitat segregation, and implications of competitive displacement. LITERATURE CITED Reed, D. F. 1995. Mountain goat numbers, distribution, and dispersal in the northern Collegiate range. Colo. Div. Wildl. Res. Rep. July, 223-234pp. Reed, D. F. 1996. Mountain goat numbers, distribution, and dispersal in the northern Collegiate range. Colo. Div. Wildl. Res. Rep. July, 255-263pp. ,. Reed, D. F. 1997. Mountain goat numbers, distribution, and dispersal in the northern Collegiate range. Colo. Div. Wildl. Res. Rep. July, 165-167pp. Prepared by _ Dale F. Reed �298 �299 Colorado Division of Wildlife Wildlife Research Report July 1999 JOB FINAL REPORT Smreof Project No. Work Packages Task No. Period Covered: C==ol=o=ra=d=o~ _ Cost Center 3430 Mammals Program W,..:....:.._-..:.;15=3:;_-=R:.....:-l=2'--_ 7120, 7210, 7610, Mammals Program Library, Publications, 8120 and 8160 Administrative, and Research Services ~2=-_ Library, Publications, Administrative, and Research Support Services July I, 1998 - June 30, 1999 Author: J. Boss, N. Wild. M. Wild. M. Miller, G. White, and B. Gill Personnel: Diane Haerter, K Larson, R M. Post Vieira, A. Dharman M Bartmann, D. C. Bowden. D. J. Freddy, M. M. Conner, ABSTRACI' Task 1 (WorkpackaJ!e 7210) During the Segment the following were accomplished: 74 2010 841 1375 2166 o Publications acquired by the Research Center Library for the use of Colorado Div. of Wildlife employees, cooperators, wildlife educators, and the public. Items of information delivered to Colorado Div. of Wildlife employees, cooperators, wildlife educators, and the public, resulting from requests and literature searches. Items of information cataloged into the electronic and card catalogues, which including duplicates and additional volumes, expanded the Research Center Library inventory to 18,420 items. Computer scanned items of information entered into the electronic catalogue for the maintenance of the circulation system of the Research Center Library. Items checked out by Colorado Div. of Wildlife employees, cooperators, wildlife educators, and the public indicating satisfaction of library services. Comments received from Colorado Div. of Wildlife employees, cooperators, wildlife educators, and the public, through a . Suggestion Box' to monitor and evaluate library patron satisfaction with library services. Task 2 (Workpackage 7120) Progress made on each Objective-during this Segment is reported below. 1. 2. 3. 4. Assisted 4 Wildlife Researchers in developing publication-ready manuscripts. Assisted Wildlife employees in creating graphics and visual materials for 46 presentations (slides, overheads, handouts, posters). Published 1998 Federal Aid Abstracts and distributed them to approximately 300 Division personnel and 150 outside agencies and libraries. Published 2 Special Reports, Nos. 73 and 74. This process involved editing; selecting and creating tables, photos, and graphics; preparing camera-ready copy; having books printed at local vendor; and �300 distributing to Division personnel and outside libraries, agencies, etc. These reports are as follows: Modeling the Population Dynamics of Bighorn Sheep: A Synthesis of Literature, No. 73 Columbian Sharp-tailed 5. Grouse Harvest and Wing Analysis: Implicationsfor Management, No. 74 Assembled, published, and distributed the FY 97-98 Wildlife Research Reports. Task 3 <Workpackage 8160) The Colorado Division of Wildlife's Foothills Wildlife Research Facility (FWRF) maintained captive animals (annual total: 165 wild ungulates of 5 species, 11 domestic cattle, and 33 prairie dogs) and facilities in support of research on chronic wasting disease (CWO) in deer and elk (and potential transmission to domestic cattle and pronghorn), deer and elk contraception, pneumonia immunization of bighorn sheep, and prairie dog research for black-footed ferret recovery. During the year, 20 additional mule deer, 10 bighorn sheep, and two elk were added to support research efforts. Thirty-three animals died or were euthanatized for health problems and an additional 12 animals were euthanatized as part of study protocols or facility management programs. Chronic wasting disease (CWO) was again the predominant mortality factor (n = 9) in cervids. A high quality of animal care and facility maintenance was provided by temporary, work-study, YCC employees, interns, and volunteers. Volunteers (including interns) contributed 846 hr (equal to about 0.4 FfE) work at FWRF. The high standard of care is in part reflected by the finding of compliance under the Animal Welfare Act during the annual QSDA APIDS inspection of FWRF. Repairs and modifications were performed to improve safety and/or function of facilities at FWRF. Task 4 <Workpackage 7610) Sixty five cases of wildlife mortalities were examined during the segment to assess cause of death. Six mountain sheep were examined, of which 4 died fromPasturella spp. pneumonia. Two pronghorn tested positive for hemorrhagic disease. Two raccoons tested positive for Canine Distemper Virus. Thirty two birds were examined that were suspected of succumbing to botulism; all 32 tested positive for botulism. Task 5 (Workpackage 7610) Consulting services were provided to 20+ individuals within the Terrestrial Wildlife Section. DEAMAN software was upgraded and revised as a Windows 95 program. A spreadsheet big game population model was developed and distributed to terrestrial biologists. Two research proposals which addressed competing causes of the mule deer decline were prepared and submitted for funding consideration. Harvest estimators were developed for several terrestrial wildlife species. A manuscript which reported on optimal allocations of manpower and fiscal resources to inventory mule deer was prepared and submitted to The Journal of Wildlife Management. Additional analyses were conducted on mule deer population data to identify causes of declining populations. Data for both deer and elk support a response of young:female rations to the previous year's male:female ratios, about 0.25 fawns: 100 does for each change of 1 buck: 100 does in deer populations and 0.28 calves per 100 cows for each change of 1 bull: 100 cows in elk populations. These effects are not adequate to compensate for the much greater decline in fawn:doe and calf:cow ratios that have been observed over the past 20 years in Colorado. Differences observed between DAUs suggests that only some of the DAUs might show an increase in young:female ratios were male:female ratios increased, and then only a small increase in young:females is predicted. There does not appear to be a threshold male:female ratio for either species where there is a drastic decline in young:female ratios. Task 6 (Workpackage 7610) All objectives were met as planned. �301 LIBRARY, PUBLICATIONS, ADMINISTRATIVE, AND RESEARCH SUPPORT SERVICES P. N. OBJECTIVES 1. Provide effective library, publication, editorial, veterinary, pen and animal maintenance, wildlife disease diagnosis and monitoring, biometrical, supervisory, and administrative services at a minimal cost by centralizing them and enhancing accountability for support services. SEGMENT OBJECTIVES Task 1 (Workpackage 7210) 1. 2. 3. 4. Acquire reference library materials requested by wildlife research staff members and cooperators. Assist wildlife research staff in conducting literature searches and in acquiring reprints of requested materials. Maintain electronic and card catalogues of all research library holdings. Develop a process for monitoring and evaluating customer satisfaction with library services. Task 2 (Workpackage 7120) 1. 2. 3. 4. 5. 6. Assist Wildlife Researchers in developing publication ready manuscripts. Assist Division of Wildlife employees in developing graphics and visual materials for oral presentations. Publish Federal Aid Abstracts and distribute to Division Personnel and outside agencies. Create camera-ready copy, print, and publish Special Reports as requested and submitted by Terrestrial Wildlife Researchers Assemble and publish the FY 97-98 version of Wildlife Research Reports. Develop a process for monitoring and evaluating customer satisfaction with publication services. Task 3 (Workpackage 8160) 1. Maintain research facilities and experimental animals in support of research on chronic wasting disease, deer contraception, and pneumonia immunization of mountain sheep. Task 4 (Workpackage 7610) I. 2. 3. Maintain systems for submitting, diagnosing, and recording on sporadic disease cases in wild animals throughout Colorado. Maintain databases and update simulation models for assimilating and analyzing data on and/or guiding management of wildlife disease problems identified through surveillance and surveys. Provide assistance in investigating and managing wildlife disease outbreaks in Colorado. Task 5 (Workpackage 7610) 1. 2. Provide ongoing consulting assistance to CDOW on game harvest surveys, terrestrial inventory systems, and game population modeling. Update DEAMAN software to run as a Windows 95 program and incorporate the graphical display of spatial data such as numbers harvested by GMU and DAU and categorical estimates of deer and elk hunter satisfaction. �302 3. 4. 5. 6. Conduct workshops as necessary to assist region personnel in the use of DEAMAN and population modeling procedures, and in the use of statistical techniques for monitoring spatial distribution and statewide abundance of wildlife populations. Maintain data analysis programs currently in use for small game harvest surveys (grouse, squirrel, waterfowl, and small game). Develop programs to input turkey (fall and spring, limited and unlimited) and furbearer (30 day permit holders) harvest data. Output data by species as estimated harvest, number of hunters, and days in the field by county, and CDOW Region. Develop a sampling protocol for estimating harvest of small game species utilizing the Harvest Inventory Program (lllP) database. Develop the necessary programming to incorporate HIP data into CDOW's small game harvest survey process. Write the code necessary to access the HIP export data, to enter harvest information, to analyze the harvest information, and report the estimated harvest by species. Harvest estimates should include number harvested, number of hunters, and days in the field by county and CDOW Region. Work with contract companies as necessary to train personnel on the system. Task 6 CWorkpackage 7610) 1. 2. 3. 4. 5. 6. Prepare PACE plans for all Mammals Program staff members. Prepare performance evaluations for all Mammals Program staffmembers. Coordinate scientific input from Mammals Program staff members into the process for implementing Amendment 14 and Senate Bill 52. Process all encumbering documents for Mammals Program staff members and maintain records of those encumbrances in the COFRS program. Update the Administrative Standard Operating Procedures Manual and distribute copies to all Mammals Program staff members. Coordinate the preparation of Federal Aid Program Narratives, Segment Narratives, Job Progress Reports, and Job Final Reports and submit required numbers of copies to the U. S. Fish and Wildlife Service. ACCOMPLISHMENTS Task 1 (Workpackage 7210) Publications Acquired in the Research Center Librarv Ballard, W. B. and A. R Rodgers, eds. 1998. Alces: a journal devoted to the biology and management of moose. Thunder Bay, Ont., Canada: Lakehead University. Alces 34(1). 259pp. Bengeyfield, W. 1990. Evaluation of an electrical field to divert coho salmon smolts from the Penstock intake at Puntledge Generating Station. White Rock, B.C., Canada: Global Fisheries Consultants Ltd. Prepared for B.C. Hydro: Environmental Resources, Vancouver, B.C. 54 leaves. Burnham, Bill, ed. 1997. The Peregrine Fund : World Center for Birds of Prey : 1996 annual report: focusing on birds to conserve nature. [Boise, ID : World Center for Birds of Prey] 20pp.. Buskirk, S. W., A. S. Harestad, M. G. Raphael, and R A. Powell, eds. 1994. Martens, sables, and fishers : biology and conservation. Ithaca, NY : Cornell University Press. 484pp. Cagney,1. 1993. Greenline riparian-wetland monitoring. Denver, CO: U.S. Bureau of Land Management. Riparian Area Management. Technical reference; 1737-8. 45pp. Carey, A. B. and C. Elliott, eds. 1994. Washington forest landscape management project - progress report. Olympia, WA: Washington State Dept. of Natural Resources. Washington forest landscape management project; report no. 1. 174pp. �303 Carey, A. B., C. Elliott, B. R Lippke, J. Sessions, C. J. Chambers, C. D. Oliver, J. F. Franklin, and M. G. Raphael. 1996. Washington forest landscape management project - a pragmatic, ecological approach to small-landscape management. Olympia, WA : Washington State Dept. of Natural Resources. Washington forest landscape management project; report no. 2. 99pp. Carlson, T. J. and A. N. Popper. 1997. Using sound to modify fish behavior at power-production and water-control facilities: a workshop: Dec. 12-13,1995: Portland State University: Portland, OR: phase II : fmal report. Portland, OR : Bonneville Power Administration. 348pp. Caughley, G. and A. Gunn. 1996. Conservation biology in theory and practice. Cambridge, MA : Blackwell Science, Inc. 459pp. Clemmer, P. 1994. The use of aerial photography to manage riparian-wetland areas. Denver, CO : U.S. Bureau of Land Management. Riparian Area Management. Technical reference; 1737-10. 54pp. Crichton, V., J. M. Peek and A. R Rodgers, eds. 1998. Alces: ajournal devoted to the biology and management of moose. Thunder Bay, Ont., Canada: Lakehead University. Alces 34(2). 261-509pp. Cvancara, V. 1998. Current references in fish research: volume 23,1998. Chippewa Falls, WI: CRFR 156pp. Douglas, C. L., T. D. Bunch, P. R Krausman, D. M. Leslie, Jr., J. J. Spillett, and J. Blaisdell, eds. [1983]. Desert Bighorn Council: 1982 transactions: a compilation of papers presented at the 26th annual meeting, April 7-9, 1982, Borrego Springs, Calif. Las Vegas, NV: Univer. of Nevada. 119pp. Douglas, C. L., P. R Krausman, and D. M. Leslie, Jr., eds. [1984]. Desert Bighorn Council: 1983 transactions: a compilation of papers presented at the 27th annual meeting, April 6-.8, 1983, Silver City, New Mexico. Las Vegas, NV: Univer. of Nevada. 46pp. Douglas, C. L., P. R Krausman, and D. M. Leslie, Jr., eds. [1985]. Desert Bighorn Council: 1984 transactions: a compilation of papers presented at the 28th annual meeting, April 5-7, 1984, Bullhead City, Arizona. Las Vegas, NV : Univer. of Nevada. 59pp. Douglas, C. L., D. M. Leslie, Jr., and T. O'Farrell, eds. [1979]. Desert Bighorn Council: 1978 transactions: a compilation of papers presented at the 22nd annual meeting, April 5-7, 1978, at Kingman, Arizona. Death Valley, CA: Desert Bighorn Council. 51pp. Douglas, C. L., D. M. Leslie, Jr., C. D. Simpson, J. J. Spillett, R Valdez, and L. D. White, eds. [1980]. Desert Bighorn Council: 1979 transactions: a compilation of papers presented at the 23rd annual meeting, April 4-6, 1979, at Boulder City, Nevada. Death Valley, CA: Desert Bighorn Council. 51pp. Douglas, C. L., L. D. White, J. J. Spillett, C. D. Simpson, R Valdez, and D. M. Leslie, Jr., eds. [1981]. Desert Bighorn Council: 1980 transactions: a compilation of papers presented at the 24th annual meeting, April 9-11, 1980, St. George, Utah. Death Valley, CA: Desert Bighorn Council. 51pp. Duda, M. D., S. J. Bissell, and K. C. Young, eds. 1998. Wildlife and the American mind : public opinion on and attitudes toward fish and wildlife management. Harrisonburg, VA : Responsive Management. 804pp. Duncan, J. R, D. H. Johnson, and T. H. Nicholls, eds. 1997. Biology and conservation of owls of the northern hemisphere: second international symposium: Feb. 5 - 9,1967: Winnipeg, Manitoba, Canada. St. Paul, MN : U.S. Forest Service. North Central Research Station. Gen. Tech. Rep. NCGTR-190. 635pp. Evans, H. E. and M. A. Evans. 1991. Cache La Poudre : the natural history of a Rocky Mountain river. Niwot, CO : University Press of Colorado. 260pp. Evans, H. E. 1997. The natural history of the Long Expedition to the Rocky Mountains: 1819 - 1820 . . New York: Oxford University Press. 268pp. Fagan, D. 1998. Canyon country wildflowers: a field guide to common wildflowers, shrubs, and frees. Helena, MT: Falcon Publishing Co. 147pp. Folzenlogen, R 1995. Birding the Front Range: a guide to seasonal highlights. Littleton, CO : Willow Press. 159pp. Garrett, G. P., ed. 1997. Proceedings of the Desert Fishes Council: Vol. XXVIII: 1996 annual symposium: 6-9 Nov. : La Paz, Baja California Sur, Mexico. Bishop, CA : Desert Fishes Council. I08pp. �304 Garrett, G. P. and AM. Sudyka, eds. 1996. Proceedings of the Desert Fishes Council : Vol. XXVII : 1995 annual symposium: 16-19 Nov. : Reno, Nev., U.S.A Bishop, CA: Desert Fishes Council. 129pp. Gebhardt, K, S. Leonard, G. Staidl, and D. Prichard. 1990. Riparian and wetland classification review. Denver, CO: U.S. Bureau of Land Management. Riparian Area Management. Technical reference; 1737-5. 56pp. Gordon, R. E., ed. 1998. 1998 conservation directory. Vienna, VA: National Wildlife Federation. 34th ed. 483pp. Greenlee, J. M., ed. 1997. Proceedings: first conference on fire effects on rare and endangered species and habitats: Coeur d'Alene, Idaho: November, 1995. Fairfield, WA : International Assoc. of Wildland Fire. 343pp. Harfenist, A, T. Power, K. L. Clark and D. B. Peakall. 1989. A review and evaluation of the amphibian toxicological literature. Ottawa, Canada: Canadian Wildlife Service. Technical report series; no. 61. 222pp. Hendrickson D. A, ed. 1995. Proceedings of the Desert Fishes Council : Vol. XXVI: 1994 annual symposium: 17-20 Nov. : Death Valley National Park: Furnace Creek, California, U.S.A Bishop, CA : Desert Fishes Council. 149pp. Hendrickson D. A and G. P. Garrett, eds. 1999. Proceedings of the Desert Fishes Council : Vol. XXIX: 1997 annual symposium: 20-23 Nov. : Death Valley National Park, California. Bishop, CA : Desert Fishes Council. 78pp. Hindley, E. 1996. Observing physical and biological change through historical photographs. Denver, CO : U.S. Bureau of Land Management. Riparian Area Management. Technical reference; 1737-13. 32pp. Hoffinan, G. L. 1999. Parasites of North American freshwater ftshes. Ithaca, NY: Comstock Publishing Associates. 2nd edition. 539pp. Johnson, S. K 1998. Acoustic target strength estimates of opossum shrimp (Mysis relicta) using a splitbeam echo sounder, M.S. Thesis, Colorado State University, Fort Collins. 109pp. Keating, J. B., Jr. 1993. The goo-positioning selection guide for resource management: a reference to select systems and techniques for obtaining and mapping natural resources or cultural data. Cheyenne, WY: U.S. Bureau of Land Management. Technical note; 389. 64pp. Kingery, Hugh E., ed. Colorado breeding bird atlas. [Denver, CO] : Colorado Bird Atlas Partnership, Colo. Div.ofWildlife. 636pp. Kinch, G. 1989. Grazing management in riparian areas. Denver, CO: U.S. Bureau of Land Management. Riparian Area Management. Technical reference; 1737-4. 44 leaves. Lee, R. M., ed. [1998]. Desert Bighorn Council : 1997 transactions: a compilation of papers presented and submitted at the 41st annual meeting: Grand Junction, Colorado: April 9-11, 1992. [S.1.]: Desert Bighorn Council. 92pp. Leonard, S., G. Kinch, V. Elsbernd, M. Borman, and S. Swanson. 1997. Grazing management for riparian-wetland areas. Denver, CO: U.S. Bureau of Land Management. Riparian Area Management. Technical reference; 1737-14. 63pp. Leonard, S., G. Staidl, J. Fogg, K Gebhardt, W. Hagenbuck, and D. Prichard. 1992. Procedures for ecological site inventory - with special reference to riparian-wetland sites. Denver, CO: U.S. Bureau of Land Management. Riparian Area Management. Technical reference; 1737-7. 135pp. Lindzey, F. G., W. G. Hepworth, T. A Mattson, and A F. Reese. 1997. Potential for competitive interactions between mule .deer and elk in the western United States and Canada: a review. Laramie, WY : Wyo. COOperativeFisheries & Wildlife Research Unit. 82pp. Lippke, B. R., J. Sessions, and A B. Carey. 1996. Economic analysis offorest landscape management alternatives. Seattle, WA: University of Washington. CINTRAFOR special paper; 21. 157pp. Lyman, R. L. 1998. White goats, white lies: the misuse of science in Olympic National Park. Salt Lake City, UT : Univ. of Utah Press. 277pp. McDonald, G., J. D. Fitzsimons, and D. C. Honeyfteld, eds. 1998. Early life stage mortality syndrome in ftshes of the Great Lakes and Baltic Sea. Bethesda, MD : American Fisheries Society. American Fisheries Society Symposium; 21. 177pp. �305 Maehr, D. S. 1997. The Florida panther: life and death of a vanishing carnivore. Washington, DC: Island Press. 259pp. Myers, L. H. 1989. Inventory and monitoring riparian areas. Denver, CO : U.S. Bureau of Land Management. Riparian Area Management. Technical reference; 1737-3. 79pp. National Research Council (U.S.). Committee on Management of Wolf and Bear Populations in Alaska. 1997. Wolves, bears, and their prey in Alaska: biological and social challenges in wildlife management. Washington, D.C. : National Academy Press. 207pp. Oedekoven, O. O. 1985. Columbian sharp-tailed grouse population distribution and habitat use in south central Wyoming. M.S. Thesis, University of Wyoming, Laramie. 58pp. Pillsbury, N. H., J. Verner, and W. D. Tietje, technical coordinators. 1997. Proceedings of a Symposium on oak woodlands: ecology, management, and urban interface issues: 19 - 22 March 1966 : San Luis Obispo, CA Albany, CA: US. Forest Service. Pacific Southwest Research Station. Gen. Tech. Rep. PSW-GTR-160. 738pp. Pister, E. P., ed. 1970. The rare and endangered fishes of the Death Valley system - a summary of the proceedings of a symposium relating to their protection and preservation: Vol. I. : held at National Park Service Headquarters: Furnace Creek, Calif. : Nov. 18 & 19, 1969. Bishop, CA: Desert Fishes Council. 18 leaves +. Pister, E. P., ed. 1971. The rare and endangered fishes of the Death Valley system: a summary of the proceedings of the second annual symposium relating to their protection and preservation: VoL II : held at Death Valley National Monument Headquarters: Furnace Creek, Calif. : Nov. 17 & 18, 1970. Bishop, CA : Desert Fishes Council. 26 leaves +. Pister, E. P., ed. 1987. Proceedings of the Desert Fishes Council: Vols. XVI - XVIII: the sixteentheighteenth annual symposia. Bishop, CA : Desert Fishes Council. 253pp. Prichard, D. 1998 (rev. ed.). Process for assessing proper functioning condition. Denver, CO: US. Bureau of Land Management. Riparian Area Management. Technical reference; 1737-9. 51pp. Prichard, D. 1994. Process for assessing proper functioning condition for lentic riparian-wetland areas. Denver, CO: US. Bureau of Land Management. Riparian Area Management. Technical reference; 1737-1l. 37pp. Prichard, D., P. Clemmer, M. Gorges, G. Meyer, and K. Shumac. 1996. Using aerial photographs to assess proper functioning condition of riparian-wetland areas. Denver, CO: U.S. Bureau of Land Management. Riparian Area Management. Technical reference; 1737-12. 41pp. Redpath, S. M. and S. J. Thirgood. 1997. Birds of prey and red grouse. London: Stationery Store. .148pp. Biddle, P., ed. [1992]. Proceedings of the fifteenth biennial pronghorn antelope workshop: Rock Springs, Wyoming: June 8-11,1992. [Laramie, WY: Wyoming Game & Fish Dept.] 151pp. Rowland, M. M., L. D. Bryant, B. K. Johnson, ]. H. Noyes, M. J. Wisdom, and], W. Thomas. 1997. The Starkey Project: history, facilities, and data collection methods for ungulate research. Portland, OR : US. Dept. of Ag., Forest Service, Pacific Northwest Research Station. General technical report; PNW-GTR-396. 62pp. Schneider, B. 1996. Bear aware: hiking and camping in bear country. Helena, MT : Falcon Publishing Co. 127pp. Siegel, A F. and C. J. Morgan. 1996. Statistics and data analysis: an introduction. New York: John Wiley & Sons. 2nd ed. 635pp. Smith, B. and D. Prichard. 1992. Management techniques in riparian areas. Denver, CO: U.S. Bureau of Land Management. Riparian Area.Management, Technical reference; 1737-6. 44pp. Standage Market Research, Inc. 1998.+Summary of project report: trends and issues in Colorado muzzleloader hunting: a profile of 1997 Colorado limited muzzleloader license applicants. Denver, CO : Standage Market Research, Inc. 11 leaves. Standage Market Research, Inc. 1998. Trends and issues in Colorado muzzleloader hunting: a profile of 1997 Colorado limited muzzleloader license applicants. Denver, CO : Standage Market Research, Inc. 56pp. Steele, M. A,]. F. Merritt, and D. A Zegers, eds. 1998. Ecology and evolutionary biology of tree squirrels / Proceedings of the International Colloquium on the Ecology of Tree Squirrels, Powdermill Biological Station, Carnegie Museum of Natural History, 22-28 April 1994. Martinsville, VA : �306 Virginia Museum of Natural History. Virginia Museum of Natural History. Special publication; no. 6. 310pp. Tarnow, K., P. Watt, and D. Silverberg. 1996. Collaborative approaches to decision making and conflict resolution for natural resource and land use issues: a handbook for land use planners, resource managers, and resource management councils. Salem, OR : Oregon Dept. of Land Conser. & Develop. 116 leaves. Theberge, J. B. and M. T. Theberge. 1998. Wolf country: eleven years tracking the Algonquin wolves. Toronto, Ont. : McClelland & Stewart, Inc. 306pp. Thompson, W. L., G. C. White, and C. Gowan. 1998. Monitoring vertebrate populations. New York: Academic Press. 365pp. Torres, S. 1997. Mountain lion alert: safety tips for yourself, your children, your pets, and your livestock in lion country. Helena, MT : Falcon Publishing Co. 123pp. Webster, D. B. and C. M. Britton, eds. 1998. Research highlights: noxious brush and weed control: range, wildlife, and fisheries management: 1998. Lubbock, TX: Texas Tech Univ. 45pp. The Wildlife Society. Northeast Section. [1998]. Northeast wildlife. S.l: Northeast Section - The Wildlife Society. Transactions of the Northeast Section: The Wildlife Society: Vol. 53.: 1996. 71pp. The Wildlife Society. Western Section. [1999]. Transactions of the Western Section of the Wildlife Society: 1998 Vol. 34. S.l: Western Section of The Wildlife Society. 68pp. White, G. C. and R A. Garrott. 1990. Analysis of wildlife radio-tracking data. New York, NY : Academic Press. 383p. Wisdom, M. J., J. G. Cook, M. M. Rowland, and J. H. Noyes. 1993. Protocols for care and handling of deer and elk at the Starkey Experimental Forest and Range. Portland, OR: U.S. Dept. of Ag., Forest Service, Pacific Northwest Research Station. General technical report; PNW-GTR-311. 49pp. Theses and Books Obtained on Interlibrary Loan Literature Searches and Information Delivered Baldo, B. A., E. R Tovey, and St. Leonards, eds. 1989. Protein blotting: methodology, research and diagnostic applications. New York: Karger. 168pp. Bjerrum, O. J. and N. H. H. Heegaard, eds. 1988. CRC Handbook ofimmunoblotting of proteins. Boca Raton, FL: CRC Press, Inc. Experimental and clinical applications, vol. II. 214pp. Breitenmoser U. and C. Breitenmoser-Wursten. 1990. Status, conservation needs and reintroduction of the lynx (Lynx lynx) in Europe. Strasbourg, France: Council of Europe. Nature and environment series; no. 45. 47pp. Brodkorb, P. 1964. Catalogue offossil birds; pt. 2 (Anseriformes through Galliformes) Gainesville, FL: Univ.ofFlorida. Bulletin of the Florida State Museum. Biological sciences; v. 8. 195-335pp. DeBano, L. F., D. G. Neary, and P. F. Ffolliott. 1998. Fire's effects on ecosystems. New York: John Wiley & Sons, Inc. 333pp. Fremont, John C. 1988 (reprint). Exploring expedition to the Rocky Mountains. Washington, D.C. : Smithsonian Institution Press. Exploring the American west. 319pp. Originally issued 1845 as: Report of the exploring expedition to the Rocky Mountains in the year 1842, and to Oregon and North California in the years 1843-44 by Brevet Captain J. C. Fremont, of the Topographical Engineers under the orders of Col. J. J. Abert, Chief of the Topographical Bureau. Gal, Y. L. and H. O. Halvorson, eds. 1998. New developments in marine biotechnology. New York: Plenum Press. Proceedings of the 4th International Marine Biotechnology Conference.held Sept. 2229,1997, in Sorrento, Paestum, Otranto, and Pugnochiuso, Italy. 343pp. Galatowitsch, S. M., and A. G. van der Valko 1994. Restoring prairie wetlands: an ecological approach. Ames, IA : Iowa State University Press. 246pp. Gardner, S. C. 1997. Movements, survival, productivity, and test of a habitat suitability index model for reintroduced Columbian sharp-tailed. M.S. Thesis, University ofIdaho, Moscow. 91pp. eireenlee, J. M., ed. 1997. Proceedings: first conference on rare and endangered species and habitats : Coeur d'Alene, Idaho: Nov. 1995. Fairfield, WA : International Association of Wildland Fire. 343pp. �307 "'" Hammond, E. L. 1990. Mark-recapture estimates of population parameters for selected species of small mammals. M.S. Thesis, Oregon State University, Corvallis. 116pp. Holdich, D. M. and R S. Lowery, eds. 1988. Freshwater crayfish: biology, management and exploitation. Portland, OR : Timber Press. 498pp. Holman, J. A. 1964 Osteology of gallinaceous birds. Quart. J. Florida Acad. Sci. 27(3):230-252. Horton, H. H., Jr. 1989. An illustrated checklist of the American crayfishes (Decapoda: Astacidae, Cambaridae, and Parastacidae). Washington, D.C. : Smithsonian Institution Press. Contributions to zoology; 480. 236pp. Hudson, G. E., R A. Parker, J. V. Berge, and P. J. Lanzillotti. 1966. A numerical analysis of the modifications of the appendicular muscles in various genera of gallinaceous birds. Am. Midland Nat. 76(1):1-73. Kansas Ornithological Society. 1979-1983. Bulletin. Manhattan, KS : Kansas Ornithological Society. vols. 30-34. Johnson, C. L. 1997. Distribution, habitat, and ecology of the Mexican spotted owl in Colorado. M.A. Thesis, University of Northern Colorado, Greeley. 78pp. Marshall, A. J., ed. 1960. Biology and comparative physiology of birds, vol. I. New York: Academic Press. 518pp. Martinez, A. M. 1983. Identification of brook, brown, rainbow, and cutthroat trout larvae. M.S. Thesis, Colorado State University, Fort Collins. 149pp. McAninch, 1:'B., ed. 1995. Urban deer: a manageable resource? Proceedings of the 1993 Symposium of the North Central Section, The Wildlife Society. S.l.: The Wildlife Society. North Central Section. 175pp.. McWherter, G. R 1991. Estimating abundance of cottontail rabbits using live trapping and visual surveys. M.S. Thesis, University of Tennessee, Knoxville. TWRA technical report; no. 91-4. 95pp. Meyburg, B.-U and RD. Chancellor, eds. 1994. Raptorconservation today. East Sussex, UK. : World Working Group on Birds of Prey and Owls. Proceedings of the NWorld Conference on Birds of Prey and Owls: Berlin, Germany, 10-17 May 1992. 799pp. Monte, M. A. 1995. Changing models of administrative decision-making: public participation and public land planning. M.S. Thesis, University of Arizona, Tucson. U.p. National Rifle Association of America. 1998. The hunter's guide. Washington, D.C. : National Rifle Assoc. of America. ???PP. Nava, J. A. 1970. The reproductive biology of the Alaska lynx (Lynx canadensis). M.S. Thesis, University of Alaska, Fairbanks. 140pp. Northrup, R D. 1991. Sharp-tailed grouse habitat use during fall and winter on the Charles M. Russell National Wildlife Refuge, Montana. M.S. Thesis, Montana State University, Bozeman. 54pp. Noss, R F. and A. Y. Cooperrider. 1994. Saving nature's legacy: protecting and restoring biodiversity. Washington, D.C. : Island Press. 416pp. Oedekoven, O. O. 1985. Columbian sharp-tailed grouse population distribution and habitat use in south central Wyoming. M.S. Thesis, University of Wyoming, Laramie. 58pp. Parker, T. L. 1970. On the ecology of the sharp-tailed grouse in southeastern Idaho. M.S. Thesis, Idaho State University, Pocatello. 14Opp. Potts, D. F., ed. 1998. Proceedings of AWRA Specialty Conference, Rangeland Management and Water Resources. Herndon, VA: American Water Resources Association. TPS-98-1. 474pp. Reed, J. M., N. Warnock, and L. W. Oring, eds. 1997. Conservation and management of shorebirds in the , western Great Basin-of North America. Norfolk, UK. : The Wader Study Group. International, wader studies; 9. 81pp. Ridgway, R and H. Friedmann. 1946. The birds of North and Middle America: Part X. Washington, D.C. : US. Gov. Printing Office. US. National Museum bulletin; 50. 484pp. Ruggles, C. P. 1984. Fish diversionary techniques for hydroelectric turbine intakes. Montreal: Canadian Electrical Assoc. Canadian Electrical Association. Research report; 149 G 339. var. paging. Schneider, J. W. 1994. Winter feeding and nutritional ecology of Columbian sharp-tailed grouse in southeastern Idaho. M.S. Thesis, University ofIdaho, Moscow. 118 leaves. Seton, E. T. 1953. Lives of game animals: an account of those land animals in America, north of the Mexican border, which are considered "game," either because they have held the attention of s,•••••• �308 sportsmen, or received the protection of law. v. 3, pt. 2. Boston, MA: Charles T. Branford, Co. 413780pp. Short, L. L., Jr. 1967. A review of the genera of grouse (Aves, Tetraoninae). New York: American Museum of Natural History. American Museum novitates; no. 2289. 39pp. Skovlin, J. M. 1991. Fifty years of research progress: a historical document on the Starkey Experimental Forest and Range. Portland, OR : U.S. Forest Service, Pacific Northwest Research Station. General technical report; PNW-GTR-266. 58pp. Sloss, M. W., R L. Kemp, and A. M. Zajac. 1994. Veterinary clinical parasitology. Ames, IA : Iowa State University Press. 6th edition. 198pp. Sureda, M. 1996. Small mammal abundance within Mexican spotted owl home ranges in the Manti-LaSal National Forest, San Juan County, UT. M.S. Thesis, Univ. of Arizona, Tucson. 117p. (Microfiche) Tarnow, K., P. Watt, and D. Silverberg. 1996. Collaborative approaches to decision making and conflict resolution for natural resource and land use issues: a handbook for land use planners, resource managers and resource management councils. [Salem, OR] : Oregon Dept. of Land Conserv. & Develop. 116 leaves. Ulliman, M. J. 1995. Winter habitat ecology of Columbian sharp-tailed grouse in southeastern Idaho. M.S. Thesis, University ofIdaho, Moscow. 119pp. Varley, N. C. L. 1996. Ecology of mountain goats in the Absaroka Range, south-central Montana. M.S. Thesis, Montana State University, Bozeman. 91pp. Vosburgh, T. C. 1996. Impacts of recreational shooting on prairie dog colonies. M.S. Thesis, Montana State University, Bozeman. 50pp. Wetmore, A. 1960. A classification for the birds of the world. Washington, D.C. : Smithsonian Institution. Smithsonian miscellaneous collections; vol. 139(11):1-37. Wisdom, M. J., J. G. Cook, M. M. Rowland, and J. H. Noyes. 1993. Protocols for care and handling of deer and elk at the Starkey Experimental Forest and Range. Portland, OR : U.S. Forest Service, Pacific Northwest Research Station. General technical report; PNW -GTR-311. 49pp. The Research Center Library staff also located and delivered approximately 2,292 individual items or free documents on request for Colorado Div. of Wildlife employees and cooperators during this segment. Maintain Electronic and Card Catalogues of all Research Library Holdings 841 is the total number of items cataloged during this period of time. This includes not only new acquisitions, but also older materials from the library collection being entered into the electronic catalog for the first time. Among the new acquisitions are Federal Aid: Job Progress Reports and manuscripts written by Colorado Div. of Wildlife Researchers and other employees. 1375 is the total number of computer scanned items added to the electronic circulation system during this period. This includes not only the above mentioned newly cataloged items, but also newly acquired serials, volumes, and items being assigned scanning numbers for the electronic circulation system for the first time. CD OW Manuscripts Published July. 1998 - June. 1999 Job Progress Reports; Federal Aid. All studies. Andelt, W. F., R L. Phillips, R H. Schmidt, and R B. Gill. 1999. Trapping forbearers: an overview of the biological and social issues surrounding a public controversy. Wildl. Soc. Bull. 27(1):114-125 Anderson, D. R, K. P. Burnham, A. B. Franklin, R J. Gutierrez, E. D. Forsman, R G. Anthony, G. C. White, and T. M. Shenk. 1999. A protocol for conflict resolution in analyzing empirical data related to natural resource controversies. Wildl. Soc. Bull. (in press) Bissell, S. J. 1998. The cultural context of illegal hunters in England and America: a historical comparison. Human Dim. of Wildlife 3(4):49-52. �309 Duda, M. D., K. C. Young, and S. 1. Bissell. 1998. Partnerships in conservation: using human dimensions to strengthen relationships between fish and wildlife agencies and their constituents. pages 268-277 in Transactions of the sixty-third North American Wildlife Conference, ed. K. G. Wadsworth. Washington, D.C. : Wildlife Management Institute. Franklin, A. B. and T. M. Shenk. 1999. The importance of parameter and variance component estimation in population viability analysis. page 29 in Population viability analysis: assessing models for recovering endangered species: program & abstracts: March 15-16, 1999: San Diego, California. Sponsored by University of California - Berkeley and The Wildlife Society: Western Section. (abstract) Freddy, D. 1., Bowden, and G. C. White. 1999. Estimating elk densities in Colorado: progress with perplexities. Western States and Provinces Deer and Elk Workshop. Salt Lake City, Utah. U.S.A. March 1999. (abstract in print) Gammonley, J. H. and L. H. Fredrickson. 1998. Breeding duck populations and productivity on montane wetlands in Arizona. Southwest. Nat. 43(2):219-227 Green, A. L., N. M. DuTeau, M. W. Miller, J. Triantis, and M. D. Salman. 1999. Polymerase chain . reaction techniques for differentiating cytotoxic and noncytotoxic Pasteurella trehalosi from Rocky Mountain bighorn sheep. Am. J. Vet. Res. 60(5):583-588. Johnson, K. H. and C. E. Braun. 1999. Viability and conservation of an exploited sage grouse population. Conser. bioI. 13(1):77-84 Jones, M. S. arid K. B. Rogers. 1998. Palmetto bass movements and habitat use in a fluctuating Colorado irrigation reservoir. North Am. J Fish. Manage. 18(3):640-648. Manfredo, M.J., H. C. Zion, S. Sikorowski, and J. Jones. 1998. Public acceptance of mountain lion management: a case study of Denver, Colorado, and nearby foothills areas. WildI. Soc. Bull. 26(4):964-970. McCarty, C. W. and M. W. Miller. 1998. Modeling the population dynamics of bighorn sheep: a synthesis of literature. Fort Collins, CO: Colo. Div. of Wildlife. Special report; no. 73. 35pp. McNeil, H. 1., M. W. Miller, 1. A. Conlon, I. K. Barker, and P. E. Shewen. 2000. Effects of delivery method on serological responses of bighorn sheep to a multivalent Pasteurella haemolytta supernatant vaccine. 1. Wildl. Dis. (in press) Merrion, D., R Standage, M. Lloyd, and L. Sikorowski. 1999. Trends and issues in Colorado muzzleloader hunting. Hum. Dim. Wildl. 4(1):70-71. Mote, K. D., R D. Applegate, J. A. Bailey, K. M. Giesen, R Horton, and J. L. Sheppard. eds. Assessment and conservation strategy for the lesser prairie-chicken (Tympanuchus pallidicinctus). Emporia, KS : Kansas Dept. of Wildlife and Parks. 51 leaves. Nehring, R B., K. G. Thompson, and S. Hebein. 1998. Impacts of whirling disease on wild trout populations in Colorado. pages 82-94 in Transactions of the sixty-third North American Wildlife Conference, ed. K. G. Wadsworth. Washington, D.C. : Wildlife Management Institute. O'Rourke, K. I., T. E. Besser, N. W. Miller, T. F. Cline, T. R Spraker, A. L. Jenny, M. A. Wild, G. L. Zebarth, and E. S. Williams. 1999. PrP genotypes of captive and fee-ranging Rocky Mountain elk (Cervus elaphus nelsoni) with chronic wasting disease. 1. Gen Virol. (in press) Seidel, J. W., B. Andree, S. Berlinger, K. Buell, A. E. Byrne, R B. Gill, D. W. Kenvin, and D. F. Reed. 1998. Draft strategy for the conservation and reestablishment of lynx and wolverine in the southern Rocky Mountains. [Fort Collins, CO: Colo. Div. of Wildlife] 116pp. Shenk, T. M. and G. Byrne. 1999. Colorado lynx recovery project: background and post-release monitoring of lynxze-introduced to the southern Rocky Mountains of southwestern Colorado. Reintro. News (in press) Shenk, T. M. and M. M. Sivert. 1999. Development of a conservation strategy for Preble's meadow jumping mouse: what can you do with little information? page 95 in 2nd International Wildlife Management Congress: program and abstracts: 28 June - 2 July 1999 : Godollo, Hungary. Sponsored by the Godollo Agricultural University and The Wildlife Society. (abstract) Shenk, T. M., G. C. White, and K. P. Burnham. 1998. Sampling-variance effects on detecting density dependence from temporal trends in natural populations. Ecolog. Mono. 68(3):445-463. �310 Sigurdson, C. 1., E. S. Williams, M. W. Miller, T. R Spraker, K. I. O'Rourke, and E. A. Hoover. 1999. Oral transmission and early lymphoid trophism of chronic wasting disease PrPres in mule deer fawns. J. Gen. Virol. (in press) Sikorowski, L., 1. Smeltzer, and M. J. Manfredo. 1998. Wildlife management in Colorado: the past, the present and predictions of the future. pages 257-267 in Transactions of the sixty-third North American Wildlife Conference, ed. K. G. Wadsworth. Washington, D.C. : Wildlife Management Institute. Swift-Miller, S. M., B. M. Johnson, R T. Muth, and D. Langlois. 1999. Distribution, abundance, and habitat use of Rio Grande sucker (Catostomus plebeius) in Hot Creek, Colorado. Southwest. Nat. 44(1): 42-48. Ward, A. C. S., D. L. Hunter, K. M. Rudolph, W. J. Del.ong, J. M. Bulgin, L. M. Cowan, H. J. McNeil, and M. W. Miller. 1999. Immunologic responses of domestic and bighorn sheep to a multivalent Pasteurella haemolytica vaccine. J. Will. Dis. 35(2):285-296. White, G. C. and T. M. Shenk. 1999. Population estimation with radio-marked animals. at The Wildlife Society 6th annual conference: Austin, Texas: Sept. 7-11,1999. (in press) Manuscripts in Review FY 1998-99 Cassirer, E. F., K. M. Rudolph, P. Fowler, V. L. Coggins, D. L. Hunter, and M. W. Miller. 1999. Evaluation of ewe vaccination as a tool for increasing bighorn lamb survival following pasteurellosis epidemics. J. Wildl. Dis. (in review) Kreeger, T. J., M. W. Miller, M. A. Wild, P. H. Elzer, and S. C. Olsen. 1999. Safety and efficacy of Brucella abortus strain RB51 vaccine in captive pregnant elk. J. Wildl. Dis. (in review) McCarty, C. W., K. P. Burnham, and M. W. Miller. 1999. A comparison of several theories of epidemic dynamics using AlC. Biometrics (in review) Miller, M. W., E. S. Williams, C. W. McCarty, T. R Spraker, T. J. Kreeger, and E. T. Thome. 1999. Epidemiology of chronic wasting disease in free-ranging cervids. J. Wildl. Dis. (in review) Miller, M. W., J. Vayhinger, S. Roush, T. Verry, A. Torres, and V. Jurgens. 1999. Drug treatment for lungworm in bighorn sheep: reevaluation of a 20-year-old management prescription. J. Wildl. Manage. (in review) Pojar, T. M., D. C. Bowden, and J. D. Madison. 1999. Pronghorn density estimates: comparison of fixedwing line transect and helicopter quadrat surveys. Wildl. Soc. Bull. (in review) Reed, D. F. 1999. A conceptual interference competition model for introduced mountain goats. J. Wildl. Manage. (in review) Reed, D. F. and J. Kindler. 1999. Snowshoe hare density/distribution estimates and potential release sites for reintroducing lynx in Colorado. Fort Collins, CO: Colo. Div. ofWildl. Division report; no. (in review) Spraker, T. R, R B. Zink, B. A. Cummings, K. I. O'Rourke, and M. W. Miller. 1999. Neuroanatomical distribution and patterns of histological lesions and immunohistochemical staining ofprion protein in mule deer (Odocotleus hemionus) with preclinical chronic wasting disease. Vet. Path. (in review) Spraker, T. R, R B. Zink, B. A. Cummings, M. A. Wild, K. I. O'rourke, and M. W. Miller. 1999. Neuroanatomical distribution and description of histological lesions and immunohistochemical staining of prion protein in mule deer (Odocoileus hemionus} and Rocky Mountain elk (Cervus elaphus nelsoni) with advanced spongiform encephalopathy. Vet. Path. (in review) Monitor and Evaluate Customer Satisfaction with Library Services During this fiscal year a 'Suggestion Box' was set up in the library for patrons to make comments. No suggestions were received. �311 Task 2 <Workpackage 7120) Progress made on each Objective during this Segment is reported below. 1. 2. 3. 4. Assisted 4 Wildlife Researchers in developing publication-ready manuscripts. Assisted Wildlife employees in creating graphics and visual materials for 46 presentations (slides, overheads, handouts, posters). Published 1998 Federal Aid Abstracts and distributed them to approximately 300 Division personnel and 150 outside agencies and libraries. Published 2 Special Reports, Nos. 73 and 74. This process involved editing; selecting and creating tables, photos, and graphics; preparing camera-ready copy; having books printed at local vendor; and distributing to Division personnel and outside libraries, agencies, etc. These reports are as follows: Modeling the Population Dynamics of Bighorn Sheep: A Synthesis of Literature, No. 73 Columbian Sharp-tailed Grouse Harvest and Wing Analysis: Implications for Management, 5. No. 74 Assembled, published, and distributed the FY 97-98 Wildlife Research Reports. Task 3 <Workpackage 8160) METIIODS AND MATERIALS Operation of Foothills Wildlife Research Facility (FWRF) during FY1999 was in support of research on chronic wasting disease in deer and elk (and potential for natural transmission of CWO to domestic cattle and pronghorn), deer and elk contraception, pneumonia immunization of bighorn sheep, and prairie dog research for black-footed ferret recovery. Animal care and facility maintenance standard operating procedures (SOP's) were followed for routine animal husbandry and facility procedures. Given this general guidance, and the direction required to meet terrestrial research needs, we performed the following tasks: Animal Maintenance General: Again this year, routine feeding and caretaking of research animals, including health observations, training, weighing, and clean-up, was performed primarily by well trained work-study and temporary employees, as well as volunteers. FWRF was inspected by USDA APHIS for compliance with federal animal welfare regulations on 27 April 1999. To maintain optimal population size of captive wildlife species for use in research, we allowed reproduction in captive female mule deer and bighorn sheep. Nutritional Maintenance Feeding protocols: Feeding protocols were as previously described (reviewed by Wild 1997). We assessed nutritional programs by determining body condition of animals at FWRF using body weight data (tractable animals) or subjective scoring (fractious animals). Health Maintenance General: We followed established FWRF protocols for the preventive medicine program (Wild 1995) and chronic wasting disease (CWO) management (Wild 1998). We continued to monitor animal health using FWRF SOP's. Dental care in prairie dogs was adopted as an additional FWRF health maintenance SOP. �312 FacilitylMaintenancelRepairsllmprovements A variety of scheduled and unscheduled maintenance and repair activities were necessary to support facility operation and ongoing research programs. We worked toward a conservation-oriented approach for facility care by undertaking preventive maintenance projects, and performing high-quality new construction and repairs to existing facilities. Facility repair and construction projects were prioritized based on animal welfare concerns and anticipated research needs. Research Projects Facility operations offered support for pilot studies and for research projects conducted by CDOW personnel and other collaborators that were initiated, conducted, or continued using FWRF animals and facilities throughout the year. Educational Contributions Facility tours and educational lectures were provided to school, university, and professional groups visiting FWRF. We emphasized the importance of maintaining captive wildlife for performing controlled experiments and the contributions made by research projects performed at FWRF. FWRF animals and facilities were also used occasionally for hands-on training for professional groups. RESULTS AND DISCUSSION Animal Maintenance General: In FYI999, temporary, work-study, YCC employees and volunteers performed the majority of tasks involving animal care and facility maintenance at FWRF. Nineteen volunteers contributed 846 hr work to FWRF. These volunteers performed primarily caretaker tasks and also assisted in weighing animals and sample collection. Contributions by volunteers represented a savings to FWRF of about 0.4 TFTE and $8333 (vs. cost of temporary employees). The animal welfare inspection by USDA APHIS found FWRF in compliance with all AW A standards. This finding highlights the high standard of animal care provided by FWRF employees and volunteers. At the close ofFYI999, FWRF maintained 23 elk, 40 bighorn sheep, 4 white-tailed deer, 38 mule deer, 9 pronghorn, 30 white-tailed prairie dogs, and 11 domestic cattle. Two adult bull elk from Rocky Mountain National Park were added to FWRF as breeding bulls. Two adult nuisance deer (one buck, one doe) from the Silt area, Colorado, were also added to the herd; however, the buck was euthanized about 2 wk after arrival due to maladjustment. In addition to this buck, another 11 animals were euthanized as part of study protocols or for population management. Thirty-three natural mortalities occurred (Table 1). Ten of these mortalities were is neonates <1 week of age. Definitive cause of death in these neonates was not determined, however, lack of maternal care and hypothermia were suspected of contributing to losses. Nutritional Maintenance Feeding protocols: Individuals of all species except mule deer maintained reasonable body condition on available diets. This is the first year that we have maintained captive prairie dogs, so body mass information on these animals is included (Fig. 1). Body mass of prairie dogs increased from July through September 1998, then remained fairly constant. Body mass of torpid animals was lower (P:S 0.013) than non-torpid animals during March and April 1999. �313 Although ad libitum amounts of alfalfa hay and a high-energy pelleted supplement (Baker et al. 1998) were available to captive mule deer throughout the year, they were in poor body condition. Poor body condition of does may have predisposed some fawns to early mortality. Lack of intake of available feeds was most likely attributable to: 1) disturbance caused by cattle in pastures (although enclosures for feeding deer only were available), or 2) unpalatability of available feeds. We addressed each of these possibilities in our attempt to resolve this problem. First, cattle will be congregated into one or two pastures (rather than spread equally throughout all pastures). Cattle will be rotated among pastures about once every month. Small pass-through gates for deer only were constructed between pastures A and B 1, and B2 and C so that deer have the option to move from the pasture that is occupied by cattle. Secondly, we reassessed type of feeds offered to the mule deer. A pelleted browser diet (Mazuri Browser Maintenance Diet) appears to be much more palatable and of similar nutritional quality (D.L. Baker, Pers. comm.); however, cost is about three times greater than our standard high energy supplement. We will assess intake and body condition of mule deer offered ad libitum quantities of the browser diet in addition to standard feedstuffs over the next several months. We will also acquire natural browse as available from a local landscape company to supplement deer diet. Health Maintenance General: Overall, captive wildlife maintained at FWRF remained healthy throughout the year. Chronic wasting disease (CWO) continued to be a significant source of mortality in captive deer. High neonatal mortality in deer may have been attributable to poor body condition of does, young does having twins, stress of captivity in deer not hand-raised, and/or unusually cool, damp weather rather than a specific etiology agent. Tooth damage in prairie dogs was a minor problem affecting about I animalImonth. Source of tooth damage was not definitely identified, but prairie dogs may lose teeth when they chew on the sides of their cage, or when they bite the studded Kevlar gloves worn during handling. When tooth damage was observed, the animal was anesthetized and examined. If the remaining incisors had become overly long, they were ground to normal length with a sanding disk attached to a high speed Dremel tool. Animals with damaged teeth were fed powdered rat chow and/or small rabbit chow pellets until the teeth grew back, as they did in all cases. Chronic wasting disease: All animals at FWRF were monitored closely for clinical signs of CWO. Tissues from all mortalities occurring at FWRF were examined histologically for evidence of CWO. Of the 13 adult mule deer that died or were euthanized with clinical illness in FY1999, eight were due to CWO (Table 1). Additionally, one mule deer that died of other causes was positive to CWO on immunohistochemical (lHC) staining of brain tissue. Brain samples from the other mule deer that died of natural causes are awaiting mc staining. Facility MaintenancelRepairslImprovements As older portions of the facility are repaired and replaced, the need for unscheduled daily repairs appears to be decreasing. Maintenance projects continue to be important for animal safety and facility function. Significant maintenance/repair/ improvement projects completed at FWRF this year included: - Replace roofs on 3 feedsheds and several animal shelters Construct one feed shelter Replace main water pump Repair plumbing in east lab and waterline to automatic waterers Construct 15 individual prairie dog cages Begin replacement of (non-electric) fencing in west deer pastures Construct two deer pass-through gates �318 SPEC IES=Deer 130 IZO • Y 110 0 u 100 n 9 : I 0 0 ......._ • • • " 90 80 70 F' e III a I SO liO II • 'to • 30 ZO 1970 1980 1990 ZOOO Figure 1. Fawns: 100 does for mule deer DAUs plotted against year. The solid line is a cubic polynomial regression, w lines providing 95% confidence intervals on the regression. SPECIES=Elk , 100 90 • Y 0 u 80 n 9 : 70 I 0 0 SO F' II III •I e • SO 'to • • • • 30 • eo 1960 • 1970 1980 •• • 1990 Ye.r Figure 2. Calves: 100 cows for elk DAUs plotted against year. The solid line is a cubic polynomial regression, with dashed lines providing 95% confidence intervals on the regression. ZOOO �319 SPEC IES=OlSer 70 • 60 H • I 50 IS • 1 o o 'to '.'. • • .....•_... . '. • • '.......•....•... .... • ....• . .. ~"': .. :--._. :"_.... I • • .....__ _ F" 30 iii " •• . • • • • • I •: • ·· • • • • • • I • • • •I EO • • I • I e • 10 o •• • • I·· • • • Y--.~--.-~-.~--.-~-.~--.-~-r~--.-~-r~--.-~-r~--.-~-r~--.-~ 1970 1980 1990 2:000 Yaitr Figure 3. Bucks: 100 does for mule deer DAUs plotted against year. The solid line is a cubic polynomial regression, with dashed lines providing 95% confidence intervals on the regression. SPECIES=EII< 90 • • 80 • H • 70 I e • 60 : 1 50 0 0 F" • • 'to e " •I 30 e 2:0 • • • • • • •• • ••I •• • • • • 10 0 1960 1970 1980 1990 Figure 4. Bulls: 100 cows for elk DAUs plotted against year. ,;f.he solid-line is a cubic polynomial regression, with dashed lines providing 95% confidence intervals on the regression. 2:000 �320 Primary Regression Analyses The decline in young:female ratios over time for both deer and elk makes the analysis of the impact of sex ratios difficult. This difficulty is caused by time trends in male:female ratios concurrently, as shown in Figures 1-4. To capture the effect ofmale:female ratios on young:female ratios requires removing the year effect from the data. Thus, models that included year and the previous sex ratio were fit to the observed data for young: 100 females, i.e., (1) where Yi . is the observed young: 100 females ratio for year t and DAU j, the 13 's are the estimated parametels, and e ij is the random error, assumed normally distributed with mean zero and constant variance across years and DAUs for purposes of estimation. For purposes of estimation, the variable year had 1985 subtracted from the year of the observation. That is, for 1984, the variable year = -1, for 1986, the variable year = 1, etc. Thus, the intercept in the model represents the level in 1985. Five different regressions were conducted, 2 for deer and 3 for elk. For deer, sex ratio was taken as the male: 100 females ratio of the December surveys on the previous year, and also the mature males (males> 1.5 yr old) to 100 females ratios. For elk, the previous December male: 100 females ratio, the December male: 100 females ratio from 2 years previous, and the previous year's preseason male: 100 females ratio were considered. The following parameter estimates were obtained for the 5 analyses. All data are from post-season surveys except for the last model where pre-season sex ratios were used, but regressed against December young:female ratios. Species c;; }ntercept 130 (SE) Year ~1 (SE) P Year Sex Ratio ~2 (SE) P Sex Ratio Deer all males lagged 1 year 59.546 (l.703) -l.138 (0.115) <0.001 0.247 (0.070) <0.001 Deer mature males lagged 1 year 60.798 (l.041) -1.149 (0.090) <0.001 0.362 (0.067) <0.001 Elk all males lagged 1 year 46.765 (l.054) -0.692 (0.061) <0.001 0.285 (0.060) <0.001 Elk all males lagged 2 years 49.286 (0.940) -0.591 (0.060) <0.001 0.l75 (0.048) <0.001 Elk pre-season all males lagged 1 year 49.092 (2.458) -0.773 (0.101) <0.001 0.021 (0.085) 0.806 The impact of the previous year's sex ratio is an important predictor of young: female ratios, even with the year effect removed from the data. For total sex ratio lagged 1 year, estimates in the above table 'imply that each increase in 1 male: 100 females results in 0.247 (deer) and 0.285 (elk) young per 100 females. The effect of the previous sex ratio is not as important as the year effect, i.e., the decline in young:female ratios over time dominates the young:female data for both species. For example, with deer, to compensate for the expected decline of 1.138 fawns:lOO does per year, the sex ratio would have to increase by 4.6 bucks: 100 does. For elk, only 2.4 bulls: 100 cows are needed to compensate for the time trend in calves: 100 cows. �321 The importance of mature males for deer is not well supported by these results. An increase in 1 mature male: 100 females results in 0.362 young per 100 females, only a small increase over the 0.247 increase for all males. For elk, all males lagged by 2 years suggests a weaker effect on calf:cow ratios than did lagging the sex ratio just one year. For elk, all pre-season males lagged by 1 year showed no effect on the calf:cow ratios during the postseason survey. The sample size for this analysis was smaller than the analyses conducted with only postharvest data, with only 60 data points available. To further understand how consequential the sex ratio effect is, consider for all males lagged 1 year that an increase in sex ratio from 10 to 40 males: 100 females only results in 7.4 additions fawns: 100 does, or 8.6 calves: 100 cows. For deer, this increase would not compensate for the loss expected over a 10-year period of these data, i.e., 11.5 fawns: 100 does. For elk, the loss expected over a 10-year period is 6.9 calves: 100 cows, so an increase of 30 bulls: 100 cows during the 10 years could have compensated for the time trend in age ratios. Consistency Across Time Because I'm concerned that the sex ratio effect could be caused by just a portion of the data, I evaluated the 2 models with sex ratio lagged 1 year further. First, I split the data into 2 groups, years 1985 and before, and years after 1985. This year was selected to split the data approximately in half, although this year also corresponds to when antler point restrictions were implemented during harvest. The following graphs display the relationship between young:female ratios and the previous year's sex ratios for DADs for each species. SPECI ES=Dear 130 • 120 y 110 a u n 100 c- g 90 I 0 0 80 •• 70 • F e .~ 60 • •• •• •• •• • <> • • • • • • • • • III • SO • 'to I a • • 30 • eo 0 10 EO -: .. f' PERIDD ••• 'to 30 Prevlau. E.rly «=198&) Sex SO 60 70 R.tla o <>,~ Racent (>198&) Figure 5. Fawns: 100 does for deer DADs plotted against the previous year's estimated bucks: 100 does. Data points are distinguished by whether after 1985, or 1985 and earlier. .~ �322 SPECIES=Elk 100 Y • 90 0 u n g • 70 I 0 0 •• 80 • • • 60 • F e 50 1'1 • I 'f0 II • 30 • ZO 0 10 20 PERIOD ••• 30 E.rly «=1985) 50 ~ ¢ ¢ Recent 60 ()1985) Figure 6. Calves: 100 cows for elk DAUs plotted against the previous year's estimated bulls: 100 cows. Data points are distinguished by whether after 1985, or 1985 and earlier. Regression analyses conducted on the partitioned data for all males lagged 1 year suggested that there was no difference in the importance of the previous year's sex ratio by time period (P> 0.583) for either deer or elk. As the graphs in Figs. 5 and 6 show, the distribution of the data between periods is not completely dispersed, yet is not completely separated either. Consistence Across DAUs. To evaluate the consistency of the relationship between the previous total sex ratio and young:females ratio, I fit the model in Eq. (1) to each DAU with at least 5 data points with sex ratio lagged 1 year for both deer and elk. In addition, I also considered sex ratio lagged 2 years for elk. The estimate of the effect of the previous sex ratio was then analyzed by looking at its distribution, and testing to see if the mean of the estimates for each species differed from zero. The following histograms provide the distribution of the estimates of the sex ratio effects for deer and elk byDAU. �323 SPECIES=Deer F"REIOIUENCY 11 10 9 8 7 6 s 3 2: -2:.5 -2:.0 -1.5 -1.0 -0.5 0.0 previous 0.5 1.0 1.5 2:.0 2:.5 Sex Ra~lo Figure 7. Distribution of the estimate of previous sex ratio on fawns: 100 does for the 30 mule deer DAUs with at least 5 years of data. The mean is -0.0419 (SE 0.137), and is notsignificantly different than zero (p= 0.762). SPECIES=Elk FREQUENCY 12: 1, 10 9 8 ., 6 5 -2:.5 -2:.0 -'.5 -'.0 -0.5 Prevlou. 0.0 Sex 0.5 '.0 '.5 2:.0 2:.5 Ratio Figure 8. Distribution of the estimate of previous sex ratio on calves: 11.00 cowssfoathe 27 elk DAUs with at least 5 years of data. The mean is 0.187 (SE 0.106), and is marginally significantly different than zero (p= 0.089). The means of the parameter estimates for neither species were near the value estimated from the overall regression. Further, the wide range of estimates obtained across DAUs suggests that the relationship between previous sex ratio and young:females is not consistent across DAUs. I would expect this kind of noise in the data, because DAU's have been and currently are managed differently with respect to sex ratios. However, I find it disconcerting that the means of the parameter estimates are not close to the value �324 predicted from the overall regression. I'm unclear on how to interpret this result. However, these results do suggest that the effects of sex ratio on age ratios is not consistent across DAUs. The following maps show the magnitude of the estimates of the previous sex ratio spatially, and thus provide some ideas about which DAUs have age ratios most affected by sex ratios. Sex Rado Coefficiont ITIIlIII!D < -0.75 (",,,,,,,,,10.25 to 0.75 !IIIlI!IlIl !:H::AAI -0.75 to -025 >0.75 ~ -025 to 025 Figure 9. Estimates of the effect of previous sex ratio on calves: 100 cows by deer DAU. No estimates were produced for units lacking pattern. D-2, D-6, D-7, D-9, D-44, D-42, D-43, D-14, D-12, D-51, D-19, D-39, D-40, and D-25 were statistically negative (P < 0.10), and the rest neutral. ~ Sox Rado Coetrlclent ammm < -0.75 ~ 0.25 to 0.75 !I!IIillIIJ ~ -0.75 to -0.25 >0.75 IZZZ'Z'l -0.25 to 0.25 Figure 10. Estimates of the effect of sex ratio lagged 2 years on calves: 100 cows by elk DAU. No estimates were produced for units lacking pattern. E-2, E-6, and E-7 were statistically positive (P < 0.10), E-32 was statistically negative (P < 0.10), and the rest neutral. �325 ~ Sex Ratio Coefficient !IIIIIIIIIl < -0.75 .,,:::::::::.0.25to 0.75 mnmm-0.75 ~>0.75 to -0.25 I'2Z2Z?'.Zl -025 to 0.25 Figure 11. Estimates of the effect of previous sex ratio on calves: 100 cows by elk DAU. No estimates were produced for units lacking pattern. E-2, E-3, and E-6 were statistically positive (P < 0.10), E-l was statistically negative (P < 0.10), and the rest neutral. Threshold Effect To evaluate whether there is a threshold point in sex ratio where young:female ratios decline more rapidly in response to sex ratio, Ifit a model that forces the age ratio to zero for a zero sex ratio: (2) where y .. is the observed young: 100 females ratio for year i and DAU j, the I' 's are the estimated param~ls, and e i . is the random error, assumed normaI1y distributed with mean zero and constant variance across yeais and DAUs for purposes of estimation. For purposes of estimation, the variable year had 1985 subtracted from the year of the observation. That is, for 1984, the variable year = -1, for 1986, the variable year = 1, etc. Thus, the intercept in the model represents the level in 1985. Three analyses were conducted. For deer, only the previous year's sex ratio was considered. For elk, both the previous year's sex ratio, and the sex ratio lagged 2 years were considered. These estimates were obtained with a non-linear regression model. Estimates of the parameters and their standard errors are given in the following table, with graphic results in the following figures. �326 130 Species 131 (SE) (SE) 132 (SE) Deer lagged 1 year 67.601 (l.216) -l.333 (0.106) 0.638 (0.249) Elk lagged 1 year 52.548 (0.909) -0.671 (0.066) 0.296 (0.174) Elk lagged 2 years 53.700 (0.898) -0.601 (0.065) 0.348 (0.182) SPEC IES=Deer 130 • 120 110 <> y D 100 u n 8 1 0 0 F <> • • 90 '> •• • 110 • • •• • •• 110 (,. • • • • TO IS ••• • • • • • • o I IS • 60 'I. 3. ZO 0 10 PERIDD ZO 30 '10 Preulou. Total Sex Ratio ••••• 16 PredIcted ••••• 86 PredIcted ••• Early «=1585) ~ Q ~ Recent (>1985) 50 60 10 ••••• 96 PredIcted Figure 12. Fawns: 100 does for deer DAUs plotted against the previous year's bucks: 100 does ratio. Dots represent years 1985 or before, and diamonds years after 1985. The 3 lines represent the predicted fawns: 100 does for the years 1975, 1985, and 1995, from high to low, respectively, and demonstrate the effect of time on the relationship. There is no evidence to support a threshold of the sex ratio for either species that causes drastic declines in age ratios until sex ratios are unreasonably. Sex ratios would have to be lower than observed during the period of the data collection for this model to predict a strong decline in age ratios. �327 .0. • .0 y U n sa · TO • • 0 110 • • • 0 ..• • • 80 D •_ • •••• • • • ••••• ~, I • • c ..• o v 0- <> <) • F" • 5. · • • G c) c- I • • -c- <, <> c- ., '10 :0. ". c- _ 1:. • 10 3. '10 50 &0 Preulau. Ta~.1 Sex Ratla P~IIIDD ••• '7• .-redlc1'ed Eerly «_lS8&) - 0::-" •• ~.,dlot.,d 0' Recant ()1S8&) - 8 ••• ,..••dlot"ed Figure 13. Calves: 100 cows for elk DAUs plotted against the previous year's bull: 100 cows ratio. Dots represent years 1985 or before, and diamonds years after 1985. The lines represent the predicted calves: 100 cows for the years 1975, 1985, and 1995 from high to low, respectively. SPECIES= Y • 100 • 0 u • 80 n 9 70 1 60 0 0 50 Elk •• •• ¢¢ 40 F e 30 m a 20 I e 8 • 10 0 0 10 20 40 30 50 60 70 80 90 lOtal Sex Ratio 2 Years Ago 75 Predicted • • • Early 1985) « = - 85 Predicted - 95 Predicted o o o Recent (> 1985) Figure 14. Calves: 100 cows for elk DAUs plotted against the bull: 100 cows ratio from 2 years previous. Dots represent years 1985 or before, and diamonds years after 1985. The lines represent the predicted calves: 100 cows for the years 1975, 1985, and 1995 from high to low, respectively. Discussion Caution must be used in interpreting these results. The data were not collected as part of an experiment to assess the importance of previous sex ratio on recruitment. Sampling ofDAUs is not random, and there is clear danger that some of the observed patterns are a result of the lack of an explicit, definitive �328 experimental design. One example of a problem that may be inherent in these data is that occasionally not all the Game Management Units (GMUs) within a DAU are sampled within the same year, so that consecutive years of data from the same DAU might represent samples from different sets ofGMUs. Such an unorthodox sampling scheme could bias the results for that DAU, depending on how similar the population dynamics of the GUMs within the DAU are, particularly if the various sets ofGMUs are managed with very different harvest strategies. The analyses presented here do not demonstrate cause-and-effect relationships, and the approach used here can never demonstrate cause-and-effect because the sex ratio of the population was not purposely manipulated to observe the impact on young:female ratios. To demonstrate that male:female ratios during breeding over the ranges observed in these data cause changes in young:female ratios would require a manipulative experiment, such has already been proposed elsewhere. Note that most studies that have manipulated male:female ratios have done so by removing or harvesting females, which not only increases male:female ratios, but also confounds the effect by decreasing density. As a result, the experimental studies of the impact ofmale:female ratios on young:female ratios have confounded the impact of sex ratio and density. For that matter, the effects of density might also be present in the analyses presented here, with lower male:female ratios actually representing the effects of lower densities, and the resulting increase in young:female ratios is because of density-dependent responses. There is a clear need for designed experiments to examine the impact of sex ratio on young:female ratios. In particular, experiments that do not confound with sex ratio and density are needed if we are to understand the impact of sex ratio on age ratios. I do not have an explanation for the similarity of the response of elk (0.285) and deer (0.247) to changes in the previous sex ratio. Because the post-hunt male:female ratio for elk does not represent the sex ratio at time of breeding, I would not have expected to see any relationship. When the sex ratio was lagged for 2 years, only a weak relationship was observed. Analysis of the pre-season elk ratio suggests no impact on calfcow ratios. Possibly this lack of an effect is because the pre-season surveys are not effective in measuring bull:cow ratios, or because the sample size is too small to demonstrate an effect. However, the strong trend in year effects was present in the pre-season analysis. I am unclear on exactly how to interpret the results for elk, but the evidence presented does not provide a strong case for increasing sex ratios to increase age ratios. Summary The data for both deer and elk support a response ofyoung:female ratios to the previous year's male:female ratios, about 0.25 fawns: 100 does per 1 buck: 100 does for deer and 0.28 calves: 100 cows per 1 bull: 100 cows for elk. These effects are not adequate to compensate for the much greater decline in fawn:doe and calf:cow ratios that have been observed over the past 20 years in Colorado. Differences observed between DAUs suggests that only some of the DAUs might show an increase in young:female ratios were male:female ratios increased, and then only a small increase in young:females is predicted. There does not appear to be a threshold male:female ratio for either species where there is a drastic decline in young:female ratios. Acknowledgments Data used in this study were provided by CDOW terrestrial biologists via the DEAMAN database. Bruce Gill and David Freddy provided comments on a previous draft. �329 Task 6 (Workpackaee 7610) 1. PACE performance plans were prepared for all 12 Mammals Program staff members by the 31 July deadline, 2. PACE performance evaluations were completed for all 12 Mammals Program staffmembers by the 31 July deadline. 3. A draft predator management policy was prepared and is in the process of revision. 4. All encumbering documents were processed using the COFRS program such that vendors were paid with 7 days of receipt of invoices. 5. The Administrative Standard Operating Procedures Manual was comprehensively updated once during the segment and continually updated thereafter as changed. LITERATURE CITED Baker, D. L., G. W. Stout, and M. W. Miller. 1998. A diet supplement for captive wild Zoo Wildl. Med. 29:150-156. Wild, M. A. 1995. Animal and pen support facilities for mammals research. Colorado Rep., WPla, 11, Jul1994 - Jun 1995, Fort Collins. Wild, M. A. 1997. Animal and pen support facilities for mammals research. Colorado Rep., WPla, 11, Ju11996 - Jun 1997, Fort Collins. Wild, M. A. 1998. Animal and pen support facilities for mammals research. Colorado Rep., Jul1997 - Jun 1998, Fort Collins. ruminants. 1. Div. Wildl. Res. Div. Wildl. Res. Div. Wildl. Res. �330 � https://cpw.cvlcollections.org/files/original/e9c48e8fe56e867c837ae60aa9a41277.pdf dd0cf230c8021a164bc3b0e75c169df0 PDF Text Text 1 Colorado Division of Wildlife Wildlife Research Report April 2000 JOB PROGRESS Smreof ~C=o=lo=r=a=do=_ .Project: W-173-R-2 REPORT _ Peregrine Falcon Investigations (Avian Research) Work Plan: __ __.2=--__ : Job _--=...1__ Job Title: - -=-P...::e~re""g.:..:ri::.:n.:..e .=..F.=alo,::c""'-on:.:....:.;R:::::e""st""o.:,.::ra:;::ti'-"'o_::.n _ Period Covered: 01 January 1999 through 30 June 2000 Author: .,...-Personnel: Ge=ra=ld~R=:..... C=ra:::,cig:>-- _ James H. Enderson, Colorado College, and Gerald R. Craig, Michael Lanzone, Terry Meyers, Trish Miller, Bret Tennis, Nicole Trahan and Marni Zaborac, Colorado Division of Wildlife ABSTRACT .The period of this report encompasses the 1999 and 2000 breeding seasons. In 1999, 92 territories were occupied by peregrines (Falco peregrinus anatum) and 60 pairs fledged 140 young for an overall productivity of 1.69 young per total pair. The rare of occupancy remained above 80%. In 2000, 113 territories were occupied and 97 pairs fledged 208 young (2.19 young per total pair). Nests were not visited and no eggshells were collected. A sample of 41 sites was monitored for the second and third years. . These sites serve as surrogates to represent the total population. ��3 PEREGRINE FALCON RESTORATION Gerald R. Craig INTRODUCTION In 1972, only half of the 22 known peregrine falcon (Falco peregrinus anatum) breeding sites in Colorado were occupied. The following year, peregrines were placed on the federal endangered species list. Over the next 2 decades, an aggressive recovery effort was implemented in cooperation with the Bureau of Land Management, U.S. Forest Service and National Park Service. Following the ban on applications of chlorinated hydrocarbon pesticides, captive propagation, fostering and hacking were used to bolster the dwindling wild population. Two hundred and twenty eight young were fostered to wild pairs and 275 young were reintroduced to vacant sites through hacking between 1976 and 1990. By 1997, the number of occupied sites had increased to 87 from a low of 11 in 1982, and the falcon was considered secure enough to remove from the State's list of endangered species. Nationally, peregrines were de-listed in 1999. This project will continue to monitor population levels for at least 5 years as mandated by the Endangered Species Act to assure that the population does not relapse. Valuable insight will be gained about nest site parameters and population dynamics as nest sites are reoccupied throughout the state. In the course of recovery efforts, Colorado obtained the most extensive collection of wild eggshells in the nation. This collection can serve as a benchmark for comparison of future shell thickness. Over the past 2 decades, more than 850 peregrines were banded and band recoveries are providing important information on movements, wintering areas and mortality factors. This project should provide essential population parameters to protect these falcons and assure that Colorado's peregrine populations remain robust. P.N. OBJECTIVES 1. Annually survey all documented peregrine breeding sites throughout Colorado to establish the presence of nesting peregrines. 2. Annually monitor a statistically significant sample of breeding pairs to document their reproductive success. 3. Annually monitor organochlorine pesticide levels. 4. Investigate population dynamics. 5. Evaluate, characterize, and protect breeding habitat. 6. Document and protect important migration and wintering areas. SEGMENT OBJECTIVES 1. Whole, nonviable eggs that are encountered during eyrie visits will be collected, preserved and submitted to the appropriate U.S. Fish and Wildlife Service approved laboratory for pesticide analysis. 2. Compile data from previous years and prepare final report of eggshell thinning and pesticide residues in Colorado peregrine falcons from 1974 through 1997. 3. Compile and analyze data on peregrine falcon recovery and population dynamics and prepare manuscripts and annual report. 4. Monitor selected peregrine falcon eyries for occupancy and productivity when funded. �4 METHODS Beginning in early April, up to 3 teams of observers (depending upon funding availability) travel throughout the state and monitor peregrine breeding behavior and nesting success. Teams are comprised of 2 individuals, at least one of which is knowledgeable of nest locations and has experience observing breeding peregrines. This assures that knowledge is passed to less experienced team members. Potential nest cliffs are usually observed from distances of ~ to Yz mile with 15-45x spotting scopes. Team members jointly keep the site under continuous observation to avoid missing often brief, but critical, behaviors. Observation periods average 4-6 hours, although some may be shortened to Yz hour if critical behaviors are documented promptly. Observations may have to be extended to 8 hours and even repeated the following day if observations are inconclusive. Generally at least 4 visits are scheduled per site; an initial visit to document the presence of a breeding pair, a second visit to document egg laying or onset of incubation, a third visit to confirm hatch of young, and a final visit to confirm the number of young fledged. Because most of the nest sites are not visible, the reproductive stage is inferred from behavior of the adults (incubation exchanges, prey deliveries, etc.). RESULTS AND DISCUSSION 1999 Survey Effort Thee teams comprised of 2 observers each were fielded for the 1999 season. Each team was assigned 20 priority territories to monitor. Among the 60 sites were 45 surrogate sites considered to be representative of the Colorado peregrine population. Selection was based upon occupancy longevity (prior to 1990), accessibility, and distribution. In addition to monitoring the priority sites for productivity throughout the season, the teams also attempted to visit all registered sites as well as survey potential cliffs throughout the state as time permitted. The teams were able to check 105 of 115 sites that were on record at the beginning of the field season. Time constraints and accessibility restricted them from visiting the 10 remaining registered sites. Six potential nest cliffs were also surveyed and two new pairs were located. Two previously undocumented pairs were also reported by other sources and occupancy was confirmed at those sites as well. By the end of the season, the number of known nesting territories within the state had increased to 119(Fig. 1) and the field teams were able to confirm occupancy at 82 of the 105 sites they visited (Fig. 2). 2000 Survey Effort Thee teams of observers were employed for the 2000 field season and conducted inventories in the same manner as the previous year. All of the team members were experienced peregrine observers, and 4 had worked on the project in previous years. The experience paid off'. All priority surrogate sites were' . monitored and through their extraordinary effort, 14 new nesting pairs were located whichincreased the' state total to 133 territories (Fig. 1). Inaccessibility and time constraints kept the teams from visiting only 3 sites. Occupancy was documented at 113 sites; the other 17 were vacant. 1999-2000 Territory Occupancy Occupancy trends have been obscured in recent years because of inability to visit all sites on record (Fig. 2). Approximately 10% of all registered sites were not checked from 1977 through 1999 and this problem will persist in the future as more sites are added and observer time remains fixed. Therefore, occupancy is best presented as the number of occupied territories per total territories checked as displayed in Figure 3. The rate of occupancy remained above the 80% level over the past 7 years and is generally considered appropriate for a stable population. In the future, it will probably be necessary to base the occupancy rate on the sample of surrogate sites that are occupied. A sampling regimen that was established in 1998 will continue to be tested. �5 140 130 120 110 l- 100 I- 90 80 .s'" en l- 70 l- l- f- I- f- I- 60 t- l- I- 50 l- I- l- I- l- 40 30 ;- 20 l- l- 10 I- l- I- l- f- l- f- f- f- l- I- f- If- f- f- l- f- lI- 0 .co N M co CD 0> e- 0> V <X) 0> ,- OJ e-- l{) <0 IX) co ,.._ co aJ co 0> ..- 0> e- en ..- 0> ,.- Years Fig. 1. Peregrine Nesting Territories on Record. 140 130 ....... --_ --_ 120 ---- --_. __ - -_ .._. __ .. _ ---- ..--_ -_ _--- ..--_ .._---_ -- -.--. __ ---- ..---- __ - -_ _ - -.-_ _-_ _ -- __ - __ _ _-_ - __ -_ - _._ _._--- --_ _ - _ 110 100 .s'" en _.-------------_.-----_.- ------ --_. _. -- 90 ... _ 80 ............. 70 60 50 -- ••••• - --_ ••• ---- ••• -_ •• _-_ •• - ••••• - - _.-.- - _ ••• _ •••• -_ •• _.- -- -. '" _ :: - _ ::::: ••• - ""'" _ ••• _ •• _ ••••••• _--_ •• _ ••• _._-- _ ••• __ • __ o ·_··_····_ - .--_ ••• _ •• - _ •• -_. - .._ _ _ _ •••• ----- _. .. -.._ __ _ ••• -- - ••• _ •• - ------- . .... _............ .. .. •• -- •••••• - • - ..- .:;::: _. mI~i _" {: ~~}~... ::::: .. '::::: _ :.-:-:-~.:.~ .i.~.~.~.~... ... :::~ . : :: ::_!i.;.I~~ •••····I··* ~ llj1~ " 40 ... ..- _.- 30 20 10 0 Years 1111 Occupied IJ Unoccupied D Unchecked I Fig. 2. Site Occupancy 1972-00. 1999 Reproduction Breeding (egg laying) pairs were present at 73 sites; non-breeding pairs occupied 6 sites (5 pairs were comprised of subadult females) and breeding could not be confirmed at lather site. Sixty pairs successfully fledged 140 young (2.33 young/pair) for an overall productivity of 1.69 per occupied site (Fig. 4). Forty-one of the surrogate sites were occupied and 36 successfully fledged 72 young. As in 1998, the ::::.. �6 100% 90% 80% 70% 60% 'E GI 0 •... GI a.. 50% 40% 30% 20% 10% -.---.--.--.--- - - -.. -- -- --------.--- -.-- - --- -.-- . Years Fig. 3. Rate of Occupancy of Monitored Sites. surrogate sites experienced a slightly higher productivity of 1.76 than the 82 sites that were visited. Hence, the surrogate sites continue to be a reasonable estimate of population productivity given the large variance that occurs in these statistics. ( 2000 Reproduction In 2000, the field teams located 97 breeding pairs. Non-breeding pairs occupied 13 sites (7 were pairs contained subadult females) and a lone adult was present at 1 site. Outcomes were not determined for 2 pairs. Seventy-seven of the 95 pairs that were monitored were successful. They produced 238 young, of which 208 successfully fledged. The 2000 breeding season experienced record productivity of2.19 young per known outcome pair. A1l41 of the surrogate sites were again occupied in 2000,38 had breeding pairs of which 32 were successful and fledged 84 young for a productivity of2.21 young per known outcome pair. These productivity values are similar to the statewide productivity. Productivity above l.25 is usually indicative of an expanding population and values above l.5 for 1998-2000 (Fig. 4) suggest that Colorado peregrines are reproductively secure. . . E22shell Condition Personnel and time limitations did not permit nest site visits to band nestlings as in pervious years and as a consequence, no whole unhatched eggs or shell fragments were collected. Or2anochlorine Residue in E22s A collection of the contents from 53 unhatched eggs acquired since 1991 still await pesticide analysis at the facilities of Colorado College. Funding was not available for the pesticide assays and the samples remain archived. �7 3.00 ...•... 2.50 , , •, '" '.".. '" GI A~ '11 1.50 t 1.00 / 1\ GI r.. .-I Ol t: . \ GI Ol '11 . 0 >< ,. l,) '" .-< '" r- M r- r- '" .-< .-< ' I . ~ J I s; '" •.. A ~ r....... ~~ ) )~ I' ~ ~ '"r- '" , • A", I \ ') ~ ..• p ~ I ~ 0.50 I \ , 11 ;l 0.00 ~ , 2.00 '" .-< eo ID r- rr- '" '".-< .-<'" .-< r-. l) ~ '" .-< '" r- 0 eo en .-< .-< '" CD CD '" ....• '" .-< ..• M CD '" eo .-< ...• '" '" eo ...• '" V eo .)a.. ID r- eo en .-< CD CD eo ....• '" .-< '" ...• '" '" 0 .-< ...• '"'" en '" '"'"...• ....• N M en en .-< )'.V )\ .••IL .....• ~ I ..• ...• '"'" '" ...• '"'" )~ ID en en .-< r- ....• '"'" eo en '" .-< '" '"'" .-< 0 0 0 cv Years ____'__w i Fig. 4. Peregrine Productivity 1d Productivity - •• - Total Productivity 1972-00. Data Compilation Field notes from 1972 through 1999 were reviewed and compiled into a comprehensive database to facilitate data analysis. Field observations from the present season will also be included. Sections were drafted on management efforts, movements, mortality and early population distribution in anticipation of a final report that chronicles Colorado's recovery program. ~(;~~k?~~~~~a~;>~~ _ PREPAREDBY: __ Ge'raid~g, GPV ��9 ,. Colorado Division of Wildlife . Wildlife Research Report ApriI2000 JOB PROGRESS ,':, REPORT . " State of: Colorado . Project: W-180-R Work Plan: _----.:1'--_ Job Title: .,-- Job __ Raptors ----'1.____ ....,.----=Ra=D=to=r'-'M::.:.=an=a""ge=m=e=n=t..=an=d-=.P..=o&.D=ul=a=ti=o=n..::.A-=.:s=s=es=s=m=e=n. _ Period Covered: 01 July 1999 through 30 June 2000 ., Author: Gerald R Craig Personnel: Gerald R Craig, Deanna Meinke, Gary Meinke, Colorado Division of Wildlife ABSTRACT A raptor nest-mapping project involving Fort Collins, Loveland, Windsor and Wellington was initiated. The . maps will provide information about sensitive raptor use areas within the growth areas of these communities ' .so they can Incorporate them into their planning process. Although records are incomplete, 89 inquiries from the public and governmental entities about raptors were logged into the office. Bald eagle nest monitoring and . mid-winter inventories continued. The numbers of wintering eagles declined and the nesting population continued to increase. Thirty-seven nesting pairs produced 45 eaglets for the highest productivity recorded for the state; To maximize resources, volunteers provided all of the field assistance on the eagle banding and nest monitoring projects, The results were comparable to paid staff. ��11 RAPTOR MANAGEMENT AND POPULA nON ASSESSMENT Gerald R. Craig INTRODUCTION Interest in raptors continues to increase and, although losses from shooting and poisoning have declined, Colorado's population growth over the past decade has reduced important wildlife habitats along the Front Range as well as inter-mountain areas. Increased human visitation and conflicting usage (e.g., running and dog exercise) are impacting existing natural areas that presently support raptors as human population in the metropolitan population increase. Their visibility and sensitivity to habitat perturbations draws raptors into land use controversies as the publics attention focuses on shrinking agricultural and open lands. Governmental agencies also include raptors among the wildlife species of concern when they address project impacts. For these resons, the CDOW receives numerous requests for information and assistance from federal, state, county and municipal governments as well as non-governmental organizations and the public. There is a need to disseminate technical and biological expertise on raptor habitat requirements, distribution, and inventory methodology to agencies and non-governmental organizations. Expertise will also be needed to assist agencies and developers as they devise mitigation measures. There is a need to provide centralized expertise to the public and CDOW field personnel. Conflicts involving raptors are often unique and may require innovative actions. Complaints vary from attacks on poultry to insomnia caused by incessant hooting of amorous owls to owls nesting in grain elevators or flowerpots. Field personnel deal with these complaints infrequently and circumstances are often unique. There is also a need to provide specialized expertise to assist District Wildlife Managers with their responses to raptor related conflicts. Basic knowledge of population levels and trends is lacking for the majority of Colorado's raptors. Some species are recently recovered (e.g., bald eagles and peregrine falcons) while others (e.g., burrowing owls and ferruginous hawks) may be declining, but limited resources preclude monitoring to assess their population trends. This project will inventory and monitor selected raptor species to develop the requisite baseline population information. P.N. OBJECTIVES The objectives of this project are to: (1) maintain a current database on important raptor nest sites and concentration areas, (2) advise federal, state and local governments on habitat requirements, recommend buffer distances and mitigation measures to assure occupancy of nests and hunting areas for raptors, (3) assist field personnel with issues concerning raptors, and (4) monitor populations of selected sensitive raptor species and those that may be declining. SEGMENT OBJECTIVES 1. In cooperation with the Habitat Section, expand and maintain a current database on the location and status of raptor nests associated with urban growth areas. Review literature and maintain an updated version of the document Recommended buffers for Colorado raptors. Evaluate results of previous efforts by land managers to preserve or enhance raptor habitats. �12 2. Respond to requests and inquiries by agencies, municipalities, developers and concerned publics to protect or enhance raptor areas threatened by urbanization or development. Review and evaluate current and past efforts that have been implemented to maintain raptor use of habitats threatened by urbanization. If protective measures are recommended, the concerned entity will be contacted to determine if the measures were implemented. Return visits should be made over ensuing years to determine if sites remain occupied after development occurs. 3. Respond to requests from field personnel for assistance on raptor related issues. 4. Monitoring of selected species will be accomplished as needed with available resources. Important sites (nests, night roosts and concentration areas) will be described, entered into the database and monitored in future years. 5. Compile data and prepare annual report. METHODS Inquiries E-mail and telephone logs were maintained on inquiries about raptors that were received from the public and private sectors. The inquires were later entered into an Excell spreadsheet for compilation and analysis. Records were also kept of meetings, lectures, and presentations where information on raptors was disseminated. Population Monitoring Bald eagle nesting efforts were monitored through annual visits to documented nesting territories. Observers consisted of Division employees (District Wildlife Mangers and Area Biologists), landowners and volunteers. Visits were also made to confirm recently reported nesting attempts. Nests were observed from sufficient distances (at least 200 yards) to avoid disturbance. Observations were timed to confirm presence of breeding pairs and later to document number and age of young. Work began in March and continued intermittently until the nests either failed or fledged young in June and July. Efforts were made to band nestlings at sites when young were 6-9 weeks of age. Banding did not occur at those locations where landowners did not want their eagles disturbed, where nests were inaccessible, or where climbing could jeopardize the nest structure. A professional tree climber experienced in handling eaglets climbed trees. The young were placed in a bag and lowered to the ground to be banded, weighed and measured. Any prey remains in and below the nest were identified and counted. US Fish and Wildfire' Service rivet style bands were placed on one leg and the other leg was marked with a red anodized band to which a short (1.5X 3.0 inch) ALLFlex® laminated vinyl tag was affixed. The anodized band and tag were embossed with similar alpha-numerics. The markers permitted identification of individuals that returned to breed without the necessity of trapping and handling them. Banding visits were timed to avoid prolonging disturbance beyond an hour. Mid-winter Eagle Counts Mid-winter bald eagle counts were conducted in accordance with methodology established in the National Wildlife Federation's Midwinter Bald Eagle Survey. Counts were conducted annually on the second weekend of January, Participants traveled standardized routes and all eagles were identified and counted. Standardized point counts were also made at roosts. Forms are returned to the Boise Office of the Biological Survey for inclusion in the national survey. I I �13 RESUL TS AND DISCUSSION Urban Raptor Nest Mapping An effort was initiated to map raptor nests in the vicinity of Fort Collins, Loveland and Windsor and Wellington growth areas. In the past, the Division has identified nest sites of species of concern (eagles, ospreys and burrowing owls), but the commoner species (red-tailed hawks, Swainson's hawks, kestrels and great horned owls) have not received attention. The following nests were identified and mapped: Species Bald Eagle Golden Eagle Osprey Red-tailed Hawk Swainson's Hawk Prairie Falcon Great Horned Owl Common Raven Number 2 8 3 8 2 I 3 I A preliminary map of nests within the Fort Collins growth area was provided to the Fort Collins Natural Resources Department for planning purposes. Requests for Information or Assistance A log was maintained of inquiries about raptors received in Fort Collins from July 1, 1999 through June 30,2000. However, due to time constraints, a number of contacts were not recorded, and the following must be considered a sample. Eighty-nine contacts were logged over the period and the source of contact follows: Contact Method Number 35 Telephone E-mail 35 Meeting 16 Lecture 2 Correspondence ~ 89 As the Division's offices received Internet capability, the number of e-mails increased toward the end of the period. The e-mails also remained on record and could be accessed for review. Phone messages were .frequently not documented because time was not taken at the end of each call to record the source and subject of the conversation. Inquiry sources were: Sources Private Parties Private Groups Federal Agencies Colo. Div. of Wildlife Colo. Div. of Parks Counties Number 11 5 14 35 I 8 Sources Municipalities Consultants Rehabilitators Newspapers Public Service Co. Developer Number 3 5 3 3 1 _1 89 �14 As expected, the greatest number of inquiries (35) originated from within the agency, since the field personnel had a history of contacting the principal investigator. Among the federal agencies, the U.S. Forest Service accounted for 12 contacts. Although not recorded, at least 6 additional inquiries came from the U.S. Fish and Wildlife Service. Geographically, most of the inquiries originated along the Front Range. The inquiry subjects are catalogued as follows: Subject of the Inquiry Nest SiteIHabitat Protection Development Impacts (subdivisions, roads, pipelines) Recreation/Climbing Disturbance Nest Reports IdentificationlNatural History/Status Inventories Artificial NestslPerches MortalitieslInjuries Rehabilitation Laws and Licensing Predation Pigeon Control Miscellaneous (feathers, mounts, photos, etc.) Number 31 11 7 8 9 2 4 4 4 5 2 2 2. 93 The majority (65%) of the inquiries related to impacts of urbanization and requests to protect, maintain or enhance habitat. Again, most inquiries were associated with the Front Range. Population Monitoring Efforts are currently underway to monitor wintering and breeding bald eagles statewide, and document the distribution of burrowing owls in Weld and Larimer counties. Breeding Bald Eagles Fewer than a dozen bald eagle nests have been reported for Colorado. In 1974 one pair was discovered in Moffat County. The landowner recalled the nest had been occupied in 1947, so it is probable that a few eagles have always bred in the state. The following chart displays the expansion over 3 decades. The Northern States Bald Eagle Recovery Team assigned a recovery goal of 10 pairs to Colorado. That level was attained in 1987 and sustained after 1990. The national population has also increased and is likely to be delisted within the year. Ongoing monitoring by Colorado will assist in meeting the mandated postdelisting monitoring program. Although the state will follow suit and delist the eagles, they remain a species of concern and should be monitored to assure the populations remain secure. Expansion of nesting pairs is occurring along the northern Front Range and northwestern portions of the state. In 1999,29 nesting territories were occupied and 25 pairs produced 40 young. The number of occupied nests increased to 37 in 2000 of which 23 pairs produced 45 young. A banding project was initiated in the late 1980s to document movements and mortalities of Colorado eaglets. Since then, bands were placed on 190 eaglets. In 1999, banding responsibilities were turned over to a volunteer team of Gary and Deanna Meinke to coordinate and they have banded 54 young over the past 2 breeding season. A standard Fish and Wildlife Service band is placed on one leg and a colored marker band and short vinyl tab is affixed to the other leg. The marker permits identification of birds at a distance. Three color-marked individuals have returned to breed in the general areas they were hatched and band recoveries of 7 others have been reported. �15 50 45 40 35 ~ ·iii 30 f_ D... g' 25 "C Q) ~ III QT /"d 20 15 N ! 7 ~ /\f 10 5 0 n n-I / i' ~m~IVII . . Years IIIIIIiilIiIITotalPairs -o-Young I Bald Eagles Nesting in Colorado Midwinter Bald Eagle Counts Between late November and the end of March a fairly large population of bald eagles winter in Colorado. It is estimated that Colorado's share of the eagles may range from 1,000 to 1,500 individuals. These birds move into Colorado from northern states and Canada. Eagles are very mobile. Wintering population in Colorado fluctuate depending upon weather conditions and upon prey availability. A national effort has been underway to inventory the wintering eagles and obtain information on population trends. Over the past 20 years, Colorado has participated in a national census conducted annually in early January. The results provide a national overview of the species population trends. Standardized routes are traveled either with vehicles or aircraft and all eagles are counted. The results are provided below: Although midwinter eagle counts were initiated in 1980, standardized routes were not finalized until 1987. The low count in 2000 is mainly attributable to failure to fly two key routes along the Yampa and White rivers. In previous years these routes have accounted for 175-300 eagles. If the same number of eagles had been present, the population levels would have at least paralleled the 1999 counts. While only the standardized routes are used for census purposes, the Division accepts all counts made during the period as a gross estimate of overall population levels. Volunteer Effort Aside from the duties of the principal investigator and assistance from the Division's Wildlife Managers and Area Biologists, volunteers conducted nest monitoring and banding. Mileage and per diem expenses were reimbursed for the eagle banding effort. Listed below are the volunteer hours contributed to this project in 1999 and 2000. �16 1000 900 800 700 600 500 400 300 200 100 0 r0::> 0::> 0::> (J) (J) (J) (J) ...- 0 0::> (J) (J) ...- (J) (J) Standardized Mid-Winter Bald Eagle Counts Hours Volunteers Project 1999 2000 1999 2000 Bald Eagle Nest Monitoring 4 16 135 550 Bald Eagle Banding 3 4 386 430 Total 7 20 521 980 Estimated Value (hrly rates of$12-$15) PREPARED BY: C;..:::l.·-,-._:_fl!__'_:~=--.:::!~L--__ Geral~alg GPV $7,024 $13,600 �17 Colorado Division of Wildlife .. . Wildlife Research Report '. April2000 JOB PROGRESS Project: -'-W::....--""1.>::,.66::<...-""'R"-_ Work Plan: __ -=1,--_: job Title: =2=.2 Migratory Bird Investigations _ M"""·""'o:::m!.::·to""'r'-'B:::.:an=d:::.in=g'-"o""-f...o.:M~a::::.lI""'a::..:rds=_.!.in..._",C"'"oc:.,::lo<.:,;ra=d""'o'_ Period Covered: Author: .; -,,- __ .Personnel: . Olterman, Job __ REPORT 01 January 1999 through 30 June 2000 -""T""'od:::.:d"-'Ao....:..:....;. S:::.:an=d~ers'-"'- _ J.Broderick, M. Szymczak; P. Creeden, V. Graham, J. Gurnber, T. Mathieson, J. Miller, J. Olterman, R. and S. Yamashita; Colorado Division of Wildlife. ABSTRACT Ducks were trapped in modified Salt Plains bait traps and banded in western Colorado from late August to early September 1999. A total of 540 mallards (Anas platyrhnchos) was banded; 228 in 1 area near Grand . Junctionand 312 in-4 areas near Cortez. The 540 bird sample consisted of68% immatures and 62% males, COLOow ~LDullll~rOO~~Imlillimr BDOW013553 ��19 PRESEASON MONITOR BANDING OF MALLARDS IN COLORADO Todd A. Sanders INTRODUCTION In 1990, the Pacific Flyway Study Committee formulated a 5-year cooperative mallard and northern pintail (Anas acuta) preseason banding program, which was endorsed by the Pacific Flyway Council and intiated in 1991. The program was designed to ascertain mallard and pintail migration and survival throughout the western U. S., including Alaska, and the Canadian provinces of British Columbia and Alberta. Following the 5th year of the banding program, analyses of band recoveries showed cohorts of mallards banded in southeastern Idaho, western Wyoming, northern Utah, and western Colorado had similar band recovery distributions. Further, trapping and banding efforts in the 4-state region were most successful in southeast Idaho and western Colorado. Consequently, these 2 regions were selected for continued banding to determine if establishment of a western mallard management unit is warranted. Banding and band recovery data also contribute toward estimating harvest rates, survival rates, and distribution of harvest for use in the development of an adaptive harvest management strategy for western mallard populations. Banding in 1999 marked the 4th year of monitor banding. P. N. OBJECTIVE Trap, band, and determine the age and sex of at least 500 mallards annually during late August to early September in the Pacific Flyway portion of Colorado. SEGMENT OBJECTIVE 1. Trap at least 500 mallards annually in the Grand Junction and Cortez areas of western Colorado during late August to early September using Salt Plains bait traps (Szymczak and Corey 1976). Ducks will be banded with U.S. Fish and Wildlife Service bands and classified according to age and sex using accepted techniques (Carney 1964, Weller 1976: 35). Banding schedules and recapture reports will be submitted to the U.S. Fish and Wildlife Services' Bird Banding Laboratory. Band recovery reports will be prepared by Colorado Division of Wildlife personnel. METHODS Trap Area and Site Selection The selection of trap sites for continued mallard banding in western Colorado were based on the number of ducks trapped per unit of effort from 1991 to 1995, and on the availability of personnel to operate banding stations. We selected 5 trap sites; Walker State Wildlife Area (WSW A) near Grand Junction (1996 to 1999) and 4 wetlands (Toten Reservoir, Nolan's Pond, Merritt's Pond, and Williamson's Pond) near Cortez (1998 to 1999). Trap sites at WSWA were located on backwater areas within side channels of the Colorado River. Trap sites in the Cortez area were on 3 small perenial ponds and a bay at Totten Reservoir. �20 Trapping Ducks were trapped in modified Salt Plains bait traps (Szymczak and Corey 1976) using whole, shelled com for bait. We checked traps daily. Mallards were the target species. We recorded the date, trap area, age, sex, and band number for each duck banded. Also, we recorded the date, trap area, and band number for each duck trapped that had been banded previuosly. RESULTS Trapping and Banding We trapped and banded a total of 540 mallards in western Colorado between 30 August and 13 September 1999 (Tables 1 and 2); 228 in 1 area near Grand Junction and 312 in 4 areas near Cortez. The 540 bird sample consisted of68% (368/540) immature birds and 62% (337/540) males. Record Keeping and Band Reporting We prepared electronic files containing data on duck banding and recaptures at the Colorado Division of Wildlife's Research Center in Fort Collins. All data was submited to the U.S. Fish and Wildlife Service's Bird Banding Laboratory on standard forms. LITERA TURE CITED Carney, S. M. 1964. Preliminary keys to waterfowl age and sex identification by means of wing plumage. U.S. Department of the Interior, Fish and Wildlife Service Special Scientific Report-Wildlife 82. 47pp. Szymczak, M.R., and 1. F. Corey. 1976. Construction and use of the Salt Plains duck trap in Colorado. Colorado Division of Wildlife, Division Report 6. 13pp. Weller, M. W. 1976. Molts and plumages of waterfowl. Pages 34-38 in F. C. Bellrose, editor. Ducks, geese and swans of North America. Stackpole Books, Harrisburg, Pennsylvania. Prepared by: ~crJd a ~/rtJ1A./j Todd A. Sanders, General Professional IV �21 Table l. Number of mallards banded by location, age, and sex in western Colorado between late August and early September 1999. . Area Site Grand Junction Walker State Wildlife Area Cortez Merritt's Pond Nolan's Pond Totten Reservoir Williamson's Pond Subtotal Total 1M Age and sex IF AM 93 73 27 6 68 27 128 221 19 7 33 15 74 147 AF Total 43 19 228 30 4 30 9 73 116 16 92 19 142 59 312 540 11 8 37 56 Table 2. Number of mallards banded annually by age and sex in the Grand Junction (GJ) and Cortez (CZ) areas of western Colorado, 1996-1999. Area Year GJ CZ 1M 1996 1997 1998 1999 Total 362 369 343 228 1302 t t 143 120 302 221 786 t No ducks were trapped 425 312 737 and banded. Age and Sex AM IF 133 116 196 147 592 68 93 202 116 479 AF Total 18 40 68 56 182 362 369 768 540 2039 ��23 ., Colorado Division of Wildlife Wildlife Research Report April2000 JOB PROGRESS State of: C=ol=o.:..:ra=d=o " Project: _ _.:W'-!........!-1:...:::6~6-...:.R::...._ _ Work Plan: __ Job Title: REPORT -"1",,,,0 __ : Job __ Migratory Bird Investigations ...:.1 __ ~C~oo~p~e~ra~ti~v~e~M~an~a=ge~m~e~n~t~P~ro~g~r~am~s _ Period Covered: 01 January 1999 through 30 June 2000 Authors: James H. Gamnionley, Michael R. Szymczak. and Todd Sanders Personnel: Alex Chappell, James H. Ganunonley, Michael R. Szymczak, and Todd Sanders, Colorado Division of Wildlife. ABSTRACT Wetland projects were reviewed and selected for funding with Wetland Initiative, Colorado Duck Stamp, and DucksUnlimited, Inc. MARSH funds. Work with the Wetland Focus Area Committees continued in , relation to wetland project development proposals. Responsibilities as Colorado's representative on Pacific Flyway Study Committee and Central Flyway Technical Committees, were fulfilled. Flyway ,'.resp~nsibilitiesincluded providing support for cooperatively funded flyway studies. Recommendations for wetland habitat improvements and/or management were provided for public and private land managers " throughout.Colorado. Presentations on wetland ecology and waterfowl identification were given at , , workshops. ' .,': -:,' BDOWD13554 ��25 COOPERA TIVE MIGRATORY BIRD MANAGEMENT PROGRAMS In 1988, the Colorado Division of Wildlife (CDOW) created the Migratory Game Bird Program Unit (MBPU) within the Terrestrial Wildlife Section. This administrative change combined all individuals having statewide responsibilities for research and management of migratory game birds. Members of the MBPU worked in concert to improve migratory bird management in Colorado. This job was created to allow team members to participate in these management programs. In November 1993, project personnel assumed additional responsibility for leading and administering the Duck Stamp wetland development program. Since 1993, personnel of the MBPU have taken on additional responsibilities within wetland programs in Colorado. In July 1996, the MBPU was dissolved with most of the responsibilities being transferred to the Avian Program. This report covers activities of the migratory bird segment of the Avian Program. P. N. OBJECTIVES 1. Continue to aid in the activities of Wetland Focus Area committees in the state, monitor the functioning of these committees, and aid in obtaining funds for proposed projects by serving as chairman of the state-wide oversight committee and state coordinator for the Intermountain West Joint Venture. 2. Advise public land management agency personnel, including area biologists and district wildlife managers, on the potential benefits to migratory birds of acquisition and/or development of wetland areas. Activities may include on-site inspection, formulating plans for the collection of biological data, recommending wetland enhancement developments, or review of development plans. Provide information on wetland habitat development to private land managers. 3. Prepare and present information programs on the principles of migratory bird and wetland management. Preparation may include literature review, construction of charts, graphs, and tables as photographic slides or posters, and rehearsal of presentations. 4. Attend flyway Technical Committee and Council meetings and flyway special workshops as assigned. Compile information on current status of Colorado's migratory bird population and hunting season results through consultation with CDOW biologists and managers. Prepare reports for presentation at _ meetings and workshops. Serve on committees as assigned QYflyway Technical Committee and/or Council' Chairman. Attend selected meetings 'in Coloradothat address 'migratory bird management programs, at which in-depth biological expertise would be ofvalue. 5. Provide methodology to migratory bird and wetland managers for sampling the biological parameters of interest. Literature review may be required to develop appropriate methodology. Parameters of interest may include breeding pairs, nesting densities, nesting success, fledging success, vegetation composition and density, invertebrate composition and density, or population survivaL" ,,', ',' 6. Assist in collecting information that will enable waterfowl and wetland managers to make decisions on population and wetland management. �26 SEGMENT OBJECTIVES 1. Use the knowledge and skills of the Federal Aid supported members of the Avian Research Program of the Colorado Division of Wildlife to facilitate wetland and waterfowl management and informational programs in Colorado. a. Monitor the functioning of Wetland Focus Area committees and aid in the design and funding of proposed wetland conservation projects by serving as Colorado Division of Wildlife representatives on the Wetland Initiative Partnership Committee and other Colorado Wetland Program committees, Intermountain West and Playa Lakes Joint Ventures of the North American Waterfowl Management Plan, and as chairman of the Waterfowl Habitat Project Review Committee. b. Attend Central and Pacific Flyway Technical Committee and Council meetings and workshops as assigned. Compile information on current status of Colorado's migratory game bird populations and hunting season results through consultation with CDOW biologists and managers. Prepare reports for presentation at meetings and workshops. Serve on committees as assigned by Flyway Technical committees and/or Council Chairmen. Attend selected meetings in Colorado that address migratory bird management programs, at which in-depth biological expertise would be of value. c. Participate in cooperative migratory bird studies sponsored by the Central and Pacific Flyway councils. d. Advise public land managers on the potential benefits to migratory birds of acquisition, development and/or management of wetland areas. Provide information on wetland habitat development to private land managers. e. Prepare and present information programs on the principles of migratory bird management. Preparation may include literature review, construction of charts, graphs, and tables as photographic slides or posters, and rehearsal of presentations. f. Provide methodology to migratory bird and wetland managers for sampling the biological parameters of interest. Literature review may be required to develop appropriate methodology. g. Assist with Canada goose trapping and banding in the upper Gunnison Basin. RESULTS The Wetland Initiative (WI), Wetland Focus Area Committees (FAC)' Joint Venture and Waterfowl Habitat Project Review Committee (WHPRC) Activities As members of the WI project planning and CDOW Wetland Program team, migratory bird researchers assisted in preparing the final grant application and developing project selection criteria; consulted on further development and refinement of wetland proposals; served on the committee to select project proposals for funding; and helped develop project development packages for some of the selected projects. Szymczak worked to facilitate delivery by Ducks Unlimited, Inc.(DU) of projects that were selected on State Wildlife Areas (SW A). Wetlands Program team members continued to communicate with all FACs in the state. Emphasis of the FACs during this reporting period was to facilitate the delivery of WI funded projects within the respective areas. ( �27 As chairman of the WHPRC, Szymczak chaired committee meetings for ranking and funding proposals submitted for the 1999-2000 and the 2000-01 funding years; informed proposal proponents of the outcome of their funding request; periodically monitored progress of project planning, construction, and money flow for new and previous years funded projects; coordinated Site Specific Agreements and fund reimbursement with the Ducks Unlimited Inc. MARSH program; served as the Project Officer for wetland development contracts formulated with Ducks Unlimited, the Bureau of Land Management, and the U. S. Fish and Wildlife Service. Flyway Technical Committee and Council Meetings In January 1999, Szymczak attended the winter meeting of the Pacific Flyway Study Committee (PFSC). Discussions at the winter meetings of importance to Colorado involved hunting season regulation packages, and developing an objective function and model sets for harvest management of western mallards. In March 1999, Gammonley attended the Central Flyway Webless Game Bird Technical Committee (CFWGBTC) and Central Flyway Waterfowl Technical Committee (CFWTC) meetings while Szymczak attended the PFSC meeting. Major items of direct relevance to Colorado included a review of mourning dove management strategies, review and approval of harvest allocation for the Rocky Mountain Population of greater sandhill cranes, review of the status of the pintail interim harvest strategy, compilation of the Four-Corners band-tailed pigeon harvest reports and recommendation for hunting seasons, discussions about the North American Bird Conservation Initiative, and review of Central and Pacific Flyway recommendations for duck harvest regulation packages under the Adaptive Harvest Management (AHM) approach. In April 1999, Gammonley attended the ARM working group meeting. A major subject of this meeting was to review analyses of the impacts on harvest resulting from changes to duck season opening and closing framework dates. The ARM working group .developed a position paper summarizing these analyses and providing recommendations, that was used by the flyway councils in subsequent discussions about frameworks. In July 1999, Gammonley attended the CFWTC and Central Flyway Council sessions while Szymczak attended the PFSC and Pacific Flyway Council meetings. The major agenda items at meetings in both flyways in July are the status of waterfowl populations, the characteristics of the most recent waterfowl hunting season harvest, and regulation proposals for the up-coming hunting season. The recommendations are forwarded through the respective Councils to the USFWS Regulation Committee. Subsequently, Gammonley and Szymczak made recommendations to CDOW regulations personnel on the structure of waterfowl hunting seasons in Colorado, and reviewed final selections for hunting season structure and regulations for migratory game birds. In December 1999, Gammonley attended the winter meeting of the CFWTC. The focus of this work session was the development of a Central Flyway strategy and guidelines for management of resident Canada goose populations. In January 2000, Sanders attended the winter meeting of the PFSC, for a work session on migratory game bird management plan revisions (pacific Population and Rocky Mountain Population of Canada geese, Four corners Population of band-tailed pigeons). Proposals for funding with the Web less Migratory Game Bird Research program were also reviewed and ranked. In March 2000, Sanders attended the PFSC meeting, while Gammonley attended the Central Flyway Webless Game Bird Technical Committee (CFWGBTC) and Central Flyway Waterfowl Technical Committee (CFWTC) meetings. Recommendations on regulations and other management issues were forwarded to the flyway Councils. �28 In May 2000, Gammonley attended the AHM working group meeting. Emphasis was on incorporating Eastern mallards into the existing ARM approach, and the potential to incorporate more information about waterfowl hunters (including measures of hunter satisfaction) into AHM objective functions. Cooperative Migratory Bird Studies The CDOW cooperates extensively with other migratory bird management agencies in the Central and Pacific flyways in conducting studies of migratory bird populations. In the Pacific Flyway, investigations are underway to: (1) define the western mallard population and establish monitoring programs to integrate the annual status of western mallards into the AHM regulatory process; (2) establish models for the continental northern pintail population that can be integrated into the AHM regulatory process; (3) document harvest distribution and harvest rates for mallards breeding in the Yukon Territory of Canada in conjunction with defining a western mallard population; and (4) monitor the annual status of the Rocky Mountain Population of greater sandhill cranes. In the Central Flyway we are providing support for (1) a large-scale mallard banding project in northern Central Flyway states; (2) banding studies of arctic-nesting snow geese, white-fronted geese, and Canada geese; and (3) studies of snow geese and habitats near Hudson Bay. In addition to funding, the CDOW provided equipment and transportation support for the mallard banding project. Gammonley assisted with an EIS public scoping meeting on alternative control methods for resident Canada geese, while Sanders assisted with the Pacific Flyway at the Coleman National Fish Hatchery in California in February, 2000. He also developed a computer program to back calculate hatching dates and to summarize wingbee data for band-tailed pigeons. Sanders also assisted with duck nest searches at Monte Vista National Wildlife Refuge. Wetland Resources and Development Szymczak visited existing and potential wetland sites and made recommendations for development andlor management, and served on the land acquisition wetland task force that evaluated possible acquisitions along the South Platte River from the standpoint of existing wetlands, water resources, potential wetland developments and recreational opportunities. Sanders and Gammonley provided suggestions and considerations for wetland developments, management, and monitoring programs on state, federal, and private lands. Gammonley assisted with development of Master Management Plans for the Mount Pleasant and Russell Lakes State Wildlife Areas, assisted Bureau of Land Management staff in duck nest searches at Hebron Wildlife Management Area, and suggested considerations for future wetland management at the site. Informational Programs Gammonley lectured on migratory game bird issues for an undergraduate wildlife course at Colorado State University, participated in a hunter education instructor workshop, completed a draft of a chapter on wildlife use of palustrine wetlands for a book "Wetland and Riparian Areas of the Intermountain West: Their Ecology and Management" being edited by staff at the University of Wyoming, and continued duties as associate editor of Wetlands, the journal of the Society of Wetland Scientists. Sanders gave a presentation on breeding habitat availability and mineral deposit use of band-tailed pigeons at the 3rd Annual Pacific Flyway Symposium. Gammonley, Sanders, and Szymczak gave presentations to CDOW trainees on waterfowl identification, wetlands conservation programs, and migratory game bird management. Canada Goose Management Activities No Canada geese were trapped and banded in the Gunnison Basin during this reporting period. �29 DISCUSSION Project personnel provide useful information in planning and evaluating waterfowl management and habitat enhancement programs in Colorado and educating land management agency personnel about the habitat requirements of waterfowl. With increasing emphasis on wetland habitat in Colorado, and the initiation of the Wetland Program within the CDOW and new funding sources such as the Wetland Initiative, wetlandrelated objectives of this job will receive more emphasis. Colorado now has 10 Wetland Focus Area Conunittees functioning in the state that will require coordination and expertise in wetland project planning. The resources provided by project personnel will insure that money raised through the Colorado Duck Stamp program or any other funding initiative will be spent in accordance with the objectives of the program. Continued participation on Pacific and Central Flyway conunittees ensures that Colorado will remain informed and involved with migratory bird issues, have input into migratory bird hunting regulations, and influence habitat programs affecting migratory game birds. Increasingly, studies on migratory birds that have flyway-wide implications are being funded cooperatively through the Flyway councils. Conducting and/or formulating surveys and banding efforts and informing management agency personnel about aspects of waterfowl and wetland ecology provides a valuable service to management agencies, the waterfowl resource and, in some cases, the hunting public. preparedbY:p*~ James H. Gammonley GPIV Todd Sanders GPIV ��31 .. -"'-" Colorado Division 'of Wildlife ,Wildlife Research Report , April 2000 JOB PROGRESS State of: REPORT, ---"'C"""o;.:..>~o::.:.rad=o::...._ _ W-166:'R Project: Work Plan~ '__ '....::2::=2,,-_ Job Title: Migratory Game Bird Investigations Job _~2==--_ ~M~i~g~rn=to~ry~G~arn~e~B~ir~d~P~u~b~lic~a=t~io~n~s _ Period Covered: 01 January 1999 through 30 June 2000 . Author: James H. Ganunonley Personnel: James H. Ganunonley, Todd Sanders, and Michael R. Szymczak, Colorado Division of Wildlife ABSTRACT ---. The following article was published: -Laubhan, M. K, and J. H. Ganunonley. 2000. Density and foraging habitat selection of waterbirds -',,breeding ill the San Luis Valley of Colorado. Journal of Wildlife Management 64:808-819. ��33 -- .Colorado Division of Wildlife Wildlife Research Report A . p"ril2000 -. JOB PROGRESS State of: - REPORT ----"C"""o:!.>~o""-ra".,d,."o'-_ Migratory Game Bird Investigations Work Plan: __ .:::.3~0 __ : Job-__ ~I _ Job Title: _--,S==<Jpl<-!nnc::' "-Cg"-'S~t""'o.l<.po><-v:..:e"'-r..:.F.>::ood=_"'R""'e""'so"_"u""r~ce~s~an=d_""L~an=d_"U""'s:>::.e_"'P-"'a=tt::::.: Sandhill Cranes in the San Luis Valley, Colorado PeriodCovered: 01 January 1999 through 30 June 2000 James R Gainmonley Author: Personnel: James H. Gammonley, Colorado Division of Wildlife, and Murray K. Laubhan, USGS ,Biological Resources Division ABSTRACT In, response to recent concerns over whether current or projected availability of waste grain and wetland habitats III the San Luis Valley (SL V) are adequate to meet the spring resource needs of Rocky Mountain Population (RMP) sandhill cranes (Grus canadensis tab ida), we initiated a study to determine food habits, body condition, and distributional patterns ofRMP sandhill cranes during their spring stopover in the SLY. We recorded the distribution of grain fields in the SLY during 1997, 1998, and 1999 in a GIS map. During 1999 and 2000, we collected waste grain samples from a total of20 fields that represented the range of post-harvest practices used on private grain fields in the SL V, and combined these results with datafr<?m 1,9fitlds sampled in 1998. Two of the sampled fields were damaged by hail and were not using normal practices. Estimates of waste grain biomass available on the soil surface of these _ fields_~"eraged252 and 332 kg/ha. _-In the remaining fields, the estimated mean biomass of waste grain was 5kgtha (95% CI 1.7-9.1 kg/ha, range for individual fields=0.1-57 kg/ha). We collected 40 sandhill cranes dlJring23 February~25 March 1999, and 20 cranes during 29 February-29 March 2000. We collected _ '¢r3rie~iti grain fields (n =23), potato fields (n = 5), and irrigated pastures and wetlands (n = 32). -~.-. Prelimi#ary analyses of esophageal contents confirm that waste grain is a major food item, but cranes also ---,. ci.>nsllll1edseveral taxa of invertebrates (beetles, spiders, snails), as well as stems, leaves, and roots of .: ".goosefoo(Cherzopodium); sedges (Carer), and various grasses. Our collections included 4 of the lesser .~subspecies (G. c: canadensis). Our collections included 4 of the lesser subspecies (G. c. canadensis). Of • ~ < tq~'ie01aiping greater sandhill cranes, males (n = 33) averaged 5.72 kg, and females (n = 23) averaged 5.17 " _.. _::>'kg.,'AIl'FUect&i'pirdswere plucked and sent to the University of Western Ontario for analysis of body .,',;<'<_nuiri~ntcc)rripc,~itiQn (lipid, protein, and mineral). Field will be completed in 2001 and laboratory work ·.,,~:>~Hbe_~mlll,etedby2002. - ruuveste<l .. -~..~.t;':'-i ::;: -f "',: ;;, :"'{'/.:':~ ·":',1 .;./;; ~.:>;'~" ,: - .:.'.- :.',' .. < •• :>~•'..;'.' .~:.,..; :>., ", .t.'~.:,:~,5.',"';:~}:·'! .-------</'?' .. "~"';'f; c. ~~:~:;:.:!~ :·.~.(~~;~:·~~i~~.::i2,.{j":.' ,.:::.::,..: · "';:'~'S'\',:,;i:: .. ,": .,r· ;' .,' .... "~' ..::..,'. .':. ,.:.',';,~ ";,'," ' . :>',: \' ��35 SPRING STOPOVER FOOD RESOURCES AND LAND USE PATTERNS OF ROCKY MOUNTAIN POPULATION SANDHJLL CRANES IN THE SAN LUIS VALLEY, COLORADO James H. Ganunonley INTRODUCTION Virtually the entire population Of Rocky Mountain Population (RMP) of greater sandhill cranes (Grus canadensis tabida) uses the San Luis Valley (SLY) of Colorado as a spring stopover area. RMP cranes in the SLV forage on unharvested grain provided on Monte Vista National Wildlife Refuge, and on waste grain in privately-owned fields. In recent years, however, fall tillage and irrigation of grain fields has become increasingly widespread in the SL V. These changes in farming practices have resulted in an unmeasured reduction in waste grain availability for RMP cranes during spring and have prompted concern over whether current or projected food supplies are adequate to meet spring nutritional demands of a target population size of 18,000-20,000. Changes have also occurred in the availability, distribution, and quality of wetland habitats in the SLY. Information on habitat use, food habits, and body condition ofRMP sandhill cranes is lacking. PROGRAM NARRATIVE OBJECTIVES 1. Determine food habits of RMP cranes on grain fields, pasture lands, and wetlands in the SL V during the spring stopover period. 2. Determine the body condition of RMP cranes in the SLV during the spring stopover period. 3. Determine the amount and distribution of waste grain available to RMP sandhill cranes during the spring stopover period. 4. Estimate the amounts of each major food item (particularly small grains) consumed by RMP cranes required to meet the cumulative energy requirements ofa population of 18,000-20,000 during the spring stopover period in the SLY, and compare these estimates to the estimates of abundance of each food item 1) during the study and 2) under projected trends in farming practices and available public land management practices. SEGMENT OBJECTIVES 1. Determine food habits of Rocky Mountain Population (RMP) sandhill cranes on grain fields, pasture lands, and wetlands in the San Luis Valley (SLY) during the spring stopover period. 2. Analyze the body condition ofRMP cranes in the SLY at arrival and immediately prior to departure in spnng. 3. Determine the amount and distribution of grains (barley, wheat) used by RMP cranes upon arrival and after departure of cranes in spring. 4. Estimate the amounts of each major food item (particularly cultivated grains) consumed by RMP cranes required to meet the cumulative energy requirements ofa population of 18,000-20,000 during the spring stopover period in the SLY. 5. Prepare annual reports. �36 RESULTS We used Natural Resource Conservation Service (NRCS) aerial photographs taken during 1997, 1998, and 1999, to identify crop type (grain or other) in 2,068 center-pivot agricultural fields throughout the SLY. The distribution of grain fields during each year was recorded in a Geographical Information Systems (GIS) map of the SLY. During 1999 and 2000, we collected waste grain samples from a total of 20 fields that represented the range of post-harvest practices used on private grain fields in the SLY. These results were combined with data from 19 fields sampled in 1998. Two of the sampled fields were damaged by hail and were not harvested using normal practices. Estimates of waste grain biomass available on the soil surface of these fields averaged 252 and 332 kg/ha, In the remaining fields, the estimated mean biomass of waste grain was 5 kg/ha (95% CI 1.7-9.1 kg/ha, range for individual fields=O.l-57 kg/ha). We also established a road survey route to monitor crane numbers and distribution. Based on weekly count totals, the survey route accounted for >90% of the RMP cranes present in the SLY. We collected timeactivity budget data on cranes in fields, unflooded pastures, and wetlands throughout the stopover period. Preliminary analysis indicates that cranes in fields and pastures spent >70% of diurnal hours foraging. We collected 40 sandhill cranes during 23 February-25 March 1999, and 20 cranes during 29 February-29 March 2000. We collected cranes in grain fields (n = 23), potato fields (n = 5), and irrigated pastures and wetlands (n = 32). Preliminary analyses of esophageal contents confirm that waste grain is a major food item, but cranes also consumed several taxa of invertebrates (beetles, spiders, snails), as well as stems, leaves, and roots of goosefoot (Chenopodium), sedges (Carer), and various grasses. Cranes also appear to readily forage on small potatoes that remain in fields following harvest. Given that >24,000 ha of potatoes have been planted in the SLY in recent years, potatoes could represent an abundant and previously unrecognized food source for migrant cranes. After removing the digestive tract contents, we determined the sex of each collected crane, and measured the culmen, tarsus, wing chord, body length, and body weight. These measurements were used to determine subspecies status of each crane. Our collections included 4 of the lesser subspecies (G. c. canadensisi. Of the remaining greater sandhill cranes, males (n = 33) averaged 5.72 kg, and females (n = 23) averaged 5.17 kg. All collected birds were plucked and sent to the University of Western Ontario for analysis of body nutrient composition (lipid, protein, and mineral). PLANS FOR 2000-2001 ~: ... We will review NRCS photographs taken during July 2000 and add the locations of grain fields to the GIS. In February and March 2001, we will continue field sampling, weekly surveys, time budgets, and collections. In October 2000 and 2001, we will conduct surveys of crane distribution for comparison with spring results. Field work is expected to be completed in fall 2001. Laboratory analysis of esophageal contents and body condition will be completed by 2002. �37 Colorado Division of Wildlife .. Wiidlife Research Report ,April 20.00, . ' .. ,- -';'-" FINAL REPORT ','"Colorado " Project...;.'',_'_....,.,..__ Work PlaIl: _,W..:...-_,1'-"6;.:::6--"-R~ ---=:;.3..;::..1--: "Job Title: ,~--, Job __ -'M~ig::.!ra-=t!:!:o:.:...ry,l......!::Gam=::::e__"B:::.!ic!.!rd::..In=..!.v_"" __ --=-I__ -"In,..,t"",egr=a=ted:>::...,;W:..:..=at~ec!.!rb::.:.ir:..::d,-,M=::an~ag=e:.:.:.m.:.:e:.:.:.nt::..,;S""tu=d.:.:ie:>< _ -, .: ," , Period Covered: 01 January 1999 'through 30 June 2000 ' ' , Autho~: iames'H. Gammonley , :Personn~l: James H. Gammonley, Colorado Division of Wildlife; Murray K Laubhan, USGS Biological .Resources Divi~,~ori ,_ , .. ; :".<'" _./" : ABSTRACT Progress during this period was centered on preparing previously reported data for publication. An interim finalreport was presented in the April 1999 progress report. A draft management plan for wetland habitats .at Russell Lakes State\Vildlife Area, presented in the April 1999 progress report, has been incorporated 'intotheColofado Division of Wildlife master management plan for the area. One major paper has been published from this study: Laubhan, M. K~ and J. H. Gammonley. 2000. Density and foraging habitat selection of waterbirds breeding in the San Luis Valley of Colorado. Journal of Wildlife Management 64:808-819. Threeadditional papers are being submitted for publication: Gammol1Jey,J. H" and M. K Laubhan. Patterns of food availability for breeding waterbirds in a Colorado ~etJ.aIld complex. Wetlands, , GammO:nley,i H.; and M. K Laubhan. Diet and foraging activity of breeding waterbirds in the San Luis " ':~.:Valley of Colorado. Journal of Wildlife Management. Laubhan; M. K; and J, H. Gammonley. ' Nesting ecology of breeding waterbirds in the San Luis Valley of , _, Colorado. Journal ofWildlife'M~ement. ��39 INTEGRATED WATERBIRD MANAGEMENT STUDIES James H. Ganunonley PROGRAM NARRATIVE OBJECTIVES 1. Map the location of wetland communities on Russell Lakes State Wildlife Area. 2. Document the hydrologic regime and water, soil and vegetation characteristics of each wetland type. 3. Identify the aquatic invertebrates associated with each wetland community, and document seasonal trends in invertebrate 'diversity, abundance and biomass. 4. Quantify the abundance, spatial and temporal use patterns, behaviors, and food habits of waterbirds in different wetland types. Relate the dynamics of endogenous lipid and protein reserves to food habits and breeding ecology. 5. Determine the seasonal wetland habitat requirements for waterbirds, and consolidate these needs into a conceptual design for an optimum wetland community. 6. Determine the water management protocol and wetland development guidelines needed to produce the optimum wetland community. Prepare a wetland development and water management plan for Russell Lakes State Wildlife Area. SEGMENT NARRATIVE OBJECTIVES 1. Determine food selection of waterbirds collected at Russell Lakes State Wildlife Area (RLSW A) by comparing food use to food availability for each species with age, reproductive status, molt intensity, size of nutrient reserves, and collection date as covariables. 2. Determine nest habitat selection and nest success for waterbirds at RLSW A in relation to habitat features (cover type, water depth, vegetation structure) and spatial attributes (e.g., distance to water, patch size) of nest locations. 3. Integrate study results into a wetland habitat management plan for RLSW A. 4. Prepare final report and publications. Prepared by: 9-<r/~ James H. Ganunonley, GP IV ��41 <; . Colorado Division of Wildlife Wildlife Research Report . April 2000 . JOB PROGRESS REPORT '. ~. . State of: =c=o~:_:o;.:_:ra=d=o'-- _ Project: -:-Work Plan: __ Job Title: .__ W~-1'-"6:..::!6~'-R~ ..:.3..:..1 __ _ Migratory Game Bird Investigations : Job _-,2=--_ ----'M=o:!!n~ito~n~· n~g"-'an=d!..:E~v~a:!;!;lu~a~t!!:io:!!n~o~f...!W~et~lan=d~D~e:::.!v.!::el!!::o:!:'p.>.!.m!!::e~nt~P!..:r'-"o~ _ Period Covered: 0 1 January 1999 through 30 June 2000 Author: James H. Gammonlev Personnel: JamesH. Gammonley and Alex Chappell, Colorado Division of Wildlife, and Matthew A. Reddy, Duck Unliinited Inc. ABSTRACT We improved and updated a database of wetland development projects in Colorado that were funded by Colorado Division of Wildlife (CDOW), Ducks Unlimited Inc. (DU), and/or the U.S. Fish and Wildlife Service Partners for Fish and Wildlife (PFW) program. We collected additional field data on 15 projects during 1999. We also began developing a companion database containing information on habitats used by ...wetland birds in Colorado. These databases will be completed during 2000-2001. We also compiled . information on historic hydrologic patterns (i.e., river flows), climatic patterns, soil types, and surrounding land use for 60 projects and for each wetland Focus Area. We developed a draft planning process for Setting biological objectives for wetland conservation at Focus Area and individual project scales. During 200()-OI, we will work with Focus Area Committees to develop biological objectives, and recommend m~>nitoringefforts needed to evaluate the su~ of conservation efforts in.relation to these objectives. -- .. $2..~ .......... 7.~G_;: Prepared by: .~ ·115"111'11 BDOW013558 ��43 Colorado Division of Wildlife •...Wildlife Research Report . April 2000 JOB PROGRESS State of Colorado, Pr~~: ~W~-~16~7~-R~ Work Plan: ---,,----,,,-1 __ : Job REPORT _ Avian Research =24-=--__ Job Title: .. ' Evaluation of Habitat Development for Ring-necked Pheasants in Eastern Colorado " Period Covered: 01 January 1999 through 30 June 2000 Auilior: Th~o~m~~~E~.~R~e~nu~·~n~ro~o~n _ ABSTRACT .There were no research results reported during this Segment. A Final Report will be written later. - , .. . ��45 ~---, Colorado Division of Wildlife Wildlife Research Report . April 2000 JOB PROGRESS REPORT State of: ~ _ _"C::..oo<.:;lo:::..:ra=do><-_ Project: -'W'""----"1'-=6...:...7....:-R..:__ _ Work Plan: Job Title: 8=--__ : Job __ -0::6 Avian Research _ :::.D:..=e..:..;ve~l::::Jopt'.!m~e::::!n~t...::o~f~a~C~o::!;n~se~rv~at~io::::.n!...:P~l~an~fo:::..:r~L~e~s~se::::.r~P~ e~-c~h~ic:::;k~en!.!!s~ _ Period Covered: 01 January 1999 through 30 June 2000 Author: Kenneth M. Giesen -~--~~~~~~~----- Personnel: Kenneth M. Giesen, Jeff Yost, Colorado Division of Wildlife ABSTRACT The Lesser prairie-chicken (Tympanuchus pallidicinctusi was petitioned for listing under the Federal Endangered Species Act in 1996. As a result, the Lesser Prairie-chicken Interstate Working Group _..~PCIWG} was formed in 1997 to address the conservation needs of this species, and several meetings were held in 1999 to complete a range-wide status assessment and conservation plan. Increased knowledge of population size and distribution, and development of habitat-based management plans to cooperatively manage sand sagebrush (Artemisia ./iiifoUa) rangelands on public and private lands were identified as priorities in Colorado for this species. A habitat management guide for landowners was completed and distributed .. Within Colorado intensive breeding surveys were conducted in 1999 resulting in 164 males, 31 females, and 31 unclassified grouse being counted on 21 active leks. This is a 30 percent decline compared .to1998 and a 50 percent decline since 1988. A cooperative agreement between several private landowners, . the Colonwo Division of Wildlife, U.~. Forest Service, U.S. Fish and Wildlife Service, and the Natural ..Resour~ Conservation Service to manage >23,000 acres for Lesser Prairie-chickens in southeast Colorado (BacaCounty}was terminated in 1999 at the landowners request. ��47 DEVELOPMENT OF A CONSERVATION PLAN FOR LESSER PRAIRIE-CHICKENS Kenneth M. Giesen INTRODUCTION The lesser prairie-chicken (Tympanuchus pallidicinctusi was petitioned for listing under the Endangered Species Act in 1996. In June 1998 the ruling by the U.S. Fish and Wildlife Service on the petition was "warranted but precluded". A multi-agency committee, the Lesser Prairie-chicken Interstate Working Group (LPCIWG) was established in 1996 to address causes for the declines in distribution and population size of this species. A review of pertinent literature suggested that the most promising approach to reversing the population declines was to restore and manage habitats used by this species. Priorities for each state included more intensive monitoring of breeding populations and working with land owners and land management agencies to manage or restore habitats, especially nesting and brood-rearing habitats, for the lesser prairie-chicken. P.N. OBJECTIVES The primary objective of this study is to monitor populations of lesser prairie-chickens in Colorado and work cooperatively with other agencies and landowners to develop a conservation plan for this species in Colorado. SEGMENT OBJECTIVES 1. Review literature on lesser prairie-chicken biology and habitat use. 2. Monitor breeding populations of lesser prairie-chickens in Baca, Prowers, and Kiowa counties in southeastern Colorado. 3. Cooperate with NRCS, U.S. Forest Service, the LPCIWG, CSU Extension Service, other agencies, and private landowners in developing and implementing conservation strategies to benefit the lesser prairiechicken. 4. Prepare annual progress report. METHODS CDOW continued to offer Cost-share on conservation practices to benefit Lesser Prairie-chickens. These practices included reseeding of cropland, planting of native mid- and tall grasses in Cropland Reserve Program fields, and modification of grazing practices using fencing and water development. Known active leks of lesser prairie-chickens were surveyed for occupancy and to obtain counts of males and females from late March through early May. Observers attempted to survey leks within 2 hours of sunrise when lek activity and numbers of birds were highest. �48 RESULTS AND DISCUSSION The annual breeding survey of Lesser Prairie-chickens in Baca, Prowers, and Kiowa counties resulted in 213 total birds being counted on 21 active leks. There were 93 birds counted on 12 leks in Baca County (including 70 males, 16 females), 100 total birds in Prowers County on 6 active leks (including 78 males and 13 females), and 20 birds on 3 leks in Kiowa County (including 16 males and 2 females). Access to leks in Cheyenne County was denied in 1999 where at least 2 active leks were counted in 1998. The number of leks and total birds counted is less than in 1998 (302 birds on 40 leks), and the populations is substantially lower from high counts in 1988 (448 birds on 35 leks). The Lesser Prairie-chicken Interstate Working Group and various subcommittees held several meetings and phone conferences to address issues concerning the decline in numbers and distribution of lesser prairiechickens and draft a regional conservation strategy. This conservation strategy (Mote et al. 1999) was completed and distributed to state wildlife departments and land management agencies in February. Within Colorado, a cooperative agreement involving habitat management and Lesser Prairie-chicken conservation on >23,000 acres of public and private lands was terminated by landowner request. Specific management activities outlined in the plan will continue without the formal agreement as funding becomes available. LITERATURE CITED Mote, K. D., R. D. Applegate, 1. A. Bailey, K. M. Giesen, R. Horton, and 1. L. Sheppard. 1999. Assessment and conservation strategy for the Lesser Prairie-chicken (Tympanuchus pallidicinctusy. PREPARED BY: Kenneth M. Giesen Wildlife Research �49 _- .. ...._, , ./ Colorado Division of Wildlife , Wildlife Research, Report April ZOOO JOB PROGRESS REPORT Smreof ~C~o~lo~m~d~o~ Project: W..:.:.....-..:..;16::..;7:_-R:.::...._---,. __ -:-- Work Plan: __ Job Title: __ _ Avian Research ~13::..__ : Job _---=-11=--_ --=E::..:v~a::.:lu:::a""ti~on:.o...:<o .••.. f_".C""o:::lu:!.:m::.:b<.!:ian=.o..S""h'_"a:::.' rp.l<.-...::ta=i..,_,le::.=d,-,G","r,-"o:.:::u""se"-,R=ei::.:n=-tr-""od=:u"-,c::.:;ti,,,,,o;o:. e"",s _..in..__ _ Western Colorado Period Covered:" 01 January 1999 through 30 June 2000 Author: Richard W. Hoffinan , Personnel: Richard W. Hoffman, Colorado Division of Wildlife; Jennifer Boisvert, University ofIdaho; and Michelle Lassige, Colorado State University. ABSTRACT The inaugural meeting of the Northwest Colorado Columbian Sharp-tailed Grouse (Tympanuchus phasianellus columbianus) Work Group was held in January 1999. Meetings have been held every month since then. The group has identified the issues potentially impacting sharp-tailed grouse in northwest -Colorado and developed objectives, goals, and conservation strategies to address the issues. Preparation of the conservation plan is underway with the first draft scheduled for completion by September 2000. Grouse Habitat Improvement Program funds were used to plant 3 shrub thickets and to collect and propagate native grass and forb seeds for future habitat restoration efforts in California Park. Ninety-nine new sharptailleks have been Iocated in Moffat and Routt counties since 1997. The number of active leks has increased from 77 to 133 and the average number of males per lek has increased from 11.9 to 19.3. Less than 10% of all known lek sites (n = 174) occur on public land. Seventy-four sharptails (30 males, 44 femalesjwere trapped on leks in CRP and mine-reclamation: lands and equipped with radio transmitters. Monitoring of these birds provided 680 macrohabitat locations. Cover types with the highest percentage of :"-use' included shrub steppe and mine reclamation. Microhabitat characteristics were measured at 141ek sites, 28 nest sites, arid 47 brood sites. Corresponding random sites were also measured for each lek, nest, and brood site. Females (2,899 ± 4,405 m) moved farther than males (408 ± 298 m) from the lek of ' capture. Hens nested on average < 2 km from the lek of capture. Mean clutch size was 10.1 eggs for first nests and 7.7 eggs for renests. Egg fertility was 95%. Hatch dates of initial nests ranged from 16 June to 8 July, Overall nesting success was 48%. At 7 weeks post-hatch, 64% of the successful hens still possessed a brood and 49% of the chicks were still alive. Between mid-June and late August, average brood size declined from9.7 to 4:4 chicks per hen. The mortality rate of radio equipped birds was 48% from 24 April ,to 31 August. Grouse breeding in CRP were less productive and suffered higher mortality than grouse '" ,breeding in mine reclamation. Data were collected to assess habitat characteristics at multiple scales sugoQuding lek .sites ~ using GIS and remote sensing techniques. These dam have not been analyzed to date. . .. . - .', . , , ��51 EVALUATION OF COLUMBIAN SHARP-TAILED GROUSE REINTRODUCTION OPPORTUNITIES IN WESTERN COLORADO Richard W. Hoffinan INTRODUCTION Use of common names and misidentification of blue grouse (Dendragapus obscurus) and sage grouse (Centrocercus urophasianus) by early explorers have made it difficult to ascertain the precise distribution of Columbian sharp-tailed grouse in Colorado (Rogers 1969, Giesen and Braun 1993). However, historical records suggest this subspecies may have occurred in at least 22 counties in western Colorado (Bailey and Niedrach 1965, Rogers 1969). Recent surveys indicate viable populations are restricted to Moffat, Routt, and Rio Blanco counties, with possible remnant populations in Mesa and Montrose counties (Giesen and Braun 1993). Similar reductions in the distribution of Columbian sharp-tailed grouse have occurred throughout western North America (Miller and Graul 1980). Factors responsible for the reduction in distribution include conversion of native rangeland to cropland, excessive grazing by livestock, vegetative succession due to fire suppression, herbicide treatments, mineral exploitation, and urban development (Meints et al. 1992, Giesen and Connelly 1993). These factors have had the most pronounced impact on nesting, brood rearing, and winter cover through loss of native grasses and deciduous shrubs (Giesen and Braun 1993). Cover types used by Columbian sharp-tailed grouse tend to be structurally and vegetatively diverse with an extensive deciduous shrub component (Meints et al. 1992, Giesen and Connelly 1993). In Colorado, Columbian sharp-tailed grouse occur in mountain shrub communities interspersed with grasslands, small aspen (Populus tremuloides) stands, and riparian zones (Giesen 1987). Serviceberry (Amelanchier spp.) is an essential element of these communities and usually grows in association with one or more of the following shrubs: Gambel oak (Quercus gambelii), common chokecherry (Frunus virginiana), snowberry (Symporicarpos spp.), and sagebrush (Artemisia spp.) (Giesen 1987). Wheat is the primary agricultural crop within the range of sharptails in western Colorado. Wheatfields may be used during late summer and fall after harvest. These fields are usually snow-covered and unavailable during winter. Much of what is known about Columbian sharp-tailed grouse in western Colorado has resulted from studies in the northwest portion of the state (Dargan et al. 1942, Rogers 1969, Giesen 1987). Little is known about sharp-tailed grouse in southwestern Colorado other than they once occurred there and may still exist in low densities on the north end of the Uncompahgre Plateau (Rogers 1969, Giesen 1985). It has been 10 years since the last intensive effort to conduct lek surveys for Columbian sharp-tailed grouse in western Colorado. Another intensive effort is needed because changes in land use practices have occurred since then including implementation.of the Conservation Reserve Program (CRP), additional mining and development activities; arid alteration of grazing practices. Perhaps the most important action in the last 10 years affecting the need for current population and distribution data has been the petition to list Columbian sharp-tailed grouse as "threatened" or "endangered" in the lower 48 conterminous United States pursuant to the Federal Endangered Species Act (Carlton 1995). This action is of special significance in Colorado because Idaho, Utah, and Colorado are the only states that allow hunting of Columbian sharp-tailed grouse and that still have adequate populations to provide transplant stock for future restoration programs. Opportunities for management of sharptails in western Colorado may be limited because much of the occupied habitat occurs on private lands. The most extensive areas of public lands within the historic distribution of Columbian sharp-tailed grouse are in southwest Colorado. The last confirmed sighting of sharptails on these lands was in 1985 (Giesen 1985). It is likely that any effort to restore sharptails in western Colorado will require a conunensurate effort to restore and protect habitat. Thus, before a reintroduction program can be implemented, the status distribution, and habitat relationships of sharptails in occupied habitats in northwest Colorado must be evaluated and management strategies formulated based on the outcome of the evaluation. �52 P. N. OBJECTIVES Objectives of this project are to (1) form a sharp-tailed grouse working group with broad citizen, community, and agency representation, and in cooperation with this group, prepare a conservation plan for Columbian sharp-tailed grouse in Colorado, (2) conduct intensive lek surveys of Columbian sharp-tailed grouse in northwest Colorado, (3) ascertain presence or absence of sharptails in historic range in southwest Colorado, (4) identify potential reintroduction sites within the historic range of Columbian sharp-tailed grouse, (5) evaluate existing habitat conditions on these sites and within currently occupied habitats, and (6) cooperate with other western states in preparing conservation strategies for Columbian sharp-tailed grouse. SEGMENT OBJECTIVES 1. Review literature pertinent to the objectives of this study. 2. Form working group and conduct regularly scheduled meetings to develop management strategies and prepare conservation plan. 3. Prepare conservation plan in collaboration with working group. 4. Prepare and monitor contracts for Columbian sharp-tailed grouse habitat improvement projects. 5. Conduct lek searches and lek counts in Moffat, Routt, and Rio Blanco counties. 6. Ascertain seasonal movements, survival, productivity, and habitat use by Columbian sharp-tailed grouse breeding in CRP and mine reclamation lands .. 7. Evaluate the potential for reintroduction of Columbian sharp-tailed grouse into previously occupied habitats using remote sensing and GIS. 8. Compile data, analyze results, and prepare progress report. METHODS, RESULTS AND DISCUSSION Segment Objective 1 - Literature on all aspects of the biology and ecology of sharp-tailed grouse was reviewed, including published and unpublished materials. Literature searches were conducted through Current Contents, Wildlife Worldwide, and the Fish and Wildlife Reference Service. Efforts were made to review draft and final conservation plans and strategies prepared by other states and to talk with the people involved in preparing these documents. Efforts also were made to review all documents pertaining to the petition to list the Columbian sharp-tailed grouse as threatened or endangered. Segment Objectives 2-3 - Three public informational meetings were held in northwest Colorado in 1998. The purpose of these meetings was to inform the public about the status, distribution, and biology of Columbian sharp-tailed grouse, introduce them to some of the issues related to management of this grouse, and form a working groupto develop a conservation plan. Thirty-six peopleattended.the meetings. . Everyone agreed we should move forward with preparation of a conservation plan.. There was general agreement we should form one working group that includes representatives from-all counties (Moffat, Routt, and Rio Blanco) within the current range of sharptails in northwest Colorado. A mailing list of 230 potential stakeholders was developed with assistance from local personnel from the Colorado Division of Wildlife (CDOW), U.S. Forest Service (USFS), Bureau of Land Management (BLM), and Natural Resource Conservation Service (NRCS). Everyone on the list was notified about the formation of the working group and invited to participate. The inaugural meeting was held in January 1999; since then, meetings have been held on the last Tuesday of every month. The group has completed the following tasks: developed a mission statement, established a population goal, identified the issues, formulated objectives, goals, and conservation actions, and developed an implementation schedule to address the issues. An outline of what should be included in the plan was approved by the work group. Preparation of the plan is currently underway with the first draft scheduled for completion by September 2000. �53 Segment Objective 4 - Limited funds were available for habitat improvement projects for this reporting period. Three shrub thickets were established on private lands in conjunction with the Wildlife Habitat Improvement Program administered by Natural Resource Conservation Service. The thickets were established immediately adjacent to CRP fields, encompassed about 0.2 ha, and included mostly serviceberry with some chokecherry, hawthorne, and skunkbush sumac. The thickets were fenced to protect the young plants from browsing by domestic and wild ungulates. Funds also were allocated for the collection and propagation of native grass and forb seed from California Park, one of the few areas where sharptails occur on public land. The plants propagated from this project will be used as a seed source for habitat restoration efforts in California Park. Segment Objective 5 - Traditionally, lek counts were designed to provide information on average number of males per lek and average number of birds per lek. These estimates, when collected consistently over long periods, were presumed to provide trends in population size. However, Kobriger (1975) concluded that lek counts have little value in measuring population size or documenting population trends of sharptailed grouse because of inconsistent lek attendance patterns within and among years. Beck and Braun (1980) likewise concluded that high variation in attendance patterns by males at leks seriously limits the utility of lek counts as a population index for sage grouse. Cannon and Knopf (1981) recommended replacing lek counts with lek surveys (i.e., number of active leks) as a trend index to prairie grouse populations. Their recommendation was based on the observation that when populations increase, males respond by forming more leks instead of increasing the average number of males on each lek. These findings suggest that lek surveys should include two components: (1) surveys of known lek sites to ascertain status (active or inactive), and (2) searches for new leks. If lek counts are conducted, the results should be interpreted with caution (Rippin and Boag 1974, Emmons and Braun 1984). Lek surveys were conducted between 27 March and 2 June and consisted of the following: (1) surveys of known lek sites to ascertain status (active or inactive), (2) searches for new leks, and (3) counts of the total birds per lek, and if possible, the number of males per lek. Accurate counts were not always possible. Birds were frequently obscured by vegetation or there was no vantage point from which to observe the entire lek. In such cases, a flush count was obtained. Also, due to the vast area that needed to be searched and the large number of leks that needed to be surveyed, most leks were checked only once. Table 1 summarizes all surveys and counts conducted since 1997. It contains the most current information on the location and status of all known sharp-tailed grouse leks in northwest Colorado and should be used as the basis for future surveys. The database includes 174 known lek sites of which 128 (101 active, 27 inactive) are in Routt County and 46 (31 active, 15 inactive) are in Moffat County. No leks have been found in Rio Blanco County, even though sharptails are known to occur there. Ninety-nine new leks were located from 1997 to .2000. During this period, the. number of active leks increased from 77 to 133 and the average numberofmales per.lek increased from 11.9 to 19,.3,. Of 77 leks counted for 3 or more years, 60 (78%) showed an increase in the average number of males per lek, 10 (13%) showed a decrease, and 5 (7%) remained stable. Only 16 of the 174 known lek sites were on public land. Seven were inactive and 11 were on State Land '.Board property with little or no access to the general public. Based on the classification of 158 lek sites, the distribution ofleks by habitat type was as follows:26% sagebrush, 25% CRP, 19% hay/pasture, 16% mine reclamation, 8% native grass/forb, 3% alfalfa, 1% wheat. 1% mountain shrub, and 1% mine spoil. �. ble 1. Columb' . -- .. - h - -.. --~. Lek Name lek . _. 2 Type' • Status tailed_. - - -- - - County - ,- - Routt K Annan's Twenty Mile 1 Routt Annan's Twenty Mile 2 Routt Annan's Twenty Mile 3 Routt Baker's Peak t rth t Colorado. 1997-2000 Land status" Status Status Status Count Count Count Count 1998 1999 2000 1997 1998 1999 2000 8 12 16 Hay/pasture private 1997 80 Road dlek _'.1 - - Habitat Type A A A A H A A A A H A A A A A A A A NC Barnes Moffat Routt K K K I I I I I I Big Elk 1 Routt N A* A A 18 26 30 CRP private Big Elk 2 Routt ' N A* A A 8 5 27 CRP private Bloomquist Routt N A* A A A 11 7 20 Hay/pasture private Buck Mountain 1 Moffat K A A A A 14 15 33 Sagebrush Buck Mountain 2 Moffat N A* A 8 3 25 18 11 22 Sagebrush private Sagebrush private 19 52 54 Native grass/forb private 16 18 Sagebrush private Sagebrush public (USFS) Sagebrush public (SLB) Sagebrush public (USFS) Unknown private A 16 9 18 10 14 CRP private 22 38 26 Sagebrush private 14 23 14 CRP private Unknown public (SLB) CRP private Buck Mountain 3 Moffat N A* A A A Burn Moffat N A* A A Calf Creek California Park 1 Routt N NC A A Routt K A I I 2 California Park 2 Routt N A* A A 10 11 California Park 3 California Park Road 1 Routt Routt N A* I 4 5 H NC NC A NC NC California Park Road 2 Routt K NC NC NC A A* A I I A A* NC 12 12 California Park Road 3 Routt N Cedar Hill Gulch Moffat K Cole Gulch 1 Moffat N A* A I 4 Cole Gulch 2 Colowyo Reclamation Moffat A* A A 13 Moffat N N County Airport Routt K Cull Reservoir Moffat N Davis Moffat N Davis 2 Moffat N Deadman Deep Creek Routt N Routt N A* A A A Dinwiddie Routt K NC I A A Dresher Reservoir Moffat Routt N Dry Creek N A* Dry Elkhead Ridge Routt Dry Fork Elkhead Routt H H Dry Fork Elkhead 2 Routt N NC 21 A* I I I I A* A A A A* 8 public (SLB) 12 Sagebrush private 18 Sagebrush private Native grass/forb private CRP private CRP private Mine reclamation private Native grass/forb private private private 33 6 18 . 22 16 Hay/pasture 12 13 CRP A* 4 CRP A* 24 10 Sagebrush Sagebrush 13 Alfalfa private 35 CRP (retired) Hay/pasture private private 6 A* I I A A A 9 A NC A A 4 A A A A A* private I private private 2 17 14 8 16 CRP private 30 29 35 Hay/pasture private 23 Sagebrush public (SLB) ..,. V1 I I �.. Dry Fork Elkhead 3 Routt N Dry Gulch 2 Routt H Dry Gulch 3 Routt K Dunckley Park Routt N Earle Routt N Eckman Park 1 Routt N Eckman Park 2 Routt N Eckman Park 2A Routt N - A* I I I I I I 9 Sagebrush Private NC Unknown private NC Unknown public (SLB) 20 Hay/pasture private A* A* A A A* A A A A* A A A A* A 12 18 32 CRP private 17 19 29 30 Mine reclamation private 18 17 51 59 Mine reclamation private 4 11 Mine reclamation - Eckman Park 3 Routt N A* A A A 17 11 13 6 Mine reclamation private Eckman Park 4 Routt N A* A A A 10 8 11 8 Mine reclamation private Eckman Park 5 Routt N A* NC A A 10 19 16 Mine reclamation private Eckman Park 6 Routt N A* A A A 11 18 21 Mine reclamation private Eckman Park 7 Routt N A* A 29 40 Mine reclamation private Eckman Park 8 Edna Mine 1 Routt N Routt N A A* A 14 30 24 Mine reclamation Mine reclamation private private Edna Mine 2 Routt N A* A 5 2 Mine reclamation private Edna Mine 3 Routt N A* A 18 22 Mine reclamation private I Elk Creek 1 Routt N A* A A A 12 9 13 Mine spoil private J Elkhead Routt N A* NC A A 10 7 Native grass/forb private Elkhead Road 1 Moffat H H I I I I private Moffat I I unknown Elkhead Road 2 I I unknown private Elkhead Road 3A Routt H A A A A Elkhead Road 3B Elkhead Road 4 Routt N A* A Moffat K I I Elk Mountain 1 Routt H I I NC NC A A A A A A 6 16 A A A A 35 31 Elk Mountain 2 Elk Mountain 3 Routt H RO'utt .. H . Roytt H. A* I I I I Energy Fuels Routt K NC A A Finger Rock Routt N A* NC NC I I Fish Creek Routt N A* A A Five Pines Mesa Routt K NC I I Elk River Cemetery Moffat K NC I NC I I I I George's Gulch Routt H A A A A Gillilands Routt H I Gnat Hill Routt N A* Green Acres Routt K I I I I I I I Fly Creek Moffat K NC A A Foidel Creek Routt H A NC I Fortification Rocks 13 9 9 8 private 7 5 CRP private unknown private 2 16 Hay/pasture private 15 22 Hay/pasture private 39 Hay/pasture private Native grass/forb private Sagebrush private private I Native grass/forb Sagebrush private i 8 8 10 11 , Sagebrush private CRP (retired) private Sagebrush public (BLM) private NC unknown private I I CRP private Sagebrush private 18 14 I I ! ! private Alfalfa 22 I CRP Sagebrush 24 I 14 3 12 ; 10 4 15 ; ! u, u, �Hayden Divide Routt H , A A A A 25 28 26 Mine reclamation private 17 14 17 CRP private 7 Sagebrush private Heidel Routt K A A A A Hicks Routt K NC NC NC A Hightail Routt N A* A A A 13 8 8 9 CRP (retired) private Hillberry Routt N A* A A A 10 22 20 31 Sagebrush private Hinkle Routt K A A A A 5 6 7 8 CRP (retired) private Hocket Routt K A A A A 8 18 22 Hay/pasture private Hoffman Routt N A* A A A 11 18 26 12 Hay/pasture private ,Homestead Routt N A* A A A 6 14 12 CRP (retired) private Routt N A* A A A 7 5 2 Mine reclamation private Horton Knoll 1 Routt K A A A A 11 25 Sagebrush private lies Dome Moffat H NC A A A 3 18 CRP lies Mountain Moffat N 25 Sagebrush Jacks Routt Jubb Moffat N N Little Buck 1 Moffat K A Little Buck 2 Moffat A Little Hunter Routt K N long Gulch Moffat Maneotis Mcinturf Mesa Homestead Ditch 7 10 A* private . private A* A 5 17 Wheat private A* A 9 9 Native grasslforb private A A A 15 19 7 Sagebrush private A A A 7 17 13 private A* NC I I Sagebrush Sagebrush K A A A A 2 11 16 Hay/pasture private Routt K A A A Hay/pasture private Moffat K NC I I I unknown public (BlM) McKinney Ranch Routt H A A A A MCR18 Middle Creek Moffat N A* A A Routt N A A Miller Routt N Milner Morapas Gas Field Routt K I Moffat H Morgan Moffat Morgan Creek Reservoir Routt Morning Crow 15 7 I 6 6 Hay/pasture private 5 8 CRP private A 7 36 Mine reclamation private A* A 3 Sagebrush private I NC I unknown private NC A A A 21 37 33 CRP private N A* A A A 44 49 53 Native grasslforb private A A 40 Sagebrush I A A 35 A* A A 30 Routt K N 4 6 CRP private private Mud Springs Routt H A A A A 20 33 CRP private Nolands Nolands 2 Moffat NC A A A A* 4 7 CRP Moffat K N private private North Giant Routt N A* A A A 31 CRP private Pelleys Moffat H NC I I I Hay/pasture private A A A* 7 private 4 7 19 17 Pilot Knob Routt N Pinnacle Mountain 1 Moffat N A* Pinnacle Mountain 2 Moffat N A* NC I 9 Pinnacle Mountain 3 Moffat N A* A A 4 6 13 A* 9 Hay/pasture 22 Hay/pasture private 13 18 Sagebrush private unknown private 16 13 Alfalfa private , Vl 0\ �Pondella Ranch Moffat N A* , A A 10 17 37 Hay/pasture private private private Postovit Routt N A* A A 10 10 16 CRP RCR 33b Routt N A* A A A 44 38 48 51 CRP 10 10 13 14 Rick's Routt N A* A A A Robinson Routt I Routt NC I I NC Rock Creek 1 K' H NC I I Rock Creek 2 Routt K A A A A Rogers Routt N A* I A A 15 Saddle Mountain 1 Routt N A* NC 5 I I I I A 12 9 CRP (retired) private Alfalfa private unknown private 21 Sagebrush private 15 CRP (retired) private Hay/pasture private unknown public (SLB) Mountain shrub private Hay/pasture private NC Sage Creek Routt H Salt Creek Routt N Schneiders Moffat H NC I I A* NC Seneca Mine 1 Routt K A A A A 15 18 20 Mine reclamation. public (SLB) Seneca Mine 2 Routt N A* A 12 21 21 Mine reclamation Sherrod-Sadelin Routt K I unknown Routt N A* I I public (SLB) private Shivers I I A NC Six Plus Routt N Slater Park 1 Routt K NC A A A 7 Smiths Routt H A A A A Smuin Gulch Gravel Pit Routt N A* A A A Smuin Gulch Oil Well 1 Routt N A* A A A Smuin Gulch Oil Well 2 Routt N A* A Soash Stokes Gulch 1 Routt Routt K A N A* A A Stokes Gulch 2 Routt N Straight Gulch Moffat N Taylors Moffat H Trapper Routt N Trapper Mine 1 Moffat H Trapper Mine 2 Moffat N Turner Creek Routt N Twentymile 1 Routt K Twentymile 2 Routt Twentymile 3 I 20 6 CRP private 14 Sagebrush private 7 7 Sagebrush public (USFS) 11 35 29 Hay/pasture private 14 15 12 16 CRP private 10 10 15 23 CRP private A 10 5 16 CRP private A A 6 7 9 Hay/pasture private A A 21 27 18 CRP private A* A 25 19 CRP private A* A A 9 23 Sagebrush private I NC NC unknown private A* A A A A A* A A A* A A A A A A N Routt Twentymile 4 Twentymile Twentymile A* NC 14 6 5 Sagebrush private 23 27 31 Mine reclamation public (SLB) 6 6 8 Mine reclamation private 15 10 30 Sagebrush private A 9 7 Hay/pasture private A* A 12 12 Hay/pasture private N A* A 14 7 Hay/pasture private Routt N A* A 17 24 Hay/pasture public (SLB) 5 Routt N 25 Sagebrush private Cliffs 1 Routt N A* I I I 3 Mine reclamation private Twentymile Cliffs 2 Routt N A* A A A 10 24 43 Mine reclamation private Twentymile Routt N A* A I A 4 3 7 Mine reclamation private Cliffs 3 NC 14 A* 26 ------ - V1 -...j �Twentymile Cliffs 4 Routt K Twentymile Cliffs 5 Routt Twentymile Cliffs 6 Routt N N Villards Moffat K Villards 2 Moffat N Warrick Pasture Routt N A* A NC A Wilderness Moffat K A A A A 5 Routt N A* A A 7 Wilson Moffat K A A 24 Wilson 2 Moffat N. Windemere Routt N Moffat H Wolf Mountain Ranch Routt N NC . A* Woods Routt Wymans 1 Routt K K A NC Wymans 2 Routt N Yampa Yellowjacket Routt N Road Routt H Yellowjacket 1 Routt H I I I I Routt K A NC A A 8 Routt N A* 10 Ranch William's Park Wiseman's 1 Yellowjacket 2 Yoast Mine Road Summary Total Established Leks New Leks Located Total Leks Total Leks Surveyed Total Active Leks Total Leks Counted Total Males Counted Average malesllek K = known A ' A* A A A 8 9 A A I 20 14 19 A* I A I 26 I A " A A I I NC NC A A A A A A A A A* A A A .A ---- 1997 1998 1999 2000 75 39 114 91 77 44 524 11.9 114 27, 141 125 94 86 1107 12,9 141 15 156 146 114 103 1646 16 156 18 174 165 133 127 2454 19.3 lek found prior to 1997; H = historic private Mine reclamation private Vl 00 private private 12 10 Sagebrush public (SLB) 27 50 Sagebrush private 21 27 Native grass/forb private 9 Native grasslforb private 3 CRP private 3 3 private unknown private . 10 18 Sagebrush 19 28 Hay/pasture CRP private private CRP private Sagebrush private Native grass/forb private Hay/pasture private Native grasslforb private Alfalfa private 17 6 20 26 5 3 5 4 7 I ! Hay/pasture Sagebrush A* A private Mine reclamation Hay/pasture A* A* Mine reclamation 12 A* A 34 ------ private ------ lek reported in Rogers, G,E, 1969. The sharp-tailed grouse in Colorado. Colo. Div, Game Fish, and Parks Tech, Pub!. 23,; = = N new lek found since 1997, 2 A active (* denotes the year the lek was first located for new leks); I inactive; NC not checked, 3 SLB State land Board; USFS United States Forest Service; BLM Bureau of Land Management = = = = = -.~. �59 Segment Objective 6 - Lek surveys conducted in northwestern Colorado since 1997 indicate that sharptailed grouse actively use CRP and post-act mine reclamation lands for breeding. Leks on these lands constituted about 42% of all known lek sites. Although CRP and mine reclamation lands appear to be important components of sharptail habitats, little is known about the use of these lands beyond the lekking period. Recent studies have associated sharp-tailed grouse use with CRP lands (Sirotnak et al. 1991, Ulliman 1995, Apa 1998, McDonald 1998), but none of these studies specifically examined how sharptailed grouse use CRP. There has been no research addressing the use of mine reclamation lands by Columbian sharp-tailed grouse, yet all reclaimed mine properties surveyed in northwest Colorado from 1997 to 2000 contained at least 1 sharp-tailed grouse lek. A study plan entitled "Ecology of Columbian Sharp-tailed Grouse Breeding in Conservation Reserve and Post-Act Mine Reclamation Lands" was prepared and approved in 1998. Data collection began in April 1999. The objectives of this study were to (1) determine year-round seasonal macrohabitat use, seasonal movements, home range sizes, and survival of Columbian sharp-tailed grouse breeding in CRP and post-act coal mine reclamation lands, (2) identify and describe microhabitat characteristics oflek, nest, and brood sites of Columbian sharp-tailed grouse breeding in CRP and post-act coal mine reclamation lands, and (3) determine reproductive success and productivity of Columbian sharp-tailed grouse hens breeding in CRP and post-act coal mine reclamation lands. Methods Birds were trapped from 9 leks in mine reclamation and 5 leks in CRP using walk-in funnel traps (Schroeder and Braun 1991). Captured birds were aged and sexed based on plumage characteristics (Ammann 1944, Henderson et al. 1967), weighed to the nearest gram with an electronic scale, and banded with a serially numbered aluminum leg band (size 12). All hens and a select number of males were equipped with a necklace mounted radio transmitter (Holohil System, model RI-2B) weighing 12-14 g, which amounts to less than 3% of the average female body weight. Monitoring of the radio-marked birds began in April. When a radio-equipped bird was located, it was encircled at approximately 20 m and 3 GPS coordinate readings were obtained. The more precise location was determined by triangulation. The following information and macrohabitat variables were recorded at each location: time, date, weather conditions, slope, aspect, elevation, cover type, nearest cover type, distance to nearest cover type, nearest road, road type, distance to nearest lek, and distance to lek of capture. Birds, other than females suspected of nesting, that were consecutively located in the same spot were presumed to be dead and an effort was made to find the carcass and retrieve the radio. The site and carcass were examined to ascertain the cause of mortality. Macrohabitat variables of the site were recorded only if it was determined that the site was the actual location where the bird was killed. During the nesting period, any female found at the same site on consecutive locations was presumed to be nesting. The suspected nest sire was encircled at a 5':'10-m radius to morepreciselylocate the nest without disturbing the bird. The nest was marked at > 10-m by attaching flagging to a conspicuous object and recording the distance and compass direction to the nest. Microhabitat characteristics were measured at all study area leks except Twentymile Cliffs #5, because of difficulties in determining its actual location. Microhabitat measurements also were obtained at paired random locations for each lek. The random site was selected by computer .•generated UTM coordinates within the study area. The site had to meet the following criteria: ~ 0.5 Ian from an active structure, in a suitable cover type, and on < 5 % slope. The plots were 20- m in radius and centered on the UTM coordinates of the random sites. For the actuallek sites, the plot center was located a random number of steps from 1-5 in a random compass direction from the center of male activity, which was identified by observing birds on the leks. Transects were extended in the 4 cardinal directions from plot center and 13 ground measurements were obtained using a sampling quadrat (Daubenmire 1959) and modified cover classes at the center, 5 -, 10-, and 20-m. Thirteen measurements of vegetation height were taken and a cover pole (Griffith and Youtie 1988) was used to acquire 26 vertical cover measurements. The line- �60 cover pole (Griffith and Youtie 1988) was used to acquire 26 vertical cover measurements. The lineintercept method (Canfield 1941) was used to measure shrub canopy cover along each transect. Measurements at the lek and corresponding random site were taken within 2 days of each other to control for phenological differences in vegetation. Microhabitat characteristics were measured at all nest sites. Paired random sites were selected by producing computer-generated random numbers corresponding to UTM coordinates within 2 km of the hen's lek of capture. The 20-m transects radiating in the 4 cardinal directions were centered in the nest bowl or on the random coordinates. Measurements were obtained at the center, 5, 10, and 20 m along each transect for a total of 16 ground cover, 32 vertical cover, and 16 vegetation height microhabitat measurements. Canopy cover along each transect was estimated using the line-intercept method. A modified, 12 x 12 in cover board (Jones 1968) placed over the nest bowl or random plot center was used to measure horizontal cover. The nest sites and associated random sites were assessed within 2 days of each other and as soon after the conclusion of nesting as possible. For brood sites, microhabitat characteristics were assessed at every fourth location for each brood hen. The site was flagged and measurements recorded within 2 days after locating the brood to allow time for the brood to move from the site. Random sites were selected by moving a random distance between 50 and 500 m in a random compass direction from the actual brood site. Measurements were identical to those recorded for nest sites. Brood and random sites were measured on the same day. Nest sites were visited after departure of the hen to determine clutch size, number of fertile eggs, and nest success. Also, if the hen was away from the nest feeding, an effort was made to find the nest and count the eggs. If the hen was killed or abandoned the suspected nest, the site was searched for any obvious and/or strong anecdotal evidence that the hen had attempted to nest. A nest was considered successful if at least 1 egg hatched. A hen was classified as successful if she hatched at least one egg regardless of the number of nesting attempts. Broods were monitored throughout the summer. Chicks were counted whenever possible. However, such opportunities were infrequent due to the secretive behavior of chicks and their tendency to hide rather than fly when young. Thus, broods were deliberately flushed at 7 weeks of age to count the chicks. A brood was classified as successful if at least 1 chick survived to 7 weeks post-hatch. Productivity was calculated as the percent of chicks surviving to 7 weeks. Results and Discussion One hundred and seventy birds (147 original captures and 23 recaptures) were captured on 13 different leks. There were 5 mortalities (4 male, 1 unknown) and 1 serious injury (female with a dislocated shoulder) sustained by birds during the trapping effort. This amounted to 3.5% of all birds handled. Serially-numbered leg bands were placed on 146 birds and radio transmitters were attached to 74 birds (30 males, 44 females). Nine radio-marked grouse (7 males, 2 females) died and 3 males lost their radios within 2 weeks of capture. In addition, the radios on 2 other males failed immediately after the birds were released. Useful information was obtained from 60 radio-marked grouse including 29 birds (9 male, 20 female) captured on CRP lands and 31 birds (9 male, 22 female) captured on mine reclamation lands. The mean weight of 46 hens was 696 g (range = 627- 819 g). Adult hens averaged 702 g (n = 34, range = 627- 819 g) and subadult hens averaged 680 g (n = 12, range = 640- 758 g). Mean weight for 93 males was 771 g (range = 687- 843 g). The mean for adults was 774 g (n = 82, range = 687- 843 g) and for subadults it was 752 g (n = 11, range = 724- 787 g). Mean weights of birds captured in this study were similar to weights recorded for Columbian sharp-tailed grouse in Wyoming and Idaho (Oedekoven 1985, Marks and Marks 1987) and were nearly identical to previous weights recorded in northwestern Colorado by Giesen (1992). Weights in Giesen's study averaged 677 g and 698 g for subadult and adult females and 741 g and 777 g for subadult and adult males, respectively. �61 Monitoring of radio-equipped birds from April to August 1999 resulted in 456 non-nestinglnon-brooding bird locations, 34 nest locations, and 190 brood locations for a total of 680 macrohabitat observations. Fourteen cover types were distinguished within the study area and birds were found in the following 10 types: Native grass/meadow Hay/pasture Shrub-steppe « 1 m) Mountain shrub (> 1 m) Mixed shrub CRP Retired CRP Mine reclamation (grass/forb) Mine reclamation (shrub/grass/forb) Deciduous forest(aspen) Cover types with the highest percentage of use by radio-equipped birds included shrub-steppe (primarily sagebrush) and mine reclamation lands. Shrub steppe was used most by non-nesting and non-brooding birds, while post-act mine reclamation cover was used most by nesting and brood-rearing hens (Table 2). Table 2. Cover type use (%) by radio-marked sharptails Habitat Type Native grass/meadow Hay/pasture Shrub-steppe Mountain shrub Mixed shrub CRP Retired CRP Mine reclamation (grass/forb) Mine reclamation (shrub/grass/forb) Deciduous forest Nest 2.9 5.9 35.3 0.0 0.0 5.9 2.9 38.2 5.9 2.9 Brood 3.7 l.6 ILl l.6 l.1 0.0 0.0 60.5 20.0 0.5 Other 2.4 0.7 32.7 3.3 4.8 7.5 9.2 23.0 16.2 0.2 Total 2.8 1.2 26.8 2.6 3.5 5.3 6.3 34.3 16.8 0;4 No locations were recorded in agricultural crops, pre-act mine reclamation lands (mine spoils), riparian areas, or coniferous forests . .Eventually the habitat data will be used to determine whether sharp-tailed grouse are using cover types in proportion to their availability within the study area. However, before this analysis can be completed, all the seasonal use data needs to be collected so the study area can be properly defined. Distances moved from the lek of capture during breeding, nesting, and brood-rearing seasons were significantly farther for females than males (P < 0.001). Females moved a mean distance of2899 ± 4405 m (SD) (n = 42, range 45 - 23,200 m), whereas males only moved a mean distance of 408 ± 298 m (SD) (n = 17, range 0 - 1330 m). Microhabitat characteristics were measured at 14 leks and 14 random locations between 11 - 28 May 1999, and at 28 nest sites and 28 random locations from 3 June to 22 July 1999. Microhabitat measurements also were obtained at 47 brood, 47 random, and 14 non-brooding bird sites (5 males, 9 females) from 19 June to 11 August 1999. An effort was made to identify all vegetation at plot sites. Trees, shrubs, and grasses were classified by species, whereas forbs were identified by genus, and species when possible. A total of 113 plant species was identified within the study area. Of these, 71 were found in native shrub steppe, 84 were found on mine reclamation lands, and only 18 species occurred within CRP. Twenty-nine initial nest sites and 3 renest sites of radio-marked birds were located in 1999. In addition, 4 nests from unmarked hens were found during the field season, of which 2 provided macrohabitat data and 1 microhabitat data. Initiation of nesting began 19 May and the final initial nest attempt was on 13 June 1999. Renest initiation occurred from 31 May to 12 June. Hatch dates of initial nests ranged from 16 June to 8 July. Eggs in 2 renests hatched on 26 June and 5 July (Figure 1). Incubation length averaged 26 days (n = 10). �62 4 rn Ol Ol Q) Em initial "'C .renests Q) - nests 3 x: o C'CS - s: s: 2 .~ .!!l rn Q) t: o ~ o+-_~ Date Figure 1. Peak hatch dates of Columbian sharp-tailed grouse nests in northwestern Colorado, 1999. Hatching dates documented in this study (mid-June to early July) were somewhat later than other documented hatch dates (range, late May to late June) (Hart et a1. 1950, Marks and Marks 1987, Giesen 1997). This was partially due to the cold, wet spring in 1999. However, the primary reason was the elevation. The study area ranges in elevation from 2100 - 2400 m and often maintains snow cover through mid-April. No other populations of Columbian sharp-tailed grouse have been studied at this high of elevation. Another possible impact of the higher elevation may be a longer incubation period (26 days). Previously reported incubation periods range from 21 to 24 days (Johnsgard 1983, UUiman et a1. 1998). Average distance moved by hens from the lek of capture to initial nest sites was 1.87 km (n = 29). Renest sites were 0.40 km (n = 3) from the leks where the hen was captured. Hens from CRP moved an average of3'.41 km (n = 11, range = 0.23-9.21 km) from the lek of capture to their initial nest site. Hens in mine reclamation only moved an average of 0.99 km (n = 18, range = 0.10-5.00 km), which was significantly less (P=0.056) than what hens moved in CRP. The longest movement was 23.2 km, which surpasses the previously reported longest movements of 20 km made by 2 Columbian sharp-tailed grouse hens in Idaho (Meints 1991). The shorter distances moved by males further supports the fidelity of male sharp-tailed grouse to lek sites, whereas females tend to venture farther from lek sites in search of suitable nesting habitat. The mean distance (1.87 km) moved by hens from the lek of capture to nest site was comparable with other research findings, which show that most hens nest within 2.0 km of the lek of capture (Oedekoven 1985, Marks and Marks 1987, Meints 1991, Giesen 1997). However, in this study, the distances moved varied depending on whether the hen was breeding in CRP or mine reclamation. The mean distance moved by hens breeding in CRP (3.41 km) was farther than most studies have documented, and the mean distance moved by hens breeding in mine reclamation lands (0.99 km) was shorter than average distances previously documented. This would suggest that most hens breeding in mine reclamation lands were able to locate nearby suitable nesting habitat, whereas CRP hens were not. Hens from both mine reclamation and CRP passed over, moved nearer, and sometimes attended other leks within the area, thus challenging past assumptions that hens nearest to a lek most likely breed at that location (Marshall and Jensen 1937, Parker 1970). The closer distance ofrenest attempts (0.40 km) to the lek of capture than initial nests (1.87 km) is contrary to that documented by Meints (1991) and Apa (1998) in Idaho. This may be due to the improvement in suitability of cover near the leks as the growing season progresses. �63 Average clutch size for 24 initial nests and 3 renests was 10.1 (range = 8 - 12) and 7.7 eggs (range = 7-8), respectively. The average clutch size of initial nests was slightly lower than clutch sizes documented elsewhere within the southern range of the Columbian sharp-tailed grouse; 10.8 in northwestern Colorado (Giesen 1987) and 10.9 (n = 127, range = 3-17) in Utah (Hart et al. 1950). Egg fertility for 17 nests was 95% (n = 160). The egg fertility rate of 4 hens breeding in CRP was 91 % (n =33) compared to 96% (n=127) for 13 hens nesting in mine reclamation. Egg fertility was within the range documented in other sharp-tailed grouse studies (86% - 99.3%) (Hamerstrom 1939, McDonald 1998). Nest success and hen success was 48% (n = 31and 33, respectively). Initial nesting attempts had a 46% success rate (n = 28) and 67% ofrenests were successful (n = 3). Nesting success was lower than rates reported in Utah (53-82%, n = 127), Colorado (62%, n = 13), and Idaho (72%, n = 25) (Hart et al. 1950, Giesen 1987, Meints 1991, respectively), but similar to rates reported from other studies in Idaho (51 %, n = 47; Apa 1998) and Washington (41%, n = 37; McDonald 1998). Hens nested in 7 different cover types identified within the study area. These. included deciduous forest (n = 1), native grass (n = 1), grazed pasture (n = 2), shrub-steppe (n = 12), CRP/retired CRP (n = 4), post-act mine reclamation (n = 10), and enhanced mine reclamation (seeded with shrubs and/or wild rye bunchgrasses) (n = 2). Seventy-four percent of all nest sites occurred within shrub-steppe and mine reclamation cover types. There were notable differences in nest success by cover type (fable 3). No successful nests were located in CRP or grazed pasture. Nests in mine reclamation were highly successful. Nests in native shrub steppe were mostly unsuccessful. This may be related to seasonal vegetation growth. The mean date of nest initiation in shrub-steppe cover was 24 May (n = 7), a week before the mean nest initiation date (1 June) for nests in mine reclamation (n = 12). Residual cover in shrub-steppe may be the most adequate, but not necessarily the best, nesting medium early in the season. Nesting cover provided by rapidly growing grasses and forbs on mine reclamation lands apparently surpasses the quality of shrub steppe as nesting habitat. Ta bl e3. Nest success by nest site cover type. Cover Type - First nest Deciduous forest Grazed pasture Shrub-steppe CRP Mine Reclamation (grass/forb) Mine Reclamation (shrub/grass/forb) Cover Type - Renest Mine Reclamation (grasslforb) 2 Success (%) 100 0 17 0 100 100 3 67 N 1 2 12 3 8 Average brood size at the beginning of the brood season was'9.7 chickslhen (n = 14). After 7 weeks, 64% of the hens still possessed a brood and the average number of chicks per hen was 4.4. Twelve hens raised their broods almost exclusively in mine reclamation lands, while 2 hens raised their broods in a combination of mine reclamation or native grass and shrub-steppe. The broods raised in mine reclamation lands averaged 4.7 chicks per hen. The 2 broods that used other habitat types in addition to mine reclamation averaged 2.5 chickslhen. The percentage of hens raising at least 1 chick to maturity was 67% (n = 12) for mine reclamation raised broods and 50% (n = 2) for broods raised in other cover types. The overall productivity (% of chicks surviving to 7 weeks) was 49%. The average brood size of 4.4 chickslhen was within the range (4.1 - 4.8) of brood sizes found in Utah and Idaho (Hart et al. 1950, Parker 1970, Marks and Marks 1987, Meints 1991). From 24 April to 31 August 1999, the mortality rate of radio -equipped birds was 48% (29 birds). Predation (44%) was the major cause of mortality. Mortality rates differed between females (39%) and males (69%), and between CRP (59%) and mine reclamation (29%) breeding habitats (Figure 2). �64 8 o Mne rec females o CRPfemales 7 (/) 6 fli;i Mne rec males CIl :E 5 t 4 ~ 0 ~ 0 'l:t: .CRPmaies 3 2 -_. 10 Figure 2. Weekly distribution of radio equipped grouse mortalities, Apr 16 - Aug 31, 1999. Although sharp-tailed grouse suffer high mortality during spring and summer (Schiller 1973, Marks and Marks 1987, McDonald 1998), the 48% mortality rate observed in this study from April through August seems excessively high. In Washington, a non-hunted population of grouse was reported to have a 47% annual mortality rate (Schroeder 1994), and in Colorado, a hunted population was documented as having 58% annual mortality (Giesen 1987). Over-winter survival can be quite variable, (29 - 86%) as documented by Ulliman (1995). Thus it is difficult to conclude whether the observed high seasonal mortality rate warrants concern about possible effects of the radio transmitters. However, mortality rates compared between the 2 breeding habitats suggest a cover type influence. The mortality rate for grouse breeding in CRP accounted for 65% of the total mortality between April and August. The preliminary results of this study indicate that Columbian sharp-tailed grouse in Colorado have found habitat changes like the introduction of CRP and mine reclamation lands attractive for breeding, nesting, and brood-rearing. However, their reproductive fitness and survival may not be equal within these cover types. Although Columbian sharp-tailed grouse are attracted to CRP for its grassland appearance, they likely suffer in terms of survival and reproductive performance due to lack of cover and vegetative diversity within CRP. Conversely, mine reclamation lands mimic or exceed the diversity and structure found in native·cover types and may benefit sharp-tailed grouse populations in northwest Colorado during-the breeding, nesting, and brood-rearing seasons. . . .~... Segment Objective 7 - A study plan entitled "Evaluation of Columbian Sharp-tailed Grouse Habitats in northwest Colorado Using Geographic Information Systems" was prepared and approved in 1999. Data collection began in spring 2000. No analyses have been completed to date. :The objective of this project is to assess habitat characteristics at multiple scales surrounding lek sites using GIS and remote sensing techniques. Data collected in this study will be used to locate and evaluate potential reintroduction sites elsewhere in Colorado. The analysis will involve 8 data layers compiled into a vector based GIS (ArcInfo, ESRI) and compared between lek sites and random sites. The base layer will consist of vegetation data created from a classified Landstat image taken 5 July 1989 (Friesen 1994). Additional layers will include roads, water features, CRP lands, mine reclamation lands, agricultural lands, land ownership (public versus private), and lek sites. ™ �65 The analysis area will consist of concentric circles surrounding the lek and random sites. The initial scale will be a 0.5-km radius around each site with each additional scale increasing by 0.5 km increments. Program Fragstats*Arc (pacific Meridian Resources) will be used to calculate the following variables within each analysis circle: patch size, sh ape, juxtaposition and composition; distance from the lek or random point to the nearest road, water feature, winter habitat, and nesting habitat; distance to nearest other lek; and nearest neighbor to winter and nesting habitats. The significance of each variable will be evaluated in a logistic regression model, increasing the scale until there are no differences in any variables between the lek and random sites. This approach should explain how grouse habitat requirements change across spatial scales (Morris 1987, Wiens 1986). LITERA TURE CITED Ammann, G.A. 1944. Determining the age of pinna ted and sharp-tailed grouse. Journal of Wildlife Management 8: 170-171. Apa, A.D. 1998. Habitat use and movements ofsympatric sage and Columbian sharp-tailed grouse in southeastern Idaho. Dissertation, University ofIdaho, Moscow, Idaho, USA. Bailey, A. M., and R J. Niedrach. 1965. Birds of Colorado, Vo1.ume 1. Denver Museum Natural History., Denver, CO. Beck, T. D. I., and C. E. Braun. The strutting ground count: variation, traditionalism, management needs. Proceedings Western Association Fish and Wildlife Agencies 60:558-566. Canfield, RH. 1941. Application of the line interception method in sampling range vegetation. Journal of Forestry 39:388-394. Cannon, R W., and F. L. Knopf. 1981. Lek numbers as a trend index to prairie grouse populations. Journal Wildlife Management 45:776-778. Carlton. J. C .. 1995. Petition for a rule to list the Columbian sharp-tailed grouse, Tympanuchus phasianellus columbianus, as "threatened" or "endangered" in the conterminous United States under the Endangered Species Act, 16 U.S.C. Sec. 1531 et seq. (1973) as amended. Biodiversity Legal Foundation, Boulder, CO. Dargan, L. M., H. R. Shepherd, and R N. Randall. 1942. Data on sharp-tailed grouse in Moffat and Routt counties. Colorado Game, Fish, and Parks Department Sage Grouse Survey, Volume 4, Denver. Daubenmire, R 1959. A canopy-coverage method of vegetational analysis. Northwest Science 33:43-64. Emmons, S. R., and C. E. Braun. 1984. Lek Attendance of male sage grouse. Journal Wildlife Management 48: 1023-1028. Giesen, K. M. 1985. Inventory of Columbian sharp-tailed grouse in western Colorado. Colorado Division Wildlife, Unpublished Report, Fort Collins. __ . 1987. Population characteristics and habitat use by Columbian sharp-tailed grouse in northwest Colorado. Final Report, Colorado Division of Wildlife, Federal Aid Project W-152-R, Denver, "~o:H)i-ado,USA. __ .. 1992.. Body mass of Columbian sharp-tailed grouse in Colorado. Prairie Naturalist 24:191-196. __ . 1997. Seasonal movements, home ranges, and habitat use by Columbian sharp-tailed grouse in Colorado. Colorado Division of Wildlife Special Report 72, Denver, Colorado, USA. _____:> and C. E. Braun. 1993. Status and distribution of Columbian sharp-tailed grouse in Colorado. Prairie Naturalist 25:237-242. _____:> and J. W. Connelly. 1993. Guidelines for management of Columbian sharp-tailed grouse habitats. Wildlife Society Bulletin 21:325-333. Griffith, B., and B.A. Youtie. 1988. Two devices for estimating foliage density and deer hiding cover. Wildlife Society Bulletin 16:206-210. Hamerstrom, F.N. Jr. 1939. A study of Wisconsin prairie chicken and sharp-tailed grouse. Wilson Bulletin 51:105-120. �66 Hart, C.M., O.S. Lee, and J.B. Low. 1950. The sharp-tailed grouse in Utah. Utah Department ofFish and Game Publ. 3, Salt Lake City, Utah, USA. Henderson, F.R., F.W. Brooks, RE. Wood, and R. B. Dahlgren. 1967. Sexing of prairie grouse by crown feather patterns. Journal of Wildlife Management 31:764-769. Jones, RE. 1968. A board to measure cover used by prairie grouse. Journal of Wildlife Management 32:29-31. Johnsgard, P.A. 1983. The grouse of the world. University of Nebraska Press, Lincoln, Nebraska, USA. Kobriger, G. D. 1975. Correlation of sharp-tailed grouse population parameters. North Dakota Outdoors 25(5):10-13. . Marks, lS., and V.A. Marks. 1987. Habitat selection by Columbian sharp-tailed grouse in west central Idaho. U.S. Department of the Interior, Bureau of Land Management, Boise, Idaho, USA. Marshall, W.H., and M.S. Jensen. 1937. Winter and spring studies of sharp-tailed grousein Utah. Journal of Wildlife Management 52:743-746. McDonald, M.W. 1998. Ecology ofColwnbian sharp-tailed grouse in eastern Washington. Thesis, University of Idaho, Moscow, Idaho, USA. Meints, D.R 1991. Seasonal movements, habitat use, and productivity ofColwnbian sharp-tailed grouse in southeastern Idaho. Thesis, University of Idaho, Moscow, Idaho, USA. Meints, D. R., J. W. Connelly, K P. Reese, A. R. Sands, and T. P. Hemker. 1992. Habitat suitability index procedure for Colwnbian sharp-tailed grouse. University Idaho Forest, Wildlife, and Range Experiment Station Bulletin 55. Miller, G.C., and W.D. Graul. 1980. Status of sharp-tailed grouse in North America. Pages 18-28 in P.A Vohs Jr. and F.L. Knopf, editors. Proceedings: Prairie Grouse Symposiwn. Oklahoma State University, Stillwater, Oklahoma, USA. Oedekoven, 0.0. 1985. Colwnbian sharp-tailed grouse population distribution and habitat use in southcentral Wyoming. Thesis, University of Wyoming, Laramie, Wyoming, USA. Parker, T.L. 1970. On the ecology of sharp-tailed grouse in southeastern Idaho. Thesis, Idaho State University, Pocatello, Idaho, USA. Rippin, A. B., and D. A. Boag. 1974. Recruitment to populations of male sharp-tailed grouse. Journal Wildlife Management 38:616-621. Rogers, G. E. 1969. The sharp-tailed grouse in Colorado. Colorado Division Game, Fish, and Parks Technical Publ.ication 23. . Schiller, RJ. 1973. Reproductive ecology of female sharp-tailed grouse (Pediocetes phasianellus) and its relation to early plant succession in northwestern Minnesota. Dissertation, University of Minnesota, St. Paul, Minnesota, USA. Schroeder, M.A. 1994. Productivity and habitat use of Columbian sharp-tailed grouse in north central Washington. Progress Report, Washington Department ofFish and Wildlife, Olympia, Washington, USA. ,. and C.E. Braun. 1991. Walk-in traps for capturing greater prairie chickens on leks. Journal of Ornithology 62:378-385. Sirotnak, J.M., KP. Reese, J. Connelly, and K'Radford. 1991. Characteristics of Conservation Reserve Program fields in southeastern Idaho associated with upland game bird and big game habitat use. Completion Report, Project W-160-R, Idaho Department ofFish and Game, Boise, Idaho, USA. Ulliman, M.J. 1995. Winter habitat ecology ofColwnbian sharp-tailed grouse in southeastern Idaho. Thesis, University of Idaho, Moscow, Idaho, USA. A. Sands, and T. Hemker. 1998. Idaho Columbian sharp-tailed grouse conservation plan (draft). Idaho Department of Fish and Game, Boise, Idaho, USA. __.J --.J Prepared by: �) 67 CDIDradODivision of Wildlife Wildlife. Research Report April 2000 . JOB PROGRESS State of Project: Colorado --'W'-'--'-1'-"6:...:..7-"-R~· _ Work Plan: _,--_..o::2!::.2__ JDbTitle: REPORT : JDb __ Avian Research ~1 __ ~A.:..:V1c.::·an=.:...;R""'e""'s<.:ea=r:..:c::.:h....:P_=u:.:::b.;:li.=.:ca:::t:::..::iD::.:.:ns=__ _ Period Covered: 01 January 1999 through 30 June 2000 Author: Th.:.=;:D=m=as=-:E=.:....:R=e::.:,m:.,::in=gt""'D::.:n _ Personnel: Clait E..Braun, Kenneth M. Giesen, Christian A. Hagen, Richard W. Hoffman, Sara J. OylerMcCance, CDIDradD Division of Wildlife ABSTRACT I .. ' The following articles were published: Commons, M.L. R.K. Baydack, and C.E. Braun. 1999. Sage grouse response to pinyon-juniper management. Pp 238-239 in S.B. Monsen and R Stevens, compilers. Proceedings: ecology and management of pinyon-juniper communities within the Interior West U.S. Dep. Agric., FDr. Servo RMRS-P-9. . Giesen, K.M. 1999. Columbian sharp-tailed grouse wing analysis: implications for management. CDID. Div. Wildl. Spec. Rep. ND. 74. 16 pp. _. and M.A. Schroeder. 1999. Population status and distribution of greater prairie chickens in CDIDradD. Pages 99-104 in W.D. Svedarsky, R.H. Hier, and N.J. Silvy, editors. The greater prairie chicken: a national look: Minnesota Agric. Exper. Sta., Misc. Publ. 99-1999, University of Minnesota, St. Paul, MN. Hagen, C.A. 1999. Sage grouse habitat use and seasonal movements in a naturally fragmented landscape, northwestern COIDradO. M.S. thesis, Univ. Manitoba, Winnipeg. 136 pp . . _. -. ... > _. RK. Baydack, and C.E. Braun. '1999. Habitat selection by sage grouse in a fragmented landscape: at which scale is preference defined? Proc. Prairie Grouse Tech. Counc. 23:Abstract. N.C. Kenkel, RL. Baydack, and C.E. Braun. 1999. Fractal-based spatial analysis of telemetry data. -Program Abstracts, Annu. Conf. The Wildlife Society 6: Ill . �68 Hoffman, R.W. 1999. Good News for grouse. Colorado Outdoors, MarchlApril2000: Johnson, K.H., and C.E. Braun. Conserv. BioI. 13:77-84. lO-13. 1999. Viability and conservation of an exploited sage grouse population. Kahn, N.W., C.E. Braun, 1.R. Young, S. Wood, D.R Mata, and T.W. Quinn. 1999. Molecular analysis of genetic variation among large- and small-bodied sage grouse using mitochondrial control-region sequences. Auk 116:819-824. Mote, K.D., RD. Applegate, J.A. Bailey, K.M. Giesen, R Horton, J.L. Sheppard, editors. 1999. Assessment and conservation strategy for the lesser prairie-chicken (Tympanuchus pallidicinctusy. Emporia, KS: Kansas Dept. of Wildlife and Parks. 51 pp. Oyler-McCance, SJ. 1999. Genetic and habitat factors underlying conservation strategies for Gunnison sage grouse. Ph.D. thesis, Colorado State Univ., Fort Collins. 162 pp. _, C.E. Braun, K.P. Burnham, and T.W. Quinn. 1999. Population genetics of Gunnison sage grouse in Colorado: implications for management. Program Abstracts, Arum. Conf. The Wildlife Society 6:160-161. __, N.W. Kahn, K.P. Burnham, C.E. Braun, and T.W. Quinn. 199. A population genetic comparison of large-and small-bodied sage grouse in Colorado using rnicrosatellite and mitochondrial DNA markers. Molecular Ecol. 8: 1457-1465. Schroeder, M.A., J.R. Young, and C.E. Braun. 1999. Sage grouse (Centrocercus urophasianus). In The Birds of North America, No. 425 (A. Poole and F. Gill, eds.). The Birds of North America, Inc., Philadelphia, PA 28pp. Svingen, D., and K.M. Giesen. 1999. Mountain Plover (Charadrius montanus) response to prescribed burns on the Comanche National Grasslands. 1. Colo. Field Ornithol. 33(4):208-212. i g/1/V1~ PREPARED BY:Jf~ Thomas E. Rerningto~/l Acting Avian Research Program Manager �69 <, Colorado Division of Wildlife Wildlife Research Report April 2000 - '- JOB PROGRESS State of: Colorado Project; ;..._ ...!.W,_-..,:.1.:::,.67.:....-..::.;R:::.__ _ Work Plan: __ REPORT Upland Bird Research . ....;2:::..:6::...__ __ : Job _----'I:::.__ __ Job Title: :....:An=al •.•. y""si"""s-"'o~f_"'U~p_"'lan=d_=B::..:i~rd"_'P=_o""'p::..:u=la=t=io=n:__T:...;r:..:::e=nd=s"-_ Period Covered: 01 January 1999 through 30 June 2000 Author: Thomas E. Remington Personnel: . Richard W. Hoffinan, Jennifer A. Nehring, Kim M. Potter, Thomas E. Remington, Colorado Division of Wildlife ABSTRACT .-The following reports were published: Hoffman, It. W. 1999. Columbian sharp-tailed grouse harvest data, northwest Colorado, 1976-99. 1999. Columbian sharp-tailed grouse lek surveys and lek counts for northwest Colorado. Unpubl. Rep., Fort Collins. 9 pp. 1999. Blue grouse Wing analyses in northwest Colorado for 1999. Nehring, J.A., and C.E. Braun. 2000. Gunnison sage grouse investigations, Poncha Pass Area, Colorado, April - December 1999. Unpubl. Rep., Colorado Div. of'Wildl., Fort Collins. 27 pp. Potter, KM., and C.E. Braun. 1999. Sage grouse investigations, Middle park, Colorado, April , December 1999. Unpubl. Rep., Colorado Div. ot-Wildl., Fort Collins. 27 pp. Remington, T.E. 1999. Sage grouse harvest report, North Park, 1999. 1999. Sage grouse harvest report, Gunnison Basin, 1999. 1999. Sage grouse harvest report, Lower Moffat and westemRoutt 1999. Sage grouse harvest report, Middle Park, 1999. j . {(:::9+- PREP..AMD B,(~ --' ... Thomas E. Renungton .-... •', A~ting Avian Research Program Manager County, 1999. ��71 Colorado Division of Wildlife Wildlife Research Report April 2000·· JOB PROGRESS State of: _--'- ....!:C~o!,!;lo~ra=do~ Project: ...!.W!--~1""-67!--~R'__ Work Plan: __ Job Title: ----=:2~8 __ : Job. REPORT _ _ Avian Research l~ __ Evaluate Population Trends of Selected Species ofNeotropical Migratory Birds in Colorado Period Covered: 01 January 1999 through 30 June 2000 Auilior: ~K~e~nn~e~ili~M~~~G~ie~se~n~_ Personnel: Mary Beth Dillon, Susan Jojola-Elverum, Kenneth M. Giesen, Colorado Division of Wildlife, Daniel Svingen, U.S. Forest Service, Mike Carter, Tony Leukering, Colorado Bird Observatory. ABSTRACT Data on Cassin's sparrow (Aimophila cassiniii breeding density on a Comanche National Grasslands study plot were collected using point-counts and will be analyzed later using program DISTANCE to calculate breeding density and compare density estimates over the breeding season. Nest searches resulted in three Cassin's sparrows nests being located; two were parasitized by brown-headed cowbirds (Molothrus ater) and all 3 were unsuccessful. Point-counts of Chestnut -collared longspurs (Calcarius ornatus) and McCown's longspur (Calcarius mccowni) were conducted on the Pawnee National Grassland to obtain estimates of breeding density and 87 nests of passerine birds were located using rope-dragging on study sites. Mayfield nest success of McCown's Longspurs was 28.1 percent on the East Willow site (27 nests) and 48 percent on the Carroll site (15 nests). Nest success of Chestnut-collared longspurs was 22.1 percent on the Carroll site (8 nests). Monitoring of selected species of passerine Neotropical migratory birds was continued in 1999 with a Division of Wildlife Contract with the Colorado Bird Observatory. A total of 309 point transects in 13 habitats was completed in 1999 and we obtained breeding density estimates on 219 bird species using Program DISTANCE. At least 325 specific locations were surveyed for colonial breeding species and limited-range species in Colorado and minimum size of breeding populations was estimated from direct counts. .'lIlioJ BDOW013564 ��73 EVALUATE POPULATION TRENDS OF SELECTED SPECIES OF NEOTROPICAL MIGRATORY BllIDS IN COLORADO Kenneth M. Giesen INTRODUCTION There is widespread concern that populations of many species ofNearctic-Neotropical migratory passerine birds have declined in the last 30 years (Robbins et al. 1986, Robbins et al. 1992). Although U.S. Fish and Wildlife Breeding Bird Survey (BBS) data monitors populations for many species, other species are not sampled adequately because of low densities, geographic distribution, or access. Many Colorado passerines are monitored annually with BBS routes, but other species are not represented or are sampled in numbers too small to ascertain population status and trend, even if one assumes BBS indices reflect actual population trends (Colorado Bird Observatory 1997). There is no statewide program for monitoring population status of most Colorado avian species, and little specific information on nest success or other measures of reproductive performance for those species apparently declining (e.g., grassland birds). A program was developed cooperatively between the Colorado Division of Wildlife, U.S. Forest Service, Bureau of Land Management, and the Colorado Bird Observatory to monitor Colorado's breeding birds using a system of distance-measured point counts (Buckland et al. 1993) stratified by habitats found in Colorado. This program (Monitoring Colorado's Birds) is designed to be able to detect a ~ 3 percent population change with a statistical significance of 0.1 and a power of 0.8 for target species in 13 Colorado habitats. Because many last 3 decades, grassland bird rates, fledging grassland avian species are reported to have undergone the largest population declines in the additional efforts will be focused on evaluation of inventory methodology for selected species and examination of their reproductive performance (i.e., nest success, parasitism success). P.N. OBJECTIVES The primary objectives of this study are to evaluate and implement a statistically reliable population monitoring program for passerine birds in Colorado, and to investigate population monitoring protocols and nesting success for selected species of grassland birds. . . SEGMENT OBJECTIVES 1. Review literature appropriate to monitoring Neotropical migratory birds and literature on ecology and biology of selected avian species. 2. Develop and implement a program to monitor trends in breeding bird abundance for Colorado's breeding bird population. 3. Select study areas and initiate research on abundance and productivity of selected species of Neotropical migratory birds in Colorado. �74 4. Monitor abundance of breeding birds, nest success, and fledging success of selected species of Neotropical migratory birds in Colorado. 5. Compile data and prepare annual progress report. METHODS Study sites for comparison of distance-measured point counts and transects were located on the Pawnee and Comanche National Grasslands, and 3 grassland species (Cassin's sparrow, chestnut-collared longspur, McCown's longspur) were selected for intensive surveys of their population densities and nesting success. Three-minute point counts were obtained at 1-2 week intervals on both sites to obtain estimates of breeding density of focal species. Points were located and marked with 30-40-cm wooden stakes and plastic flagging. Consecutive points were 200 m apart on the Pawnee site and 300 m apart on the Comanche site. The distance to each bird was recorded using a laser rangefinder, or occasionally, by pacing or ocular estimate. Line transect surveys were conducted on the same study plots as the point counts. Transect surveys were conducted by walking slowly along the marked point-count route and measuring distance and compass direction to each detected bird. As with the point counts, distance was measured using a laser rangefinder, and occasionally by ocular estimate or pacing. Data will be compiled at a later date and analyzed using Program DISTANCE (Buckland et at. 1993). Nests were located by dragging a 30.5-m rope along transects to flush birds from nests. A few nests were found incidental to other activities. Nests were marked by a short wood stake 10 paces from the nest and located relative to grid stakes placed on the area. Nests were checked at 3-4 day intervals, and nest success was calculated using the methods of Mayfield (1961, 1975). Literature concerning monitoring of avian populations and detecting population trends was reviewed and discussed with personnel of the Colorado Bird Observatory (CBO) and other agencies involved with bird monitoring in Colorado. A distance-based point count methodology (Buckland et al. 1993) was used to design a program for population monitoring of avian species in Colorado's major habitats. The protocol was to select transects of 15 points each in 30, randomly selected, vegetative stands for all 11 habitats in Colorado. Uncommon species were recorded on points and on transects between points. Colonial species and "special techniques" species were surveyed using various methodologies (Leukering, pers. comm.). RESUL TS AND DISCUSSION Point counts for Cassin's sparrow on the Comanche National Grassland were conducted at weekly intervals from 15 June to 13 July 1999 along the same 6.0 km transect (20 points). The number of Cassin's sparrows detected ranged from 53 on 22 June to 22 on 8 July and was markedly affected by weather conditions, especially wind. The number of detections totaled 198 (avg. 2.0/point per survey, Table 1). Only 3 nests of Cassin's sparrow were located using the rope dragging technique. Two were parasitized by brown-headed cowbirds and none was successful in producing young. On the East Willow pasture on the Pawnee National Grasslands point-count transects were conducted on 11 May, 25 May, 8 June, and 22 June for McCown's longspurs. The number of detections ranged from 64 to 90 on the 20-point transect and averaged 3.6 per point. One line transect survey was attempted on 13 May but was terminated due to the high number of McCown's longspurs displaying at the same time and the inability to ascertain detection distance and direction for multiple birds at the same time. Further, it was apparent that double counting of individuals was a problem as individual birds moved throughout their territories. �75 Nest searches on 20 May, 4 June, 17 June, 24 June, 8 July, and 15 July resulted in 27 McCown's longspur nests being located, in addition to 3 lark bunting (Calamospiza melanocorys) nests, 2 homed lark (Eremophila alpestris) nests, and 2 mountain plover (Charadrtus montanus) nests. The number of eggs in the McCown's longspur nests ranged from 2 to 5 (avg. 3.46). Mayfield egg success was 0.73 and fledging success was 0.39 for an overall Mayfield nest success of 0.28 for the McCown's longspur. 1. Summary of Cassin's sparrow point-count data on the Comanche National Grasslands, 1999. 0-25m 26-50m 51-75m 76-IOOm IOI-125m 126-150m 151-175m I 76-200m 200+m 15 Jun 4 9 7 6 6 4 2 2 1 22 Jun 1 8 9 6 15 4 3 1 6 30 Jun 0 3 4 9 15 4 6 1 0 8 Jul 0 3 5 8 2 2 0 1 1 13 Jul 0 1 9 6 7 6 5 4 2 Total 5 24 34 35 45 20 16 9 10 Point-counts for chestnut-collared longspurs on the Carroll Pasture were conducted on 19 May, 2 June, 15 June, and 30 June. The number of chestnut-collared longspurs ranged from 24 to 41 on the 20-point transect and averaged 1.75 per point. As on the East Willow pasture, the number of simultaneously displaying birds made it difficult to conduct line transect surveys without double counting individuals. Only 8 chestnut-collared longspur nests were found on the Carroll Pasture during searches on 18 May, 6 June, 16 June, 24 June, 30 June, 6 July, and 13 July. Mayfield egg success was 0.30, fledging success was 0.73, and overall Mayfield nest success was 0.22. There were also 15 lark bunting nests located on this pasture (Mayfield nest success 0.19), as well as 15 McCown's longspur nests (Mayfield nest success 0.48), and 8 homed lark nests (Mayfield nest success 0.30). The results of the statewide monitoring program for all non-game birds in Colorado is attached to this report and provides specific methodologies and protocols for this effort as well as the preliminary results. LITERA TURE CITED Buckland, S. T., D. R. Anderson, K. P. Burnham, and 1. L. Laake. 1993. Distance sampling: estimating abundance of biological populations. Chapman & Hall, London, England. Colorado Bird Observatory. 1997. 1996 reference guide to the monitoring and conservation status of Colorado's breeding birds. Colorado Bird Observatory and Colorado Div. Wildlife, Denver. Mayfield, H. 1961. Nest success calculated from exposure. Wilson Bull. 73:255-261. Mayfield, H. 1975. Suggestions for calculating nest success. Wilson Bull. 87:456-466, �76 Robbins, C. S., D. Bystrak, P. H_ Geissler. 1986. The breeding bird survey: its first 15 years, 1965-1979. Resource Pub. 157. Washington, D.C. Fish and Wild!. Serv., U. S. Dep. Inter: 133-159. Robbins, C. S., 1. R. Sauer, and B. G. Peterjohn. 1992. Population trends and management opportunities for neotropical migrants. Pages 17-23 in D. M. Finch and P. W. Stengel (eds.). Status and management ofNeotropical migratory birds. U. S. Dep. Agric. Forest Serv., Rocky Mountain Forest and Range Exper. Sta., Gen. Tech. Rep. RM-229. Fort Collin, CO. 422 pp. PREPARED BY: ~~ Kenneth M. Giesen Wildlife Researcher �77 Monitoring Colorado's Birds: Report for the 1999 First Full-effort Year Submitted by Tony Leukering and Rich Levad Colorado Bird Observatory 13401 Picadilly Road Brighton, CO 80601 Submitted to Ken Giesen Colorado Division of Wildlife 317W. Prospect Ft. Collins, CO 80526 Chris Schultz U.S.D.A. Forest Service 15 Burnett Ct. Durango, CO 81301 Ron Lambeth U.S. Bureau of Land Management 624 Yucca Rd. Grand Junction, CO 81503 31 March 2000 Abstract In 1999, Colorado Bird Observatory, in conjunction with itsfunding partners - Colorado Division of Wildlife, U.S.D.A. Forest Service, and U.S. Bureau of Land Management - conducted the pilot effort of. the full-scale breeding-bird monitoring plan, as delineated by Leukering and Carter (1998). We conducted an intensive literature search to compile all known breeding localities for all of Colorado's colonially-breeding waterbirds and a small suite of limited-range species. We augmented the literature search with an extensive effort to contact biologists, land managers, birders, and other people with information on breeding locations of these species. During the course of the 1999 field season, we visited a large number of these to determine how many sites were still active and, if so, the number of breeding individuals at each. In addition, we obtained data from various contacts on the activity and colony size of many more colonies. After 1998's successful pilot transect season (Leukering 1999; three habitats: Aspen, Ponderosa Pine, and Spruce-Fir), we attempted to initiate the transect protocol in an additional ten habitats in 1999. Though we encountered many difficulties, this year's effort was an incredible success, as the habitat-based transects �78 provided excellent data on 94 breeding species (coefficients of variation of :sSO%) and solid data on an additional 54 breeding species (coefficients of variation between 50% and 100%). This total of 148 is 62% of all regularly-occurring Colorado breeding species. Future years' data should be even better. Establishing 300 new transects in 1999 proved to be an impossible task for a number of reasons. About the most recalcitrant hindrance was the difficulty we encountered in finding appropriate tracts in which to place transects due to the numerous errors in habitat allocation in the Colorado GAP data set. This is the GIS information that we used to randomly select transect locations in both years (1998 and 1999). This proved to be a difficulty in 1998, but we only conducted transects in three habitats that year; having these difficulties with ten habitats caused uncountable lost field days and resulted in incomplete sample size for a number of habitats. Additionally, we were unable to obtain the services of enough qualified field workers to physically conduct all the work required. This was due to two reasons: 1) we had a very small pool of qualified applicants from which to select (for reasons unsure) and 2) we had to wait until contracts were in place before hiring, thus we lost a few of the qualified applicants to other positions before we started the hiring process. Because the Monitoring Colorado's Birds project is personnel-intensive, it is imperative that we be able to conduct hiring in January and February before a large number of qualified applicants have already signed on to other projects. Introduction Colorado Bird Observatory (CBO) initiated efforts to create and conduct a Colorado-wide effort to monitor breeding-bird populations in 1995. In 1997, after review by statisticians and Colorado Division of Wildlife (CDOW) biologists, we redesigned the program (Monitoring Colorado's Birds (Leukering and Carter 1998» and conducted a small, pilot effort in 1998 on three habitats (Leukering and Carter 1999). With the success of the 1998 effort, we expanded field work in 1999 to include all originally-allocated habitats and special-species efforts - in effect, a full-scale pilot effort. This report delineates effort and results of the 1999 field season and provides recommendations and suggestions for changes to be incorporated in 2000. Methods We used three methods: point transects, colony counts, and censussing, to obtain population data for all of Colorado's breeding-bird species. Point transects-We established transects of 15 point counts in each of30 randomly-selected stands in each of 11 habitats. Using the Colorado GAP data set, we numbered all publicly-owned stands of the habitats in Colorado and randomly selected 60 from each habitat.' We then randomly selected 30 of those in which we established point transects. In a few instances, selected stands were not the indicated habitat or access across private land was denied, so we discarded them and randomly selected a replacement from the original set of randomly-selected stands. Most replacement stands were randomly selected from all stands, regardless of ownership. We selected all Grassland transects randomly from all stands with only seven falling on public lands. Each transect was conducted by one observer using protocol established by Leukering (1998). The observer located the selected stand on the ground and ran the transect along a randomly-selected bearing. It was usually impossible to run the entire transect along the random bearing, as stand boundaries, property boundaries, and physical obstructions forced turns in the transect direction. When this happened, the observer randomly turned right or left perpendicular to the random bearing, subsequently alternating perpendicular directions if additional turns were necessary. In some stands, the narrowness of the stands predicated the location and bearing of the transects. �103 Colorado Division of Wildlife Wildlife Research Report April 2000 JOB PROGRESS State of: Project: Work Plan: __ Job Title: _;C~o~l~o~ra~d~o REPORT _ W-167-R ---=:2.:::.,.9 __ : Job __ Avian Research ....!.,_ __ --=D;..:;is:::.:t~ri"""bu:;t:.:.;io"""n:.....:an=d:....:r~ep,r;:.;r'_"od=uc=_=t:.:...;iv:...:::e....::s:=ta=tu::,:s::.,_o"",f:... _ Period Covered: 01 January 1999 through 30 June 2000 Auilior: =K=e=nn=e=th==M=.~G=I=·e=se=n~ _ Personnel: Mary Beth Dillon, Susan Jojola-Elverum, Kenneth M. Giesen, Colorado Division of Wildlife, Daniel Svingen, U.S. Forest Service ABSTRACT Mountain Plover (Charadrius montanus) use of native shortgrass rangeland, agricultural fields, and shortgrass rangeland managed with prescribed burning was evaluated with surveys during spring migration and monitoring of nesting on the Pawnee National Grasslands and surrounding private lands in Weld County, and on the Comanche National Grasslands in Baca County, Colorado. Three United States Forest Service (USFS) pastures on the Comanche National Grasslands were burned on March 10 and counts of plovers on these pastures peaked during the first 2 weeks of April. Forty-five plover nests were located on these pastures (and no nests on control pastures) with 23 (51 percent) being successful (Mayfield nest success 4l.7 percent). Surveys were conducted in April in Weld County and 150 Mountain Plovers were observed during 332 miles of transects. The number of plovers detected on private lands was proportional to land ownership, as was proportion of birds seen on agricultural fields and shortgrass rangeland. Eighteen nests were located (9 on rangeland, 9 in agricultural fields) of which 11 hatched (61 percent). Mayfield nest success was 30.1 percent and was higher on agricultural fields (55 percent) than on native rangeland (14 percent). Radio transmitters placed on 22 nesting plovers (13 in Baca County, 9 in Weld County) were lost or failed before fledging of broods so no estimate of brood survival after hatch was possible. Numbers of post-breeding plovers observed on staging areas in Baca County peaked between July 15 and August 3, with most plovers migrating by early September. ��105 DISTRIBUTION AND REPRODUCTIVE STATUS OF MOUNTAIN PLOVERS Kenneth M. Giesen INTRODUCTION The mountain plover (Charadrius montanus) is known to breed in the eastern Colorado counties of Weld, Bent, Cheyenne, Baca, and Kiowa; a small population also breeds in Fremont and Park counties with scattered individuals in other localities (Bailey and Niedrach 1965, Andrews and Righter 1992, Kuenning and Kingery 1998). Historically mountain plovers were considered numerous in eastern Colorado (Graul and Webster 1976), but numbers have declined over the last 2-3 decades (Knopf 1991). It is estimated the population is declining at an annul rate of 3.7 percent. The primary cause appears to be loss of wintering habitat in California and conversion of native prairie grasslands to cultivation on breeding grounds. In Colorado, the mountain plover is currently classified as a Species of Special Concern and has been petitioned for listing under the federal Endangered Species Act. Studies by Graul (1975) and Knopf and Rupert (1996) have described the plover's natural history and breeding requirements on the short grass prairies of eastern Colorado. Although the species has declined throughout its range, the recently completed Colorado Breeding Bird Atlas suggests that mountain plovers may be more widespread that previously known (Kuenning and Kingery 1998). Knopf (1996) recommended the use of prescribed burning to enhance attractiveness of native prairie for mountain plovers. Because the U.S. Forest Service initiated prescribed burning for mountain plover habitat improvement on the Pawnee and Comanche National Grasslands several years ago there is a need to evaluate plover response to these treatments, especially regarding breeding density and nest success. SEGMENT OBJECTIVES The primary objectives of this study are to (1) develop and evaluate a statewide monitoring program to identify distribution, population trends, and changes in productivity for Mountain Plover, and (2) investigate contributions of burned grasslands and fallow cultivated fields to plover productivity. P.N. OBJECTIVES 1. Monitor plover breeding densities, nest success and chick survival on prescribed burns on the Comanche National Grasslands and evaluate the effectiveness of that management practice. Up to 3 designated prescribed bum areas will be inventoried a year prior to the burn, the year of the bum and a year following the bum. 2. Up to 30 plovers nesting within the study sites will be trapped, banded and equipped with transmitters. Movements of broods will be tracked so that mortality, dispersal and habitat preferences can be documented. 3. Monitor breeding densities, nest success and chick survival on fallow, cultivated fields. Fallow cultivated fields will be surveyed by the investigators, DWMs, and Area Biologists for the presence of �106 nesting plovers. Nests will be monitored to document success and productivity. A sample of up to 30 nesting adults will be trapped and treated as described in 2 (above) to monitor movements and attrition of broods. 4. Compile data and prepare annual report. METHODS We conducted roadside transects in northern Weld County, on and adjacent to the Pawnee National Grasslands, to document relative abundance of Mountain Plovers (Charadrius montanus) on rangeland and agricultural fields (wheat stubble, fallow fields). Transects were conducted using vehicles traveling g 50 kmIhr. Within suitable habitat (short grass rangeland, prairie dog colonies, fallow fields), observers stopped the vehicle every 0.8 km and scanned for 3-5 minutes. When plovers were observed, habitat at the site was recorded and the location marked on a topographic map. A Pawnee National Grassland map was used to ascertain land ownership (public vs. private). Three pastures on the Carizzo Unit of the Comanche National Grasslands (7B, 8E, 13D), Baca County, were burned by U.S. Forest Service personnel on March 10, 1999 to enhance breeding habitat for Mountain Plovers. We conducted surveys for plovers on these pastures 3-5 times weekly during spring migration (late March - early May) using 4WD vehicles to traverse parallel north-south transects 0.2-0.3 km apart. When plovers were observed, the location, time, and number of birds was recorded on 7.5minute topographic maps. Double counting of individual flocks was minimized by recording both flock size and location precisely and taking care not to flush plovers. Surveys were conducted alternately in mornings and afternoons, and in each instance, were typically completed within 3-5 hours. We compared plover use of burned and unburned pastures by conducting additional transect surveys from 0630 to 1100 h on 15 and 21 April on the 3 burned pastures, on 2 pastures burned in March 1998 (7A, 14G), and on 3 pastures scheduled for burning in spring 2000 (41, 5B, and 14M). Starting on the east side of each pasture, we drove north-south transects spaced at 0.4 km and stopped every 0.4 km. At each stop, the observer exited the vehicle and used binoculars to scan in all directions for three minutes. Location and numbers of plovers observed withing 0.2 km were documented. We searched for plover nests with 1-2 observers in vehicles (primarily 4WD pickups) on the Comanche National Grasslands and 1 observer in an ATV in Weld County. Parallel transects 50-100 m apart were driven at 15-20 kmIhr on study pastures (Comanche N. G.) and in potential habitat in Weld County while scanning for plovers. A few nests in Weld County were located incidental to other activities. When plovers were observed we examined the site for eggs or a nest scrape. Nests were usually marked with a 3 x 20-cm wood stake placed 10 paces from the nest, and a wire flag placed 15 paces from the nest. We monitored nests from vehicles 1-3 times weekly until hatch or nest failure. Nest success was ascertained from observation of adults with young at the nest or observation of tiny eggshell fragments at the nest after expected completion of incubation (Mabee 1997). We estimated nest success after the methods of Mayfield (1961, 1975). Incubation period was estimated at 29 days (Graul 1975). Selected adults were trapped on nests and fitted with miniature radio transmitters « 2.0 gms) glued to their backs to monitor movements and survival of dependent juveniles. Efforts were made to locate radio-marked adults 1-2 times weekly and document numbers of surviving juveniles. �107 RESUL TS AND DISCUSSION Weld County A total of 150 Mountain Plovers was observed during 332 miles of surveys in April (0.45 plovers/mile) which included approximately 1184 quarter sections of potentially suitable habitat. Because transects were completed in April, many plovers observed were likely migratory rather than resident breeding birds. Most plovers were observed in short grass rangeland (N = 127) with fewer being observed in green winter wheat (N = 17) or in wheat stubble or fallow fields (N = 6). More plovers (N = 112; 74.7%) were seen on private lands than on public (USFS) lands (N = 38) with private lands comprising 80.3 % of the area surveyed. Because these surveys were conducted in April, some plovers observed were likely migrants and did not breed in the areas where observed. Regardless, the data indicate no selection for public lands over private lands, nor any gross selection for specific habitat type as long as it met criteria of having short vegetative cover and bare ground. If these data are indicative of habitat preferences for breeding plovers, agricultural lands, especially fallow fields, may comprise a significant portion of potential breeding habitat in northeastern Colorado as was hypothesized by Shackford et al. (1999). Eighteen Mountain Plover nests were located in Weld County. Nine of these were in rangeland and 9 in agricultural fields. Clutch size observed was 1 egg (1 nest), 2 eggs (2 nests) or 3 eggs (15 nests). Because the clutches of 1 and 2 eggs were not observed early in incubation some eggs may have been lost prior to our recording clutch size. Gross nest success was 61.1 percent (11 of 18 nests hatched), while Mayfield nest success was 30.1 percent based on 176 nest-days of observation and a 29-day incubation period. Overall nest success on agricultural fields (7 of 9) was higher than that observed on native short grass rangeland (4 of 9). Mayfield nest success on agricultural fields and native rangeland was 0.55% and 0.14%, respectively. However, one additional nest in a fallow field may have been destroyed by cultivation if the nest location had not been marked and identified to the landowner when he was cultivating the field. The average hatch date of 11 successful nests was 8 June (median 5 June, range 2 June - 26 June) indicating an average nest initiation date of 6 May. We attached radio transmitters to nine plovers (5 nesting on rangeland, 4 nesting on fallow fields) prior to hatch of their clutches. Transmitters fell offwithin a few days of capture on 4 plovers, and three plovers lost their transmitters within 10 days post-hatch. Two remaining plovers were tracked through the fourth week post-hatch before signals were lost. One of these birds had moved 15 miles before the signal was lost. Since the fledging age is 33-36 days (Graul 1975, Miller and Knopf 1993) we believe both these birds had lost their broods. Because no other plovers were marked, we were unable to ascertain brood survival from any of the adults for which we had located nests. Other methods for attaching transmitters to plovers need to be investigated. Baca County Weekly surveys of the three burned pastures resulted in maximum counts of 44,44, and 107 plovers in pastures 7B (473 ha), 8E (194 ha), and 13D (210 ha), respectively, between 10 and 15 April (Table 1). This period represented the peak of spring migration for Mountain Plovers in Baca County. Although plovers were not banded or individually marked, it appeared that migrating plovers remained on pastures for only a short period before resuming migration. Surveys in April of pastures burned in 1998 (7A., 14G) resulted in only 1 pair of plovers being observed on approximately 794 ha, and those were associated with a prairie dog (Cynomys ludovicianus) colony. Because of vegetation growth the effect of burning was short-lived. Encouraging heavy grazing following the burn might result in suitable plover breeding habitat in subsequent years. Similar surveys of pastures scheduled for prescribed burning in 2000 (41, 5B, 14M) totaling 718 ha did not detect any plovers. �108 Table 1. Peak numbers of Mountain Plover counted during weekly surveys on the Comanche National Grasslands, Baca County, Colorado, 1999. Weekly Period Pasture 7B Pasture 8E Pasture 13D Total Mar. 21- Mar.27 Mar. 28 - Apr. 3 Apr. 4 - Apr. 10 Apr. 11 - Apr. 17 Apr. 18 - Apr. 24 Apr. 25 - May 1 35 Not counted 44 22 17 6 1 Not counted 40 44 10 11 76 Not counted 89 107 42 33 112 Not counted 173 173 69 50 Mating was first observed on 15 April and backdating of hatched clutches (N = 23) indicated the mean date of clutch initiation was 24 April. A total of 45 nests was located, most (N = 33) were found in Pasture 13D. There was no positive correlation between pasture size and number of nests located suggesting breeding habitat and nest density may be related to landscape or topographic features external to the size of burned pastures (Svingen and Giesen 1999). Clutch size was 3 eggs in 42 nests and 2 eggs in 2 nests. One unusual nest contained 3 Mountain Plover eggs and 3 Killdeer eggs and was being incubated by a Killdeer (Jojola-Elverum and Giesen, in press). The mean hatch date was 27 May (median 26 May, range 15 May - 19 June) with 12 of23 (52.2 percent) successful nests hatching between 22 and 28 May. Overall nest success was 51.1 percent (23 of45) with Mayfield nest success being 41.7 percent based on 740 nestdays of incubation (Table 2). Most nest failure was due to predation with only 1 instance of nest abandonment suspected. Table 2. Nest success of Mountain Plover on the Comanche National Grasslands, Baca County, Colorado, 1999 Pasture Size (ha) n Nests n Successful 7B 473 10 4 0.265 8E 194 2 0 0.000 13D 210 33 19 0.504 Nests Mayfield Nest Success Thirteen plovers were trapped on nests prior to nests hatching in an attempt to use telemetry methods to document movements of broods and survival of young. Unfortunately, most radios were removed by the plovers within a week of hatch and no radios remained on plovers after 7 June, thus no data on brood movements or fledging rates was obtained. Telemetry indicated adult plovers with broods remained within the burned pastures at least until radios failed or fell off. Results of surveys on both burned and unburned pastures in April, and documentation of nesting plovers on burned pastures indicates prescribed burning of appropriate rangeland may be beneficial to Mountain Plovers (Knopf 1996). Although long-term data are not available concerning productivity of plovers on rangeland managed by prescribed burning, overall nest success rates (51 %, Mayfield 42%) do not indicate these areas are a population "sink". However, additional information on fledging rates is needed to quantify productivity. Counts of post-breeding plovers on approximately 30 parcels of privately owned agricultural land (32-130 ha) suitable for plovers (fallow cultivated fields, burned wheat stubble) indicated few plovers «20/day) were observed prior to 12 July. Between 15 July and 3 August up to 440 plovers were counted/day on the �109 same fields. Counts of plovers declined thereafter until early September when no plovers could be located on these fields. Plovers appeared to select certain fields for their post-breeding staging areas and avoid other nearby, and seemingly similar, fields. It is possible that fields selected contained a higher density of available invertebrates and were selected on the basis of foraging efficiency, rather than for other characteristics. LITERA TURE CITED Andrew, R. A, and R. Righter. 1992. Colorado Birds. Denver Mus. Nat. History, Denver. Bailey, A M., and R 1. Niedrach. 1965. Birds of Colorado. Denver Mus. Nat. History, Denver. Graul, W. D. 1975. Breeding biology of the Mountain Plover. Wilson Bull. 887:6-3l. _, and L. E. Webster. 1976. Breeding status of the mountain plover. Condor 78:265-267. Jojola-Elverum, S. and K. M. Giesen. 2000. Killdeer parasitizes Mountain Plover nest. Wilson Bull. 112: in press. Knopf, F. L. 1991. Status and conservation of mountain plovers: the evolving regional effort. Report of research activities, National Ecology Research Center, U. S. Dep. Inter., Fish and Wildl. Servo Fort Collins, CO. _. 1996. Mountain Plover (Charadrius montanus). A Poole and F. Gill (editors). The birds of North America, No. 21l. The Academy of Natural Sciences, Philadelphia, PA, and the American Ornithologists' Union, Washington, D.C. _, and J. R. Rupert. 1996. Productivity and movements of mountain plovers breeding in Colorado. Wilson Bull. 108:28-35. Kuenning, R R, and H. E. Kingery. 1998. Mountain Plover. Pp. 170-171 in H. E. Kingery, editor, Colorado Breeding Bird Atlas. Colorado Bird Atlas Partnership and Colorado Division of Wildlife, Denver. Mabee, T. J. 1997. Using eggshell evidence to determine nest fate of shorebirds. Wilson Bull. 109:307313. Mayfield, H. F. 1961. Nesting success calculated form exposure. Wilson Bull. 73:255-261. _. 1975. Suggestions for calculating nest success. Wilson Bull. 887:456-466. Miller, B. J., and F. L. Knopf. 1993. Growth and survival of Mountain Plovers. J. Field Ornithol. 64:500506. Shackford, 1. S., D. M. Leslie, Jr., and W. D. Harden. 1999. Range-wide use of cultivated fields by Mountain Plovers during the breeding season. J. Field. Ornithol. 70:114-120. Svingen, D., and K. Giesen. 1999. Mountain Plover (Charadrius montanus) response to prescribed burns on the Comanche National Grasslands. J. Colo. Field. Ornithol. 33:208-212. PREPARED BY: ~ Kenneth M. Giesen Wildlife Researcher � https://cpw.cvlcollections.org/files/original/73c86cbbca040efde9c89c6307d3a582.pdf 48595c755af20f2e03c7e2656ee55f1e PDF Text Text 1 Colorado Division of Wildlife Wildlife Research Annual Report July 2000 JOB PROGRESS State of Project No. Work Package No. __ Task No. REPORT Colorado Cost Center 3430 W-153-R-13 --=0:...::6-"-6=.2 1 Mammals Program _ Preble's Meadow Jumping Mouse Conservation· Develop Conservation Plan for Preble's Meadow Jumping Mouse Period Covered: July 1, 1999 - June 30,2000 Author: Tanya M. Shenk and Gary C. White ._-_._--_._--_ ._--------. ABSTRACT The second field season of a three year study on the demography and movement patterns of Prebles meadow jumping mouse (Zapus hudsonius preblei) was completed. A total of 175 individual mice were captured on stream-side transects at three study sites (60 mice at Maytag Property, 60 mice at PineCliff Ranch, and 55 mice at Woodhouse Ranch) during the 1999 field season. These captures included 34 mice PIT-tagged in 1998 (14 at Maytag Property, 12 at PineCliffRanch, and 8 at Woodhouse Ranch). These mice were captured over three different trapping sessions with an effort of 19,222 trap-nights. Most adult mice captured exhibited evidence of active reproductive behavior, either pregnancy, lactation, or enlarged genitalia. We were able to locate one potential maternal nest. Juvenile mice were not captured during the June trapping session. Juvenile mice were captured at all three sites during both the July and September trapping sessions. Natural mortality factors documented during both years include predation by house cats, garter snakes, rattlesnakes, yellow-bellied racers, bullfrogs, weasel, and fox as well as accidents by drowning and road kill. In 1999 mice were also found in torpor throughout the summer, often in very exposed areas frequently resulting in death by either exposure or predation. Preliminary analysis of the movement data collected on radio-collared PMJM in 1999 support results found in 1998. In particular we were able to document again in this second field season that PMJM exhibit (1) greater use of upland habitats than previously assumed, (2) general site fidelity to both daytime nesting sites and nighttime feeding sites, (3) seasonal shifts in movement patterns, and (4) use of both perennial and intermittent tributaries adjacent to the' capture drainage. Detailed vegetation maps for all three study sites were created through intensive field mapping. These maps will be used in conjunction with the movement data to further refine habitat use information. Thirteen possible hibernacula were located in 1999. Three sites were located at Woodhouse Ranch and ten possible hibernacula at Pine Cliff Ranch. In contrast to 1998 data, these potential hibernation sites were located closer to the stream. However, at both Pine Cliff and Woodhouse Ranches stream banks rise out of the floodplain in closer proximity to the stream than at Maytag Property where the potential hibernation sites located in 1998 were located long distances from the stream center. Vegetation characteristics of the 21 sites located during both 1998 and 1999 are similar to �2 other hibemacula described for PMJM. By cooperating with other researchers PMJM densities (mice/km of stream) were estimated from nine study areas during June 1998 and June 1999, providing a total of 15 study area x year combinations. With these limited data available, 68% of the variation in PMJM density was explained by a model that includes riparian shrub and tree cover (halkm stream). These results suggest that habitat quality ofPMJM can be predicted by the riparian shrub and tree cover available on a site. Preliminary results from the demography, movement and distribution studies of PMJM have suggested the need to continue with research questions currently being addressed. However, the preliminary results also suggest further research be conducted on (1) use of upland and riparian habitat by PMJM, (2) refining water requirements ofPMJM (i.e., do they require stream habitat or are wetland areas sufficient?), (3) range-wide distributional boundaries (e.g., elevation restrictions), and (4) investigating areas of potential sympatry or hibridization with the western jumping mouse (Z princeps). The demography and movement studies were modified for the summer 2000 field season to further investigate upland use and water requirements, including a habitat modification experiment. TIlls report includes results from the first two years of a multi-year project to follow individually marked PMJM through time. Collection of more data, and more years of data, will improve our ability to evaluate demographic parameters, movement patterns, and habitat use and evaluate how they vary across space and time. �3 DEVELOP CONSERVATION PLAN FOR PREBLE'S MEADOW JUMPING MOUSE Tanya M. Shenk and Gary C. White P. N. OBJECTIVE Conservation of Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. SEGMENT OBJECTIVES FY99-00 1. Estimate abundance, over-summer survival, annual survival, Preble's meadow jumping mouse at three study sites. 2.. Evaluate movement data of Preble's meadow jumping mouse study sites. 3. Refine and prioritize needed research components to develop Preble's meadow jumping mouse. 4. Develop study plan to evaluate meta-population dynamics in funded in FYOO-01). 5. Prepare a Federal Aid Job Progress Report. and population age and sex structure of as it relates to landscape features at three sound strategies for conservation of Preble's meadow jumping mouse (to be PERFORMANCE INDICATORS FY99-00 1. 2. 3. 4. Second year survival rate estimates for three populations of PMJM. Second year abundance and density estimates for three populations ofPMlM. Second year movement patterns for three populations of PMlM. Study plan for PMlM meta-population study. INTRODUCTION On May 12, 1998 the U.S. Fish and Wildlife Service (USFWS) published a final rule in the Federal Register (63 FR 26517) to list Preble's meadow jumping mouse (Zapus hudsonius preblei) as 'threatened' under the Federal Endangered Species Act (ESA) of 1973~ as amended. Recovery goals for Preble's meadow jumping mouse (PMJM) should work towards the sustainabilityprotection, and restoration of Z. h. preblei populations and habitats on both private and public lands to provide the spatial, genetic, and demographic structure needed to promote long-term species viability and provide species management flexibility. Recovery efforts for the subspecies will be most effective if reliable information is available on the basic ecology of the subspecies and this information used to design recovery efforts such as Habitat Conservation Plans. A review of studies conducted on PMJM shows that there is insufficient information to fully address defining fange-wide ecological requirements, limiting factors, limits of species tolerance, or population status (Shenk 1998). Most work to date has focused on geographic distribution (presence or absence of Z. h. preblei), taxonomy, and habitat descriptions of sites where mice have and have not been captured. For PMlM in particular, information on dispersal, habitat use, and population dynamics is most needed to identify minimal ecological requirements of the subspecies. Monitoring, by way of estimating demographic parameters (such as survival, abundance, and reproduction), of a single population over time provides an opportunity to document demography of a �.......... --------------_ 4 species, estimate temporal fluctuations in demography, and gain insights into the temporal variation inherent in demographic parameters. Monitoring of multiple populations over time and over multiple geographic locations provides the opportunity to gain further understanding of population processes and detaches time effects from spatial effects .. Population monitoring activities do not constitute scientific experiments, in the spirit of manipulation of salient ecological variables, however, replication of monitoring activities for natural populations over long periods of time and in diverse geographic locations can lead to insights into population processes (Cook and Campbell 1979). These insights can then be translated into hypotheses useful for predicting changes in population demography resulting from either natural perturbations (e.g., flooding events) or anthropogenic modifications (e.g., gravel mining). Experimentation would then be required to test these hypotheses and establish cause and effect. Movement and dispersal pattern information will be key to any conservation strategy designed for PMlM. Documenting daily and seasonal movement patterns of PMJM will provide information on habitats used by the mice and on the relative configuration and juxtaposition of these habitats. Configuration of habitats include vegetation and size of the areas used. Juxtaposition includes relative locations of different habitats used such as nest sites, feeding areas, and movement corridors connecting those areas. Key dispersal factors to document for PMJM include (1) which segment of the population disperses, (2) when do they disperse, (3) through what habitat do they disperse, (4) how far will individuals disperse (i.e., what is the maximum distance that separates adjacent populations) and (5) how critical is dispersal (both into and out of a population) to the persistence of a given population. Areas of suitable habitat must provide requirements to survive throughout the life cycle. These requirements must provide necessities for both the active period and hibernation periods. During the active period suitable habitat must provide requirements for daily survival, reproductive activities (breeding, nesting, and rearing of young to independence), and dispersal. The hibernation period requires sufficient food supplies to assure fat storage prior to hibernation and suitable hibernacula. Habitat providing all seasonal and life cycle requirements mayor may not occur in a single contiguous area. If not in a contiguous area, habitat patches must occur in a mosaic of usable areas where suitable corridors exist for seasonal movement among sites. The habitat matrix within the range of Z. h. preblei is mixed grasslandsadjacent to the Colorado Front Range along the Piedmont and along the base of the Laramie Mountains in Wyoming and extends to the Colorado plains. Within this matrix., PMJM occur along stream drainages that contain patches of suitable vegetation. Suitable habitat appears to have at least two major components, The first component is a supply of open water, at least in part of the active season. Secondly, areas where PMJM has been found have dense cover. Based on studies of Z. h. preblei and Z. hudsonius elsewhere, Z. h. preblei apparently occurs mostly in undergrowth consisting of grasses, forbs, or both in open wet meadows and riparian corridors, or where tall shrubs and low trees form an overstory and provide adequate cover (Armstrong et al. 1997). Meadow jumping mice are widespread in abandoned grassy fields, but are often more abundant in thick vegetation along ponds, streams, and marshes or in rank herbaceous vegetation of wooded areas (Whitaker 1963). The mouse does not appear to have an affinity toward any single plant species but instead favors sites that are structurally diverse and provide adequate cover and food throughout its life cycle. PMJM are typically not found in upland areas away from riparian habitats but are most often captured where either ground water becomes visible as either seep springs or as main water channels (M. Bakeman, T. Ryon, personal communication) suggesting a dependence on open water, at least during their active periods. PMlM have been trapped in natural riparian areas as well as areas altered by anthropogenic influence including ditches and wetlands adjacent to interstate highways, cement-lined ditches with tall cover, ditches along driveways and moderate road. use, and moderate cattle grazing. If PMlM occurs as metapopulations in the classical sense of a set of local populations linked by infrequent dispersal then habitat includes not just one area of suitable habitation but also areas suitable for nearby mouse populations. These suitable areas must also be linked by dispersal habitat. If the mice are �5 dependent on dense riparian habitat for dispersal as well as for areas to reproduce, persistence of discrete populations would require a mosaic of suitable discrete riparian patches interconnected with dispersal corridors of similarly dense riparian vegetation. If mouse populations function in a source (populations . where growth rate ~ 1) and sink (those populations where growth rate < 1, maintained through immigration) system, it will be critical to identify and protect those populations serving as sources. Thus, for a source-sink population critical habitat will include those areas that support source population, dispersal habitat to sink areas will be less critical. If local mouse populations are functionally discrete, such a mosaic of interconnected areas of suitable habitat would provide a buffer for local, and source, populations against deleterious stochastic events by providing the opportunity for local population failures to be 'rescued' by immigration from other populations. The primary objective of this study is to investigate spatial and temporal variation in the demography and movement patterns of PMlM. Demographic parameters to be estimated include survival, reproduction, temporary emigration, immigration, population structure, and density. These demographic parameters will be estimated from individually marked animals from geographically distinct populations. To evaluate spatial differences in the demography of PMJM, populations were selected from three sites that provided a variety of habitat matrices available to the mouse. To begin to understand PMJM movement, dispersal, and habitat use we monitored radio-collared mice at three different study areas .. Study areas were selected to cover a variety of habitat configurations to evaluate spatial variation in movement patterns of PMlM. Multiple years of conducting the study at those same sites will provide an estimate of the temporal variation in demography and movement patterns of PMlM. STUDY SITES Demography and movement patterns ofPMlM are being evaluated for three populations. All three populations selected were located in areas where PMJM had previously been found. To evaluate spatial differences in the demography and movement patterns of PMJM, populations were selected on sites that provided a variety of habitat matrices available to the mouse. The first site selected, Colorado Division of Wildlife (CDOW) Maytag Property, has one primary water source available to the mouse. This water source is East Plum Creek. The second site, PineCliffRanch, provides both a tributary (Garber Creek) and a main stem drainage (West Plum Creek). This provided the opportunity to investigate whether PMlM will use upland areas to move from one drainage to another or if they are restricted to only moving along riparian corridors. The third study site, CDOW Woodhouse Ranch provides an area containing a tributary (Indian Creek) and a series of ponds and irrigation ditches scattered throughout the property. The configuration at Woodhouse Ranch provided an even greater opportunity to investigate how much the mice use upland areas or if they restrict their movements strictly to riparian corridors. By replicating the same methodologies at each of these three unique habitat matrices we can begin to estimate spatial variation in nightly and seasonai movements of PMlM. . i. OBJECTIVES Objectives of the demography study are to: 1. Estimate abundance and density ofPMlM for each of three study populations. 2. Estimate over-summer, over-winter, and annual survival ofPMJM at each of three study populations. 3. Estimate temporary emigration ofPMJM at each of three study populations. 4. Estimate immigration of marked PMlM back into each of three study populations. 5. Estimate reproduction ofPMlM at each of three study populations. 6. Evaluate the affect of weight, sex., age, abundance (i.e., density dependent response), and habitat features such as stream reach, vegetation composition and density on survival, reproduction, abundance, temporary emigration, and immigration of marked animals back into three study populations of PMlM. �6 7. Estimate age and sex ratios ofPMJM at each of three study populations. The PMJM movement study is designed to describe nightly and seasonal movement patterns ofPMJM and to describe habitats used by PMJM. These movement patterns will be described for three different study areas to evaluate spatial variation, and over three different years at the same three study areas to evaluate temporal variation. Specific objectives include: 1.' Describe nightly movements of PMJM. Evaluate difference in nightly movements as they relate to sex, age, and habitat available to the animals. 2. Describe 30-day (or life of the radio transmitter) interval movements ofPMJM. Evaluate difference in 30-day (or life of the radio transmitter) interval movements as they relate to sex, age, and habitat available to the animals. 3. Describe seasonal movements of PMJM. Evaluate difference in seasonal movements as they relate to sex, age, and habitat available to the animals. 4. Describe habitats where mice occur: movement corridors, end point descriptions (i.e., movement from what to what), and landscape features (connectivity with other riparian areas and other habitats used). 5. Estimate the mean amount of time PMJM spend in each available habitat. 6. Evaluate spatial and temporal variation in movement patterns ofPMJM. The objective of the habitat use study is to identify and refine habitat requirements of Z. h. preblei, including hibernation sites, and to determine if they influenced any of the deinographic parameters that will be estimated in this and complementary studies (see Shenk and Sivert 1999a, 1999b). METHODS See Shenk and Sivert (1999a, 1999b) for field methods. RESULTS Demography Study Trapping and PIT-tagging Effort A total of 186 individual PMIM were captured from the three study areas during the three 1998 trapping sessions (fable 1: 73 at Maytag Property, 77 at PineCliffRanch, and 36 at Woodhouse Ranch). These mice were captured over three different trapping sessions with an effort of 17,330 trap-nights. Every PMJM captured was PIT -tagged, providing each mouse with a unique, permanent, life-time identification marker. A total of 175 individual mice were captured on stream-side transects at three study sites during the 1999 field season including 34 mice PIT-tagged in 1998 (fable 1). These mice were captured over three different trapping sessions with an effort of 19,222 trap-nights. Other small mammals captured during the trapping sessions included Peromyscus spp., Mexican wood rat (Neotoma mexicana), house mouse (Mus musculus), vole (Microns spp.), western harvest mouse (Reithrodontomys megalotis), hispid pocket mouse (Chaetodipus hispidus) ,and shrew (Sorex spp.), See Shenk and Sivert (1999a) for details of 1998 field season. Reproduction During both 1998 and 1999, most adult mice captured exhibited evidence of active reproductive behavior, either pregnancy, lactation, or enlarged genitalia. We were able to locate one potential maternal nest in 1999. Juvenile mice were not captured during either June trapping sessions. Juvenile mice were captured at all three sites during both the July and September trapping sessions in both years. No estimates of reproduction could be made. �7 Survival Survival rate was estimated using the mark-recapture estimator for Pollock's Robust Design (see Kendall et al. 1995, 1997 and Kendall and Nichols 1995 for detail) in Program MARK (White and Burnham 1999). The robust design is a combination of the Cormack-Jolly-Seber (CJS)(Cormack 1964, Jolly 1965, Seber 1965) live recapture model and the closed capture models. The best fitting model combined data from all three study sites over all three trapping sessions in 1998. Survival was estimated as 0.75 (se = 0.033) for a five week period in 1998. Extrapolating this survival rate across the summer results in an over-summer (June 1- October 5, 18 weeks) survival rate of 0:355 (se = 0.056). No estimates of over-winter or summer survival in 1999 have been completed. Temporary emigration and immigration Temporary emigration rate and immigration was estimated using the mark-recapture estimator for Pollock's Robust Design in Program MARK. The best fitting model combined data from all three study sites over all three trapping sessions for these two parameters. Temporary emigration was estimated as 0.64 (se = 2.21), immigration was estimated as 0.45 (se = 89.94). Age and Sex Ratios As expected, no juvenile PMJM were captured during either June trapping session as it was too soon after hibernation for any litters to have been born and/or if born the juveniles would still be restricted to their nests. Juvenile mice were captured during both the July and September trapping sessions at all three sites in both years. Juveniles as small as 109 were captured. Sex ratios may be biased towards males, however further analyses need to be completed to confirm this hypothesis (Table 1). Density ~ Mark-recapture analysis was used to estimate abundance ( N) at each of the three study sites (Maytag Property, Woodhouse Ranch, and Pine Cliff Ranch) for each of the three trapping sessions in 1998 and for June 1999 (Table 2) using Program MARK (White and Burnham 1999) and Program CAPTURE (Otis et al. 1978, White et al. 1981). These abundance estimates and length of the trapline were used to estimate an unadjusted density of mice per kilometer of stream stretch. However, mice that typically don't use the area of stream stretch where the traps were placed could be trapped because they were attracted to the area by the artificial food source. Including these mice in the density estimates for the stream stretch covered by the trapline would artificially increase density estimates. Therefore, an adjustment parameter was estimated to calibrate density estimates for a more realistic estimate of PMJM per kilometer of stream length. Without the adjustment densities would be inflated. In collaboration with other PMJM researchers (M. Bakeman, C. Meaney, T. Ryon, and R. Schorr) population estimates (if) for a specified length of stream were determined with capture-recapture techniques using Program MARK. (White and Burnham 1999) and Program CAPTURE (Otis et al. 1978, White et al. 1981) for nine study areas over two years for early season trapping (June) only. To make these estimates comparable across study areas with unequal lengths of stream reaches trapped, mouse density (mice!km of stream) must be computed. However, traps tend to attract mice from some unknown N distance, so that naive density estimate of divided by stream length is biased high. To.remove this bias, radio-tracking data were used to estimate the proportion of tiine (P) radio-collared mice spent within the original trapline once the traps were removed. Data from six study areas (Table 3) provided by this study and the study conducted by T. Ryon at Rocky Flats were used to estimate this correction factor (P) for population estimates from linear traplines or grids. Only these study areas had radio-collared mice available from which to estimate the correction factor. Corrections were applied to all study areas with the function relatingp to trapliIie length (L) developed from these data. �8 The Michaelis-Menton model was fit to the observed p value: where L, is the length of the trapline for grid i, Nonlinear least squares were used to minimize the sum of squared Ej values to obtain an estimate of the unknown parameter, BSW (i.e., boundary strip width). Observations were weighted by the number of radio-collared mice providing the observed value of Pi. Because the variance of a proportion is a function of the value of the proportion, a logistic transformation was used to stabilize the variances of the Ei across the range of the Pi values: L, + 2BSW 1 - ----- L. I + Li 2BSW Predicted values of p (j3) are obtained from either fitted model for a particular trapline length (L) by substituting the estimate of BSW into the predictive equation: L A P = L + 2BSW· Estimates ofBSW for the two estimation methods are shown in Table 4, predicted values are shown in Table 3, and the fit of the two fitted equations is shown in Figure 1. The parameter estimate from the logistic transformation model was used to correct the population estimates, i.e., L A P = L + 2x41.5446 . This model was preferred because of the variance stabilization provided by the logistic transformation. However; as shown in Figure 1, the difference in the predictions of the two fitted equations is not biologically important. To obtain an adjusted number of mice on a trapline or trapping grid, the estimated population on the grid (if) from mark -recapture estimation is multiplied by the correction (ft) from the above equation to give the adjusted number: The variance of s; is computed as the variance of a product using the delta method: �9 where = var(ft) 4L 2 var(BSW) (L + 2BSW)4 Similarly, density (mice per unit of length of stream) is computed as the adjusted population size divided by the length of the trapline or the original trapline or grid population estimate over the adjusted length of the trapline: Nadf 15 = 15 is computed The variance of 15 = var(Nad) p, a trapline function of = L + 2BSW ~2 = 1 ~ var(N) + (L + 2 *BSW)2 SE(BSW)2 N ~ var(BSW) (L + 2 *BSW)4 . with SE(BSW) given in Table 4 .: As an example of the variance of length of 500m results in SE(.D) p = using the delta method: L2 Note that V"ar(BSW) L = ~var(.D) = 0.02696. Confidence intervals on the fitted are shown in Figure 2, computed as P±1.96xSE(.D). An alternative approach to computing confidence intervals on work better than the ±L96SE(ft) p that we would have thought would intervals shown in Figure 2 is to compute the confidence interval on the logit(p), and then back-transform the interval endpoints: 1 1 + 1 exp[ -(logit(.D) - 1.96SE(logit(.D)))' where SE(logit(.D)) = SE(BSW) . BSW 1 + exp[ -Vogit(.D) + 1.96SE(logit(.D))] = 9.1676/41.5446 = 0.22067 for the data presented here. After ... . computing confidence intervals by both methods, the results were only negligibly different, with the . maximum difference in the interval lengths being only 0.00111 over the range of L from 100 to 200Om. Riparian Vegetation Cover Data Total area of riparian shrubs, trees, herbaceous cover, non-vegetated, and open water were computed for each study area from riparian vegetation maps (Table 5). These riparian maps were developed from photo interpretation of infrared aerial photography at a resolution of 1:24,000. Areas of riparian vegetation were calculated when patches of vegetation were greater than 24 m wide. To estimate area of each riparian vegetation classification for each study area, stream length for each study area was delineated on the riparian vegetation map. A 200-m buffer was then generated on either side of the study area stream reach. Within this 200-m buffer total area (ha) of patches with dominant vegetation classified as riparian �10 herbaceous, riparian shrub, riparian tree, non-vegetated, and open water were calculated using ArcInfo (ESRI 1987). No ground truthing of vegetation classification was performed. Statistical Analysis ofPMlM Density and Riparian Vegetation Cover The adjusted density and areas of riparian vegetative cover per krn of stream (fable 5) were used to assess the relationship between PMJM density (mice/km of stream) and riparian vegetation cover. Estimates for multiple years for study areas were treated as separate estimates so that the year-to-year variation in PMIM density could be evaluated. An ANOVA ofPMJM density for study area and year effects suggested no differences between years (1998: = 35.08, SE = 8.85; 1999: = 31.04, SE = 6.29; P = 0.511) or across study areas (P = 0.245). The variation in PMJM density across study areas is what we want to explain with riparian vegetation cover differences across study areas. Individually, none of the vegetation cover variables were important predictors ofPMIM density (fable 6). When model selection following Burnham and Anderson (1998) was performed across all models involving the riparian shrub, tree, herbaceous, non-vegetated, and open water cover (ha!km stream ) variables, the best AlCc model was shrub and tree cover (fable 7) explaining 68% of the variation in PMJM density. The next best model included open water as well as shrub and tree cover, explaining 71 % of the variation ofPMIM density. The combination of riparian shrub and tree cover appears to be an important set of predictors for PMJM density based on the study areas included in this analysis. The parameter estimates for the best AlCc model are shown in Table 8, and a residual plot in Figure 3. All riparian vegetation cover variables have positive slope parameters, indicating a positive influence on PMIM density over the range of the vegetation cover variables considered in this analysis. x x Movement Study Radio-tracking effort . A total of 125 radio-collars were put on mice over the three trapping sessions in summer 1998 (Table 9: 48 at Maytag Property, 47 at PineCliffRanch, and 30 at Woodhouse Ranch). A total of 62 females and 63 males were radio-collared over the 1998 field season. A total of 138 radio-collars were put on mice over the three trapping sessions in summer 1999 (fable 9: 51 at Maytag Property, 52 at PineCliff Ranch, and 35 at Woodhouse Ranch). A total of 57 females and 81 males were radio-collared and tracked in 1999. Daily movements The general daily movement pattern throughout the active season was for mice to become active at dusk, remain active through the night, and to return to a day nest location at dawn. Most of our tracking was confined to night time movements; however we did make a greater effort in 1999 to confirm the mice were primarily in day nests during daylight hours. This was true for most daytime observations, however on occasion mice were active in the daytime as well. Percent ofPMIM locations, as determined from radio-tracking, at each of three distance categories were calculated for each study site and the latter two radio-tracking sessions in 1998 (see Shenk and Sivert 1999b for details). The majority oflocations at Maytag, PineCliff, and Woodhouse were within 46m of stream center for both. tracking periods (July-August and September-October). For all areas and tracking periods except Maytag during the last tracking session, 90% of PMIM movements were within 91m of either side of the center of the capture drainage. Mouse movements at Maytag during September-October resulted in 32.6% of the locations being> 91m from the stream center. Similar patterns were observed for 1999. . During nightime activity periods more than one mouse would often be found foraging in the same area. These foraging areas were often the same areas used by the same mice on numerous consecutive nights �11 Seasonal movement Field methods used in June 1998 did not allow us to document accurate June movement patterns for that year. The distribution of mouse locations in 1998 at Maytag Property for the July-August tracking session is different from the distribution of PMJM locations for the September-October tracking session. In 1998, the pattern shifted from heavy use along East Plum Creek to the north of the trapline to a more concentrated distribution either side of the trapline. The more concentrated use of the areas east and west of the trapline also resulted in locations being further from the center of the creek. The northern end of the trapline is largely vegetated by willows with the remainder of the trapline more diversely vegetated. In 1999 we were able to document movements for all three tracking sessions. Greatest movement away from the center of the stream occurred in the August session. In 1998, mouse movements were more concentrated along the drainages at PineCliff Ranch during the September-October tracking sessions than during the July-August tracking sessions. Vegetation along both West Plum Creek and Garber Creek along the trapline was primarily willow. In 1999, greatest movements from either stream center were documented during the August tracking session. At Woodhouse Ranch, the distribution of mouse locations also shifted between tracking sessions. Mouse movements during June 1999 were the most diverse of all tracking sessions. Mouse movements were more concentrated along the southern half of the trapline in September-October as compared to the more even distribution of mouse locations recorded for July-August in both 1998 and 1999. The northern end of the trapline is dominated by willows, the middle and southern end of the trapline is in more diverse riparian habitat. In September 1999 we had fewer mice radio-collared at each of the three sites because we were not able to trap mice. We assume the majority of the mice had already hibernated by the time we started our last trapping effort on September 9, 1999. Over the two-year study, 27 PMIM were captured, radio-collared, and tracked for more than one tracking session (fable 10) . Other mice were captured during more than one trapping session but radio collars were not replaced either because all the radio collars had been put on other mice or because the physical condition of the animal was such that we decided not to replace the radio-collar. Such conditions included mice who had significant hair worn off the neck from the previous radio collar. In no incidence did we find open wounds caused by radio collars. Of the 27 mice radio-collared for more than one session, we have only analyzed the mice captured in 1998. Of those, four provided information to evaluate seasonal changes in areas and habitats use between the July-August tracking period and the September-October tracking period: One mouse, a male from Maytag was observed only once during the September-October tracking session, providing no information concerning seasonal changes in movement patterns. Thus, changes in areas used by individual mice between June and either the July-August or September-October tracking session are not summarized here. A female at Maytag exhibited a seasonal movement shift, small sample size prohibited any evaluation of the movement patterns of the Maytag male (n = 1 in September-October). There does not appear to be any shift in areas used by the male tracked during both latter sessions at Pine Cliff Ranch, however, sample sizes were small (n = 17, 14). Two females at Woodhouse Ranch exhibited a shift in areas used between July-August and September-October. However, caution should be used in interpreting these results because of low sample sizes in the latter tracking session (n = 14,6). Annual variation in movement patterns Preliminary comparisons of mouse movements from 19981:0--1999 for tracking sessions August and September were completed. Mouse movements during August tracking sessions were similar at Maytag Property. More mice were captured on the southern half of the Maytag Property in 1999 and this is reflected in the greater number of locations in the southern half of the property for that year. The same area located outside a 100m strip from the stream center was used both years during this tracking session and was comprised primarily of a seep area. August movements at Pine cliff Ranch were further from the �12 stream in 1999. Mice were also found using an ephemeral stream in August at PineCliff Ranch in 1999. This stream was dry at the time. Similar mouse movements were observed at Woodhouse Ranch over both August tracking sessions although movements were more concentrated in the north in 1998 and more concentrated in the south in 1999: Temporal comparisons of mouse movements in the September tracking sessions are not entirely comparable because fewer mice were followed in the 1999 September tracking session because we were not able to capture mice. We assume a large portion of the mice in 1999 were already hibernating when we started our September trapping and radio-collaring session. We did not observe the greater movements away from the stream center at Maytag in 1999 that we observed there in 1998, however we probably missed most of the movements to hibernation sites. Movement patterns at PineClifffRanch and Woodhouse Ranch were similar for both September tracking sessions with movements being more concentrated near the stream. The same feeding hotspot at Woodhouse Ranch was used both years in September. Habitat Use Detailed vegetation maps were created using Global Positioning Systems. Combining these vegetation maps and locations of mice we will be able to describe in further detail habitat use. These analyses are not yet completed. Nest sites Numerous daytime nest sites were located at all three study sites from both years. Most nest sites were made of tightly woven vegetation, located on the ground. However, we also located underground daytime nests in summer 1999. Nest material included leaves, grass, and small sticks. There was only one small entrance hole on each of the nests found. When observing mice in their nests the mice would peer out of the hole and remain motionless as long as we were present. One possible maternal nest was located in 1999. This possible maternal nest was underground, made of densely packed grasses with the burrow lined as well. There also were several locations at each of the study sites where adult females returned. to repeatedly. Each of these sites were in patches of extremely dense vegetation or underneath a large downed log. Hibernacula Potential hibernation sites were located by noting stationary radio-telemetry signals over repeated nights. These signals were coming from underground with no evidence of predation or slipped. collars. Eight potential hibernacula were located, at least one at each study site in 1998. In 1999 a total of 13 potential hibernation sites were located. Three were located at Woodhouse Ranch (1 female, 2 males) and 10 sites at PineCliffRanch (2 females, 8 males). In contrast to 1998 data, these potential hibernation sites were located closer to the stream. However, at both Pine Cliff and Woodhouse Ranches stream banks rise out of the floodplain in closer proximity to the stream than at Maytag Property where the potential hibernation sites located in 1998 were located long distances from the stream center. Vegetation characteristics of the eight sites located during both 1998 and 1999 are similar to other hibernacula described. for PMJM. Hibernation sites are considered. potential until the sites can be dug up to confirm a hibernation chamber. One male mouse was dug up at Pine Cliff Ranch in 1999 to confirm the site as a true hibernation location and to gain information on the hibernation nest and chamber. The mouse was in the chamber and was not aroused when the earth was dug out around him. Once we finished our assessment of the nest the soil was put back in place, leaving the mouse in the chamber. The remaining hibernation sites will be dug up in June 2000 so as not to disturb other hibernating mice. We hope to confirm these sites as hibernation sites then and will document features of the hibernation sites at that time. �13 Mortality factors Mortalities factors of radio-collared mice in 1998 included predation by rattlesnake, garter snake, fox, and house cat. Four probable predations were also noted and identified by finding tightly crimped (i.e., no possibility of the mouse having slipped the collar over its head) radio-collars lying on the ground. Two accidental deaths were documented, a road kill and drowning. Mortality factors also included trapping andlor handling mortalities and unknown causes. In 1999 mortality factors also included predation by bullfrogs, weasel, and yellow-bellied racer (Table 11). Nine mice also died once they entered a state of torpor above ground. These mice either succumbed to cold or were predated on. Fecal analysis The amount of bait found in each of the fecal samples was quantified as either 100%, abundant (> 70%), trace « 10%) or 0%. The following summaries are made after eliminating all samples of 100% bait, and then only classifying the fecal sample contents other than bait. Fecal analyses indicate a seasonal shift in diets. The most common item found in the fecal samples during the June trapping session at all three sites for 1998 were arthropods. This was true for three of the five June samples collected at Maytag, eight often June samples at PineCliff, and four of five June samples collected at Woodhouse. Other common items included endogenous fungus (1) and seed (1) at Maytag; endogenous fungus (2) at PineCliff,; and Poa (1) at Woodhouse. During the July 21-August 4, 1998 trapping session the most common items in the fecal samples at Maytag were arthropod (2), endogenous fungus (1), pollen (1), and Carex (1). During the same trapping period, the most common items in five of eight samples collected at PineCliff were endogenous fungus, the other three samples having a majority of arthropod (1), moss (1), and pollen (1). The two samples from Woodhouse during this trapping session were composed primarily of either mushroom or seed. On August 13, 1998, nine samples were collected during an extra trapping session (on the same trapline as the other trapping sessions occur) with the most common items in the fecal samples being moss (4), endogenous fungus (3) or pollen (2). All three samples collected from PMJM captured at a back drainage on August 23, 1998 at Woodhouse Ranch were primarily fungus. The September 1998 trapping session at Maytag yielded samples with majority fecal contents of arthropod (2), moss (2), pollen (2), endogenous fungus (1), and seed (1). Ten samples from PineCliff during September 1998 had majority fecal contents of arthropod (3), endogenous fungus (3), and seed (3). Eight of nine samples taken from Woodhouse contained primarily arthropods, with one a trace of seed. A total of329 fecal samples were collected during the 1999 field season. Analyses of these samples have not been completed. We also collected 16 stomach samples from dead mice. Analyses of these stomach samples are not yet completed. DISCUSSION Information on the population dynamics of PMJM is necessary to determine which areas and habitats support viable populations. To begin to evaluate the viability of a population information on key demographic parameters must be obtained. Conducting studies on individually marked animals provides the greatest insight on the demography of a population. In general, estimated sex ratios from this study are comparable to those found by Armstrong et al. (1997) who reported an overall sex ratio for all captured PMIM of51.6 males: 48.4 females; approximately 86.0% of captures were identified as adults. There is a possible male sex bias at PineCliff Ranch. However, small sample sizes and possible trapping biases by sex may explain the discrepancy. Density estimates were expected to increase as the summer progressed to account for the birth pulses in late June, and late July-August. In 1998, this was the trend observed at Maytag. PMJM densities at Woodhouse Ranch increased from June to July but did not increase further during the September trapping session. Highest densities occurred at PineCliff, where the vegetation is primarily willow and both a �14 mainstern and tributary were used by mice. Lowest densities occurred at Woodhouse Ranch where the riparian vegetation has fewer willow but was dense with other riparian vegetation. Vegetation at Maytag provided some areas of dense willow with the remaining areas being of moderate density of riparian vegetation. The lower density of PMJM reported for Woodhouse Ranch might be explained by the composition and densities of other small mammals at that site. Woodhouse Ranch had the highest captures of both house mice and voles of the three sites studied. Defining summer as June 1 - October 5, over-summer survival was estimated as 0.36 (se = 0.056) over all three study sites. Meaney et al. (1999) report a one-month summer survival rate of 78%. Extrapolating Meaney et al.'s (1999) estimate over our summer period (- four months) would result in a similar oversummer survival rate estimate of 36%. Prior to studies conducted in 1998 no information existed on survival rates for populations of Z. h. preblei although Whitaker (1963) reported a 67% loss of Z. hudsonius over hibernation. Temporary emigration and immigration rates were estimated but both had extremely high variances associated with those estimates. Thus these estimates cannot be used, with any confidence, to provide information on movements of mice into and out of these populations. Very little new information was gained during this study on reproductive parameters of PMIM. Most adult mice captured exhibited evidence of active reproductive behavior, either pregnancy, lactation, or enlarged genitalia. Juvenile mice were not captured during the June trapping session. Juvenile mice were captured at all three sites during both the July and September trapping sessions. Given that breeding peaks appear to occur in early to mid-June and August with a possible third litter in September (Whitaker 1963) this was not unexpected and agrees with previous observations (Meaney et al. 1996, 1997, PTI 1996a, M. Bakeman unpublished data, T. Ryon unpublished data) Preliminary analysis of the movement data collected on radio-collared PMIM in 1999 supported results found in 1998. In particular we were able to document again in this second field season that PMIM exhibit (1) greater use of upland habitats than previously assumed, (2) general site fidelity to both daytime nesting sites and nighttime feeding sites, (3) seasonal shifts in movement patterns, and (4) use of both perennial and intermittent tributaries adjacent to the capture drainage. Jumping mice of the genus Zapus are true hibernators, spending much of their lives in hibernation. Meadow jumping mice spend approximately 7 months (-210 days) per year in hibernation (Quimby 1951) whereas estimates for Z. princeps indicate that some populations (e.g., in the western mountains of Utah) spend up to 300 days per year in hibernation (Cranford 1983). Jumping mice hibernate in underground burrows (Quimby 1951, Whitaker 1963). They are excellent burrowers and create their own hibemacula. Meadow jumping mice are generally solitary hibernators, however, there have been occurrences of more than one mouse found in a single hibernaculum. Eight possible hibernacula were located during 1998. Five of the eight mice using these possible hibernacula traveled ~ 90 meters from the center of their typical September night time locations. Of the 13 possible hibernacula located in 1999, three sites were located at Woodhouse Ranch and ten possible hibernacula at Pine Cliff Ranch. In contrast to 1998 data, these potential hibernation sites were located closer to the stream. However, at both Pine Cliff and Woodhouse Ranches stream banks rise out of the floodplain in closer proximity to the stream than at Maytag Property where the potential hibernation sites located in 1998 were located long distances from the stream center. Vegetation characteristics of all the potential hibernacula located over both years are similar to other hibemacula described for PMIM. One confirmed hibernaculum, located on Rocky Flats Environmental Technology Site, used by Z. h. preblei has been located (Armstrong et al. 1997). This site was 9m above a creek bed (Walnut Creek); it had a thick cover of chokecherry (Prunus virginianat and snowberry (Symphoricarpos spp.), the mouse was found in a leaf litter nest 30cm beneath the ground in coarse textured soil (Armstrong et al. 1997). Four possible hibernacula were located by tracking radiotelernetered mice at the U. S. Air Force Academy in fall 1997. These sites are located 7, 12,29, and 31m from a creek bed (R. Schorr, unpublished data). There was no consistency among sites in aspect. Three sites were in vegetation dominated by coyote willow (Salix exigua), one site was in vegetation dominated by snowberry and mullein (Verbascum thapsus). However, all four hibemacula appeared to be below �15 coyote willows. The eight sites located during this study and the four U. S. Air Force Academy sites were not disturbed to protect any hibernating mice and therefore are only possible hibernacula because there is no confirmation a mouse actually hibernated there. Confirmation of a true hibernaculum cannot be made until a chamber, or nest is located. The other explanation for these collar locations might be either locations of radios discarded by the mice or dead mice carried underground by a predator. Location or more hibernation sites was limited primarily by normal battery failure of the radio transmitters before mice went into hibernation. Prior to the 1998 field season, natural mortality factors reported for Z. hudsonius included only insufficient fat storage prior to hibernation (Whitaker 1963), predation (Whitaker 1963, Poly and Boucher 1997, R. Schorr unpublished data) and cannibalism (Sheldon 1934). Other assumed natural mortality factors for Z. h. preblei included starvation, exposure, and disease. Natural mortality factors documented during this study included predation by house cats, garter snakes, rattlesnakes, yellow-bellied racers, bullfrogs, weasel, and fox as well as accidents by drowning and road kilL In 1999 mice were also found in torpor throughout the summer, often in very exposed areas frequently resulting in death by either exposure or predation. Use of ephemeral drainages was observed at both Maytag Property (in l 998) and Pine Cliff Ranch (in 1999), Two mice moved away from East Plum Creek at Maytag Property in September 1998 and focused their movements -300 meters from their previous locations, up a dry drainage dominated by upland grasses and gambel oak. These two mice continued to use this new area and were not observed again near East Plum Creek for at least two weeks. Normal radio-failure (i.e., batteries failed after -4 weeks) after this time did not allow us to determine if the mice ever returned to East Plum Creek or if they hibernated in their new location. Use of the ephemeral drainage at Pine Cliff Ranch occurred in the August tracking session. The high percentage of arthropods and endogenous fungus found in the fecal samples provides a new aspect to evaluating PMlM habitat requirements. When considering habitats used by PMJM, or in trying to predict habitats that might be suitable for the subspecies, consideration should be given as to whether those habitats could support the arthropods and fungus apparently being selected for by the mouse. Arthropods are a food source high in protein and fat which would benefit mice emerging from hibernation and mice preparing for hibernation. Whitaker (1963) reported a 67% loss of individuals over hibernation and that average body mass of individuals emerging from hibernation was greater than the average for mice entering hibernation. Because no mice are known to store food in their hibernacula, this indicates that the lighter individuals died during hibernation and only those entering with higher masses survived. All the energy they use during hibernation and the periodic arousals (the energetically most expensive part of hibernation) must be the fat they carry into hibernation (B. Wunder, personal communication). The ability to put on sufficient fat for overwinter survival during hibernation is a critical factor in the life history of these mice. Thus, appropriate and sufficient food sources must be available to the mouse to meet these nutritional requirements. The apparent seasonal shift in mouse movements between the July-August and September tracking sessions may be a result of diet switching. The broader diets suggested by the fecal analyses from samples collected during July and August possibly represent both a Wider availability of suitable foods and the ability ofPMlM to exploit these resources. It might also suggest a need to exploit these other resources to provide the mouse with necessary food requirements for breeding. Because mice tracked during each session were not generally the same mice there is a possibility these apparent movement shifts might only be different areas used by different mice. The probability of this being the case is small for two reasons. The first is the low density of mice in the stream (Shenk and Sivert 1999). Given that, the proportion of mice followed during each session is a high proportion of the mice in the population. Thus, each subset of mice should be representative of the population. Secondly, evidence from the four mice followed during both the latter tracking sessions also exhibited movement shifts between sessions. The combination of shifts in both general mouse movements, individual mouse movements, and diet provide strong circumstantial evidence that PMlM may be selecting for or require specific seasonal diets. If �16 this is the case, all these requirements must be considered and provided for to ensure conservation of the subspecies. A detailed literature search and further studies need to be conducted to investigate the possible implications of these dietary patterns. In comparing movement patterns from 1998 to 1999 we were able to evaluate annual variation in daily and seasonal movement patterns of PMJM. General patterns that emerged from the comparison of the two years included (1) similar areas being used by the mice over both years, (2) greater use of upland habitats than previously assumed, (2) general site fidelity to both daytime nesting sites and nighttime feeding sites, (3) seasonal shifts in movement patterns, and (4) use of both perennial and intermittent tributaries adjacent to the capture drainage. This study looked at PMlM movements at only three sites. These sites were selected based on known presence of PMlM. Other sites, perhaps because of different habitat configurations or sites of poorer quality may require mice to move further from the creek or may allow mice to remain closer to the creek in order to obtain all life requirements. However, these study sites were specifically selected to address spatial variation in movement patterns of PMJM due to different spatial configuration and juxtaposition of habitats. And although all study sites were different they could all be considered within the general description of what is consider typical habitat for PMlM. It should also be noted that we generally trapped along the creek. Thus, mice captured and subsequently radio-collared and followed were those mice that use the area near the most prominent drainage. If there are mice that do not regularly use the main channel and thus were not available for capture there would be a bias in the data towards fewer observations 300 feet away from the creek. To test for such a bias, traps were placed 300 m from the stream center at Maytag and Woodhouse Properties during suminer 1999 as part of a Master's Project conducted by a student ay Colorado State University. Capture of mice in transects placed away from the stream lends further support for use of habitats further from the creek than initially thought. Mice captured 300 meters from the stream moved near their capture site and between the capture site and the stream. The prevailing idea of what constitutes quality PMlM habitat is amount of riparian shrubs. Therefore, we focused our analysis on describing the relationship between PMIM densities and amount of various riparian vegetation cover, including riparian shrubs. This analysis is not meant to identify all critical components ofPMlM habitat. For example, ifa necessary component ofPMlM habitat is open water, but not necessarily in any large amount, this analysis would not detect this relationship. The best model describing the relationship between PMlM density and riparian vegetation cover included shrub and tree cover, the next best model included open water as well as shrub and tree cover. Tree cover may also reflect shrub and herbaceous cover, because the tree canopy likely may be covering these other understory vegetation classes. Thus, results from this analysis appear to support the hypothesis that riparian shrubs are an important component ofPMlM habitat. However, because the study areas included in this analysis were not randomly selected from all possible PMJM habitats, the conclusions reported here must be treated with some caution. To truly assess the relationship between PMJM density and riparian vegetation would require estimating densities from a random selection of areas presumed to have suitable PMlM habitat. Analysis of how the riparian vegetation characteristics at these study sites relates to their PMlM densities would then provide an unbiased estimate of the relationship. With the limited data available, 68% of the variation in PMIM density is explained by a model that includes riparian shrub and tree cover (ha!km stream), as identified by the vegetation mapping techniques used in this study. These results suggest that habitat quality ofPMIM can be predicted by the shrub and tree cover available on a site. This report includes results from only the first two years of a multi-year project to follow individually marked PMIM through time. Further analyses of these data, collection of more data, and more years of data, will continue to improve our ability to evaluate demographic parameters estimates and movement patterns ofPMlM and of how they vary across space and time. Preliminary results from the demography, movement and distribution studies of PMIM have suggested the need to continue with research currently �17 being addressed in these three studies. However, the preliminary results also suggest further research be conducted on (1) use of upland habitat by PMlM, (2) refining water requirements ofPMJM (i.e., do they require stream habitat or are wetland areas sufficient), (3) range-wide distributional boundaries (e.g., elevation restrictions), and (4) investigating areas of potential sympatry or hibridization with Z. princeps (western jumping mouse). The demography and movement studies will be modified during the summer 2000 field season to further investigate upland use and water requirements. Acknowledgments Additional data on PMlM densities for the vegetation analysis were provided by Mark Bakeman, Rob Schorr, Carron Meaney, and Tom Ryon. Bruce Lubow provided helpful comments on the first drafts of the adjusted PMJM density correction method, and computed the estimates ofPMJM population size for several of the study areas included in this analysis. Seth McClean with CDOW provided the riparian vegetation data. Mark Bakeman and Tom Ryon provided useful comments on earlier drafts of the vegetation analysis. Literature Cited Armstrong, D. M., M. E. Bakeman, A Deans, C. A Meaney, and T. R. Ryon. 1997. Conclusions and recommendations in: Report on habitat findings on the Preble's meadow jumping mouse. Edited by M. E. Bakeman. Report to USFWS and Colorado Division of Wildlife. Burnham, K. P., and D. R. Anderson. 1998. Model selection and inference: a practical informationtheoretic approach. Springer-Verlag, New York, New York, USA 353pp. Cook, T. D. and D. C. Campbell. 1979. Quasi-experimentation: design and analysis issues for field settings. Houghton-Mifflin, Boston. Cormack, R. M. 1964. Estimates of survival from the sightings of marked animals. Biometrika 51:429438. Cranford, J. A 1983. Ecological strategies ofa small hibernator.jhe western jumping mouse Zapus princeps. Canadian Journal of Zoology 61:232-240. ESRI. 1987. ARCIINFO users manual. Environmental Systems Research Institute. Redlands, California, USA Jolly, G. M. 1965. Explicit estimates from capture-recapture data with both death and immigration stochastic model. Biometrika 52:225-247. Kendall, W. L., and J. D. Nichols. 1995. On the use of secondary capture-recapture samples to estimate temporary emigration and breeding proportions. Journal of Applied Statistics.22:751-762. Kendall, W. L., K. H. Pollock, and C. Brownie. 1995. A likelihood-based approach to capture-recapture estimation of demographic parameters under the robust design. Biometrics 51:293-308. Kendall, W. L., J. D. Nichols, and J. E. Hines. 1997. Estimating temporary emigration using capturerecapture data with Pollock's robust design. Ecology 78:563-578. Krutzsch, P. H. 1954. North American jumping mice (genus Zapus). University of Kansas Publications, Museum of Natural History 7:349-472. Meaney, C. A, N. W. Clippinger, A. Deans, and M. OShea-Stone. 1996. Second year survey for Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. Report prepared for the Colorado Division of Wildlife. Meaney, C. A, A Deans, N. W. Clippinger, M. Rider, N. Daly, and M. O'Shea-Stone. 1997. Third year survey for Preble's meadow jumping mouse (Zapus hudsonius preblei) in Colorado. Report prepared for the Colorado Division of Wildlife. Meaney, C. A, A. Ruggles, B. Lubow, N. W. Clippinger, and A. Deans. 1999. Preliminary results: Second year study of the impact of trails on small mammals and population estimates for Preble's meadow jumping mice on City of Boulder Open Space. Report for Greenways Program, City of Boulder Transportation Agency, Region 8. Department, City of Boulder Open Space, and Environmental Protection �18 Otis, D. L., K. P. Burnham, G. C. White, and D. R. Anderson. 1978. Statistical inference from capture data on closed animal populations. Wildlife Monograph 62: 1-135. Poly, W. J., and C. E. Boucher. 1997. Record ofa creek chub preying on a jumping mouse in Bruffey Creek, West Virginia. Brimleyana 24: 29-32. PTI Environmental Services. 1996a. Preble's Meadow Jumping Mouse Study at Rocky Flats . Environmental Technology Site, Annual Report 1996. Final. Rocky Flats Environmental Technology Site, Golden, Colorado. PTI Environmental Services. 1996b. Preble's Meadow Jumping Mouse Study at Rocky Flats Environmental Technology Site, Spring 1996. Final. Rocky Flats Environmental Technology Site, Golden, Colorado. Quimby, D. C. 1951. The life history and ecology of the jumping mouse, Zapus hudsonius. Ecological Monographs 21:61-95. Seber, G. A. F. 1965. A note on the multiple recapture census. Biometrika 52:249-259. Sheldon, C. 1934. Studies on the life histories of Zapus and Napaeozapus in Nova Scotia. Journal of Mammalogy 15:290-300. Shenk, T. M. 1998. Conservation assessment and preliminary conservation strategy for Preble's meadow jumping mouse (Zapus hudsonius preblei). Colorado Division of Wildlife FY1997-98 Annual Report. Shenk, T. M. and Sivert M. 1999a. Temporal and spatial variation in the demography of preble's meadow jumping mouse (Zapus hudsonius prebleiy. Colorado Division of Wildlife. Annual Report 1999. Shenk, T. M. and Sivert M. 1999b. Movement patterns of Preble's meadow jumping mouse (Zapus hudsonius preblei) as they vary across time and space. Colorado Division of Wildlife. Annual Report. Whitaker, J. 0., Jr. 1963. A study of the meadow jumping mouse, Zapus hudsonius (Zimmerman), in cental New York. Ecological Monographs 33:3. White, G. C., D. R. Anderson, K. P. Burnham, and D. L. Otis. 1982. Capture-recapture and removal methods for sampling closed populations. LA-8787-NERP, Los Alamos National Laboratory, Los Alamos, New Mexico, USA. 235 pp. White, G. C. and K P. Burnham. 1999. Program MARK: survival estimation from populations of marked animals. Bird Study 46 Supplement: 120-138. Table 1. Total number of individual Preble's meadow jumping mice (Zapus.hudsonius preblei) captured in 1998, new individuals captured in 1999, recaptures in 1999 from 1998, and total number of individual mice caEtured in 1999 at three studi: sites. Total individuals captured 1998 Site New captures 1999 Recaptures in 1999 from 1998 Total individuals captured 1999 F M U Total F M Total F M Total F M Total Maytag Property 34 38 1 73 19 27 46 4 10 14 23 37 60 PineCliff Ranch 25 52 0 77 19 29 48 4 8 12 23 37 60 Woodhouse 17 18 1 36 17 30 47 3 5 8 20 35 55 TOTAL 75 108 2 185 55 86 141 11 23 34 66 109 175 �19 Table 2. Stream reach abundance estimates (N) for Preble's meadow jumping mouse (PMJM) from three sites in Douglas County, Colorado for three traEEing sessions in 1998 and for June 1999. 95% Confidence Interval Trapping Session Site Maytag Maytag Maytag PineCliff PineCliff PineCliff Woodhouse Woodhouse Woodhouse Maytag Woodhouse PineCliff June98 July98 September98 June98 July98 September98 June98 July98 September98 June99 June99 June99 if seeN) Lower Upper 18 31 44 30 17 51 11 20 20 35 32 28 2.7 11.4 1.8 5.7 0.9 3.3 3.8 5.2 3.1 3.8 3.2 2.1 15 20 42 24 16 48 7 15 17 24 56 52 21 42 46 36 18 54 15 25 23 39 69 61 Trapline Length (m) Adjusted Density (pMJM/krn) 550 608 494 490 504 504 510 516 574 1130 504 503 26.2 41.3 69.5 47.7 26.3 78.9 16.8 30.7 27.8 31.2 62.6 56.5 Table 3. Data on the proportion of locations (P) of Preble's meadow jumping mouse locations within the area defined by parallel lines running through either end of the trapline, sorted into ascending order by traEline length. Site Year Session Meanp Weighted Fit Logistic Trapline No. SE(P) Length Mice Online Weighted Fit Walnut Creek 1999 May/June 5 0.946 0.0040 0.761 0.796 325 A-upper Woodhouse 1999 June 24 0.629 0.0843 0.800 0.831 409 Woodhouse 1999 July 409 19 0.902 0.0532 0.800 0.831 Woodhouse 1999 1.000 Sept 6 0.0000 0.800 0.831 409 Walnut Creek 1999 May/June 0.308 490 1 0.827 0.855 Lower Maytag 1998 Sept 16 0.954 0.0385 0.829 494 0.856 Walnut Creek 1999 May/June 0.828 0.830 500 1 0.858 B-series PineCliff 1999 12 0.841 0.0670 0.831 0.858 June 503 PineCliff 1999 Sept 503 15 0.924 0.0464 0.831 0.858 PineCliff 1999 July 22 0.899 0.831 503 0.0372 0.858 PineCliff 0·.0750 1998 July 14 0.805 0.831 504 0.858 PineCliff 1998 16 0.671 0.0964 0.831 Sept 504 0.858 Woodhouse 1998 12 July 0.817 0.0712 0.835 0.861 516 Woodhouse 1998 Sept 14 0.883 0.0291 0.849 0.874 574 Maytag 1998 July 608 16 0.719 0.0744 0.856 0:880 Maytag 1999 11 0.917 Sept 1130 0.969 0.1664 0.932 Maytag 1999 July 1130 26 0.908 0.0364 0.917 0.932 Mavtag 1999 June 7 0.873 0.917 0.932 1130 0.1339 �20 Table 4. Estimates of boundary strip width (BSW) for nonlinear weighted least squares fit and nonlinear weighted least sguares fit with the logistic transformation. Method Estimate of BSW SE of estimate Nonlinear weighted least squares 51.1241 10.1033 Nonlinear weighted least squares with logistic transformation 41.5446 9.1676 Table 5. Data used to evaluate the relationship between Preble's meadow jumping mouse density and riparian vegetation cover. Adjusted Area (ha/km) Adjusted SE Stream InvestiStudy Area Year Density Adjusted Length HerbaNonOpen gator Tree (mice/kin) Density Shrub (km) ceous Vegetated Water Shenk Maytag 1998 32.52 5.84 0.2619 2.4643 2.8571 1.3452 2.0357 0.84 Shenk Maytag 1999 29.04 3.05 0.1549 1.8239 1.9296 1.0000 1.7817 1.42 Shenk Pinecliff 1998 57.35 18.39 5.4324 3.3649 1.2432 0.0000 1.5946 0.74 Shenk Pinecliff 1999 53.79 5.52 5.4324 3.3649 1.2432 0.0000 1.5946 0.74 Shenk Woodhouse 1998 15.52 3.14 2.7600 0.0000 1.7200 0.0000 0.0000 0.50 Shenk Woodhouse 1999 48.52 3.52 2.7600 0.0000 1.7200 0.0000 0.0000 0.50 Bakeman Dirty Woman Cr. 1998 20.46 7.29 2.9190 1.3646 0.2987 0.0861 0.0354 3.95 Bakeman Dirty Woman Cr. 1999 6.89 2.11 1.3769 1.8396 0.4403 0.0000 0.0522 2.68 Bakeman Castle Rock 1999 41.70 26.91 3.8086 0.1584 0.2145 1.0066 1.1683 3.03 Schorr Monument Cr. 1998 67.08 13.34 5.9363 0.0000 0.4968 0.8535 0.0000 1.57 Schorr Monument Cr. 1999 28.75 8.52 5.9363 0.0000 0.4968 0.8535 0.0000 1.57 Meany South Boulder Cr. 1998 48.80 10.60 0.1640 27.4270 3.8629 0.0000 0.1933 4.45 Meany South Boulder Cr. 1999 34.50 7.75 0.1640 27.4270 3.8629 0.0000 0.1933 4.45 Ryon Walnut Cr. 1999 5.10 3.34 0.1231 1.0865 0.0416 0.0000 1.0266 6.01 Ryon Rock Cr. 1998 3.80 0.36 0.3815 0.9505 0.0000 0.0786 0.0219 14.13 Table 6. Results of linear regressions of Preble's meadow jumping mouse density (micelkm stream) predicted by riparian vegetation cover (ha/km stream ) for 15 study area and year combinations from Table 5. Variable Shrub Herbaceous Tree Non-vegetated Total Cover Open Water Intercept Slope P R2 30.22 0.63 0.638 0.018 31.79 0.06 0.656 0.016 30.80 0.67 0.495 . 0.037 31.51 2.18 0.729 0.01l 31.23 0.06 0.596 0.022 35.84 -2.35 .0.469 0.041 -_" �21 Table 7. Sununary of AlCc model selection for the riparian vegetation cover (haIkm stream) variables Shrub, Tree, Herbaceous, Non-Vegetated, and Open Water predicting Prebles' meadow jumping mouse density (mice/km stream). AlCc, !:1-AlCc, and Akaike Weights follow definitions in Burnham and Anderson (1998). Akaike R2 !:1-AlCc AlCc Variables in Model Weights 83.5040 0.0000 0.6831 0.59584 Shrub Tree 86.6162 3.1122 0.7143 0.12570 Shrub Tree Open Water 87.4205 3.9165 0.08408 0.6986 Shrub Tree Non-Vegetated 88.1465 4.6425 0.6836 0.05848 Shrub Tree Herbaceous 89.0245 5.5205 0.5421 0.03770 Shrub Herbaceous 89.3639 5.8599 0.6569 0.03182 Shrub Herbaceous Open Water 90.9029 7.3989 0.6198 0.01474 Shrub Herbaceous Non-Vegetated 91.1034 7.5994 0.3216 0.01333 Shrub 9l.8340 8.3300 0.7258 0.00925 Shrub Tree Herbaceous Open Water 92.3015 0.7171 8.7975 0.00732 Shrub Tree Non-Vegetated Open Water 92.8597 9.3557 0.7064 0.00554 Shrub Tree Herbaceous Non-Vegetated 93.5258 10.0218 0.3819 0.00397 Shrub Open Water 94.1043 10.6003 0.6810 0.00297 Shrub Herbaceous Non-Vegetated Open Water 94.5170 0.3396 1l.0130 0.00242 Shrub Non-Vegetated 95.4143 1l.9103 0.0957 0.00154 Tree 96.2135 0.0462 12.7095 0.00104 Open Water 96.3496 0.0375 12.8456 0.00097 Herbaceous 96.3621 12.8581 0.0367 0.00096 Non-Vegetated 0.3828 98.1688 14.6648 0.00039 Shrub Non-Vegetated Open Water 98.5520 0.1358 15.0480 0.00032 Tree Non-Vegetated 98.7044 15.2004 0.1270 0.00030 Tree Open Water 98.8480 15.3440 0.7345 0.00028 Shrub Tree Herbaceous Non-Vegetated Open Water 0.1059 99.0626 15.5586 0.00025 Herbaceous Non-Vegetated 0.1035 99.1022 15.5982 0.00024 Tree Herbaceous 15.6879 0.0982 99.1919 0.00023 Herbaceous Open Water 99.8370 0.0585 16.3330 0.00017 Non-Vegetated Open Water 0.1459 103.0432 19.5392 0.00003 Tree Non-Vegetated Open Water 0.1358 19.7144 103.2184 0.00003 Tree Herbaceous Non-Vegetated 0.1296 103.3266 19.8226 0.00003 Herbaceous Non-Vegetated Open Water 0.1273 103.3666 19.8626 0.00003 Tree Herbaceous Open Water 108.8576 25.3536 0.1470 0.00000 Tree Herbaceous Non-Vegetated Open Water Table 8. Parameter estimates for best AlCc model explaining PMlM density (mice/km stream length) from . vegetative cover variables (haIkm stream length). Standard Variable t statistic Estimate Pr> I~ Error 0.892 Intercept 7.1047 0.14 0.9853 Shrub <0.001 7.2342 l.5339 4.72 Tree 0.003 10.1308 2.7381 3.70 �22 Table 9. Number of Preble's meadow jumping mice radio-collared at each study site during each trapping session for each ~ear. 1999 1998 Site Session Trap nights Females Males Trap Nights Females . Males Total Total 2604 6 10 12 742 1442 1921 1651 3 7 6 7 7 7 5 10 17 12 13 1215 2114 8 3 9 5 3 125 19222 57 81 Maytag Maytag Jun Jul 1949 2370 9 11 5 5 14 16 Maytag Sep Jun Jul Sep 3256 10 5 6 4 2 8 18 3191 4341 18 13 16 5 PineCliff PineCliff PineCliff Woodhouse Woodhouse Woodhouse 1095 1603 1544 1525 Jun Jul Sep TOTALS 2420 1568 8 7 13 7 12 3 4 6 17330 62 63 18 23 13 10 12 16 24 16 13 6 138 Table 10. Preble's meadow jumping mice radio-collared and tracked for more than one tracking session. Location of capture, sex, identification, and number of locations for each mouse during each tracking session is noted. 1998 1999 Site Sex PIT-tag # Jun Jul-Aug Jul-Aug SeE-Oct SeE-Oct 60 24 Maytag F 4141f87C76 Maytag Maytag Maytag Maytag Maytag Maytag Maytag Maytag Maytag Maytag Maytag PineCliff PineCliff PineCliff PineCliff PineCliff PineCliff Woodhouse Woodhouse Woodhouse Woodhouse Woodhouse Woodhouse Woodhouse Woodhouse Woodhouse F F F F M M M M M M M F M M M M M F F F F M M M M M 41412B4A2D 413E296246 4141276D78 4140753620 4141241836 4140754707 4135354E37 41357B5AOE 414130663A 41411E6879 4141445465 4141554B28 41413D4C04 4141144535 413E02020F 414037184D 4141714141 41413E610B 41413E7B07 41414C721F 41412A2508 4141611413 4141074013 4141252066 41412D3509 41406AI049 103 105 73 93 . 82 1 19 38 144 7 76 3 40 61 13 1 70 37 49 11 65 18 78 66 44 15 21 52 21 45 37 14 5 16 63 42 46 59 75 81 6 63?NC ?NC 9 74 38 22 87 12 9 6 47 39 84 93 -,,- 64 �23 Table 11. Causes of mortality of Preble's meadow jumping mice from Maytag Property, PineCliffRanch, and Woodhouse Ranch collected during the 1998 and 1999 field season (June l-October 31). 1999 1998 Cause of Mortality Predation Bullfrog Garter snake Rattlesnake Yellowbellied racer Weasel Fox House cat Raptor Unknown Exposure Drowning Road Kill Handlingffrap Unknown TOTAL Total Maytag PineCliff Woodhouse Maytag PineCliff Woodhouse 0 0 0 5 0 0 5 0 0 1 1 0 0 2 1 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 0 1 0 0 1 1 0 0 0 0 0 1 0 0 1 0 0 0 1 0 0 0 1 1 2 4 19 0 3 2 7 5 2 0 0 0 3 2 4 9 0 1 0 0 0 2 3 1 0 0 0 0 0 1 4 0 2 0 2 1 9 1 0 2 5 3 6 17 8 4 8 24 13 17 74 1 ~----~~------------------------. _. .._.• .__ a---- • ---------.----------------------------------------------- • 0_9 --------------------.----_._-.----------------------------------------= 0.8 • -------------------------------------------------_-------------------------------------------------------------------_.- 0.7 • • ~--------------------------~I 0.6 0.5 • Observed Data -----------------------------Weighted Least Squares - 0.4 0.3 Weighted Least Squares Logistic +---+---j----I----j---+----f----t--t----+------i 200 400 800 600 Trapline Length 1000 1200 Figure 1. Observed estimates of p with the fit of the weighted nonlinear least squares model, and the logistic transformed nonlinear least squares model. �24 1 ~"-. ---{--------------------. 0.9 0.8 ~ .-9 0.7 ~ 0.6 0.5 0.4 o 500 1000 1500 2000 Trap1ine Length (m) Figure 2. Predicted p with 95% confidence intervals as a function of trap line length in meters. 20 -y-------------, 10 • Maytag .a • Walnut Creek • Rock Creek 0 ~ .•.....• .C~ouse • South Boulder Creek ·PinecJ.ij· • Maytag • Pinecliff • Dirty Woman Creek " • South Boulder Creek • Dirty Woman Creek I::I'.l ~ -10 -20 -30 • 1-onument Creek • Monument Creek • Woodhouse +--+--!--+-+--~-+--_1__I__+__+__+__+_~ o 10 20 30 40 50 60 70 PMJM Density (micelkm) Figure 3. Residuals for the best Alec model explaining PMlM density (mice/km stream length) from the standardized vegetation cover variables shrubs and trees (ha/km stream length). �25 Colorado Division of Wildlife Wildlife Research Report July 2000 JOB PROGRESS ~W~-1~5:.=::3....:.-R~-~1.=..3_ Work Package No. __ TMkNo. Cost Center 3430 Colorado State of Project No. REPORT ~0~66~3::..._ ~1 _ _ Mammals Research Program Kit Fox Conservation Kit Fox Augmentation Study Period covered: July 1, 1999 - June 30,2000 Author: T.D.!. Beck Personnel: T. Beck, G. Byrne, CDOW; E. Everett, B. Miller; Denver Zoological Foundation. ABSTRACT Relative abundance of kit fox prey surveys were developed M a written protocol. However, administrative problems prevented conduct of the late-summer 1999 field surveys. Spring 2000 surveys which had been planned were deferred because of organizational changes within CDOW relative to management of threatened and endangered species. The new organizational unit is to develop a prioritization process for allocating resources among species and habitats and it WM believed prudent to defer further field work pending this process since active kit fox augmentation had not begun. Surveys of historic kit fox dens in Uncompahgre Valley did not find evidence of pup production in 1999 and no active kit fox dens were located in spring 2000. Data on historic range rehabilitation projects in the shadscale desert communities of western Colorado WM compiled. ��27 KIT FOX AUGMENTATION STUDY Thomas D. L Beck SEGMENT OBJECTIVES I. Compare relative abundance of potential kit fox prey between Uncompahgre and Colorado river valleys. 2. Capture as many kit fox as possible in Uncompahgre Valley for DNA work and pup dispersal. 3. Develop a kit fox reintroduction plan for Mesa County. METHODS AND MATERIALS Sampling protocols were developed for comparing relative abundance of small rodents and cottontail rabbits at 2 sites in each valley. Small rodent surveys were to use a 200-trap concentric trapping web. Snap traps were selected rather than live trapping because of concerns about hantavirus prevalence in these areas. Because such trapping removes captured individuals, trapping was to be conducted during late summer; presumably the time of highest animal density. Thus it would be unlikely to create a local depleted zone. Museum collections from these areas of Colorado are scarce so all trapped animals would have been saved for museum collections. Rabbit abundance was to be measured by a combination of mark-resight estimation and spotlight counts. Hopefully the spotlight counts could be indexed to population estimations to provide a long-term monitoring index. Rabbit population monitoring would be conducted in early-spring; presumably the time oflowest animal abundance. Selection of only 2 sites per valley probably would not accurately represent the variation to be expected throughout the valley. However, this was the most that could be done with manpower and budget. Weekly precipitation records were collected from 6 stations maintained by NOAA in the Uncompahgre and Colorado river valleys in Colorado (Montrose, Olathe, Delta, Grand Junction, Fruita, Loma) for the growing season (March-October) for the period 1970-1998. Not all stations had data for each year. Historic kit fox dens were visited in July and August 1999 to survey for adult fox presence and pup .production. Historic dens and den areas were visited in April 2000 to examine for kit fox use. Records compiled by Ron Kufeld, CDOW retired, were searched to document all range restoration projects -conducted in the low elevation desert valleys. This data base was believed to be complete for the period 1960-1996. During other field activities, visits were made to most of the range project sites to see if any noticeable differences could be seen from surrounding desert vegetation. RESUL TS AND DISCUSSIONS Because of administrative delays in allocating budgets and temporary FTE' s, the trapping webs could not be set and run prior to the graduate student having to attend classes. Thus the late-summer 1999 prey studies were deferred to spring of 2000. During late-fall 1999 the Colorado Div. of Wildlife altered its organizational structure and created a new section to deal with Species Conservation; specifically species �28 currently on either a federal or state list of threatened or endangered species or a species in decline with potential for listing. Thus lead role in kit fox restoration shifted to this new section. Discussions among administrators resulted in the decision to defer further field studies until the new section could develop a prioritization process for addressing the species conservation needs. Thus prey abundance surveys were deferred during spring 2000; awaiting the development of the new section priorities. All material collected during our kit fox research surveys has been copied to the new section. Den site surveys in summer of 1999 did not produce any evidence of pup production. There was only one active den found in spring 1999 and for the third consecutive year no pups were produced at this den. No active dens were located in spring 2000 in a survey of historic den sites. Extensive surveys were not conducted in areas where we had previously failed to find active kit fox dens based on the rationale that this declining population will not produce significant dispersal. Based on the lack of finding any active dens, no trapping operations were conducted in 2000. Earlier surveys based on trapping and infra-red photography clearly indicate that trapping rarely produced kit fox captures when field surveys could not find evidence of their presence prior to trapping. The growing season precipitation data was examined to see if any seasonal patterns emerged. The variability within a year and among sites was extremely high and no patterns by year or area were apparent. This is likely a reflection of the nature of the summer precipitation events, which are part of a regional monsoon pattern dominated by afternoon thunderstorms. These storm events are quite local in scope yet capable of delivering 2-3 em of rain in less than an hour. Casual surveys of range rehabilitation sites indicated no difference from surrounding areas. Thus, when developing augmentation plans no special accord needs to be given to these historic management sites. No formal augmentation plan was developed pending results from the new organizational unit on priorities. However, the material needed to do so is on file. Contacts have been made with 2 surrounding states (Utah and Arizona) for sources of kit fox should augmentation move forward. Representatives of both states were confident of being able to provide kit foxes in adequate numbers. �29 Colorado Division of Wildlife Wildlife Research Annual Report July 2000 JOB PROGRESS State of Project No. Work Package No. __ REPORT Colorado Cost Center 3430 W-153-R-13 Mammals Research Program ....::;0-=-67.:....;O=--- Task No. 1 Lynx Reintroduction _ Post-Release Monitoring of Reintroduced Lynx Period Covered: July 1, 1999 - June 30, 2000 Author: Tanya M. Shenk Personnel: Gene Byrne, Rick Kahn, Dave Kenvin, Jim Olterman, Scott Wait, Whitey Wannamaker, Margaret Wild, Dave Younkin ABSTRACT In an effort to reestablish a viable population oflynx (Lynx canadensis) in Colorado, 41 lynx were reintroduced into southwestern Colorado in 1999 and an additional 55 lynx released in Spring 2000. Release protocols were evaluated by closely monitoring each lynx released in 1999 through radiotelemetry. Number of mortalities and causes of each mortality were documented. With this new information, release protocols were modified in an effort to release each lynx with the highest probability of survival. Three different release protocols were used in 1999. Differences in release protocol included the length of time animals were kept in the Colorado holding facility and timing of the release. Percent mortality due to starvation within six months of release date decreased with each modification of release protocols (75% under Protocol 1, 11% under Protocol 2, 0% under Protocol 3 except for female lynx released pregnant). Of the 55 lynx released in 2000,41 were released in April (protocol 2) and 14 were released in May (Protocol 3) following a minimum of three weeks in the Colorado holding facility. Of the tota196lynx released, 63 are being followed on a regular basis within Colorado, six male lynx have not been located since October 1999, one lynx possibly slipped a collar, and 26lyux are known to have died. Known mortality factors included starvation (7), gunshot (3), vehicle collision (3), trauma (2), predation (1), and disease (1). Cause of death could not be determined for nine mortalities. Initial dispersal movement patterns of the lynx released in 1999 were extremely variable. Dispersal habitat used by the lynx released in 1999 were also highly variable, from high elevation Engelmann spruce/subalpine fir to Nebraska agricultural lands. Lynx released in 2000 have remained closer to their release site and fewer have been observed using atypical lynx habitats. Through snow-tracking efforts (221 snow-tracking days) in 1999 and 2000 we have located 147 kills, 371 beds, and 132 scats. Of the 147 kills, 75% were of snowshoe hare (Lepus americanus), 23% were pine (red) squirrel (Tamiasciurus hudsonicus), and the remaining 2% were made up of other mammals and birds. We collected 132 scat samples that will be analyzed later for content. No reproduction has been documented to date, however whether or not lynx have bred and had litters at some point remains unknown. �30 �31 POST-RELEASE MONITORING OF REINTRODUCED LYNX Tanya M. Shenk P. N. OBJECTIVE 1. Post release monitoring of lynx released in Colorado. SEGMENT 2. 3. 4. 5. 6. 9. FY99-00 Estimate first year survival rates of lynx reintroduced to Colorado. Identify first year mortality factors of lynx reintroduced to Colorado. Describe first year movement patterns of lynx reintroduced to Colorado. Refine and prioritize needed research components to develop sound management strategies for lynx in Colorado. Prepare a Federal Aid Job Progress Report. PERFORMANCE 1. 2. 3. 4. 5. 6. 7. 8. OBJECTIVES INDICATORS FY99-00 Survival estimates of lynx reintroduced to Colorado. Summary of mortality factors of lynx reintroduced to Colorado. Description and analysis of habitats used by lynx reintroduced to Colorado. Description of movement patterns for lynx reintroduced to Colorado. Reproduction estimates of lynx reintroduced to Colorado. Evaluation and modification of release protocols for reintroducing lynx. Sites selected for second year release of lynx Refinement and prioritization of needed research components to develop sound management strategies for lynx in Colorado. Report on first year release oflynx in Colorado: movement patterns, habitat use, survival and reproduction INTRODUCTION In an effort to reestablish a viable population of lynx (Lynx canadensis) to Colorado, 41 lynx were reintroduced into southwestern Colorado in the spring of 1999 and an additional 55 lynx were released in Spring 2000. Monitoring of these lynx is crucial to evaluating the progress of this lynx reintroduction effort. The monitoring program will also provide information and data critical to improving release techniques to ensure the highest probability of survival for each individual lynx released in future years of the Colorado effort, and perhaps in other reintroduction efforts. The post-release monitoring program for the reintroduced lynx has two primary goals. The first goal is to obtain regular locations of released lynx. From these locations we will be able to determine how many lynx remain in Colorado and their locations relative to each other. Given this information and knowing the sex of each individual we will be able to assess the feasability of these lynx to form a breeding core from which a viable population might be established. Also from these data we can describe general movement patterns and habitats used. The second primary goal of the monitoring program is to estimate survival of the reintroduced lynx and, where possible, determine cause of mortality of reintroduced lynx. �32 Additional goals of the post-release monitoring program for lynx reintroduced to the southern Rocky Mountains include refining descriptions of habitat use and movement patterns, determining food habits, and obtaining information on reproduction. When the lynx establish home ranges that encompass their preferred habitat, more emphasis will be placed on refining descriptions of movement patterns and habitat use. Lynx is currently a species listed as threatened under the Endangered Species Act (ESA) of 1973, as amended (16 U. S. C. 1531 et. seq.)(U. S. Fish and Wildlife Service 2000). As a listed species, information specific to the ecology of the lynx in its southern range such as habitats used, movement patterns, mortality factors, survival, and reproduction in Colorado will be needed to develop recovery goals and conservation strategies for this species specific to its southern range. Thus, an additional objective of the post-release monitoring program is to develop conservation strategies relevant to lynx in Colorado. OBJECTIVES The initial post-release monitoring of reintroduced lynx will emphasize five primary objectives: 1. Assess and modify release protocols to enure the highest probability of survival. 2. To obtain regular locations of released lynx to describe general movement patterns and habitats used by lynx. 3. Determine causes of mortality occurring in reintroduced lynx. 4. Estimate survival of lynx reintroduced to Colorado. 5. Estimate reproduction of reintroduced lynx. Three additional objectives will run concurrently or become active after lynx become established in an area that encompasses their movements. These objectives include: 6. Better refine descriptions of habitats used by reintroduced lynx. 7. 8. Better refine descriptions of daily and overall movement patterns of reintroduced lynx. Describe food habits and prey of reintroduced lynx. The data collected during the post -release monitoring will be analyzed to evaluate habitat use, movement patterns, reproduction and survival. These data will be used to further the knowledge about habitat requirements for this species in the southern Rocky Mountains. Thus, the final objective for the postrelease monitoring plan is to: 9. Refine habitat protection recommendations and conservation strategies based on information collected from released lynx. STUDY AREA Five areas throughout Colorado were evaluated as potential lynx habitat (Byrne 1998). Criteria investigated in these five areas for comparison were (1) relative snowshoe hare densities (Reed at al., unpublished data), (2) road density, (3) size of area, (4) juxtaposition of habitats within the area, (5) historical records oflynx observations, and (6) public issues. Based on results from this analysis, the San Juan Mountains of southwestern Colorado were selected as the.release area for reintroducing lynx. Ten release sites within the San Juan Mountains were selected based on land ownership and accessability during time of release for the 41 animals released in 1999. Of the 55 lynx released in Spring 2000, 45 were released at Rio Grande Reservoir and ten lynx were released at three sites west of the Continental Divide. Based on current locations of the majority of the released lynx, the core research area remains in the southern San Juan Mountains, however lynx may need to be captured from areas of non-suitable habitat in Colorado and adjacent states. �33 METHODS Assessment of Release Protocols A total of 41 lynx were released in 1999 at selected areas in the San Juan Mountains of southwestern Colorado. Prior to release each lynx was examined and age, sex, and body condition determined. Each lynx was fitted with a Telonics'Pt VHF radio-collar for post-release monitoring. The collars were also equipped with a mortality switch that activates if the collar remains motionless for a period of four hours or more. Specific release sites were selected based on land ownership and accessability during times of release. Lynx were transported from the holding facility to the release site in individual cages. Release site location was recorded in Universal Trans Mercator (UTM) coordinates and identification of all other lynx released at the same location, on the same day, was recorded. Behavior of the lynx on release and movement away from the release site was documented. Monitoring of the survival and mortality factors (see below) of each lynx was used to modify release protocols in 1999 in an attempt to release each lynx with the highest probability of survival. Release protocols for the 55 lynx released in 2000 were developed from survival, mortality factor, and movement pattern data obtained from lynx released in 1999. Documenting Movement Patterns To obtain regular locations of released lynx to determine general movement patterns and habitats used by reintroduced lynx a combination of satellite, aerial and ground radio-tracking were conducted. Locations and general habitat descriptions of each location were recorded and mapped for all locations. All 41 of the lynx released in 1999 were monitored from the air through radio-tracking. Frequent flights (three times a week) were critical during the initial post-release periods because of the greater likelihood of dispersal and mortality in reintroduced carnivores. Every effort was made to locate every lynx during each flight during this period. Sixty days from the date of the last release, aerial locations of the radio-collared lynx were to be determined two times per week for the remainder of the life of the transmitters. Flights were also conducted three times per week, weather permitting, to locate lynx during the snow-tracking field season (December through April) to aid in the snow-tracking efforts. When possible at least one observer flew with the pilot to become familiar with the terrain, to operate the radio telemetry receiver, and to record the global positioning system (GPS) locations of the lynx. Generally, the pilot circled a strong telemetry signal and then bisected the circle activating the GPS unit when approaching directly overhead. The date and time of the beginning and ending of the flight, the time each collar was located, the UTM coordinates for each animal located, general weather conditions, primary overstory vegetation type, and name of the personnel were recorded. All locations were entered into a database for mapping and data analysis. Fifty-one of the 55 lynx released in spring 2000 were fitted with Sirtrack'Y dual VHF/satellite transmitter collars. The remaining four lynx were fitted with TelonicsThi VHF collars identical to those used on lynx released in 1999. Each dual collar weighed 137-156 grams. The satellite component of each collar is programmed to be active for 12 hours per week. The 12-hour active periods are staggered throughout the week, with approximately seven collars being active each day of the week. Signals from the collars allow for locations of the animals to be made via Argos, NASA, and NOAA satellites. The location information was processed by ServiceArgos and distributed daily to the Colorado Division of Wildlife through e-mail messages. Both the VHF and satellite transmitter in the dual collar has a mortality switch which is triggered by four or more hours of stationarity. Determining Causes of Mortality To determine causes of mortality occurring in reintroduced lynx every effort was made to locate and retrieve carcasses of dead lynx as soon as possible. When a mortality signal (75 ppm vs 50 ppm for the Telonics'O' VHF transmitters, 20bpm vs 40bpm for the SirtrackThi VHF transmitters, 0 activity for �34 Sirtrack ™ PIT) was heard during either satellite, aerial or ground surveys, the location (UTM coordinates) was recorded. Ground crews located and retrieved the carcasses. The immediate area was searched for evidence of other predators and the carcass photographed in place before removal. Additionally, the mortality site was described, habitat associations, and exact location were recorded. Any scat found near the dead lynx that appeared to be from the lynx was collected. All carcasses were transported immediately to the Colorado State University Veterinary Hospital for a post mortem exam. Lynx carcasses were not frozen but kept cool. If carcasses were already frozen due to field conditions, this was noted on the field form. The objectives of the post-mortem examination were to 1) determine the cause of death and document with evidence, 2) collect samples for a variety of research projects, and 3) archive samples for future reference (research or forensic). The gross necropsy and histology were performed by, or under the lead and direct supervision of a board certified veterinary pathologist. At least one research personnel from the Colorado Division of Wildlife involved with the lynx program was also present. In general, the protocol followed standard procedures used for thorough post-mortem examination and sample collection for histopathology and diagnostic testing. Some additional data/samples were routinely collected for research, forensics, and archiving. Other data/samples were collected based on the circumstances of the death (e.g., photographs, video, radiographs, bullet recovery, samples for toxicology or other diagnostic tests, etc.,). The CDOW retained all samples and carcass remains with the exception of tissues in formalin for histopathology, brain for rabies exam, feces for parasitology, external parasites for ID, and other diagnostic samples. Estimating Survival Survival rates of lynx reintroduced to Colorado will be estimated using the Kaplan-Meier method with staggered entries (Pollock et al. 1989). Documenting Habitat Use and Hunting Behavior More refined descriptions of habitats used by reintroduced lynx were obtained through snow-tracking of animals. Data were collected on habitats used, daybed and hunting bed locations, and travel corridors. Hunting and feeding behavior information was also collected by documenting prey taken, prey chases, relative abundance of prey (tracks and sightings), and use of carrion. Snow-tracking was conducted during February-May, 1999 and beginning again in November, 1999 through April 2000. Locations from the aerial-tracking were used to help ground-trackers "locate lynx tracks in the snow. One or more persons working together conducted the snow-tracking surveys. Snowmobiles, where permitted, were used to gain the closest possible access to the lynx tracks without disturbing the animal. From that point, snowshoes were used by the tracking team to reach the tracks. Once tracks are found, the ground crew back-tracked the animal. Back-tracking avoided the possibility of disturbing the lynx by moving away from the animal rather than toward the animal. However, monitoring of the lynx through radiotelemetry was also used to assure that ground crews stayed a sufficient distance away from the lynx in the event the lynx might double back on its tracks. If the lynx began to move in response to the observers, the observers retreated. If the lynx began to move and the movement did not appear to be a response to the observers, the crew continued to follow and record locations, habitats used, and behavioral information for as long as possible. Locations oflynx tracks were recorded using a Garmin XL12 GPS and 7.5 topographic map. Habitat descriptions included overstory and understory vegetation and seral stage. Locations and behavioral observations that could be interpreted from the tracks (e.g., chases, scent marking) were recorded. These data will be used for mapping and spatial analyses and analyzed to make inferences on how different habitats are used, frequency of use, daily movement patterns, hunting areas, daybed locations, den sites, and travel corridors. Data will also be used to document any changes in habitat use as animals begin to settle into a home range. 0 �35 An attempt was made to locate tracks from all lynx. However, first priority was given to locating any animal that appeared to be consistently in the same location from aerial surveys. Such stationarity may indicate an injured, starving, or otherwise traumatized animal. Data on hunting behavior was collected by location of kills, food caches, chases, and through scat analysis. Prey from attempted and successful hunting attempts were identified by either tracks or prey remains. Information from scat analysis will also provide information on foods consumed. Scat samples were collected wherever found, recording location and individual lynx identification. Only part of the scat was collected, the remainder was left where found so as not to interfere with the possibility the scat was being used by the animal as a territory mark. Comparisons of food composition and percent occurrence will be made within and among individuals. Analyses of temporal, spatial, and individual differences will be conducted to provide information on feeding ecology of reintroduced lynx in the southern Rocky Mountains. Estimating Reproduction Reproductive status of all female lynx was determined prior to release through radiographs. All females known to be pregnant or thought to possibly be pregnant on release were monitored closely from their release through the following August to determine reproductive success. Females remaining within a limited area immediately after release through August were located and observed to look for accompanying kittens or a den site. Females that had been released in 1999 and were alive in spring 2000 were monitored for proximity to males during breeding season and for site fidelity to a given area during the denning period of May and June. Each female lynx from the 1999 releases were directly observed in summer 2000 over 3-5 different visits to look for accompanying kittens or evidence of denning. Locations of both males and females released in 1999 were evaluated during March and April 2000 to document proximity of males to females in an attempt to determine if breeding could have occurred. RESULTS Assessment of Release Protocols A total of 41 lynx were released in Colorado in 1999 under five different release protocols (Table 1). The initial release protocol called for the immediate release of females once they passed veterinary inspection in Colorado. Males were to be held for a period of weeks until females established a territory, and then males were to be released near female territories. Four animals were released in early February, however, three of these died of starvation within six weeks of their release and the fourth was recaptured and returned to the holding facility where she recovered and was later re-released (Table 2). Reevaluation on the condition of animals released under the first protocol suggested that these animals may not have been in optimal physical shape when released. Therefore, a second release protocol was initiated whereby lynx were held at the Colorado holding facility for a minimum of three weeks and fed high quality diets to encourage weight gain. Most lynx gained considerable body weight while in captivity (Wild 1999). Nine lynx were released under this second protocol (Table 2). Of these nine lynx, one juvenile female died of starvation seven weeks after release. After the starvation death of the first lynx under the second protocol, a third release protocol was developed that called for releasing all subsequent lynx in the spring after a minimum stay in the holding facility of at least three weeks (Table 1). A spring release would assure the lynx were released when prey was most abundant (i.e., young of the year would be most abundant and hibernating and migratory prey would be available). Twenty lynx were released under this protocol (Table 2). Additionally, six females were released under this third protocol that were known to be pregnant (Protocol 3P) and two that were possibly pregnant (3P?). No lynx reintroduced under Protocol 3 died of starvation within six months postrelease (Table 2). However, two of the six lynx released when pregnant died of starvation within six months post-release. �36 An assessment of the fates of each lynx under all five release protocols used in 1999 led to release protocols for lynx released in 2000. Release protocols 2 and 3 resulted in the fewest post-release (up to six months after release date) starvation mortalities (Table 2). The common element in both protocols was increased captivity time in the Colorado holding facility. The single starvation mortality for lynx released under Protocol 2 in 1999 was also the only juvenile released under that protocol and the only animal released in February (the other eight Protocol 2 lynx were released in March 1999). Thus, all lynx released in 2000 were released under either Protocol 2 or 3 but not before April 1. Because of the high percentage of starvation mortalities in females pregnant on release (Table 2), we also attempted to avoid reintroducing lynx that were known to be pregnant. This was best accomplished by trying to have animals captured for the reintroduction effort in Canada and Alaska prior to their breeding season. Afover.nentJ>afterns Through extensive aerial and satellite tracking, we continue to search and locate 63 of the 70 lynx with collars on and assumed to be alive (one lynx has presumably slipped her collar). We have 1206 satellite locations for 49 of the 51 lynx fitted with dual collars (2 satellite collars never worked after the lynx were released) and 1122 aerial VHF locations for all 96 reintroduced lynx. Six males from the 1999 releases have not been found since at least 1 October 1999. Possible reasons for not locating these six males include (1) long distance dispersal, beyond the areas currently being searched, (2) radio failure, or (3) destruction of the radio (e.g., run over by car). We continue to search for all missing lynx during both aerial and ground searches. Last known locations for each of the 70 lynx assumed to be alive are presented in Figure 1. Initial dispersal movement patterns of the lynx released in 1999 were extremely variable. Dispersal habitat used by lynx released in 1999 has been highly variable, from high elevation Engelmann spruce/Subalpine fir to Nebraska agricultural lands. However, numerous travel corridors have been used repeatedly by more than one lynx, possibly suggesting route selection based on olfactory cues. Dispersal movement patterns of lynx released in 2000 were much less than those observed by lynx released in 1999. Most of 2000 releases have remained within an area encompassed by 100 km radius circle from the release locations. Most movement away from this core area has been to the north (Figure 1). We currently have six lynx using areas near Interstate 70. Survival and Afortality Factors Of the 96 lynx released, 26 mortalities have been recorded to date (Table 3). From the 1999 releases (41 animals) we have had 22 known mortalities (6 from starvation, 8 unknown, 3 gunshot, 2 hit by car, 2 trauma, and 1 predation). We have six missing males. We are following 13 of the lynx from the 1999 releases on a regular basis. From the 2000 releases (55 animals) we have four known mortalities (1 hit by car, 1 disease, 1 starvation, 1 unknown) and one animal that possibly slipped her collar. We are following the remaining 50 animals on a regular basis. Of the total seven confirmed starvation deaths, three were associated with animals released in less than ideal body condition and two were lynx less than one year old. Percent mortality due to starvation decreased with each modification of release protocols (75% under Protocol 1, 11% under Protocol 2, 0% under Protocol 3). Necropsy results for lynx BCOOF3, a female released on April 2, 2000 near Creede, Colorado indicated she died from pneumonic plague. The lynx was in fairly good condition, there was some abdominal fat, no muscle wasting, and the bone marrow had fat in it. The only gross lesion was an acute fibrinous pneumonia (i.e., lung infection of short duration). The lynx had probably only been sick a few days before it died. The carcass was recovered near her release site. Plague was diagnosed by flourescent antibody test and isolation of Yersinia pestis from lung and spleen samples. �37 Recaptures Three lynx have been recaptured and subsequently re-released since their initial release. Lynx BC99F6 was released in 1999 under Protocol 1. Her behavior and incidental sightings by the public suggested the lynx was in poor condition. We trapped her using a Tomahawk™ live trap baited with rabbit. She was recaptured the first night (March 25, 1999) we set the trap. On capture, we found she was severely emaciated. We anesthetized her with Telezol (2 mg/kg) and returned her to the Colorado holding facility. She was rehabilitated through diet. The lynx gained weight steadily and was re-released on May 28, 1999. She was hit by a car on Interstate 70 on July 19, 1999. Necropsy results indicated she was in excellent body condition at her time of death. Lynx AK99M9 was released on May 12, 1999 and recaptured on March 24,2000. Field observations by the lynx monitoring crew suggested that the lynx was severely emaciated. Live-trapping the lynx failed, so the lynx was darted with Telazol (3 mg/kg) using a Dan-Inject CO2 pistol. Physical examination revealed severe emaciation (6 kg). The lynx was returned to the Colorado holding facility and rehabilitated through diet. The lynx gained weight steadily and was re-released on May 3,2000. Lynx AK99F2 was released on May 7, 1999 and recaptured on April 18, 2000. Field observations by the lynx monitoring crew suggested that the lynx was emaciated. She was live-trapped with a Tomohawk™ live trap with one nights effort. On capture, we found she was emaciated. We anesthetized her with Telezol (2 rug/kg) and returned her to the Colorado holding facility. She was rehabilitated through diet. The lynx gained weight steadily and was re-released on May 22,2000. Habitat Use and Hunting Behavior February 1999-May 1999 Through snow-tracking, we were able to document habitat use, daily movement patterns, and hunting behavior of the earlier released lynx. Snow-tracking of lynx began shortly after the first release, Feb. 6, and continued until May 15, 1999. Although we tried to continue beyond May 15, efforts beyond this date did not yield any information because of either the lack of snow in the areas where the lynx were, or the snow conditions were too difficult to track in (hard, crusty, patchy). Because the majority (28) of the lynx were first released under Protocol 3, after May 6, the snow-tracking effort focused on the 13 lynx released prior to this date, under Release Protocols 1 and 2. Approximately 114 km of lynx tracks were followed. These tracks were from 11 different lynx, with kilometers tracked for any individual varying from 1 to 31 kilometers (Table 4). Two lynx (one female and one male) from Release Protocols 1 and 2 were never snow-tracked because we were either not able to locate the animals or because when we did locate them we could not readily access where they were. Daybeds and hunting beds were each located for eight of the lynx. Prey chases or kills were found for four lynx, scat samples were collected from five lynx, and possibly from a sixth. From the kills found and from initial examination of the scat samples, the lynx fed on snowshoe hare (Lepus american us), pine (red) squirrel (Tamiasciurus hudsonicus), and waterfowl. All the snow-tracking effort was conducted on nine lynx released under Protocols 1 and 2. Any lynx released under Protocol 3 were released too late to track. November 1999 -April 2000 Ground crews tracked 13 of the lynx released in 1999 during this period (Table 4). Two other lynx were being located during this time but were not in snow. A total of 139 kills or chases were located, 75% were snowshoe hare, 23% were pine (red) squirrel, and the remaining 2% were made up of other mammals and birds. We collected 115 scat samples that will be analyzed for content. Lynx released in 2000 were released too late to snow track. �38 Reproduction Six lynx released under Protocol 3 in 1999 were known to be pregnant (Table 1, Release Protocol 3P). Two other females may have been pregnant, the radiographs were suggestive but inconclusive (Table 1, Release ProtocoI3P?). Three of the six lynx known to have been pregnant on release in 1999 died within two months after release. Two starved and one was killed on the road (Table 2). Long distance movements and lack of stationarity in the movement patterns of the other three lynx known to have been pregnant on release in 1999 suggests these females did not have young with them by July 1999. Of the two females that might have been pregnant, movement patterns were not suggestive of a female rearing young. It is not known if any other females bred and/or had young once released, however no females snow-tracked November 1999 through April 2000 had young with them. From radiographs taken of the 35 females released in 2000, one female was known to be pregnant and three were possibly pregnant. Movement patterns suggest that none of these females have kittens with them as of July 2000. There were seven females released in 1999 that were alive during the Spring 2000 breeding season. All seven females were in close « 5 km) proximity to a male during the breeding season and could have bred. The seven females were monitored closely for stationary movement patterns, indicative of denning, from May-July 2000. Ground trackers also walked in on all seven females for visual observations on a minimum of three occasions and two females were visited on five occasions. No kittens were observed. However, the question of whether they successfully bred or had kittens at some point in 2000 is unknown. One of these females has since died and three others have made movements of over 100 km. Although we are confident none of the six live females have kittens at this time, for further confirmation we will snow-track each of these females as soon as they are in areas with fresh snow to check for kitten tracks. Beginning in March 2000 both male and female lynx began to exhibit extensive movements (> 100 km) away from areas they had used throughout the winter. For example, female (AK99F3) moved from the area near Grizzly Gulch she used throughout the winter to the Wolf Creek Pass area, a straight line distance of approximately 255km (Figure 3). Male YK99M3 moved from the area near the Climax mine which he had used throughout the winter to Taylor Mesa, a straight line distance of approximately 270km (Figure 4). Such movements by both females and males put them in close « 5 km) proximity to a lynx of the opposite sex. Two isolated males did not move during March or April and thus were not in close proximity to a known female during reeding season. This was a male that had used the area in and adjacent to the northwest comer of Rocky Mountain National Park and a male that used the area around Cuchara, Colorado throughout the winter. DISCUSSION Monitoring of lynx reintroduced to southwestern Colorado is crucial to evaluating the progress of the lynx reintroduction. Monitoring of these released lynx provides information and data necessary for improving release techniques to ensure the highest probability of survival for each individual lynx released in future years, and perhaps in other areas. Lynx is currently a species listed as threatened under the ESA. Information collected on the progress of the lynx reintroduction program, including habitats used, movement patterns, mortality factors, survival, and reproduction, could also be used to help develop recovery goals and conservation strategies for this species specific to its southern range. Three release protocols were used in the reintroduction of lynx to Colorado in 1999. Release protocols were modified as new information became available from monitoring the released lynx through radio-telemetry and snow-tracking. Each modification of the release protocols decreased the percent of animals dying from starvation. The primary element in later, more successful release protocols was an increased time in captivity at the Colorado holding facility. Increasing the amount oftime lynx were held in �39 the Colorado holding facility provided each lynx with an opportunity to increase body weight and acclimate to the climate, elevation, and local conditions of the envirorunent they would be released into. Although most lynx were housed in individual pens, with a few sharing a pen with one other lynx, the holding facility also allowed the lynx to hear and smell each other throughout this acclimation period. Such contact may have provided time for social interactions to occur. Such social interactions. may improve the likelihood these animals could form a breeding population. Post-release monitoring provided preliminary information on habitat use specific to Colorado that might later be used to refine habitat protection and management recommendations specific to Colorado. However, caution must be used in interpreting the information collected to date on habitats used by the introduced lynx. The aerial locations and snow-tracking results do provide some information but may also reflect behavior of displaced animals. General observations to note may be repeated use by multiple lynx of certain travel corridors and lack of use of tundra areas for any length of time. Both these habitat use characteristics have been noted for naturally occurring lynx populations. Preliminary data collected on kills suggests the reintroduced lynx are feeding on their preferred prey species, snowshoe hare and pine (red) squirrel in similar proportions as those reported for northen lynx during lows in the snowshoe hare cycle (Aubry et al., 1999). Caution must be used in interpreting the proportion of identified kills. Such a proportion ignores other food items that are consumed in their entirety. Through snow-tracking we have evidence that lynx are mousing and several of the fresh carcasses have yielded small mammals in the gut on necropsy. Nearly all the scat samples collected have been found through snow-tracking efforts and thus are representative of winter diet only. However, the summer diet of lynx has been documented to include less snowshoe hare and more alternative prey than in winter (Mowat et al., 1999). The extreme movements observed by both females and males in March and April 2000 may have been related to breeding behavior. March and April are the natural breeding periods for northern lynx (Tumlison 1987). We do not know if any of the females bred or had kittens but we are fairly sure that no female has kittens at this time. With only seven females from the 1999 releases in the wild in spring 2000 it was not unexpected that there might not be successful reproduction in 2000. During the summer of 2000, some lynx that were released in 1999 and had been faithful to a given area have made large movements away from these areas. Extensive summer movements away from areas used throughout the rest of the year have been documented by native lynx in Wyoming and Montana (Squires and Laurion 1999). Proposed monitoring and research include continued aerial radiotelemetry to document current locations and movement patterns, documentation of mortalities and causes of death, use of snow-tracking to document habitat use and hunting behavior, and further assessment of snowshoe hare densities in the state. The habitats used by the lynx will continue to be identified, mapped, and analyzed. These data will be used to further the knowledge about habitat requirements and preferences for this species in the southern Rocky Mountains. This information will be used to identify other blocks of potential habitat located throughout the Southern Rocky Mountains and evaluate conflicts that might jeopardize the recovery of lynx in Colorado. If conflicts are identified, such information can be used to develop conservation strategies and recommend land management strategies to mitigate them. ACKNOWLEDGMENTS The Colorado lynx reintroduction program and post-release monitoring is a large project involving many people. John Mumma, former director of the Colorado Division of Wildlife was instrumental in the implementation of the program. Rick Kahn of the CDOW is the program leader. Many CDOW biologists, researchers, wildlife managers and other personnel are involved in the program and or have advised us in the development of the monitoring protion of the program including Bill Andree, Tom Beck, Gene Byrne, Bruce Gill, Dave Kenvin, Todd Malmsbury, Jim Olterman, Dale Reed, John Seidel, Scott Wait, Margaret �40 Wild. The Lynx Advisory Team members from outside the CDOW include Steve Buskirk, Jeff Copeland, Dave Kenny, Steve King, John Krebs, Brian Miller, Gary Patton, Jerry Mastel, Kim Poole, Rob Ramey, Rich Reading, John Weaver, and Mike Wunder. We thank Susan and Herman Dieterich of the Frisco Creek Wildlife Rehabilitation Center for the care and maintenance of the lynx while being held in Colorado. The aerial post-release monitoring has been conducted by state pilots including Dell Dhabolt, Jim Olterman, Matt Secor, Whitey Wannamaker, and Dave Younkin. Ground field crew members include Bob Dickman, Chris Parmater, Jake Powell, and Jennifer Zahratka. Jon Kindler and Anne Trainor of CDOW were most helpful in preparation of the maps. Funding has been provided by Vail Associates, Turner Foundation, Great Outdoors Colorado (GOCO), and the Colorado Division of Wildlife. LITERATURE CITED Aubry, K. B., G. M. Koehler, and J. R. Squires. 1999. Ecology of Canada lynx in southern boreal forests. in Ecology and Conservation of Lynx in the United States. General Technical Report for U. S. D. A. Rocky Mountain Research Station. University Press of Colorado. Byrne, G. 1998. Core area release site selection and considerations for a Canada lynx reintroduction in Colorado. Report for the Colorado Division of Wildlife. Mowat, G., B. G. Slough, and S. Boutin. 1996. Lynx recruitment during a snowshoe hare population peak and decline in southwest Yukon. Journal of Wildlife Management 60:441-452. Mowat, G. and B. G. Slough. 1998. Some observations on the natural history and behaviour of the Canada lynx, Lynx canadensis. Canadian Field Naturalist 112: 32-36. Mowat, G. ,K. G. Poole, and M. O'Donoghue. 1999. Ecology oflynx in northern Canada and Alaska. in Ecology and Conservation of Lynx in the United States. General Technical Report for U. S. D. A. Rocky Mountain Research Station. University Press of Colorado. Nava, J. 1970. The reproductive biology of the Alaska lynx. M.S. Thesis University of Alaska, Fairbanks. Pollock, K. H., S. R. Winterstein, C. M. Bunck, and P. D. Curtis. 1989. Survival analysis in telemetry studies: the staggered entry design. Journal of wildlife management 53: 7-15. Poole, K. G., G. Mowat, and B. G. Slough. 1993. Chemical immobilization of lynx. Wildlife Society Bulletin 21: 136-140. Seidel, J., B. Andree, S. Berlinger, K. Buell, G. Byrne, B. Gill, D. Kenvin, and D. Reed. 1998. Draft strategy for the conservation and reestablishment of lynx and wolverine in the southern Rocky Mountains. Report for the Colorado Division of Wildlife. Slough, B. G. 1999. Characteristics of Canada lynx, Lynx canadensis, maternal dens and denning habitat. Canadian Field Naturalist 113:605-608. Squires, J. R. and T. Laurion. 1999. Lynx home range and movements in Montana and Wyoming: preliminary results. in Ecology and Conservation of Lynx in the United States. General Technical Report for U. S. D. A. Rocky Mountain Research Station. University Press of Colorado. Tumlison, R. 1987. Mammalian Species: Felis lynx. American Society of Mammalogists. U. S. Fish and Wildlife Service. 2000. Endangered and threatened wildlife and plants: final rule to list the contiguous United States distinct population segment of the Canada lynx as a threatened species. Federal Register 63, Number 58. Wild, M. A. 1999. Lynx veterinary services and diagnostics. Job Progress Report for the Colorado Division of Wildlife. Fort Collins, Colorado. �41 Table 1. Release protocols for lynx released in southwestern Colorado in 1999. Protocol 1 Description Release females as soon as they pass veterinary inspection in Colorado. Release males once females appear to have settled into an area. 2 Release males or females after they have been held in Colorado holding facility for a minimum of 3 weeks. During this holding period, the lynx were fed high quality diets to encourage weight gain, assuring each lynx would be released in optimal physical condition. Such a minimal holding period also provided an opportunity for the lynx to acclimate to the climate, elevation, and local conditions of the environment they would be released into. Although most lynx were housed in individual pens, with a few sharing a pen with one other lynx, the holding facility allowed the lynx to hear and smell each other throughout this acclimation period. Such contact may also have provided time for social interactions to occur. 3 All lynx to be kept in the holding facility for not only the minimal three week period but until spring. A spring release would assure the lynx were released when prey was most abundant (i.e., young of the year would be most abundant and hibernating prey would be available). Coupled with the minimum holding period of three weeks, these lynx would also be released when in optimal physical condition and after a period of acclimation to their new surroundings. 3P Pregnant females released under Protocol 3. 3P? Possibly pregnant females released under Protocol 3. Table 2. Summary of number of lynx released under each release protocol and numbers of lynx mortalities six months post-release for lynx released into southwestern Colorado in 1999 and four months post-release for lynx released in 2000. 2000 1999 Number released Protocol 1 2 3 3P 3P? Total Mortalities 6 months post-release (n, %) Number released Female Male Starvation Other Female Male 3 3 8 1 6 12 19 0,0% 0,0% 3,15% 2,33% 0,0% 5,12% 0 25 6 1 3 35 0 16 4 6 2 22 3,75% 1,11% 0,0% 2,33% 0,0% 6,14% 20 Mortalities 4 months post-release (n, %) Starvation Other 1,2% 1,10% 0,0% 0,0% 1,1% 3,7% 0,0% 0,0% 0,0% 3,5% �42 Table 3. Release and mortality information for lynx released into southwestern Colorado in 1999 and 2000. Mortality Information Release Information Animal ID Sex Age Date Site Protocol Date Cause of death BC99Ml M 8mo 2/4/99 Goose Creek 1 2124/99 starvation BC99F9 F 2+ 2/3/99 Goose Creek 1 2126/99 starvation BC99F7 F 3+ 2/3/99 Goose Creek 1 3/16/99 starvation BC99F8 F 9mo 2/20/99 Red Mtn Creek 2 4/10/99 starvation AK99F4 F 1-2 5/7/99 Sand Bench 3p 6/13/99 starvation AK99M23 M 1-2 5/14/99 Love Lake 3 6/18/99 shot BC99F6 F 2+ 2/4/99 Goose Creek 1 7/19/99 hit by car AK99F17 F 2-3 5/10/99 First Fork 3p 7122/99 hit by car AK99F8 F 5+ 5/10/99 First Fork 3p 7/30/99 starvation AK99F18 F 1-2 5/14/99 Love Lake 3 8/25/99 trauma, emaciation AK99FI0 F 10mo 5/12/99 Lemon Res 3p 9/13/99 unknown, not starvation BC99M2 M 4+ 3/19/99 Red Mtn Creek 2 10/20/99 unknown, not starvation AK99F27 F 10mo 5/14/99 Love Lake 3 10/31/99 shot AK99M6 M 5 5/13/99 Vallecito Res 3 11/16/99 shot AK99F15 F 2-3 5/14/99 Love Lake 3 11/24/99 blunt trauma YK99F4 F 4-5 5/13/99 Vallecito Res 3 1/25/00 predation, emaciation AK99Mli M 2-3 5/12/99 Lemon Res 3 1129/00 unknown BCOOF3 F 1 4/2/00 Goose Creek 2 5/24/00 pneumonic plague YKOOM5 M 10mos 4/2/00 Beaver Meadows 2 5/25/00 starvation YK99F3 F 2 5110/99 First Fork 3 6/7/00 unknown, not starvation YK99M6 M 3 5/13/99 Vallecito Res 3 6/19/00 unknown AKOOF4 F 10mos 5/22/00 Rio Grande Res 3 6/19/00 slipped collar? AK99F13 F 10mo 5/12/99 Lemon Res 3 6/22/00 unknown YKOOF17 F 1 4/17/00 Rio Grande Res 2 7/29/00 unknown, not starvation BC99MI0 M 3-4 3/19/99 Red Mtn Creek 2 8/2/00 unknown AK99F25 YKOOF6 F F 10mo 2 5/7/99 4/2/00 Sand Bench Rio Grande Res 3 2 8/10/00 8/17/00 unknown, not starvation hit b~ car Table 4. Habitat use and hunting behavior as described by summarizing kills, mousing activity, territory marks, hunting beds, day beds, and chases for each lynx tracked. Total number of days tracked to date and number of scat samples collected tracking field season. are also summarized. Data presented here are for the 1999-2000 snow- Tracking Period No. of lynx tracked Kills Beds Scats Tracking Days Feb 99 - May 99 11 8 71 17 84 Nov 99 - Apr 00 13 139 300 115 137 �I I --~--~: -----I --------.~,. ..... -: ... -~: - ..._ ... --' _ ',...- - __ . --.-~ .' \ -- - -r-I 'l' i I_ ----- t\ ----I ' , __ 1_--~l, r--"--' .J I ., .:::: .,,_ ~/--.,_I • I --. -"_ -"1 !NEWMEXlCO r· i \ I i /--------I N -- \ -, -t;':;/ . --- -1 __ ~ _ ___J '._j ,,L_i ~------ , I \ ,i \. \__ I W*E s I i Figure 1. Most recent locations, as of August 17, 2000, of the 70 lynx known or assumed to be alive from the 96 lynx reintroduced to southwestern Colorado in 1999 and 2000. Black triangles indicate last known VHF location, black circles indicate last known satellite location. Black lines are Colorado highways, grey lines are county boundaries. Each location is identified with the animal code. "" IJ.J �--_._- .. - . .1 ! I-_J l I I 1 ! - -v- --.-. --~-- ! L__ __ I------·L---·--·-l ..._.. - ..------ ~ ~ * r- l I--- .. _j - - ~----- .... ,. ---- '·1 i .. ..L;. •. .,.' c ( -- ... _--.,"" I I i J: s:l ., .'. ,~~! ------ .. ~-.--\-:.; .~ --.::;l' r= f I i -·1 - -T:E-::=O I I • I ) --..__j , , I ? ?~ <; Figure 2. Locations of all 26 known lynx mortalities from the 96 lynx reintroduced to southwestern Colorado in 1999 and 2000. Different symbols indicate different causes of death: starvation (.), hit by car (+), predation (*), disease (A), gunshot (*), and unknown (?). Dark lines within the Colorado border are highways, grey lines are county lines. �r---------- NEBRASKA WYOMING ·--T---- ----;r-------- '-" '( " J: ~ ::::> NEW MEXICO s Figure 3. Movements oflynx female AK.99F3 from her release to August 2000. Smallest circles are oldest locations with circles increasing in size as date becomes more recent. Largest circle is most recent location. Black lines are Colorado highways, grey lines are county boundaries. v. """ �~ WYOMING NEBRASKA I'· ~ I ~ ::> ·~E NEW MEXICO Figure 4. Movements of lynx male YK99M3 from his release to August 2000. Smallest circles are oldest locations with circles increasing in size as date becomes more recent. Largest circle is most recent location. Black lines are Colorado highways, grey lines are county boundaries. 0\ �47 Colorado Division of Wildlife Wildlife Research Report July 2000 JOB PROGRESS Smreof Project No. Work Package No. TMkNo. C==ol=o~rad~o REPORT _ ----!W~-1'_.:::5..:::.3~-R~-~1...!..4 _ ----'O::,.o6:...,:.7.,:::.0 ~2 Mammals Research _ Lynx Conservation _ Lynx Veterinary Services and Diagnostics Period Covered: July 1, 1999 - June 30,2000. Author: M. A. Wild. Personnel: T. Shenk, S. Dieterich, H Dieterich, T. R. Spraker, D. H Gould. ABSTRACT Fifty-six lynx (34 adult females, 19 adult males, 2 juvenile females, 1juvenile male) were received from British Columbia, the Yukon, and Alaska during January - April 2000. Lynx arrived in generally good condition with the exception of one female lynx that required euthanasia due to an extensive foot lesion from trapping. All lynx were anesthetized (TelazoI2-3 mglkg 1M) for physical examination and identification at least twice during the 2:3 week acclimation period at the captive holding facility. During captivity lynx maintained or improved body condition. Using abdominal radiographs, we diagnosed one female M pregnant and three others were suspicious. Seven lynx had lesions to 1-2 toes that required treatment. One of these lynx also had two fractured metacarpal bones that required splinting. All recovered and were released in Spring 2000. One lynx from the 1999 release WM recaptured in March 2000. The lynx WM emaciated but no other abnormalities were determined. He WM released in May 2000 after rehabilitation. Fifteen mortalities were recorded in free-ranging lynx during FY2000. Eight of the carcasses were in advanced stages of decay (or only the collar WM found) and cause of death could not be determined; however, in three of these cases we were able to rule out starvation M cause of death. Two lynx died from gunshot wounds, two from trauma, and one from predation. One trauma case and the predation case were in poor body condition. One male kitten died from starvation about 7 weeks postrelease. A female yearling died of pneumonic plague in Hinsdale County, Colorado. �48 �49 LYNX VETERINARY SERVICES AND DIAGNOSTICS Margaret A. Wild P. N. OBJECTIVES 1. Provide veterinary care and diagnostic services for reintroduced lynx. SEGMENT 1. OBJECTIVES Provide veterinary care and diagnostic services for reintroduced lynx. METHODS AND MATERIALS Lynx were trapped in British Columbia (BC) and the Yukon, Canada, and Alaska then transported by truck and/or airline to holding pens at the Frisco Creek Wildlife Hospital and Rehabilitation Center, Del Norte, Colorado. Care of lynx during captivity was based on the Husbandry and Management Protocol (Attachment 1 of Addendum A). Post-release, diagnostic and forensic services were provided to support ongoing research and law enforcement efforts. Carcasses were transported to the Colorado State Diagnostic Laboratory, Fort Collins, for examination by a board certified veterinary pathologist and a wildlife veterinarian. Complete post-mortem examination was conducted following the Necropsy Protocol (Attachment 2 of Addendum A). RESUL TS AND DISCUSSION Results of the 1999 lynx reintroduction are summarized in Addendum A. An additiona156lynx were received January - April 2000. Lynx appeared healthy on arrival with no transportation-related injuries. Captive management and husbandry techniques were similar to those reported in 1999; however, all1ynx were held at least 3 weeks for acclimation and fattening prior to release. Releases occurred in early April through late May. Daily clinical assessment suggested that lynx adjusted well to confinement in our isolated holding facility. Daily feed intake was not quantified this year however consumption appeared similar to the average 10-15% of body weight/day over a weekly basis that was observed in 1999. Health Assessment We anesthetized each lynx with about 2.5-3 mg/kg body weight Telazol 1M for initial examination and sample collection (as described in Attachment 1 of Addendum A). One lynx (BCOOFI7) was euthanized upon initial exam due to extensive foot lesions. The other 55 lynx were anesthetized again with about 2-2.5 mg/kg Telazol 1M for placement of a radiocollar just prior to release. In general, lynx arrived in good condition. Upon arrival, females from British Columbia (BC; n = 9) and The Yukon (n = 20) averaged 8.86 kg (SE 0.28) and 8.72 kg (SE = 0.22), respectively. At examination prior to release, body weights had increased to 10.23 kg (SE = 0.19) and 10.28 kg (SE= 0.13) for BC and Yukon females, respectively (Fig. 1). Adult females from Alaska (n = 4) averaged 9.82 kg (SE = 0.67) at arrival and 1l.91 (SE = 0.63) prior to release; however, one of these was known pregnant and three were suspected to be pregnant. Body weight change was less marked in male lynx. Upon arrival, adult males from BC (n = 9), The Yukon (n = �50 6), and Alaska (n = 4) averaged 12.2 kg (SE 0.53), 10.64 kg (SE = 0.32), and 11.23 (SE = 0.32) respectively. At examination prior to release, body weights were 12.93 kg (SE = 0.45), 12.13 kg (SE= 0.17), and 13.27 (SE = 0.63) for BC, Yukon, and Alaska males, respectively (Fig. I). One male kitten from The Yukon weighed 6.2 kg on arrival and 7.87 kg prior to release. Two female kittens from Alaska averaged 6.89 kg (SE = 0.34) on arrival and 9.17 kg (SE = 0.08) prior to release. Using radiographic examination of the growth plates of the distal radius and ulna as an indicator of age (Nava 1970), we determined that three juveniles were received (YK00M5, AKOOF1, and AKOOF4). Apparently, the body length measurements that we collected in 1999 to distinguish kittens from adults proved useful in field evaluation of lynx. and minimized the number of kittens that we received. This year we also adopted a new approach for estimating age of lynx. based on percent pulp cavity in the canine tooth. This method has been used to age canine teeth collected from lynx. carcasses (K. Poole, pers. comm.), but we attempted to collect the information in situ using skull radiographs. Evaluation of the method is underway. Several injuries to lynx. were found. Eight lynx. had fractured toes or toes previously amputated by veterinarians in BC or The Yukon. Damage to the foot of one of these lynx. (BCOOF17) was severe and involved the middle three toes on a front paw. Due to the poor prognosis, we euthanized this lynx. at the time of initial veterinary examination. Of the remaining seven lynx, three lynx. had one toe amputated (one of these lynx. also had two fractured metacarpal bones that required splinting), two lynx. had one toe amputated and an additional toe damaged, one lynx. had two toes amputated, and one lynx. had two fractured toes that were ankylosed but were not treated. Five other lynx. had minor lacerations on the legs or face that healed without complications. Recapture On 24 March 2000, we recaptured a lynx. (AK99M9) released in 1999. Field observations by the lynx. monitoring crew suggested that the lynx. was severely emaciated. Live trapping the lynx. failed, so we darted the lynx. with Telazol (3 mglkg) using the Dan-Inject CO2 pistol. Physical examination revealed severe emaciation (6 kg). It can be assumed that the lynx would have died ifnot recaptured. Blood work was unremarkable with the exception ofa low titer to Toxoplasmosis; however, this was unlikely the cause of the abnormality. No underlying disease condition was found that would explain the debilitation. The lynx. responded well to supportive care at the holding facility and was released in May 2000. Post-mortem Examination Fifteen mortalities were recorded in free-ranging lynx. during FY2000 (Table I; one of these was included in Addendum A as well). Eight of the carcasses were in advanced stages of decay (or only the collar was found) and cause of death could not be determined; however, in three of these cases we were able to rule out starvation as cause of death based on the presence oflipid in bone marrow. Two lynx. died from gunshot wounds, two from trauma, and one from predation (apparently from a bobcat). One trauma case and the predation case were in poor body condition. One male kitten died from starvation about 7 weeks post-release. A female yearling died of pneumonic plague. The carcass was recovered in Hinsdale County, Colorado. Necropsy findings showed an acute fibrinous pneumonia. Plague was diagnosed by fluorescent antibody test and isolation of Yersinia pestis from lung and spleen samples. After this diagnosis, we retrieved bone marrow samples to test for plague in six other lynx. that had died from unknown causes in 1999 and 2000. Fluorescent antibody test was negative on all of these cases. Plague in rodents is not uncommon in southwestern Colorado. This lynx. most likely consumed a rodent or rabbit infected with plague. �51 LITERA TORE CITED Nava, J. 1970. 141pp. Table The reproductive l. Summary of mortalities biology of the Alaska lynx. in free-ranging M.S. Thesis. Univ. Alas., Fairbanks. lynx during FY 2000. Date' Animal ID Sex Age State of Carcass Cause of Death 08/26/99 09/15199 10/22/99 AK99F18 AK99FI0 BC99-02 AK99F27 AK99M6 AK99FI5 YK99F4 AK99MII BCOOF3 YKOOM5 YK99F3 AKOOF4 YK99M6 AK99F13 YKOOF17 F F M F M F F M F M F F M F F 2 2 4 1 5 3 5 3 1 0 3 0 4 1 2 Good Poor Poor Good Good Good Good Minimal remains Good Good Poor Collar only Collar only Minimal remains Poor Trauma, emaciation' Unknown, not starvation' Unknown, not starvation' Shot Shot BIWlt trauma Predation, emaciation Unknown Pneumonic plague Starvation' Unknown, not starvation' Unknown Unknown Unknown Unknown' 11/03/99 11117/99 11/24/99 01126/00 01/29/00 05123/00 05/23/00 06/08/00 06/15/00 06/15/00 06122/00 07/29/00 I 2 Date mortality confirmed, date of death may have been earlier. Plague negative on FA of bone marrow 14 -r 12 -f- lO - 8 - 6 - 4 - 2 - ~ r-m!!i; • -11111111 ::::!:~~~~ 11111111 0 BC YK Females Fig. 1. Initial body weight (stippled) reintroduced in 2000. AK BC YK AK Males and body weight at release (solid) of female and male lynx �52 ADDENDUM A Status Report on the Health of Lynx Reintroduced to Colorado by Margaret A. Wild, DVM Colorado Division of Wildlife With additional information from Herman Dieterich, DVM, DACVS and Susan Dieterich Frisco Creek Wildlife Hospital and Rehabilitation Center For the 30-31 August 1999 CLAWSILAT Meeting GOAL To assure optimal health of lynx reintroduced into Colorado. OBJECTIVES Objectives of the health program were: l. 2. 3. 4. To To To To protect the health of wild and domestic animals in Colorado. optimize health of lynx released into Colorado. optimize welfare oflynx during transport and holding. provide diagnostic and forensic services in the case of mortalities. MATERIALS AND METHODS Lynx were trapped in British Columbia (BC) and the Yukon, Canada, and Alaska then transported by truck and/or airline to holding pens at the Frisco Creek Wildlife Hospital and Rehabilitation Center, Del Norte, Colorado. Care of lynx during captivity was based on the Husbandry and Management Protocol (Attachment 1). Post-release, diagnostic and forensic services were provided to support ongoing research and law enforcement efforts. Carcasses were transported to the Colorado State Diagnostic Laboratory, Fort Collins, for examination by a board certified veterinary pathologist and a wildlife veterinarian. Complete post-mortem examination was conducted following the Necropsy Protocol (Attachment 2). PRELIMINARY RESULTS AND DISCUSSION Captive Management and Husbandry Forty-two lynx were received at the holding facility between 29 January and 14 April 1999. All appeared healthy on arrival with no transportation-related injuries. Individual lynx were held at the facility for varying lengths of time based on assigned release protocol. Lynx from BC were held <1-7 wk (median 6.5 wk), from the Yukon 3 - 10 wk (median 8 wk), and from Alaska 3 - 6 wk (median 5) prior to release. Daily clinical assessment suggested that most lynx adjusted well to confinement in our isolated holding �53 facility. However, at least three lynx never adjusted to confinement; two paced the pen causing erosions on the palmar and plantar surfaces of the pads (AKFX99 and AKF499) and one appeared distressed and remained in relatively poor body condition (AKFI599). For other animals, clinical observations suggesting acclimation to confinement were supported by daily feed intake and body weight data. Lynx consumed on average about 10-15% of their body weight/day over a weekly average (Fig. 1) despite the fact that highly palatable feed items were not offered ad libitum. Lynx often fasted rather than consume less palatable items such as the prepared feline diet, fish, or beaver and engorged themselves on highly palatable items such as wild rabbits (up to >2 kg/day). Most lynx gained considerable body weight while in captivity (Fig. 2). In 1999, our initial plan was to hold lynx in captivity for as short a period as possible. We assumed that the animals would be distressed in confinement and would have a greater chance of survival if released immediately to the wild. However, we found that most lynx appeared to adjust to confinement without distress. Further, this period in confinement was likely beneficial to the animals in giving them time to acclimate to Colorado and to gain body weight prior to release. Health Assessment We anesthetized all lynx at least once for examination, and most were anesthetized again for placement of a radiocollar prior to release. We used 3-6 mg/kg body weight Telazol 1M to perform 84 anesthetic events on 42 lynx. Of the 79 anesthesias induced with a single injection, mean induction time was 4.3 min (SE 0.2 min; range 2-10 min). Five other anesthesias required supplementation and mean induction time was extended to 16.2 min (SE 7.2 min; range 7-29 min). Full recovery was achieved in all cases on average 91 min (range 27-188 min) after initial Telazol injection. All parameters stayed within acceptable limits; however, lynx receiving 5-6 mg/kg were markedly deeper and oxygen saturation frequently dropped to 70 80%. Supplemental oxygen was administered in these cases. In general, lynx arrived in good condition. Lynx from the Yukon were in markedly better body condition than those from other sites (Fig. 2). Using radiographic examination of the growth plates of the distal radius and ulna as an indicator of age (Nava 1970), we determined that at least six juveniles were received (two from BC, one from the Yukon, and three from Alaska). Total length of the two juveniles from BC was markedly less than that of other lynx from BC (juveniles 78 em, range in adults 87-97.5 em). The juvenile from the Yukon was correctly identified using the body length criterion of Slough (1996); the juvenile was 90.2 em while the adults were all well above Slough's 90.5 em eut-offfor adults (range 96105 em). Identification of juveniles from Alaska was more difficult, likely in part because evaluation occurred later in the year. Three juveniles were identified based on lack of closure of the growth plate; these lynx ranged from 86-89.5 em in length. Lynx classified as adults ranged from 89.5-106.5 em, Given these findings, body length can likely be used as an indicator to classify juveniles; however, site-specific cut-offpoints for body length will be required (i.e., while Slough's classification is appropriate in the Yukon, it appears less reliable for BC and potentially Alaska). We also used radiography to diagnose advanced pregnancy (>45 days) in lynx. Six lynx (AKF299, AKF399, AKF499, AKF899, AKFI099, AKF1799) had fetuses with calcified skeletons present and two other lynx had radiographic evidence suggesting earlier pregnancy (AKF599, YKF599). Of these pregnant females, three have died as of 28 August 1999 (one starvation, one hit-by-car, one undetermined cause). Five lynx had lesions or fractures of one to three toes that required medical treatment or surgical amputation. Two lynx (BC99-8 and AKF499) had minor infections on one toe of a front foot that were treated medically. These animals recovered fully prior to release. Two lynx had fractures and/or freeze �54 damage to two toes on a front foot, with one toe amputated from each lynx (AKFI099, AKF1899). The remaining lynx had fractures and freeze damage to three toes on a hind foot that required amputation of all three toes (BC99-15). Of the three lynx released with toes amputated, two are alive and one has died (cause of death unknown, although animal was slightly emaciated and had traumatic injuries) as of 28 August 1999. Sample collection, treatments, and identification were as described in the Husbandry and Management Protocol; however, eartags were placed only in lynx from BC. An incidental finding observed in the majority of lynx from the Yukon and Alaska were pinpoint to 2 mm slightly raised lesions on the oral mucosa, primarily the ventral surface of the tongue. Biopsy samples were submitted to Colorado State Diagnostic Laboratory for histopathology. Results indicated the plaques were caused by a slightly thickened epidermal layer, with no specific lesions present. Otherwise, lynx remained healthy in captivity with the exception of one adult male (YKM 199) and one adult female (BC99-06). Lynx YKM199 exhibited paresis and muscle weakness of about 1 wk duration. Results of diagnostic procedures were unremarkable, with the exception of apparently enlarged kidneys on radiographs. The lynx was euthanized 29 April 1999 and a thorough post-mortem examination was performed. No gross or histologic lesions were found (kidneys were normal despite radiographic interpretation). Kidney lead levels were normal. Serology for FelvlFIV, CDV, FIP and Toxoplasma gondii were negative. Rabies and parasitology exam were negative. The cause of paresis in this lynx was undetermined; however, infectious diseases that could have affected other lynx in the holding facility were excluded from the list of possible causes. Lynx BC99-06 was diagnosed with an abdominal hernia after recapture. The lynx was initially released under protocol 1, then recaptured in a debilitated, emaciated state. The lynx was nursed back to health on a diet slowly increasing in amount. Although no adverse effects were observed during the rehabilitation, an abdominal hernia developed. The rent in the abdominal wall was surgically corrected without complication. Post-mortem Examination In addition to the one lynx euthanized and examined prior to release, as of 28 August 1999 we have examined 10 mortalities (Table I). Starvation was diagnosed as the cause of death in five lynx based on extreme emaciation as evidenced by lack of depot fat, muscle atrophy, and serous atrophy of bone marrow. Three other lynx that died acutely (one shot, two hit-by-car), were in good body condition. Cause of death in two lynx was undetermined. One lynx (AKF899) had undergone severe autolysis and dehydration prior to examination. No internal organs or bone marrow remained for examination. Another lynx (AKFI899) was in a moderate state of autolysis, however, a complete examination was still possible. Gross examination of this animal revealed slight emaciation and traumatic injuries. Histologic examination of tissues is pending. All carcasses have been tested for rabies and internal and external parasites. Rabies examination has been negative in all cases. Incidental findings on parasitology have included Trichinella, Trichuris, coccidia, Toxascaris, Taenia, giardia, and rodent fleas. LITERATURE CITED Nava,1. 1970. The reproductive biology of the Alaska lynx. M.S. Thesis. Univ. Alas., Fairbanks. 141pp. Slough, B. G. 1996. Estimating lynx population age ratio with pelt-length data. Wildl. Soc. Bull. 24:495499. �55 Attachment 1 Husbandry and Management of Captive Lynx at Frisco Creek Wildlife Hospital and Rehabilitation Center Husbandry Lynx will be held singly or in pairs in holding pens (about 5 m2) with attached nest boxes (about 0.7 m2) at the Frisco Creek Wildlife Hospital and Rehabilitation Center. Two units, each with 10 pens, are available. Pens are fully enclosed by fencing and have concrete floors. Straw or hay bedding will be placed in the nest box as well as in two additional piles in the pen. Pens will be cleaned daily. Fresh water, snow, and feed will also be provided ad libitum daily. Feed remaining in the pen after 24 hr will be inspected for quality and discarded if rejected by the animal or if unfit for consumption. Quantity of feed provided and removed will be measured daily. Type of feed will vary, but will consist primarily of domestic, cottontail, and jackrabbits, deer and elk meat, and poultry. Additional feeds may include a prepared feline diet, fish, wild birds, and small mammals (excluding plague vector species, i.e., prairie dogs). Head, kidneys, and urinary bladder will be removed from domestic rabbits prior to feeding to prevent transfer of Encephalitozoon cuniculi. Health and behavior of lynx will be observed twice daily. Individual animal records will be maintained and abnormalities reported to attending veterinarians. Site Security Animal housing units will remain under double lock except when access is required for animal care or other approved activities. Access to the holding pens will be limited to caretakers and visitors with approval from the Colorado Division of Wildlife (CDOW) Program Manager. Alarms and other security devices may be installed as needed. Handling When handling or transportation is required, lynx will be placed in their individual nest box or alternatively, a skykennel. If required, the lynx will be transferred from their nest box to a custom squeeze cage where they can be hand-injected with an anesthetic or other treatments. Lynx will be anesthetized by, or under the direct supervision of, a veterinarian with 5 mg/kg Telazol, 1M. Temperature, pulse, respiration rate, oxygen saturation, and anesthetic depth will be assessed throughout the anesthetic event. A record for each anesthetic event will be completed (see attached). Supplemental oxygen will be provided by mask as needed. Lynx will not be returned to their pens until recovered from anesthesia. Examination and Animal Identification Lynx will receive a brief visual examination by a veterinarian upon arrival at the holding facility. Abnormalities will be noted and appropriate care provided. After 1-7 days rest, each lynx will be anesthetized for thorough examination. We will obtain a body weight and morphometric measurements, assess body condition, estimate age (based on tooth wear), take a photograph of the face, collect hair and whole blood for DNA analysis (R.Ramey, University of Colorado, Boulder) and blood for archiving, administer penicillin prophylactically (22,000 U/kg, sq, perform a thorough physical examination, and apply needed treatments and markings for identification. The physical examination will include close inspection of the condition of toes, claws, and teeth. We will use radiology to augment diagnosis of . injuries, identify juveniles (based on lack of closure of the growth plate of the distal radius and ulna), and confirm late pregnancy. Other diagnostic samples will be collected as needed if abnormalities are found. If �56 written documentation is not available to confirm that required anthelmintic treatments have been provided, we will administer praziquantel (5 mg/kg sq, ivermectin (0.3 mg/kg sq, and topical dusting with carbaryl. Lynx will be identified with two subcutaneously placed transponders (one on dorsal midline, the other in the intermandibular area) and an eartag. A radiocollar will be placed on each lynx about 2-14 days prior to release. If additional anesthesias are required for placement of radiocollar, treatment, etc., determination of body weight and other required procedures may be repeated as well. Treatment and Euthanasia Medical treatments will be provided by, or under the direct supervision of, the attending veterinarians as needed. Surgery will be performed by a board certified veterinary surgeon. If euthanasia is required, ideally it will be performed with the lynx under anesthesia using an overdose of barbiturates. A diagnostic necropsy will be performed on all mortalities by a board certified veterinary pathologist. �57 ANIMAL CAPTURE RECORD Date: / / 99 Name of investigator(s): M Sex: 0 L nx Species: Age:__ F 0 ~~~------------------------------ Est 0 Actual 0 Weight Est _ Actual 0 0 Body Condition: Animal No. 0 0 Excel Good ------Fair 0 Poor 0 Purpose of Capture: ~P.:,_re=--,.:..re=-/:..:e-=a:..:s-=e:-e:.:x..:.::a~m.:.:.:.... -=Location of Capture: _::_D,.:..ie::...:t_::_e.:_:ri_::_c.:_:h:..:'s_:H_:_:_o.:_:ld::...:in::-'9"'--'-P_:e_:_n.:_:s'-=:--.,:-:-:Ambient Temperature: Weather Conditions: _ _ _ Drug Administration Time Drug Method Dose Location Ivermectin Praziquantel Penicillin Event Times Immobilized: Finish Procedures: Start Procedures: Recovered: Vital Signs Temp Time Pulse Respiration Results of Physical Exam: Body Measurements: Samples Taken: D Blood-EDTA Blood-BIT H ir BodvLenath Tail Lenath D em em D D em cm D Collar Freq Markings Transponder Number em Ear Tag Number Location Color Left Right Comments: Colorado Division of Wildlife Revision Date: 08124100 �58 Attachment Protocol for post-mortem Contacts: 2 sampling of lynx Dr. Margaret Wild (CDOW) Dr. Dan Gould (CSVDL) All information releases and contact with press need to be through the CDOW. Margaret Wild will coordinate. The objectives of the post-mortem examination are to: 1) determine the caus'e of death and document with evidence, 2) collect samples for a variety of research projects, 3) archive samples for future reference (research or forensic). The gross necropsy and histology will be performed by, or under the lead and direct supervision of, a board certified veterinary pathologist. Preferably, Drs. Margaret Wild and/or Tanya Shenk of the Colorado Division of Wildlife (CDOW) will also be present. In general, the protocol will follow standard procedures used for thorough post-mortem examination and sample collection for histopathology and diagnostic testing. Some other data/samples listed below will be routinely collected for research, forensics, and archiving. Additional data/samples may be collected based on the circumstances of the death (e.g., photographs, video, radiographs, bullet recovery, samples for toxicology or other diagnostic tests, etc.). Procedures: 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. Remove the radio collar (animal ID will be the number written on the collar) Determine the body weight and note general body condition Note condition of the claws and teeth Collect external parasites and submit for ID If the carcass is in good condition, please skin and save the pelt (freeze) Remove and save PIT tags (about 1.5mm x 12mm rod-shaped transponder--one under chin, the other on dorsal midline between the scapulae) Conduct necropsy Collect samples of all major organs including long bone, muscle, spinal cord, etc. (formalin) Collect samples of all major organs including long bone, muscle, spinal cord, etc. (freeze) Collect a 3cm x 3cm section of muscle (freeze) Note condition of bone marrow in femur or other long bone Collect a sample of vas deferens and testis (bag with a moist gauze and refrigerate) Bag gut contents that could be used to determine diet (freeze) Collect a fecal sample and submit for parasitology Collect brain. Submit half for rabies exam; save other half in formalin After examination, place the skull in one bag, the pelt in another bag, and other remains in a separate bag (freeze) The CDOW will retain all samples and carcass remains with the exception of tissues in formalin for histopathology, brain for rabies exam, feces for parasitology, external parasites for ID, other diagnostic samples. �59 Table 1. Mortality oflynx reintroduced to Colorado, February-August 1999. Animal ID Sex Age Protocol Date of Death Weeks Out Cause of Death YKM0199 M M 2 N/A 04/29/99 N/A Undetermined 0 1 02/24/99 3 StaIvation BC99-09 BC99-07 F 2 1 02/26/99 3 StaIvation F 3 F M 0 1 2 03/16/99 BC99-08 AKM2399 StaIvation StaIvation 1 3 06/18/99 6 7 4 AKF0499 F F 1 2 2 3 06/12/99 StaIvation 3 3 3 3 07/19/99 5 7 BC99-01 BC99-06 AKF1799 AKF0899 AKF1899 F F F 5 2 04/10/99 Shot HBC HBC Undetermined Trawna, emaciation 11 12 15 07/24/99 07/31199 08/26/99 1600 1400 Of) .._... -(1) 1200 ~ s:: 1000 ..Q 800 0 600 ·ca ~ 400 200 0 3/10 3/17 3/24 1-'- 3/31 Males -;;- 417 Females 4/14 4121 A Pregnant 4/28 515 1 Fig. 1. Average daily feed intake of captive lynx reported weekly from 10 March through 5 May 1999. �60 16 14 12 00 ~ '-" 10 </l ~ ::E 8 ;;.-. 6 "'0 0 o::l 4 2 0 Be AK YK 10 Arrival Wt [] Release Wt Fig. 2a. Body weight of adult male lynx from British Columbia (BC; n (YK.; n = 7 arrival, n = 5 release), and Alaska (AK; n = 7). I = 5 arrival, n = 3 release), Yukon 12 10 00 ~ '-" ~ 8 00 .a3 ~ "'0 6 ;;.-. 0 o::l 4 2 0 Be AK YK I 0 Arrival Wt [] Release Wt I Fig.2b. Body mass of juvenile female lynx from British Columbia (BC, n = 1), Yukon (YK., n = 1), and Alaska (AK, n = 3). �61 14 12 -. ~ '-" <Il ~ 10 8 >. 6 a:l 4 -0 0 2 0 BC-O BC-P YK-P YK-O AK-O AK-P Fig 2e. Body mass of adult female lynx from British Columbia (BC; open n = 4 arrival, n = 1, release; pregnant n = 0), Yukon (YK; open n = 2 arrival, n = 1 release; pregnant n = 1), and Alaska (AK; open n 3; pregnant n = 7). 6 5 ~ 4 (1.) 1 3 2 1 0 Shot Starve Trauma Cause of Death IIIII\/Iale III Female I Fig. 3. Lynx mortality by cause through August 1999. HBC = �62 4 3 ,_ ~ .D e i 2 1 o Feb mStarve Mar Apr .Unk May (]llShot Fig. 4. Lynx mortality by month through August 1999. Jun IIIHBC Jul Aug • Trauma �63 Colorado Division of Wildlife Wildlife Research Report July 2000 JOB FINAL REPORT State of. Project No. ___;C::::;o~l~o~rad=o___; _ Cost Center 3430 -'W..:,_-....:1=5=3_:-R""--..,:.1=-3 _ Mammals Research Program Work Package No. __ Task No. ---"'0=67.:....;0"-~3 _ _ Lynx Conservation Assessing Abundance of Snowshoe Hares Period Covered: July 1, 1999 - June 30,2000 Authors: D. F. Reed Personnel: D. FReed, G. Byrne, T. Shenk, 1. Kindler, P. Albers, T. Black, H. McNally, S. Znamenacek, K. Buell, S. King, G. Patton, J. Zahratka, L. Uphham, B. Andree, J. Hicks,°L. Green, K. Wright, B. Heicher, P. Jones, D. Homan, R. Adams, S. Wait, D. Kenvin, B. Woodward, D. Boyer, K. Cordove, B. Ochs, T. McLean, A. Bellotti, K. Schrollt, G. Cross, F. Kelffner, G. Madison, L. Dwyer, D. Swanson, D. Brownne, F. Pusateri, M. Wunder. ABSTRACT Data on hare density for this report were derived from pellet surveys using methods describe by Krebs et al. 1987. Samples were distributed among 5 blocks selected by biologist who believed them to be suitable for sustaining lynx populations. Sample sizes of plot arrays ranged from 62-239 with the most plot arrays occurring in block 5 (n = 239). Density calculations yielded 0.44 hares per ha for block 5 and lower densities for the remaining 4 blocks. Hare pellet densities were highest in white fir habitats; followed by limber pine and Englemann spruce habitats. Pellet densities were lowest in Gambel's oak and ponderosa pine habitats, but these were also habitats which were not intensively sampled. The most relevant results in this report are estimates of snowshoe density, the type of overstory in which .the highest densities occurred (Engleman Spruce, based on those with adequate sample size) and where the highest densities occurred (National Forests/wilderness areas and counties in Block 5; Figs. 4-8, 9, 10, 12 in Appendix 1). These indicate marginal densities for supporting reintroduced lynx according to the literature and, professionals working with these species. Granted that most of the work and most of the ...professionals working in the field are in the northern boreal forests where snowshoe populations cycle dramatically. Poole (1995) reported that concomitant with the first full winter of low hare density in the Northwest Territories (91-92), all resident lynx died or dispersed in a 135-km2 study area. Marginal habitat conditions for hares probably result in a scarcity of prey and may explain the relatively large home ranges (and hence their behavioral adaptations toward dispersal) of the lynx (Koehler 1990, Roloff 1997). Demographic characteristics of lynx, if any successfully establish home ranges, may be representative of lynx populations along the southern periphery of their range where habitat conditions are marginal for snowshoe hares. �64 �65 SNOWSHOE HARE DENSITY /DISTRIBUTION ESTIMATES AND POTENTIAL RELEASE SITES FOR REINTRODUCING LYNX IN COLORADO D.F. Reed P.N. OBJECTIVES Analyze, summarize, and organize snowshoe hare survey data into a comprehensive final report for release to the public (Division Report No. 20). SEGMENT OBJECTIVES 1. Analyze, summarize, and organize snowshoe hare survey data into a comprehensive final report for release to the public (Division Report No. 20). RESULTS Results of the analysis of snowshoe hare pellet sampling are detailed in the attached Appendix 1. Division staff contracted with West, Inc. to evaluate and review the adequacy of both the methodology and the analysis of these data and that report is contained in Appendix 2 �66 �67 APPENDIX 1. SNOWSHOE HARE DENSITYIDISTRIBUTION ESTIMATES AND POTENTIAL RELEASE SITES FOR REINTRODUCING LYNX IN COLORADO Dale F. Reed FOREWORD Native wildlife are valuable resources in Colorado. Such values are manifested in several ways. Some people enjoy seeing or photographing non-hunted species or get satisfaction from merely knowing that they exist as important components of the ecosystem. During the 1980s and 1990s interest increased in the concept of ecosystem management and in maintaining or enhancing biodiversity. Furthermore, interest in enhancing the numbers of or in reintroducing species such as the lynx (Felis lynx) gained momentum in the late 1990s. To that end, a reintroduction oflynx was planned for the winter and spring of 1999 and a prerequisite to such an effort should involve examining the distribution and abundance of its principal prey, the snowshoe hare (Lepus americanus). TABLE OF CONTENTS INTRODUCTION. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1 METHODS . Krebs Plots Habitat Attributes at Plot Sites Model Probability Surface Potential Release Sites RESULTS AND DISCUSSION Krebs Plots Habitat Attributes at Plot Sites Model Probability Surface Potential Release Sites CONCLUSIONS/SUMMARY ACKNOWLEDGMENTS LITERATURE CITED TABLES 1 Number of points and plot arrays 2 Wyoming and Colorado hare. densities 3 Probability of suitable hare habitat FIGURES 1 GAP Vegetation Classification and Survey Blocks 1-5 2 Random Sample Points Generated for Krebs plots 3 Krebs Plot Points Completed 4 Map of Snowshoe Hare Density by National Forest 5 Graph of Snowshoe Hare Density by National Forest 6 Map of Snowshoe Hare Density by County 7 Graph of Snowshoe Hare Density by County 8 Map of Snowshoe Hare Density by Block . . . . . . . . . . . . . . . . . . . . . . . �68 9 Graph of Snowshoe Hare Density by Block . 10 Snowshoe Hare Density by Block Using "Old" Equation . 11 Snowshoe Hare Density by Elevation . 12 Snowshoe Hare Density by Primary Overstory . 13 Snowshoe Hare Density by Forest Structure . 14 Snowshoe Hare Density by Primary Understory : . 15 Snowshoe Hare Density by Understory Density . 16 Snowshoe Hare Density by Slope . 17 Snowshoe Hare Density by Aspect . 18 Snowshoe Hare Density by Domestic Grazing . 19 Model Probability Surface for Blocks 1-5 . 20 Predicted Areas with Greater than 80% Probability of Suitable Hare Habitat in Block 5 . APPENDICIES A Krebs Pellet Plot Field Form . B Selected Data from Block 1 . C Selected Data from Block 2 D Selected Data from Block 3 E Selected Data from Block 4 . F Selected Data from Block 5 . G Genus-species Understory Abbreviations in Fig. 14 . H Summary by Hare Densities per Block, Primary Overstory, National ForestlWilderness Areas, and by County . �69 SNOWSHOE HARE DENSITYIDISTRIBUTION ESTIMATES AND POTENTIAL RELEASE SITES FOR REINTRODUCING LYNX IN COLORADO INTRODUCTION Based on a renewed interest in the lynx and the possibility of reintroducing the animal in Colorado, four agencies including the U.S. Forest service, National Park Service, U.S. Fish and Wildlife Service, and Colorado Division of Wildlife prepared a draft strategy for the conservation and reestablishment oflynx (Felis lynx) in the southern Rocky Mountains (Seidel et al. 1998). This draft outlined generally two methods for assessing snowshoe hare (Lepus americanus) habitat use or presence and abundance or population trends as recommended by Weaver (1997). The first method was winter track surveys and the second was counts of snowshoe hare pellets (fecal droppings). Tracks in the snow have long been used to note habitat use and the presence of animals (Forrest 1988, Halfpenny 1986). However, only careful approaches or methods may yield definitive density or population estimates (Becker et al. 1998). A winter track survey was completed during the winter months of 1998 and reported by Byrne (1998). . Counts of snowshoe hare fecal pellets have been used to estimate population trends or density (Angerbjorn 1983, Krebs et al. 1987, Wolff 1982). Angerbjorn (1983) estimated density in Sweden from counts of square-shaped quadrats of 0.1 m2 and obtained log-log regression: log (pellets) = 1.02 log (hare density) + 0.44, where hare density is per hectare (ha). He found a close relationship between pellet counts and hare numbers (r2 = 0.91). Conversely, Wolff (1982: 141) indicated that It ••• pellet counts can be used to indicate habitat use or population trends, but not to estimate population numbers." Krebs et al. (1987) found a relationship between mean pellet counts and mean hare density on 50 quadrats of 0.155 m2 (5.08 x 305 em). A modified version of his equation, log., (hares/ha) = ([log)o (pellets) - 0.2359] x 2.303) + 0.2773, was used by Slough and Mowat (1996). Since then, the equation has been modified several times. The purpose of this document is to report on snowshoe hare pellet counts, the results of estimating hare densities across major habitat types and associated selected habitat characteristics, and an approach for estimating hare habitat as it relates to providing the primary prey for reintroduced lynx. METHODS Krebs Plots Five major core areas were delineated for the Kreb's pelletplot counts (Fig. 1). The blocks were selected based on large contiguous parcels ofland that appeared to have suitable habitat (oak, aspen, coniferous forest type or a mixture of these types). The vegetation data is from the Colorado GAP Analysis project (Thompson et al. 1993). Natural or manmade barriers, such as Interstate 70, further define the block boundaries. Small isolated blocks of suitable habitat such as Greenhorn Mountain, the Roan Plateau, Sangre de Cristos, and the Uncomphagre Plateau were not considered in the hare data collection and the GIS maps. Random points for each of the five blocks were generated using a program modified after Zack (pers. com., 1. Zack) which created a grid for each vegetation type with random numbers between 0 and 100,000. A sub-program checked the number of cells containing identical values, and aggregated these until a total was reached that was equal or greater than the required number of random points. The centroids of the cells with values lower than this cut-offvalue were then plotted as an ArclInfo coverage. �70 The number of random points generated across the vegetation types (Fig. 2) were about 200 for blocks I, 4, and 5, and 100 for blocks 2 and 3 (in some cases random point program generated a few extra points). Initially, every 4th random point ploted on map (BLM Surface Mangement Status 1:100,000 scale) overlays were selected to insure wide distribution and randomness in case all points could not be completed. According to the protocol used, once the location was determined by use of a GPS unit (matching UTM coordinates generated for a given point as listed on the overlay) or map orienteering, a random bearing (0360 deg) was chosen, and a rectangular array of 10 Krebs plots was set (A), each 30 m apart (rectangular array = 30 m x 120 m, with 5 plots per side in direction and then in opposite direction of the bearing). Additionally, a second array (B) was selected at a randomly chosen bearing and distance (10-190 m inclusive) from the completion point (about 30 m from the beginning of the first array) of the first array (A). Hence, pellet counts were made for two fairly close areas near each randomly selected point. These pellets were tallied as either new or old and their means calculated. New and old pellets were differentiated by color, lighter tan being judged "new" (estimated from current season), and darker gray being judged "old" (estimated from before current season). The mean number of pellets (2 sets of 10, i.e. n = 20 plots for each randomly selected point [Fig. 3]) was entered into a published equation to produce an estimated number of hares per hectare. This equation was a modified version of Krebs et al. (1987) (log., hares/ha = {[(0.8127 x log., pellets) - 0.2359] x 2.303} + 0.2773, as in Slough and Mowat 1996). Later, the equation was modified (per. com., K. Poole, 9 Nov 98). Still later, it was modified again (per. com., C. Krebs, 29 Sep 98 and 24 Nov 98). This third iteration, 10& (hareslha) = (0.888962 x 10& [pellets] - 1.203391) x 1.567026, was recommended by Charles Krebs (per. com., C. Krebs, 24 Nov 98) and the one ultimately used for the overall analyses. It was based on his review of his data (collected samples from the Yukon: 1976-1996, n = 86, calculated from annual pellet counts in June and average densities of hares from the previous spring and autumn estimated by markrecapture including adding a correction for bias [SprugeI1983]). Habitat Attributes at Plot Sites Categories other than those related to the plot locations (Name of Property, Game Management Unit [GMU], County, Lynx Survey Block Number, General Location Description, Land Status, and UTM coordinates) recorded on the field forms (Appendix A) included: - elevation - overstory (primary, secondary, and tertiary) - vegetative structure (e.g. mature, old growth) - understory vegetation (primary, secondary, and tertiary) - understory vegetation density (estimated visual obstruction of vegetation from 10m in front of a vertical 1 x 6 ft [0.30 x 1.83 m] white panel, marked off in ft2 [0.30 nr'], where highest value of 6.0 = no visual obstruction, and to be recorded with photograph) - slope - aspect - overstory and understory distribution (patchy or continuous) - domestic grazing presence or absence. Model Probability Surface A logistic model was used to generate a continuous surface representing the probability of any geographic location supporting snowshoe hare habitat (pers. com., M. Wunder). The results from the Krebs plot effort were examined spatially using chi-squared analysis to produce parameter estimates for the model. �71 Parameter values were estimated for each of vegetation (primary cover type), elevation, slope, and aspect using the Krebs plot coverage, the vegetation coverage from the GAP analysis project, and a digital elevation model (DEM) for the study area. The DEM was used to generate separate grids for elevation, slope and aspect. Parameter estimates with P-values less than 0.05 were then used in the logistic model, which was then run using the pertinent grids. The model equation follows: 1/I+exp(Ao + Aa * aspect + Ae * elevation + Av * vegetation), where Ao is the overall parameter estimate, Aa is the estimate value for aspect, Ae is that for elevation, and Av is that for vegetation. Release Sites Initially, release sites were selected based on the model probability surface, but for practical reasons of access and managing the media and number of observers during the releases, private land areas abutting potential lynx habitat or proximity to the Weminuche Wilderness were given high priority. Potential sites were inspected and coordinated with landowners and caretakers where appropriate. RESUL TS AND DISCUSSION Krebs Plots Paired pellet plot arrays and single plot arrays were completed at 303 and 5 randomly selected points, respectively. Sample sizes of the number of plot arrays ranged from 62-239 (Table 1, Appendices B-F) with the most plot arrays (n = 239) completed in block 5 (Appendix F). Distribution of hare densities by National Forest (Figs. 4-5), by county (Figs. 6-7), and by block (Fig. 8), suggest that block 5 had the highest hare density. Density calculations yielded 0.44 hares per ha for block 5 and lower densities for the remaining blocks (Fig. 9). This mean of hares per ha in block 5 results from using the latest regression equation from Krebs as discussed under Krebs Plots in Methods. The initial calculation using an. earlier equation (Slough and Mowat [1996]) yielded a mean of almost 1.2 hares per ha (Fig. lO). Using different equations and possibly using them incorrectly yields substantially different results. Also, initial analysis of block 5 data using a SAS program used in the Northwest Territories (per. com., K. Poole, 14 Sep 98) was corrected as were the analyses of data from 2 areas over 2 years in Wyoming, where lynx were present (per. com., K. Poole, 9 Sep 98; per. com. T. Laurion, 18 May 99) (Table 2). Our calculations using the latest regression equation from Krebs (per. com., C. Krebs, 27 Oct 98 and 24 Nov 98), matched these same values (0.817, 0.856, 0.694, and 1.429, respectively) (Table 2). Thus, using the most current accepted calculations, the Wyoming areas having lynx present, had hare 'densities ranging from 0.817-1.429, similar to some other densities reported (0.73/ ha, Dolbeer and Clark 1975; 1.401ha, Lima 1998; 2.401ha, Meslow and Keith 1968), but subtantially lower than others (12.3/ha, Baileyet al. 1986), and all higher than that considered the lower "edge" for suppporting lynx (0.5/ha; C. , .Krebs, per. comm., 24 Nov 98). Dolbeer and Clark's (1975) 0.73 hares per ha might be considered a model for habitat in Colorado, but another study using 175 permanent plots, reports lower densities (0.053/ha in lodgepole pine and O.068/ha in spruce/fir) (Shivery 1997). This causes concern that the density of our 0.44 hares per ha in block 5 may not be sufficient as a primary prey base to support lynx. Part of the problem, however, is that our data were collected randomly across 11 primary overstory types. The Wyoming data were collected from plots selected in coniferous forest types. �72 It is generally accepted that some of the primary overstory types (probably the 3 non-conifer types [aspen {Populus tremuloides}, willow {Salix spp}, and Gamble's oak {Quercus gambelii} or mixed mountain shrub], ponderosa pine (Pinus ponderosa), and bristlecone pine (Pinus aristata) might be expected to have low hare densities. Apparently this was the case based on our small sample sizes. Most of our samples were taken in 6 conifer types described below. Hence, the problem remains - we apparently have a relatively low density of hares if compared to the Wyoming data, but is it comparable? Given that Wyoming's plots were taken non-randomly in optimal habitats, the extent to which optimal habitat considerations could induce bias is unknown. It is possible that if we had used our method of selecting random samples in Wyoming that their means would have been lower. Similarly, it is possible that if we had sampled randomly in "optimal habitats" (whatever those are in Colorado) that our means would have been higher. In either case, how much lower, or higher, is simply unknown. This probably limits the utility of comparing these 2 sets of data. Habitat Attributes at Plot Sites Attributes from the plot locations as prescribed by data collection (Appendix A) across densities, included elevation (Fig. 11), 11 primary overstory or canopy types, - Lodgepole pine (pinus contorta) - Englemann spruce (Picea engelmannli) - Subalpine fir (Abies lasiocarpa) - Douglas fir (pseudotsuga menztesli) - Ponderosa pine - Limber pine (pinus flexilis) - Bristlecone pine - White fir (Abies concolor) -Aspen - Willow - Gamble's oak or mixed mountain shrub (Fig. 12), structure (Fig. 13), primary understory vegetation (Fig. 14), vegetation density indices (Fig. 15), slope (Fig. 16), aspect (Fig. 17), and grazing (Fig. 18). The relationships between hare density and some of these attributes, i.e. elevation, structure, primary understory, aspect, and grazing, yielded relatively predictable results. Forest types preferred by hares in Colorado occur at higher elevations (Fig. 11). Limber pine and white fir show the highest hare densities, but with very small sample sizes these densities are probably not representative (Fig. 12). Conversely, Englemann spruce had a sample of 220, lodgepole pine 108, subalpine fir 64, and Douglas fir 48 (Fig. 12). Willow had only one sample where it was considered the primary overstory and is not included in Fig. 12. For structure, most of the plots completed were in mature or old growth. The small sample of sapling/pole (n = 49 of 606 records, with only about 50% of the 49 having any pellets) and the even more negligible sample sizes ofGrasslForb and Shrub/Seedling, essentially yield zero densities (Fig. 13). Selected understory species appear to be associated with higher hare densities than others (Fig. 14), but another measurement, the vegetation density indices, may suggest that the amount and height of the understory is limited (Fig. 15 and 376 photographs appended to data records). Most of the samples (245 and 97 of 428 records) occurred in the 5.0-6.0 and 4.0-4.9 categories, respectively, where little understory obscured the panel (Fig. 15). The value for <l.0 is unlikely to be representative. Other workers have suggested that predation on snowshoe hares may be heaviest in habitats that have little understory (Sievert and Keith 1985), that snowshoe hares avoided open understories and greater understory �73 density provided both escape and thermal cover (Litvaitis et al. 1985), and that height of understory was important (Wolfe et al. [1982] found a strong correlation between hare use and cover densities at heights of 1.0-2.5 m above ground, and that vegetation types in which cover densities above snow level [1.0-1.5 m] were at least 40%, accounted for 85% of winter use by hares). Hare density shows a bimodal response to slope (Fig. 16), but upon inspection of photographs it appears that at least one field person consistently over-estimated degree of slope, probably skewing these data. In relation to aspect, hares probably avoided the warmer, dryer, southern exposures (Fig. 17). Hare density means from plots judged to have domestic grazing or not (Fig. 18) were predictably no different (0.358 vs 0.342, respectively). How recent or the extent of any grazing was not estimated. Judging vegetation distribution for overstory and understory as patchy or continuous (Appendix A) was largely dependent on "scale" and was therefore not analyzed. Similarly, "Old" pellets were not analyzed because their age or persistence could not be determined. Generally, these pellets occurred in low numbers and were noted in 402 of613 records. Cottontail (Sylvilagus nuttalli) and jackrabbit (Lepus townsendi) pellets were noted in a low number of records and were not analyzed. Some overlap or abutting of habitat boundaries might be expected between hares and mountain cottontails, but doubtfully with jackrabbits. One could conclude, that if you counted jackrabbit pellets, you were in the "wrong" (i.e., non-hare) habitat. Model Probability Surface The probability values from the model were naturally distributed across five range classes (pers. com., M. Wunder). These were represented as zero percent probability, one to twenty percent, twenty to eighty, eighty to ninety-three and ninety-three to one hundred percent probability (Fig. 19). The most relevant results include the areas that were modeled as greater than 80 percent probability of hosting snowshoe hare habitat. Block 5 included the largest surface area and proportion of area that was modeled greater than 80 percent probability. Areas within a radius of3.2 km from potential lynx release sites that were predicted with greater than 80% probability to provide suitable snowshoe hare habitat were delineated (Fig. 20). The distribution and amount of the area is shown in Table 3. It should be noted, however, that raw values for area may not be the best (and certainly is not the only) criterion for drawing conclusions about the suitability of an area for snowshoe hare or lynx. We did not develop any method for examining the distribution of this area for spatial context. If and when this issue is addressed in the future, it should be acknowledged that there are many varied factors involved with such an analysis. Release Sites Hence, one of the sites north of the Weminuche Wilderness, Humphreys (Goose Creek), was ultimately chosen for the initial release February 3-4, 1999. Similarly, other sites northwest, north, south, and southwest of the Weminuche Wilderness Area, were ultimately chosen for later releases in March. Release sites in Block 5 located northwest, north, and east (n = 4) and south and southwest (n = 4) of the Weminuche Wilderness were initially described using a predicted probability surface of snowshoe hare habitat (per. comm., M. Wunder) and examined using the following criteria: - ownership, public and private lands where permission for egress was obtained - access, where at least snowmobiles could be used to gain access - seclusion, where public access is generally not available - habitat near release site should have a range of 1,000-10,000 ha of continuous or at least only lightly broken conifer forests �74 - connectivity, where forest segments are sufficiently contiguous to allow for traveling lynx to avoid open areas The 4 sites northwest, north, and east of the Weminuche Wilderness Area were ranked as follows: - Humphreys (Goose Creek) - Red Mountain Ck - Rio Grand Reservoir, and - Big Meadows Similarly, the 4 sites located south and southwest of the Weminuche Wilderness Area were ranked as follows: - Weminuche Valley - Beaver Meadows - Endlich Mesa, and - Middle MtnlBear Ck CONCLUSIONS/SUMMARY The most relevant results in this report are the estimates of snowshoe hare density, the type of overstory in which the highest densities occurred (Engleman Spruce, based on those with adequate sample size) and where the highest densities occurred (National Forests/wilderness areas and counties in Block 5; Figs. 4-8, 9, 10, 12, and Appendix H). These indicate marginal densities for supporting reintroduced lynx according to the literature and professionals working with these species. Granted that most of the work and most of the professionals working in this field are in the northern boreal forests where snowshoe hare populations cycle dramatically. Poole (1995) reported that concomitant with the first full winter oflow hare density in the North West Territories (91-92), all resident lynx died or dispersed in a 135-km2 study area. Marginal habitat conditions for hares probably result in a scarcity of prey and may explain the relatively large home ranges (and hence their behavioral adaptations toward dispersal) of the lynx (Koehler 1990, Roloff 1997). Demographic characteristics of lynx, if any successfully establish home ranges, may be representative of lynx populations along the southern periphery of their range where habitat conditions are mariginal for snowshoe hares. ACKNOWLEDGMENTS We especially thank Pam Albers, Travis Black, Heath McNally, and Steven Znamenacek for the completion of most of the Krebs plots. Other agency field assistance came from Steve King, Rocky Mountain National Park; Gary Patton, U.S. Fish and Wildlife; Kit Buell and Jennifer Zahratka, U.S. Forest Service; and Lee Upham and crews, Bureau of Land Management. Field work by Division personnel included Bill Andree, Jim Hicks, Larry Green, Kevin Wright, Bill Heicher, Paul Jones, Doug Homan, Rick Adams, Scot Wait, Dave Kenvin, Dale Reed, Gene Byrne, and Brent Woodward. Gene Byrne organized and coordinated the field work. Work by volunteers included Don Boyer, K. Cordova, Brett Ochs, Travis McLean, Amy Bellotti, Kathrin Schrott, Gretchen Cross, Francis Kleffner, Griffin Madison, Lynn Dwyer, and Dawson Swanson. Geographic Information Systems (GIS) support and cartography provided by Jon Kindler. Dawn Brownne assisted with data entry. Francie Pusateri provided advice and converted Visual dBase files to EXCEL. Mike Wunder, Natural Heritage Program, generated the model probability surface and associated figures. We also thank Tom Beck, Rich Reading, and Tanya Shenk for reviewing the manuscript. Work reported here was funded mostly by Division GoCo funds and from funds from Vail Associates. �75 LITERA TURE CITED Angerbjorn, A 1983. Reliability of pellet counts as density estimates of mountain hares. Finn. Game Res. 41:13-20. Bailey, T. N., E. E. Bangs, M. F. Portner, 1. C. Malloy, and R 1. McAvinchey. 1986. An apparent overexploitated lynx population on the Kenai Peninsula, Alaska. 1. Wildt Manage. 50:279-290. Becker, E. F., M. A Spindler, and T. O. Osborne. 1998. A population estimator based on network sampling of tracks in the snow. 1. Wildl. Manage. 62:968-977. Byrne, G. 1998. A Colorado winter track survey for snowshoe hares and other species. Colo. Div. Wildt 35pp. Dolbeer, R A, and W. R Clark. 1975. Population ecology of snowshoe hares in the central Rocky Mountains. J. Wildt Manage. 39:535-549. Forrest, L. R 1988. Field guide to tracking animals in the snow. Stackpole Books, Harrisburg, PA 185 pp. Halfpenny, 1. 1986. A field guide to mammal tracking in North America. Johnson Publishing Co., Boulder, CO. 161 pp. Koehler, G. M. 1990. Population and habitat characteristics oflynx and snowshoe hare in north central Washington. Can. 1. Zoo!. 68:845-851. Krebs, C. 1., B. S. Gilbert, S. Boutin, and R Boonstra. 1987. Estimation of snowshoe hare density from turd transects. Can. J. Zool. 65:565-567. Lima, S. L. 1998. Nonlethal effects in the ecology ofpredatorprey interactions. BioSci.48:25-34. Litvaitis, 1. A., 1. A Sherburne, and J. A Bissonnette. 1985. Influence of understory characteristics on snowshoe hare habitat use and density. J. Wildl. Manage. 49:866-873. Meslow, E. C., and L. B. Keith. 1968. Demographic parameters ofa snowshoe hare population. 1. Wildt Manage. 32(4):812-834. Poole, K. G. 1995. Spatial organization ofa lynx population. Can. 1. Zool. 73:632-64l. Roloff, G. 1., and J. B. Haufler. 1997. Establishing population viability planning objectives based on habitat potentials. Wildl. Soc. Bull. 25:895-904. Seidel, 1., B. Andree, S. Berlinger, K. Buell, G. Byrne, B. Gill, D. Kenvin, and D. Reed. 1998. Draft strategy for the conservation and reestablishment of lynx and wolverine in the southern Rocky Mountains. U.S. Forest service, National Park Service, U.S. Fish and Wildlife Service, and Colorado Division of Wildlife. 115pp. Shively, K. 1997. Snowshoe hare density in Vail, Colorado; implications for lynx reintroduction. Unpub. Rep.: 1-7. Sievert, P. R, and L. B. Keith. 1985. Survival of snowshoe hares at a geographic range boundary. J. Wildl. Manage. 49:854-866. Slough. B. G., and G. Mowat. 1996. Lynx population dynamics in an untrapped refugium. 1. Wildl. Manage. 60:946-961. Sprugel, D. G. 1983. Correcting for bias in log-transformed allometric equations. Eco1. 64:209-210. Thompson, T. G., W. A Reiners, and D. L. Schrupp. 1993. Colorado gap analysis vegetation classification (draft). Unpub. Rpt. Weaver, J. L. 1997. Reconnaissance oflynx habitat in Colorado. Wildl. Conserv. Soc.IOpp. Wolff, 1. O. 1982. Snowshoe hare. Pages 140-141 in CRC handbook of census methods for terrestrial vertebrates. Edited by D E Davis. CRC Press, Boca Raton, FL. Wolff, 1. 0., N. V. Debyln, C. S. Winchell, and T. R McCabe. 1982. Snowshoe hare cover relationships in northern Utah. 1. Wildl. Manage. 46:662-670. �76 Table 1. Block numbers, number of points generated, points completed (2 plot arrays for each point with 5 exceptions where only 1 array was completed), and plot arrays completed (10 plots for each array). Block No. of Points No. of Points No. of Plot Arrays Generated Completed (2 x points completed) 1 2 3 4 5 Total 200 100 100 200 200 800 57 31 33.5 64.5 119.5 305.5 114 62 67 129 239 611 Table 2. Mean number of hare pellets per Krebs plot and calculated hare densities per ha in 2 areas over 2 years in Wyoming where lynx occurred and in Block 5 of this study. Wyomin~ Colorado Area DUBOIS97 DUBOIS98 MERNA97 MERN98 BLOCK 5 Pellets/plot Hareslha 1.86 0.82 (0.64-1.05)h 1.96 0.86 (0.67-1.09) 1.55 0.69 (0.54-0.89) 3.49 1.43 (1.11-1.84) 0.94 0.44 (0.35-0.56) "Data from T. Laurion, calculations by K. Poole. h95% CI. Table 3. Study areas or blocks, areas with greater than 80 percent probability of providing suitable hare habitat, and the proportion of the study areas involved. Study ArealBlock Area (ha) with> 80% Probability of Proportion (%) of Study Area Providing Suitable Hare Habitat 1 2 3 4 5 Total 460,753 97,920 110,233 587,903 658,258 1915067 20.3 11.7 15.7 21.4 24.1 93.2 �o , 1· •••\ _. - 1 • '- I - - --------,-- - ,, - I - , 1 , '- I -- 77 �-..J 00 * Ft~. z Sample Point ##••••,/ Study Area BoundAly , /' -,' ' County Boundary /\j' I) , __ .1 IntenrtAte Highway ,.,. 100 . IJI--. ~ o 40 Mileo 80 ,, : ,, , , 1 , ,- - - - - - - - - - , - - - - - - - - - -1- - - - - -1- - -,- - - - - - , ,, , r-~----------~---~------, \ , I' \ r" ""," , •... \ .. ,,1 * , '** ~~ r---*-*-*-l y * _"-.".( ** *, * * ~ '* * :, ** '* ,* *** , ** : ' ,, : '* * :* '* *' * * "***,* ** *--- -"'-_!' -- -~*-~- - --;- -~ - --:"t , * *f.. ,* ** **, * *: Al{llII1osa ***, "'*' 1 I : r I .: I I ,:,: I I 5 • Durango, .; , * * * •••.•••.....................•. '-- I * 'lit I I • 1Jc*: ~~ *~ **: 11'< ** '.* .••. d *:"C I ' I -- .•.. ... , \ , "'", , / I , '-, I '\ \ ) , r I( ,'" ,, I '.J , 1- _ --~-----I-- -- �**' ,, ~___..,.. t .f '~ * 1 *,,,_ i ,* * *\. ,'* '* ) ~\* * * * "..,', ' .• ,-,*_, _"*'. *, ~-- L. ".'•.......••: .:.. •••••••• .• ~ ,,--*-;t. **:*': .: r -------------, ------ *., ~*" : ----- .. - - _"":. ~ -- , • * lr , .!!' --..---"" ' *.• *'*,* - - - " " * -, * * o ' ': , • •.. : • I ---------I_ : * "t -----1 I, ,'_ ••.J,_I" * \ * * J-.- -L, '* *,...~.... \ . +,« *, G - - _m _ .,•.•••••. *: • * ** *~ , * ~ ... " ~Vista t:.••.• : -, :: , 'Y,/ I" * *' ~ -* -* - - ;rl I , A\. ",*" ( *, * ~'\ * *, * , * * ,* I ·. • 5 I I I I I 1i* '* """ "" ..................••....... ... • - '/It , - - - - I ,-_.-- ,, , : 'r--------------, *_....... :',' . : .., • * - - - ':" \, ,,, 1... """ • .: ', • : ** \ d • " *: "'- ,, , , / I I 1,-/"" , ..:__~ : ••. , r' i~ " I \ "'\ ------1------------"" ........__.__ I 1- - ,, - - - - - - - - - -1- - - - - - : v c, , : •.. yt, ......•••. \ J' _ " •• •• ~ *** *: ,* *, __..t '4t* ,* ** * .•. ,*-----~-------""'r ""* * ~*' ** .• * ' • _, * r' ,a =; " it •, Durango f , 1 " , \.* '-'I! ,)..._1;:', *** * "-,;'. ..,.\ *. * .., '*--*J!.., .,..**,' ''* * *', '*, ' ("""\*" 1- *. - *. I r-: - " ,, c....*, : * \ __•• ~•••••••••• L __ 100 , , C,,** \ .. --', 11"., . : *: ____ ...,' *i!'* ,*:t * ,'* * f!*- - - - - - - -..., * '* . *' '* I I I ~. _I : = - - - - - - - -, 11" • , -_.1_----------: , 1"* * ' ...• " ~ontrose' • .' •• .!.•••••••••••• • :tlI_,;) . . -l[J:~~~~':~~L~~~~ r ~~~~~ " .~ *, -*- - * JI .; • I--'----A- ~ Highway ••• ,, -;.- - - --li;tt- - - --' '" / /' ,/ ' Coum:y Boundary /'vf' cIntemato , 1""../ ,*- Study Area Boundary ----~------------ JI_ --- ".~.,/ : ,, G'~, s Samplo Point * ~ • •• - .: ",r';f,. 'r: -- -- i -,",. ....._ ~ ',_ * \, * *. * ,, J j_ ,'. •• - •. '.'''': * I. ': *,*. ' ~ •• "t._• -.••"·-11-;\:. ~ -- - • * •• J __ ,-- - - - - - - - -- ~ SNOWSHOE HARE DENSITY SA --- , >- / __ , \ •••. , '\ ", , - " ' ~...J 1I _ -_....f I' ' _ , _ I ~-- _ \ / Al!.llII1osa ) '( , -..Jl ~ �80 E-< ~ ..J f£ ~ ~ ~ >~ b rIl ~ ~ ~ rIl s ~ ~ rIl "<I; ~ 0 1, :xl ~ ~ Hl ~ 0 "'l ~ J ~~ ~$ ~ 0 :xl :xl • d 0 -. ::r- 0 0 I~~!I i:.I cJ7 '-i:::: 1~~~ :xl ~ o 0 I) ~ - --------1 I _ ..1_ ---"'---1 I - - - - - - 7-- , ~-- , , �:::t: II) (i; en ,S "C~, ..• CD zr II) o o ~ o t-.> o w o :,:,.. o 0'1 o m o -.J !=> ex> !=> co ARAPAHO NF BUFFALO PEAKS WILDERNESS GRAND MESA NF ~ GUNNISON NF LAGARITA WILDERNESS AREA CD Al CO CD :::I: D) (i; 6' RIOGRANDE ~ ~ ROCKY MOUNTAIN NATIONAL PARK (i; en ..• NF ~ t/) ~ C" ROOSEVELT NF ,.. '< o (i; l> m CD :::s CD en en C ROUTT NF t/) r+ SAN ISABEL NF SANJUAN NF UNCOMPAHGRE NF WEST ELK WILDERNESS WHITE RIVER NF --.-, LS> VI 18 �00 N SNOWSHOE HARE DENSITY BY COUNTY III0.3 - 0.399 H~ 0.001 -.099 HaIesiha IIIOA- 0.499 H~ o Harealha 0.1 - 0.199 Hareslha • >0.5 H~ 0.2 - 0.299 Hareslha e XlI ~ _ h'~,~ ~ . 40 o Mila ~---------'f'~ I I , I - - .!_ - - ~ - - - - - - -I I ,----------- I :- , 1 I I I ,, - - - - - - - - -1- - - - -- ,>-, ,, I ,- , I I I \ \ , 'r ,'" , I r '.J , 1- --,------ _ I - - - - - - - 1- - - - - ...• _ - - - - - - - - - - - �0 0 0 0 0 0 0 0 0 0 0 0 0 N ~ 0 0 0 0 0 0 0 m 0 en 0 0 0 0 N 0 0 0 ::I: ..• I» CD (I) 1j CD ..• =rI» :I: ., SU CD C (I) ::l en ~ o o s::: :l ~ 0" '< o o C ::l ~ " -.() ~3 £8 �84 ci V I a 0 illill o ~--, I ~-- ,. I T - - -- - •.. - n=> I I \ l-/' , _--, , r , , - j ~.... - - r ...;.., _\.).... -\, - _'- - - - - - �85 0.60 0.50 0.40 ra .J::. •... Q) Q. (/) 0.30 Q) •... ra ::c 0.20 0.10 0.00 Fig. 9. Mean hare density (wi SO) by block (samp~e sizes 10. bars) . Mean hare density by block using Kreb's oriqinal 10). 5 4 3 2 Fig. same as in Fig. equation (sample sizes above �86 f'. I ~. 11 2.5 --0- 2 co ~ :0 :E. 'II Q) ::- 1.5 'iii c Q) 0 e co I 0.5 o 8,000 - 8,999 7,000 - 7,999 <7,000 FT. 9,000 - 9,999 10,000 -10,999 11,000 - 11,999 >=12,000 Eltvation Ranges 1.2000 1.0000 0.8000 Hares per ha 0.6000 0.4000 0.2000 0.0000 Z W 0.. 0.. w Z W (f) « ~ 0 () W ...J I(f) a: 0::: Q; (f) lL W u:: :s (9 ::J 0 0 ~ 0.. ..J « III ::J (f) W :>C W 0.. w ..J 0 a:: ~ 0 0.. ill (9 0 0 ..J « Z W « (f) III 0 (f) ..J :::2; « (9 Z Zw «(.) :::2;::J wo::: ..Jo.. 0::: ~(f) ill ill 0 Z 0 0.. III Fig. 12. Hare density by primary overstory (sample sizes above bars). 0::: u:: w I- I ~ W ~ 0.. 0::: W III ~ ..J �87 Hare Density by Forest Structure 1.6 1.4 1.2 ....,. ro ~ </) e to ::c ~ 0.8 </) c III a ~ 0.6 ::c 0.4 0.2 o SHRUB/SEEDLING GRASS/FORB SAPLING/POLE ~orest MATURE STAND OLD GROWTH Structure 08 III a. 06 </) ero ::c 04 0.2 o AMAL Fig. J 4 ARCO Hare density ARUV by selected CA spp primary JU spp understories (number abbr. see Appendix QUGA of records G). SHCA SYAL n = 10 or >, and pellets SYOC n = 1 or >; VACA VASC �88 0.5 0.4 Hares per ha 0.3 0.2 0.1 o 5.0-6.0 4.0-4.9 3.0-3.9 2.0-2.9 1.0-1.9 < 1.0 I i Fiq, 15. Vegetation density indices by hare density. 2 '" L: Vi ~ ~'" .~ <JJ c: 1.5 '" ~ Cl '" J: <10 10·19 20 - 29 30 - 39 40 - 49 Slope Range (degrees) 50 - 59 60 - 69 >= 70 �89 Hare Density by Aspect (Area 5) 1.8 1.6 1.4 i!' '(ii a; 0.8 c ~ ." J: 0.6 0.4 0.2 o NORTH NORTHEAST EAST SOUTHEAST SOUTH SOUTHWEST Aspect 0.3 ro s: 0.25 Vi '" ;;; s: .~ c '" 0 '" 0.2 e '" J: 0.15 Fig. 18. Absence or presence of grazing (lst and 2nd bar, respectively) versus hare density. WEST NORTHWEST FLAT �Fig. 1Cf. Study area-wide view showing predicted probability surface of snowshoe hare nabitat, study area boundaries, roads, and municipality boundaries. Variable Ao Aa As Ae Av Parameter P 15.6638 0.2300 0.0641 ·0.7106 ·1.4909 0.0001 0.0503 0.8573 0.0001 0.0130 " . . ....•, . ~... . ..... -. \0 I~"':'·-,,' o - ;/ t-. /,,/ .',"Q.. ._, ,./.;"f '._" \ .:~ ./ = Model Surface 1/1+exp(Ao+Aa·Aspect+Ae*Elevation+Av·Vegetation) ',-// Highways Study Areas Predicted Probability _0 _ 1-20% ~ 20 - 80at10 ~ '--180-93% '-93·100% . ¥)) ":n: . --....·r·· 'i\? ,/' ./ f· I.. • ........._ •••...... L ..--.L..c Snowshoe hare data used in this map were collected by the CDOW, USFS, NPS, USFWS, biologists and numerous volunteers. The data were compiled and provided by CDOW staff The predictive model was constructed by CNHP staff ~tk .-~ \ _H"i'nn.ul"4 •.. ~ :/~, .;\ o 100 COi01l~ ---_.__ ...- ._.._- .... - '" ....._- ------- ''".\c' .--; ._~__ _._ 100 _-_.__ .. ___l. ,.~ .:/:., . __ • . _ ~ 200 Kilometers .... -- ...'. ----_ .. __ "*' . �Fig. 20. Areas within a radius of 3.2 km from potential lynx release sites that are predicted with greater than 80% probability to provide suitable snowshoe hare habitat. • Potential release sites o 80-1 OO%Probability within two mile radius _0 Wilderness and Research Natural Areas Predicted Probability of Suitable Habitat _1-20% I '.'.; 120-80% ==:J 80-93% --. 93-100% I I Name Endlich Mesa Rio Grande 1463 Reservoi ------ic-----;::;::---.~ Humphreys_ Weminuche I Hectares "' Valley Beaver Meadows Big Meadows , ---~~~ .. §_~_J ~ ."" ':t"i 469 2108 i 921 . Transfer Pk. - --_ Fourmile TH -------------- Middle Fork Piedra 794 ------ --. 656 East Fork Piedra 431 Plumtaw 120 _ ..... _ .... __ .._ ... ---- w.' ·B·~ !J.,. N __ ~.~ o ~~~~~~~~~ s 40 Kilometers .._~ ~ . C'D!0It1JP ------------_._---------- ~ ~ \0 •.... �92 Col0.'0d 0 Di VISIon - - of Wldrfle- Krebs Pellet Plot Survey Form Krebs Plot # Observer( s): Slope: (in Degrees) Name of Property GMU: Lynx Survey Block # (Actual) County: Start Location from? 0 General Location Description: 0 GPS Map 0 0 Aspect: 0 Vegetation DistJibution: Land Status Code: OIerstory UTMY: UTMX: Zone: Understory 0 0 12 13 Datum: A Apf~d,"x -NAD27 \M3S84 NAD83 0 0 0 0 Patchy Patchy Continuous Continuous KREBS PLOT DATA: ntail ~~1j~Cottc New Old J 0 0 0 feet Ja~bbit New Old 1 - - - - -I 2 - - - - 3 I -- - - ---- - - - - - - 0 4 - - - - - - - GPS 0 5 - - - 6 - - - - 7 - - - 8 - - 9 - 10 _ other Primary Overstory: Secondary Overstory: Tertiary Overstory: Vegetative stn.icture Class: GraSSl/FOrb Shrublseedling SaplingJPole Mature Old Growth Understory Vegetative Type: Common Name 1 (Primary) 2 (Seoondary) 3 (Tertiary) Grazed 0 Yes Krebs Form --I -i - --- - - - -- - - - - - - - -I - - - - ---- , Overstory Type: 0 7-8-98 other - Bevation (MSL): From: Tapa - . Rat 0-5 Moderate 6-20 Moderately Steep 21--40 Steep 41 degrees or more 0 NE 0 NW 0 SE C SW N S E W 0 it ; 0 0 0 0 lime: Date: Density Index 0 Yes 0 No Photo Taken? - -- LJ I -I -, C r~ ,. L..; Land Status Codes: ,-- C ~e IrF Private - P U.S. F. S.-F BLM-B State;Land Board - S OOW-D National Pali< - N other - 0 (Explain) No Comments (Continue on back if needed): Overstory Veg, Type Codes: Lodgepole Pine - L ~mann SP'1!ce- S pine Fir- F ~lasFir-D erosa Pine - P Umber Pine - X Bristleoone Pine - B WtiteFir-W Asoeo-A Wllow-Y Gamble's Oak or _C mixed mountain Shrub - -- �KREBSPL LSBNUM FC·21·A FC·21·B FC·95-A FC·95-B FC·97·A FC·97·B FC·101·A FC-101·B FC-178-A FC·178-B VA·62·A VA·62·B VA·132·A VA·132-B VA·144·A VA·144-B VA·149-A VA·149-B VA·150-A VA·150-B VA·155-A VA·155-B VA·161·A VA·161·B VA·163-A VA·163-B EP-4O-A EP-4O-B VA·164·A VA·164-B VA·135-B VA·153-A VA·153-B VA·135-A WA·77·B WA-81·A WA·81·B WA-73-A WA·73-B WA·75-A WA·75-B VA·166-A VA·166-B DW·13-A DW·13-B DW-49-A ,- UTMX 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 446555 446510 448682 448660 428211 428210 425389 425240 450931 450870 388866 389100 394594 394760 371270 371161 406232 406200 393331 393518 406653 406432 405438 405303 403862 403935 453565 453580 397240 397326 392830 373296 373376 392913 403816 407484 407495 406978 407057 405250 405123 406867 407088 449594 449594 434487 UTMY 4488703 4488640 4492238 4492180 4490776 4490840 4489448 4489480 4505757 4505920 4373370 4372950 4424300 4424351 4410075 4410046 4405739 4405872 4405295 4405428 4394020 43~.3960 4386943 4386987 4384857 43&4789 4442252 4442167 4381807 4381750 4419021 4399796 4399898 4419011 4528065 4521979 4522070 4538343 4538167 4531982 4532088 4376295 4376415 4374855 4374855 4398253 SN2 PRIMOVE SN1 SN8 SN3 A ppwd 2 F S S S S S ix (3 SN9 SN10 1 1 1 I L L S S A L F F L S L L L A S S F S F L F F F F L L L S S L L L A L L A A S 2 6 4 1 5 3 1 L .•. ' 1 0 1 1 1 1 1 1 1 1 4 1 1 5 3 . 1 2 1 21 2 1 1 7 '2 4 1 1 1 2 1 .1 1 1 1 1 1 1 2 1 1 1 3 1 3 2 2 10 1 1 4 1 16 4 3 4 1 . Page 1 of3 2 4 1 r . KREB'S N HARE DEN$- , MEAN 1 0.4 0.2 0.1 0.1 0 0.3 0 0 0.9 1.2 0 0 0.1 0 0.1 0 0 0.1 0.2 0.1 0.1 0.2 0 0 0 0 0.6 0.1 0.5 0 0.6 0.9 2.6 1.2 0 0.1 0.1 0.1 0.1 0.7 0.3 0 0.5 2.1 2.5 0.9 ·0.55929 -0.80394 -1.0486 . -1.0486 0 -0.66083 0 0 -0.27306 -0.17152 0 0 -1.0486 0 -1.0486 0 0 ·1.0486 ·0.80394 -1.0486 -1.0486 -0.80394 0 0 0 0 -0.41617 -1.0486 -0.48052 0 -0.41617 ·0.27306 0.101391 -0.17152 0 -1.0486 -1.0486 -1.0486 -1.0486 -0.36176 -0.66083 0 -0.48052 0.026007 0.087548 ·0.27306 0.275876311 0.1570576~ 0.08941368E 0.08941368E ( 0.21836012', ( ( 0.53326447' 0.67372666! ( ( 0.08941368( ( 0.089413681 ( ( 0.089413681 0.1570576! 0.089413681 0.089413681 0.1570576! 1 I I I 0.3835557: 0.08941368' 0.33073167 0.3835557 0.53326447 1.26296421 0.67372666 0.08941368 0.08941368 0.08941368 0.08941368 0.43474840 0.21836012 0.33073167 1.06171345 1.22334116 0.53326M7 w �UTMX KREBSPL LSBNUM DW-49-B DW-50-A DW-50-B DW-53-A DW-53-B DW-56-A DW-56-B DW-145-A DW-145-8 DW-157-A DW-157-S EP-30-A EP-30-B EP-33-A EP-33-8 EP-118-A EP-118-B EP-38-A EP-38-B EP-180-A EP-180-B EP-9-A EP-9-B EP-31-A EP-31-B EP-26-A EP-26-B EP-6-A EP-6-B EP-27-A EP-27-B EP-1-A EP-1-B EP-4-A EP-4-B EP-28-A EP-28-B EP-24-A EP-24-B EP-32-A EP-32-B EP-l1-A EP-ll-B EP-8-A EP-8-B EP-36-A 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 434500 426188 426270 435584 435605 454053 453998 451826 451826 461484 461435 444784 444662 451484 451463 451984 452014 450684 450679 450158 450001 442308 442240 445384 445432 452684 452726 437509 437449 438284 438338 427480 427373 422217 422068 425884 426471 431484 431517 432284 432314 425062 424950 433478 433573 429484 UTMY 4398210 4396472 4396380 4388779 4388725 4379173 4379265 4410001 4410131 4390954 4390970 4470479 4470409 4460879 4460801 4448579 4448550 4448279 4448511 4414204 44U203 4465163 4464949 4470179 4470199 441.a879 4478919 4471239 4471169 4476679 4476732 4472833 4472806 4470980 4470689 4474379 4473939 4482579 4482788 4461379 4461296 4464125 4464089 4451783 4451435 4452379 PRIMOVE SN1 S S S S S S S L L L L S X X X F F F F L F S F L L L L S S S S L L S S S L S S L S L L L F L SN3 SN2 3 1 SN5 SN4 SN6 SN7 SN8 SN9 SN10 1 MEAN 3 1 1 1 2 1 8 1 1 1 1 1 1 2 3 7 1 0 1 0 1 1 3 5 1 2 1 2 4 1 1 4 1 1 3 8 1 1 2 2 4 3 1 (..,:) 1 2 1 1 1 1 1 1 2 1 2 2 4 1 1 1 4 0 1 1 2 2 0.2 0 004 1 1 1 2 1 1 1 2 1 1 1 2 1 2 3 1 1 0.1 0.3 004 3 1 .1 1 14 0 0.1 0 0.9 1 0.3 0.1. 004 1 2 1 ~ 004 1 4 4 1 1 KREB'S N HARE DENSI!, 0.8 0.5 0.1 1 0.1 0.5 0.8 0.3 0.6 0.3 1 2.1 1.5 1.6 1 0.1 2 0 1 2 1 3 2 1 1 3 1 1 1 1 1 2 204 2 1 16 1 3 Page 20f3 0.1 0.1 0.2 0.1 1.2 0.2 1.6 0.1 0 0.3 0 0 0 0 ~- -0048052 OA8458460S 0.33073167t -1.0486 -0.23587 -1.0486 0.08941368E 0.58093962~ 0.08941368E -0048052 0.33073167t OA8458460~ -0.31463 -0.31463 -0.66083 -0041617 -0.66083 -0.23587 0.026007 -0.09275 -0.06998 -0.23587 -1.0486 -0.55929 -0.80394 0 -0.55929 0 -1.0486 0 -0.27306 -0.23587 -0.66083 -1.0486 -0.55929 -1.0486 -0.66083 -0.55929 -1.0486 -1.0486 -0.80394 -1.0486 -0.17152 0.073139 -0.80394 -0.06998 -1.0486 0 -0.66083 0 0.218360121 0.38355571 0.218360121 0.58093962~ 1.06171345< 0.807690771 0.85118668~ 0.58093962~ 0.08941368E 0.275876311 0.1570576S ( 0.275876311 ( 0.08941368E ( 0.53326447< 0.58093962: 0.218360121 0.08941368E 0.275876311 0.08941368E 0.21836012', 0.275876311 0.08941368E 0.08941368E 0.1570576S 0.08941368E 0.67372666: 1.183420101 0.1570576S 0.85118668: 0.08941368E ( 0.21836012~ ( 0 ( 0 0 ( ( �KREBSPL LSBNUM EP-36-B EP-35-A EP-35-B 8S-103-A 8S-103-B S8-107-A S8-107-B S8-111-A 8S-111-B SS115-A 88-115-B WA-n-A SS-37-A S8-37-B VA-11-A VA-11-B VA-12-A VA-12-B VA-153-A VA-153-BVA-154-A VA-154-B UTMX 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 429442 434484 434201 406384 408211 386984 387083 413184 413242 406384 406348 403884 401384 401418 366284 365940 387284 387580 373284 373100 367984 368100 UTMY PRIMOVE SN1 4452678 L 4454279 L 4454346 L 44603798 4482059 S 44693798 44694728 4466179 L 4466143 L 4460379 L 4460391 L 4529079 F 4450579 S 44504248 4409979 A 4409800 S 4380379 S 4380600 S 4~.9874 S 4399720 S 43993798 4398800 S SN2 SN3 SN5 SN4 SN6 SN7 SN8 SN9 8N10 1 1 3 2 1 1 1. 2 2 1 1 1 1 1 2 1 1 2 1 1 1 1 1 2 1 1 1 3 3 2 KREB'S N HARE DENSIl MEAN 3 5 1 4 2 2 1 1 0 0 0 0 0.1 0 0 0 0.1 0 0 0.3 0 0.3 0 1.1 0.7 0.2 1.6 0.9 0.6 0.3 0 0 0 0 -1.0486 0 0 0 -1.0486 0 0 -0.66083 0 -0.66083 0 -0.20223 -0.36176 -0.80394 -0.06998 -0.27306 -0.41617 -0.66083 ( ( ( ( 0.08941368( ( ( ( 0.089413681 1 1 0.21836012 I 0.21836012 I 0.62772865 0.43474840: 0.1570576! 0.85118668! 0.53326447· 0.3835557; 0.21836012 '.'), \0 VI Page 3 of 3 �UTMX KREBSPL LSBNUM CG-12-A CG-12-B CG-7B-A CG-7B-B SS-21-A SS-21-B S5-24-A S5-24-B S5-29-A SS-29-B S5-31-A S5-31-B S5-33-A S5-33-B SS-37-A S5-37-B S5-91-A S5-91-B VA-41-A VA-41-B W-5-A W-5-B W-13-A W-13-B W-53-A W-53-B W-81-A W-81-B W-82-A W-82-B S5-35-A SS-35-B S5-65-A S5-65-B WA-45-A WA-45-B CR-73-A CR-73-B CR-46-A CR-46-B CR-49-A CR-49-B WA-85-A WA-85-B S5-66-A S5-66-B 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 323599 323546 322068 322094 367299 367408 351299 351255 355299 355435 363099 363097 361899 361730 342699 342599 366082 365986 345207 345219 336521 336407 331599 331657 345899 345745 347099 347250 331599 331514 345099 345191 355899 355886 346477 346543 330499 330582 310699 310548 313599 313790 333799 333714 352650 352523 UTMV PRIMOVE SN1 4516720 4516636 4518850 4518851 4470720 4470747 4451320 4451400 4439720 4439703 4437620 4437718 4432620 4432676 4429320 4429389 4467220 44q7128 4426619 4425591 4533917 453.3875 4513120 45,.3062 F L F F A A L L L L L L L L L L A A L L L W F F 4509020 4508942 4514220 4514433 4514120 4513955 4430920 4430965 4475620 4475833 4521319 4521281 4530820 4530639 4518920 4518948 4518520 4518470 4482873 4492646 4468880 4468800 L A A F A L F A S S S S A L S S S S A A A A SN8 SN3 SN2 A ffQM-cL·x SN9 C. SN10 MEAN 1 1 1 2 5 1 :.! 1 1 1 1 1 1 2 3 2 41 1 1 13 2 1 4. .' ~ 2 1 1 4 1 6 3 9 2 2 1 2 1 4 2 3 1 9 1 11 2 2 9 2 2 1 1 1 9 2 1 1 2 2 4 1 3 2 3 2 - Page 1 of 2 -------- !--- KREB'S N HARE DENSI~ 0.1 0 0.2 0.1 0 0 1.7 0 0.4 0 0 0 0.1 0.1 0.5 0 0.9 2.7 0 0 5.4 2.2 0.1 1.1 0 0 1.3 1.4 0 0 0 0 0 0 0.4 0.1 0 0 0.1 0 0 0 0.5 1.2 0.2 0 -1.0486 0 -0.80394 -1.0486 0 0 -0.04858 0 -0.55929 0 0 0 -1.0486 -1.0486 -0.48052 0 -0.27306 0.114712 0 0 0.359367 0.042427 -1.0486 -0.20223 0 0 -0.14326 -0.11711 0 0 0 0 0 0 -0.55929 -1.0486 0 0 -1.0486 0 0 0 -0.48052 -0.17152 -0.80394 0 0.089413686 C 0.15705769 0.089413686 C C 0.894176106 C 0.275876311 C C C 0.089413686 0.089413686 0.33073167e C 0.533264474 1.302302811 C C 2.287532027 1.102623281 0.089413686 0.627728653 0 0 0.719011504 0.763647967 C C C C C C 0.275876311 0.089413686 C C 0.089413686 G C 0 0.330731678 0.673726665 0.15705769 n �KREBSPLLSBNUM SS-69-A SS-69-B WA-1-A WA-1-B WA-17-A WA-17-B CR-SO-A CR-50-B CR-83-A S5-25-A SS-25-B WA-61-A WA-61-B WA-86-A WA-9-A WA-9-B UTMX 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 355658 355565 334299 334285 361099 361201 303378 303492 304699 359099 359206 353099 353099 345895 365099 365120 UTMY 4461691 4461595 4540420 4540385 4497920 4497971 4514532 4514404 4508720 4450020 4450122 4493520 4493520 4491620 4528020 452]890 ..., PRIMOVESN1 S S S L F F S S A L L S S A L L SN3 SN2 SN4 SN5 SN6 SN7 SN8 SN9 SN10 MEAN 1 2 1 1 1 1 1 2 1 2 3 L:. ', :~ 1 KREB'S N HARE DENSIT 0 0.1 0.2 0.3 0 0.1 0 0.1 0 0 0 0 0.2 0.1 0.3 0.3 0 -1.0486 -0.80394 -0.66083 0 -1.0486 0 -1.0486 0 0 0 0 -0.80394 -1.0486 -0.66083 -0.66083 ( 0.08941368E 0.1570576S 0.218360121 ( 0.08941368E C 0.08941368E C C C ( 0.1570576S 0.08941368E 0.218360121 0.218360121 ,~ "'.lI 10 Page 2 of 2 -.....) �KRE8SPL LS8NUM UTMX G8-9-A GS·9-8 GS·13-A G8-13-8 GS·57·A GS·57·B G8-60-A GS·60-8 GS·61·A G8-61·B GS·65-A GS·65-8 GS·67·A GS·67·8 GS·93-A G8-93-B GS·98-A GS·98·B M·25-A M·25-B M·71·A M·71·B M·73-A M·73-B M·1·A M·1·B M·74-A M·74-B M·77·A M·77·B M-79-A M·79-B VA·19-A VA·19-B ME·29-A ME·29-B ME·33-A ME·33-8 ME-85-A ME·85-B ME·26-A ME·26-B ME-81·A ME-81·B ME·21·A ME·21·B 3 .3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 323290 323297 276237 276431 274782 274773 296592 296642 297492 297601 290492 290300 276892 276966 311992 311826 290592 290500 321592 321584 298192 298256 287292 287246 308292 308099 313192 313305 324792 324962 323992 324030 329547 329434 319157 319160 290592 290682 313870 313932 314992 315015 313492 313407 314792 314733 UTMY 4412469 4412272 4395669 4395598 4408689 4408444 4403498 4403428 4402598 4402735 4399198 4399250 4396587 4396513 440lj098 4405172 4397498 4397850 4442998 444;2917 4457798 4457786 4455598 4455648 4448798 4448695 4454298 4454069 4452690 4452684 4450598 4450468 4416879 4416904 4435523 4435606 4432198 4432190 4440307 4440419 4441498 4441563 4448198 4448093 4446098 4446242 SN3 SN2 PRIMOVE SN1 SN4 SN9 F F A fpo.M.d 0 0 MEAN tx D 1 S S S S S S S S A 4 1 4 2 , 5 3 8 1 1 3 1 3 0 L .:. 7 1 S S S S S S A A A A A A A A L L A AA . F F F A F A F S S F A S S SN10 1 1 1 1 1 2 3 3 10 13 1 .3 1 3 1 5 5 1 1 4 5 2 1 1 27 23 1 1 1 2 2 2 1 3 1 2 2 2 5 1 1 1 1 1 1 1 1 1 6 1 1 2 1 1 1 _. 1 2 Page 1 of 2 \0 00 KREB'S N HARE DENSm 0.1 0 0 0 0.2 0.5 0.6 0 0.9 1.3 0 0.4 0 0 0.9 0 0.1 0.5 0.3 1.3 1.3 0 0.7 0 0 0 0 0 4 4 0 0 0.6 0.4 0 0.1 0.1 0.1 0.5 1.6 0.3 0.2 0.3 0.1 0.1 0.3 -1.0486 0 0 0 ·0.80394 ·0.48052 ·0.41617 0 ·0.27306 -0.14326 0 -0.55929 0 0 -0.27306 0 -1.0486 -0.48052 -0.66083 ·0.14326 -0.14326 0 -0.36176 0 0 0 0 0 0.253441 0.253441 0 0 -0.41617 -0.55929 0 ·1.0486 -1.0486 -1.0486 -0.48052 -0.06998 ·0.66083 ·0.80394 ·0.66083 -1.0486 -1.0486 -0.66083 0.08941368€ C C C 0.15705769 0.330731678 0.383555?e C 0.533264474 0.719011504 C 0.275876311 C C 0.533264474 C 0.08941368E 0.33073167E 0.218360121 0.719011504 0.719011504 C 0.434748402 C C C C C 1.79242672: 1.79242672: c C 0.383555n 0.275876311 C 0.08941368E 0.08941368E 0.08941368E 0.33073167t 0.85118668~ 0.218360121 0.1570576~ 0.218360121 0.08941368E 0.08941368E 0.218360121 �KREBSPLLSBNUM GS·53-A GS·53-B GS-S9-A GS·S9-B GS-17·A GS-17-B GS-41-A GS-41·B GS-42·A GS-42·B GS-43-A GS-43-B ME-45-A ME-45-B ME-46-A ME-46-B ME.a9-A ME·89-B GS-49-A ME-3Q.A ME-3Q.B UTMX 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 315807 315741 303319 303421 283207 283246 319767 319706 320543 320495 320477 320499 295792 295928 291692 291663 295600 295532 268390 325792 325748 UTMY PRIMOVESN1 4412384 F 4412364 F 4391172 S 439142S S 4400690 S 4400732 A 4422156 F 4422201 F 4421864 F 4421868 F 4421531 F 4421391 F 444J)Jl98 S 4420845 S 4420398 S 4420254 S 441~32 A 441~957 A 4415600 A 4433698 F 4433677 L SN2 SN3 SN4 SN5 SNS SN7 SN8 SN9 2 2 8 1 2 1 1 0 1 8 2 8 1 SN10 MEAN 1 3 7 5 3 3 1 10 14 4 3 0 1 2 1 3 3 7 1 3 2 5 1 3 3 4 4 1 1 1 3 1 1 5 3 5 2 8 3 6 2 1 4 1 1 .. 1 1 6 ---- KREB'S N HARE DENS In 1.S 0 0.1 0.3 0.3 0 3.5 3.2 1.6 3.3 2.3 2.1 0 0 0 0 0 0 0.1 0.1 0.6 -0.06998 0 -1.0486 -0.66083 -0.66083 0 0.20631 0.17468 -0.06998 0.185541 0.058117 0.026007 0 0 0 0 0 0 -1.0486 -1.0486 -0.41617 0.851186685 0 0.089413686 0.218360121 0.218360121 0 1.60808792 1.495133686 0.851186685 1.532996908 1.143186215 1.061713454 0 0 0 0 0 0 0.089413686 0.089413686 0.38355578 \0 \0 Page 2 of 2 �KREBSPL LSBNUM PA-12-A PA-12-B GU-82-A GU-82-B GU-86-A GU-86-B SA-14-A SA-14-B SA-64-A SA-64-B CA-25-A CA-25-B CA-27-A CA-27-B CA-30-A CA-30-B CA-32-A CA-32-B CA-38-A CA-38-B CA-46-A CA-46-B CA-119-A CA-119-B CA-132-A CA-132-B CA-141-A CA-141-B GU-61-A GU-61-B GU-80-A GU-8O-B GU-82-A GU-82-B GU-86-A GU-86-B GU-89-A GU-89-B GU-91-A GU-91-B GU-92-A GU-92-B PA-19-A PA-19-B PA-56-A PA-56-B 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 UTMX UTMY PRIMOVE SN1 308071 42730550 308068 ·42730080 372071 4301755 L 371956 4301612 L 370471 4289955 L 370421 4290005 L 370371 4230655 P 370487 4230576 P 374471 4235455 L 374271 4235529 L 291683 4359468 A 291929 435~523 A 292637 4358699 A 292438 4358853 A 260735 4356491 A 260659 4356410 A 282106 4352554 0 282058 4352570 0 293676 4335150 A 293766 4335189 A 253361 4321074 A 253422 4320992 A 283084 4357784 S 283109 4357727 S 307856 4337843 D 307950 4337810 D 313455 4326347 D 313440 4326279 0 412171 4308255 P 412146 4308040 P 398571 4312655 P 398581 4312777 P 372071 4301755 L 371956 4301612 L 370471 4289955 L 370421 4290005 L 394371 4280155 L 394552 4280140 L 372671 4274155 L 372600 4274150 L 364671 4269255 D 364685 4269700 D 292071 4292555 F 292034 4292573 F 290071 4291055 A 290001 4290862 A SN3 SN2 SN8 3 0 2 4 18 2 1 0 0 2 6 0 1 0 0 0 0 0 2 1 A 2 0 0 0 SN10 MEAN 3 p P Q.M. d ,.,)( [ 0 SN9 0 0 1 5 12 4 0 0 0 0 1 3 2 0 0 0 0 0 0 0 3 (. .:. 1 6 1 1 5 1 1 2 1 1 6 3 1 2 1 2 3 3 1 6 7 2 2 1 1 1 4 3 18 2 1 6 1 2 1 2 1 1 2 1 2 2 4 1 5 12 4 33 1 3 1 2 1 1 -- Page 1 of 3 4 1 1 3 1 2 3 1 KREB'S N HARE DENSI"§ 0.6 -0.41617 0.6 -0.41617 1.1 -0.20223 3.2 0.17468 1.9 -0.00932 0.7 -0.36176 0.2 -0.80394 0.4 -0.55929 0 0 0 0 0 0 0 0 0.3 -0.66083 0 0 0 0 0 0 0 0 0.1 -1.0486 0 0 0 0 0 0 0 0 0.5 -0.48052 1.2 -0.17152 1.7 -0.04858 0.7 -0.36176 0 0 0.6 -0.41617 0.8 -0.31463 0.2 -0.80394 0.9 -0.27306 0.8 -0.31463 1.1 -0.20223 3.2 0.17468 1.9 -0.00932 0.7 -0.36176 4 0.253441 0.3 -0.66083 0 0 0 0 0 0 0 0 0.6 -0.41617 0.2 -0.80394 0 0 0 0 0.38355578 0.38355578 0.627728653 1.495133686 0.978772092 0.434748402 0.15705769 0.275876311 0 0 0 0 0.218360121 0 0 0 0 0.089413686 0 0 0 0 0.330731678 0.673726665 0.894176106 0.434748402 0 0.38355578 0.484584609 0.15705769 0.533264474 0.484584609 0.627728653 1.495133686 0.978772092 0.434748402 1.792426725 0.218360121 C C 0 C 0.38355578 0.15705769 0 0 �KREBSPLLSBNUM SA-65-A SA-65-B SA-69-A SA-69-B SA-67-A SA-67-B SA-100-A SA-l00-B SA-l02-A SA-102-B SA-104-A SA-l04-B SA-107-A SA-l07-B SA-108-A SA-108-B SA-109-A SA-109-B SA-110-A SA-110-B VA-1-A VA-l-B VA-2-A VA·2-B CA-33-A CA-33-B LE·3-A LE-3-B CA-133-A CA-133-B LE-24-A LE·24-B LE·116-A LE·116-B VA·1·A·2 VA-111-A VA·113-A CA· 17·A· CA·17·B CA·123-A CA·123-B GU·52·A GU·52·B GU·157·A GU·157-B GU·153-A UTMX 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 . 4 4 4 4 4 4 4 4 4 4 4 4 4 4 373071 373233 390271 390070 368013 367745 370371 370242 374271 374148 362171 362268 368771 368724 365571 365543 351871 351662 351530 351524 337267 337329 351577 351544 302269 302330 436930 436945 245402 245436 350683 350801 360122 360057 337271 356671 357371 296459 296390 315656 315720 327571 327548 348771 348674 365371 UTMY PRIMOVESN1 4235355 A 4235373 A 42300550 4229851 0 4231272 0 4231280 A 4246555 P 4246541 L 424Q655 L 4245672 L 4241955 L 4241864 L 4236155 L 42~6361 L 42347550 4234940 S 4231455 A 4231491 0 4229214 0 42292970 4380741 L 4380577 L 4374462 L 4374495 L 4351331 4351244 A 3397400 S 3397800 S 4335106 A 4335245 F 4370138 L 4369892 A 4371050 S 4370936 S 4380755 L 4379855 L 4378955 S 4373781 S 4373800 S 4351061 L 4350930 F 4296255 A 4296315 A 4303555 S 4303630 S 4309155 L SN3 SN2 SN4 SN5 .~ SN7 SN9 SN8 SN10 MEAN 1 1 2 1 0 7 2 4 1 2 1 1 1 1 1 2 4 (_ 1 .. ' 4 1 13 2 3 2 1 2 1 1 1 2 5 2 11 1 1 1 5 1 1 0 1 1 5 1 4 0 1 2 6 2 1 1 1 3 1 3 4 1 3 1 1 1 Page 2 of 3 .'"" SN6 3 KREB'S N HARE DENSlr 0 0.2 1.6 0.3 0 0 0 0 0.1 0.1 0.1 0.2 0.3 0.4 0.4 0 0.3 0,6 2.4 1.4 0.1 0.8 0 0.1 0 0 0 0 1.1 0 0 0 '0 0.5 1.6 0.1 0 0.4 0.1 0.7 0.2 0 0 0 0 0 ( 0 -0.80394 0.1570576~ 0.85118668~ -0.06998 -0.66083 0.218360121 ( 0 ( 0 ( 0 ( 0 -1.0486 0.08941368E -1.0486 0.08941368E -1.0486 0.08941368E -0.80394 0.1570576~ -0.66083 0.21836012', -0.55929 0.27587631', -0,55929 0.27587631' ( 0 0.21836012' -0.66083 -0.41617 0.38355571 0.073139 . 1.18342010' 0.76364796; -0.11711 -1.0486 0.08941368( -0.31463 0.48458460~ ( 0 0.08941368( ·1.0486 ( 0 ( 0 ( 0 ( 0 0.62772865: -0.20223 ( 0 ( 0 ( 0 I 0 -0.48052 0.330731671 0.85118668! -0.06998 -1.0486 0.089413681 I 0 ·0.55929 0.27587631 0.089413681 ·1.0486 0.43474840: ·0.36176 0.1570576! ·0.80394 1 0 1 0 I 0 1 0 •.... 0 - �KREBSPLLSBNUM UTMX UTMY SN2 PRIMOVESN1 SN4 SN3 SN5 SN6 SN7 SN8 SN9 SN10 MEAN o KREB'S N HARE DENSIT N GU·153-B LE·9-A LE·9·B LE·73-A LE·73-B LE·74·A LE·74-B LE·78-A LE·78-B LE·128-A LE·128-B LE·129-A LE·129-B LE·144-A LE·144-B PA-18-A PA·18-B PA·22·A PA-22-B PA·20-A PA·20-B PA-21·A PA·21-B PA-186-A PA-186-B PA·187·A PA-187·B SA·106-A SA·106-B LE·29-A LE-29-B LE·120-A LE·120-B C0-148-B LE-5-A LE-5-B CR-83-B 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 365558 412871 412756 377871 377906 370877 370998 407171 407263 373571 373292 368871 369228 371171 371104 320871 320822 288076 288225 293771 293804 294371 294368 290271 290372 288471 288585 360209 360219 346471 346471 350771 350771 761178 380700 380520 304199 4309087 4317355 4317207 4334155 4334211 4333181 4333200 4318255 4318427 4347255 4346979 434.7155 4347084 4324855 4324853 4304655 4304619 4266178 4266146 4275055 4275216 4270255 4270108 4271155 4271208 4270055 4270239 4237602 4237494 4357055 4357055 4354155 4354155 4148888 4361960 4362240 4509001 L P P L L S S S S F F F F L L S S S A F F S S S S S S S S L L L L S S S A 1 1 1 1 1 1 3 2 1 1 L.:·~ 1 1 1 6 1 1 1 2 2 2 1 1 1 1 1 1 1 1 3 1 2 1 1 2 1 1 1 8 5 1 1 -------- 2 4 1 1 1 1 1 1 ------- --------- Page 3 of 3 !t. 1 1 2 ------ 3 1 1 7 0.1 ·1.0486 0 0 0 0 0.1 ·1.0486 0.2 ·0.80394 0.3 -0.66083 0.2 -0.80394 0.1 -1.0486 0.3 -0.66083 0.2 -0.80394 0 0 0 0 0.1 -1.0486 0.5 -0.48052 0.7 -0.36176 0.1 -1.0486 0 0 0.2 -0.80394 0.7 -0.36176 0.3 -0.66083 0.4 -0.55929 0.5 -0.48052 0.2 -0.80394 0.4 -0.55929 0.5 -0.48052 0.2 -0.80394 0.3 -0.66083 0.2 -0.80394 0 0 0 0 0 0 0 0 . 0 0 2.3 0.058117 0.2 -0.80394 0.2 -0.80394 0 0 0.089413681 ( ( 0.089413681 0.1570576! 0.21836012 0.1570576~ 0.08941368€ 0.21836012· 0.1570576! ( ( 0.08941368( 0.330731671 0.43474840; 0.089413681 ( 0.1570576! 0.43474840; 0.21836012· 0.27587631· 0.330731671 0.15705W 0.27587631· 0.330731671 0.1570576! 0.21836012· 0.1570576! ( ( ( ( ( 1.14318621! 0.1570576! 0.1570576! ( �KREBSPL LSBNUM UTMX AN·152·A AN·152·B AN·156-A AN·156-B AN·157·A AN·157·B AN·158-A AN·158-B AN·160-A AN·160-B AN·168-B AN·168-A C0-14-A C0-14B C0-187·A C0-187·B DE·8-A DE·8-B DE·13-A DE·13-B DE-49-A DE-49-B DE·54-A DE·54·B DE·89-B DE·89-A DE-146-A DE·146-B DU-30-A DU·30-B DU·192·A DU·192·B MO-3-A MO-3-B M0-28-A M0-28-B M0-29-A MO-29-B M0-64-A MO-64-B M0-66-A M0-66-B DU·186-A DU·186-B DU·190-A DU·190-B 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 370326 370357 367132 367087 369726 369772 366889 366884 371126 371201 375941 375935 747483 747553 748557 748500 364326 364304 370832 370796 378026 377952 368766 368729 364807 364826 335236 334987 319412 319396 251960 251911 277574 277510 249194 249047 258872 258968 277527 277463 277150 277120 254345 254522 265782 265683 UTMY PRIMOVE SN1 4145971 S 4143036 S 4142726 D 414~692 D 41~'O671 S 4140586 S 4139791 S 4139726 S 4137071 S 4137124 S 4105024 S 4104968 S 4146928 A 4146893 A 4145794 A 4145810 A 4189271 D 4189201 D 4151461 S 4151379 S 4205671 S 4205646 S 4158215 P 4158145 P 4203561 B 4203671 B 4152177 S 4152211 S 4135828 0 4135981 0 4134560 P 4134607 P. 4234397 A 4234334 A 4221691 0 4221649 0 4220445 A 4220443 A 4231212 D 4231125 D 4228274 S 4228172S 4147354 0 4147273 P 4138467 D 4138570 D SN2 SN3 1 3 1 1 4 4 2 1 0 1 0 0 0 0 1 2 3 2 2 6 5 0 0 3 0 1 0 AfP~d,·x 1 5 2 4 3 F 1 6 23 2 1 1 1 0 0 SN9 1 3 11 10 2 2 0 0 4 0 0 0 0 0 1 2 3 3 1 0 0 1 1 1 0 1 0 o SN8 0 1 2 0 2 0 0 4 0 0 0 1 0 1 2 5 I 1 0 0 0 11 5 0 0 0 19 0 1 1 0 0 2 2 0 0 2 1 SN10 4 7 5 2 7 5 14 2 12 0 0 1 1 1 2 13 7 0 0 0 0 3 0 6 1 0 0 0 2 0 1 0 .t1..l', Page 1 of 6 KREB'S N HARE DENSIT1 MEAN 0.1 0.1 2.5 5.1 1.5 0.7 3.5 2.5 0.4 0.7 2.5 3 0 0 0 0 3.5 1.3 0.2 1.3 0.4 0.2 0.4 0.7 0.1 0.1 0.2 0 0 0 0 0 0 0 0 0 0 0 0 0 0.4 0.8 0 0 0 0 ·1.0486 ·1.0486 0.087548 0.339192 ·0.09275 ·0.36176 0.20631 0.087548 ·0.55929 ·0.36176 0.087548 0.1519 0 0 0 0 0.20631 -0.14326 -0.80394 ·0.14326 -0.55929 -0.80394 -0.55929 -0.36176 -1.0486 -1.0486 -0.80394 0 0 0 0 0 0 0 0 0 0 0 0 0 ·0.55929 ·0.31463 0 0 0 0 0.08941368E 0.08941368E 1.22334116L 2.18369704E 0.80769077~ 0.43474840; 1.6080879; 1.22334116' 0.27587631' 0.43474840; 1.22334116' 1.41873187' ( ( ( ( 1.6080879: 0.71901150' 0.1570576! 0.71901150, 0.27587631 0.1570576' 0.27587631 0.43474840: 0.089413681 0.089413681 0.1570576' I , 0.27587631 0.48458460 0 �KREB8PLL8BNUM UTMX UTMY 8N2 PRIMOVE8N1 8N4 8N3 8N5 8N6 8N8 8N7 8N9 8N10 KREB'S N HARE DENS IT '0 MEAN .j::. DU-193-A DU-193-B M0-2-B M0-2-A M0-60-A M0-60-B M0-61-A MQ.61-B MQ.65-A MQ.65-B MQ.68-A MQ.68-B MQ.79-A MQ.79-B MQ.179-A MQ.179-B DE-144-A DE-144-B AN-167-A AN-167-B DU-33-A DU-33-8 M0-4-B M0-4-A M0-26-A MQ.26-8 MQ.72-A M0-72-B MQ.77-A MQ.77-8 MQ.80-8 MO-8O-A 8A-1-A SA-1-B 8A-39-A SA-39-8 SA-41-A 8A-41-8 8A-69-A 8A-69-8 8A-76-A 8A-76-8 8A-78-A 8A-78-8 81-96-A 81-96-B 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 318011 318116 268285 268326 284257 284212 280720 280683 286117 280095 281510 281561 242213 242282 241604 241618 373926 373904 379909 379825 268426 268396 271352 271426 265626 265631 276226 276156 271726 271755 251688 251626 '337326 337333 332426 332433 339126 339218 331926 332041 341126 341115 327073 327039 267926 267859 4134176 P 4134283 P 42a~607 A 4238671 A 4241267 A 4241345 A 42343788 42344388 42290708 42291198 42255408 42255788 42116638 42117468 4222825 A 4222749 A 4152871 8 4152938 S 41050458 4105078 8 4136471 P 4136238 P 4232051 A 4232071 A 4235571 0 4235639 0 4222071 8 42220858 4215171 8 4215157 8 42112068 4211171 8 4244871 F 4244929 A 4230371 L 4230234 L 4235071 0 4234944 0 4224571 L 4224576l 4217171 8 4217057 8 4213018 F 4213095 F 4196571 8 419~596 8 3 1 1 0 4 3 4 1 3 1 2 0 6 2 1 4 6 7 2 8 2. 2 0 1 1 2 0 1 1 1 3 0 7 2 14 8 0 1 1 3 1 3 1 0 1 4 16 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 6 5 1 0 0 0 0 0 1 2 2 2 3 4 2 1 1 1 9 0 1 0 2 1 - -- -- 1 2 3 3 13 1 10 7 Page 2 of 6 <. 1 1 0 9 0 8 11 2 0 1 1 6 3 1 0 5 6 0 2 6 3 5 L.:' 4 13 23 1 3 2 7 12 6 2 4 2 8 0 1 5 1 7 0 3 1 1 0 0 0 2 0 0 0 0 2 2 2 6 1 1 1 1 5 5 2 2 0 3 7 7 0 5 9 0 0 1 1 2 0 2 3 0 0 0 0 0 0 0 0 0 0 0 0 3 2 1 2 1 2 3 2 2 3 4 6 1 0 0 0 0 0 0 1.2 2.1 0.4 0.4 0.9 1.1 2.6 2.2 0 0 2.9 5.3 0.7 0.9 0 0 0 0 0 0 3.5 2.2 2 3.2 3.3 0.9 1.2 0 0 0 0 0.3 0.2 1.8 1.1 1.4 1.4 1.9 1.9 3.1 0 0 0 0 0 0 -0.17152 0.026007 -0.55929 -0.55929 -0.27306 -0.20223 0.101391 0.042427 0 0 0.139934 0.35277 -0.36176 -0.27306 0 0 0 0 0 0 0.20631 0.042427 0.008786 0.17468 0.185541 -0.27306 -0.17152 0 0 0 0 -0.66083 -0.80394 -0.0284 -0.20223 -0.11711 -0.11711 -0.00932 -0.00932 0.163474 ( ( ( ( ( ( 0.67372666~ 1.06171345L 0.275876311 0.275876311 0.53326447l 0.62772865~ 1.26296421 ; 1.10262328 ' ( ( 1.38017557( 2.25304338~ 0.43474840; 0.53326447< ( ( ( ( ( ( 1.6080879: 1.10262328 . 1.02043701 ~ 1.49513368( 1.53299690! 0.53326447' 0.67372666: ( ( ( ( 0.21836012 0.1570576! 0.936694251 0.62772865: 0.76364796'. 0.76364796'. 0.97877209: 0.97877209: 1.45704811." �KREBSPLLSBNUM SI·102·A SI·102·B SI·107·A SI·107·9 SI·120-A SI·120-9 DE·111·A DE·111·9 DE·112·A DE·112·B DE-138-A DE·138-B SI-SO-A SI-50-9 SI-86-A SI-86-9 SI·110-A SI·110-9 SI-113-A SI-113-9 SI·122·A SI·122-B SI·126-A SI·126-9 DC·140-A DC·140-B DC-141-A DC-141-B SI-12-A SI-12-9 SI·185-A SI·185-9 DU·147·A DU·147·9 DU-145-A DU-145-B DU-151·A DU-151-B DU·154·A DU-154·B DU·189-A DU·189-B SI·123-A SI·123-9 DE·132·A DE-132-B UTMX 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 260726 260809 274226 274147 317826 317901 344226 344198 350326 350264 351826 351853 298726 298652 308226 308294 293526 293451 294626 294582 316126 316056 319126 319118 761926 761750 761426 761350 247730 247800 253126 253110 261419 261451 245026 244980 248713 248830 281790 281712 257530 257150 254400 254369 334612 .334624 UTMY PRIMOVESN1 41Qa271 0 4193273 0 4188771 S 4188681 S 4171771 S 4171806 S 4185671 A 4185621 A 4183271 S 4183331 S 4157371 S 4157311 S 4201771 S 4201741 S 4206871 S 4206850 S 4186371 A 4186413 A 4182671 S 4182594 S 4171271 S 4171236 S 4168071 S 4168086 S 4156371 F 4156280 F 4156271 F 4156350 F 4156170 A 4156200 A 4155871 P ·41557300 4150956 F 4150964 S 4152171 0 41522600 4146127 D 4146020 0 4144654 A 4144508 A 4140571 0 4140565 0 41,70897 A 4110937 0 4165099 S 4165200 S SN4 SN3 SN2 SN5 SN6 SN7 SN8 SN9 SN10 2 7 7 1 2 2 4 2 1 1 8 7 1 8 4 5 2 4 8 1 3 6 4 6 3 5 1 8 1 5 8 1 16 4 6 18 1 1 2 2 5 6 3 1 5 5 5 10 3 1 2 3 3 5 1 3 2 14 3 3 9 5 2 8 1 4 5 7 2 2 1 2 10 5 9 3 1 3 12 7 26 7 11 3 4 4 7 3 2 5 3 1 2 1 8 2 7 11 12 2 1 4 1 1 3 4 4 9 14 6 7 6 19 1 7 6 4 19 5 3 1 1 3 2 1 3 1 2 3 3 1 11 1 1 1 2 1 1 3 1 1 1 1 3 4 I 2 8 1 1 Page 3 of6 KREB'S N HARE DENSIT'I MEAN 4 2 0.4 0.7 0.7 3.3 0.9 1.6 1.4 2.1 4.3 3.2 1.6 2.4 3.6 7.3 2.8 4.5 0 0 1.4 2.4 2 3.8 1.9 3.3 0 0 0 0 3.8 1.7 0.3 0.4 1.8 0.5 0 0 0 0 0.1 0.3 0.9 0.4 0 0.2 0.1 1.4 ·0.55929 ·0.36176 ·0.36176 0.185541 ·0.27306 -0.06998 -0.11711 0.026007 0.278968 0.17468 -0.06998 0.073139 0.216253 0.465777 0.127548 0.295014 0 0 -0.11711 0.073139 0.008786 0.235337 -0.00932 0.185541 0 0 0 0 0.235337 -0.04858 -0.66083 -0.55929 -0.0284 -0.48052 0 0 0 0 -1.0486 -0.66083 -0.27306 -0.55929 0 -0.80394 -1.0486 -0.11711 0.275876311 0.434748402 0.43474840, 1.53299690f 0.53326447~ 0.85118668E 0.76364796i 1.0617134~ 1.90093784 i 1.49513368E 0.85118668E 1.18342010', 1.645330m 2.92264992' 1.34136943: 1.97248834~ ( ( 0.76364796; 1.18342010' 1.02043701\ 1.71924109: 0.97877209: 1.53299690! ( { { 1 1.71924109 0.894176101 0.21836012 0.27587631 0.93669425· 0.33073167 0.08941368 0.21836012 0.53326447 0.27587631 0.1570576 0.08941368 0.7636479~ �KREBSPLLSBNUM SI-121-A SI-121-B DC-136-A DC-136-B AN-163-A AN-163-B AN-198-A AN-198-B AN-199-A AN-199-B AN-200-A AN-200-B AN-208-A AN-208-B AN-210-A AN-210-B DU-149-A DU-149-B AN-18-A AN-18-B AN-21-A AN-21-B AN-23-A AN-23-B AN-20-A AN-20-B AN-31-A AN-31-B AN-37-A AN-37-B AN-38-A AN-38-B AN-56-A AN-56-B AN-153-A AN-153-B C0-15-A C0-15-B CO-148-A DC-118-A DC-118-B DC-119-A DC-119-B DC-100-A DC-100-B DC-103-A UTMX 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 302378 302370 763726 763840 339226 339400 329626 329550 331126 331200 331426 331550 339626 339610 336626 336381 315226 315200 374826 374869 379849 379852 363426 363412 348526 348459 346626 346544 389326 389319 358726 358634 372726 372684 337526 337416 755508 755418 761083 763232 763158 763333 763262 746543 746470 751667 UTMY 4171441 4171435 4161571 4161650 4127771 4127800 4128671 4128600 4126371 4126400 4125971 4126000 4107771 4107779 4105271 4105136 4147571 4147900 4114271 4114352 4108329 4108424 4096771 4096848 4112171 4112107 4102771 4102762 4100371 4100270 4096271 4096289 4112271 41123320 4145471 4145472 4143632 4143617 4148897 4173781 4173814 4171687 4171605 4192851 4192824 4191077 PRIMOVESN1 S S S S 0 F P P P P P P SN2 SN4 SN3 SN5 2 1 2 SN6 SN7 SN8 3 SN9 4 1 3 2 SN10 MEAN 1 8 6 0 0 (...:'. P P 0 0 A A A A A A A A 2 1 1 0 0 P P S S 0 F F A A S S S S S S S S 3 8 2 8 2 2 1 3 6 1 5 3 7 11 3 5 1 2 5 6 3 2 Page 4 of 6 2 1 4 1 2 KREB'S N HARE DENSI~ 1 0.2 1.1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0.4 0 0 0 0 0 0 0 0 0 0 0 0 0 1.1 0 0 0.6 2.1 0 0 2.4 0 0 0.5 0.6 1 1.6 0 -0.23587 -0.80394 -0.20223 -0.23587 0 0 0 0 0 0 0 0 0 0 0 0 0 -0.55929 0 0 0 0 0 0 0 0 0 0 0 0 0 -0.20223 0 0 -0.41617 0.026007 0 0 0.073139 0 0 -0.48052 -0.41617 -0.23587 -0.06998 0 0.580939625 0.15705769 0.627728653 0.580939625 0 0 0 0 0 0 0 0 0 0 0 0 0 0.275876311 0 0 0 0 0 0 0 0 0 0 0 0 0 0.627728653 0 0 0.38355578 1.061713454 0 0 1.183420101 0 0 0.330731678 0.38355578 0.580939625 0.85118668 - �" , KREBSPLLSBNUM DC·103-B DC· 124-A DC·124·B DC·128-A DC·128-B DE·142·A DE·142·B DN·98-A DN·98-B DN·9g.A DN·9g.B DU·35-A DU·35-B DU·l94·A DU·l94·B DU·195-A DU·195-B DU·197·A DU·197·B DU·201·A DU·201·B DU·204·A DU·204·B DC·9·A DC·9·B DC·97-A DC·97-B DC·101-A DC·101-B DC·104-A DC·104-B DC·108-A DC-108-B SI·51·A SI·51-B SI-87·A SI·87·B 51·90-B SI·90-A SI·106-A 51·106-B SI·10g.A SI·109·B SI-114-A SI-114-B SI·11·A UTMX 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 751699 760062 760116 756231 756142 341726 341679 358626 358674 359126 359063 299926 299861 274126 274131 265726 265795 304326 304386 298826 298965 303126 303137 211626 211613 233026 232894 229826 229820 233926 233885 232Q26 232125 293005 293126 293826 293746 271226 271226 238966 238966 243126 243560 259926 260012 288426 UTMY 4191024 4169573 4169674 4166799 4166761 4154871 4154753 4195171 4195215 4194671 4194666 4130371 4130316 4132771 4132686 4132271 4132317 4130571 4130495 4124971 4125051 4119771 4119857 4181971 4181996 4196471 4196480 4193471 4193380 4190971 4190960 4188571 4188570 4199502 4199371 4206271 4206331 4203471 4203471 4189716 4189971 4187271 4187052 4181871 4181977 4157071 SN2 PRIMOVESN1 S D D P P 5 S S S 5 S P P P P P P P P D D P P A A S S 5 S S S S S S S S S 5 5 S S S S S S A SN4 SN3 9 1 5 1 6 1 1 3 5 4 SN5 3 1 7 9 SN6 2 5 5 SN7 1 3 8 5N9 5N8 4 8 1 8 SN10 MEAN 5 1 2 9 6 7 2 ~ .' 0 0 7 2 3 0 6 9 0 5 2 1 3 1 3 2 5 5 2 24 10 3 5 5 3 1 3 2 12 7 1 8 1 3 3 0 6 7 0 5 5 3 3 1 3 5 2 1 1 1 4 1 8 31 8 15 1 6 4 2 9 8 7 7 2 2 7 3 4 1 9 3 5 1 5 7 1 3 9 8 2 2 3 4 2 6 , 5 1 2 1 4 2 4 3 6 4 7 1 2 - Page 5 of 6 KREB'S N HARE DEN51T'l 0 0 0 0 0 0 2.3 1.9 2.8 3.5 2.8 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 2.1 3.4 1.6 3.1 4.7 2.2 1.7 2.2 1.9 2.4 2.1 2.2 1.6 2.5 3.3 0 0 2.4 1.8 0 0 0 0 0 0 0 0.058117 ·0.00932 0.127548 0.20631 0.127548 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.008786 0.026007 0.196078 ·0.06998 0.163474 0.310363 0.042427 ·0.04858 0.042427 -0.00932 0.073139 0.026007 0.042427 -0.06998 0.087548 0.185541 0 0 0.073139 -0.0264 0 0 0 0 0 0 0 1.143186215 0.978772092 1.341369433 1.60808792 1.341369433 0 0 0 0 0 C C C C C C C C C 1.02043701 ~ 1.06171345~ 1.570645831 0.85118668~ 1.45704818~ 2.04344557! 1.10262328 . 0.89417610( 1.10262328 . 0.97877209: 1.18342010' 1.06171345< 1.10262328 0.85118668~ 1.22334116, 1.53299690! ( ( 1.18342010' 0.93669425! .E -...l �KRE9SPLLS9NUM SI-11-9 SI-125-A SI-125-9 SI-129-A SI-129-8 SI·133-A SI-133-9 SI-134-A SI-134-8 UTMX 5 5 5 5 5 5 5 5 5 288420 276100 276250 264326 264400 287326 287150 277726 277550 UTMY 4156972 4167900 4167850 4166571 4166500 4164071 4164100 4162971 4163000 PRIMOVESN1 A A A 0 0 S S 0 0 SN2 SN4 SN3 SN5 SN6 SN7 SN8 2 7 SN9 SN10 5 7 3 3 1 1 9 2 2 9 1 c.. •• ., Page 6 of 6 - KRE8'S N HARE DENSITY o 00 MEAN 3 0.2 1.2 0.7 0.6 1.1 0 0.5 0.2 1 -0.80394 -0.17152 ·0.36176 ·0.41617 ·0.20223 0 -0.48052 -0.80394 -0.23587 ----- 0.15705769 0.673726665 0.434748402 0.38355578 0.627728653 0 0.330731678 0.15705769 0.580939625 ------ �109 Appendix G Genus-species Abbr. Name Common Scientific AMAL ARCO Arnica ARUV QUGA SHCA SYAL SYOC VA spp VASC cordifolia uva- Arctostaphylos ursi CA spp JU spp Serviceberry Amelanchier alnifolia Juniper gambelii Gambel's Oak Buffaloberry Shepherdia canadensis Symphoricarpos albus Symphoricarpos occidentalis Snowberry Snowberry spp Blueberry scoparium Broom Huckleberry Vaccinium Vaccinium Kinnikinnik spp Juniperus Quercus Arnica Sedge Carex spp . Heart-leaved �110 Appendix ------1--- -- ------ SUMMARY BY FOREST I H ------- --- --- SAMPLE __ -----SIZE 32 12 21 63 7 89 31 22 83 16 92 45 4 94 ARAPAHO NF BUFFALO PEAKS WILDERNESS GRAND MESA NF GUNNISON NF LAGARITA WILDERNESS AREA RIOGRANDE NF ROCKY MOUNTAIN NATIONAL PARK ROOSEVELT NF ROUTTNF SAN ISABEL NF SANJUAN NF UNCOMPAHGRE NF WEST ELK WILDERNESS WHITE RIVER NF I I I I SUMMARY BY AR~ I AREA AREA AREA AREA AREA HARES PER HA .. _------- 95% CI 1 2 3 4 5 SAMPLE SIZE HARES PER HA 114 62 67 129 239 0.29 0.27 0.41 0.28 0.56 0.16 0.14 0.22 0.16 0.35 0_3142 0_5089 0.1046 0.3259 0.8712 0_6679 0.2744 0_2158 0.2114 0.2815 0.3546 0.7032 0.1352 0.2632 0.2200 0.1900 0.3000 0.2100 0.4400 I SAMPLE SIZE SUMMARY BY PRIMARY OVERSTORY I I ASPEN BRISTLECONE PINE DOUGLAS FIR SUBALPINE FIR LODGEPOLE PINE GAMBLE'S OAK PONDEROSA PINE ENGLEMANN SPRUCE WHITE PINE LIMBER PINE I - -~, -- 105 2 48 64 108 16 41 220 1 3 , ••_-- -r- - -- --- -- ~-~~-=-j -------- -- -------- ------_-- -------- ------- ----- ~~~~~~~1A' ~~~-:--= CHAFFEE CLEAR CREEK CONEJOS DELTA DOLORES EAGLE GARFIELD GILPIN I GRAND GUNNISON HINSDALE JACKSON LA PLATA LAKE LARIMER MESA MINERAL MOFFAT MONTEZUMA I OURAY PARK PITKIN RIO GRANDE RIO BLANCO ROUTT I SAGUACHE SANJUAN SAN MIGUEL SARATOGA SUMMIT I 0.1558 0.0894 0.3351 0.3301 0.2981 0.0056 0.0737 0.5163 1.1026 0.7466 I SUMMARY BY COUNTY ---- HARES PER HA -- -- --- _-_. - - SAMPLE SIZE ----HARES _. .. _- _-- -_----_ 1----... _. .--- --.--.. -- ---- ---------- . --~ 33 -:;:- c------- 6 8 17 2 18 35 36 2 31 59 18 8 28 8 36 8 18 6 14 16 8 16 14 21 52 45 24 2 2 14 -------.-- . - - --------_._- PERHA 0_0438 0.1659 0.6117 0.3635 0.3642 0 0.6749 0.2941 0.3096 0.3010 0.1955 0.3163 0.9531 0.0769 0.3114 0.1931 0.2840 0.2152 0.7937 0.0874 0.0000 0.2864 0.4260 0.1393 0.9766 0.3316 0.1942 0.4143 0.5622 1.1828 0.1877 0.0888 �III Review of the habitat assessment study of snowshoe hares by the Colorado Division of Wildlife . Shay Howlin Lyman McDonald West, Inc. 2003 Central Avenue Cheyenne, Wyoming 82001 (307) 634-1756 February 14,2000 In our examination of the issues surrounding the snowshoe hare pellet collection in 1998 we have identified 4 major concerns in the use of this data. Here we attempt to clarify these concerns and offer our analysis of their severity as they relate to the use of the pellet data as an index to saowshoe hare abundance. CONCERNSOFREVffi~ • Failure to validate or accurately duplicate Krebs' protocol This criticism is based on the premise that the division of wildlife (DOW) snowshoe hare pellet study intended to use the linear models developed by Krebs et al. (1996) to estimate snowshoe hare density based on the number of snowshoe hare pellets. Krebs' model for estimating the density of hares was developed in Canada with different snowshoe hare ecology and habitat. And the field protocol used by Krebs to develop the density estimation formula specified the removal of pellets from plots at a set time period before counting pellets. The DOW study protocol differed from Krebs' protocol in that they did not clear the study plots of old pellets before conducting pellet surveys. The biologists conducting the surveys for the DOW were trained to age pellets and could identify last year versus current year pellets. In our view, the use of current year pellets can provide an index to abundance of snowshoe hares. The analysis of the DOW data does not need to validate or use Krebs' formula if only an index to abundance is attempted. • Failure to follow DOW survey protocol The DOW generated a random sample of survey points in each of the five study areas. The number of random points generated for the survey was roughly proportional to the size of the area. Recognizing the logistical problems associated with visiting all the random points, the DOW field crews reduced the number of plots-to visit by selecting every fourth point from the original random sample (call this subset the reduced sample). DOW and BLM biologists surveyed additional points from the original random sample when convenient. The resuiring dataset contains points in the reduced sample visited by the field crews, and points in the original sample visited by biologists opportunistically on the way to other points or other work. Since the resulting sample contains a �112 disproportionate number of points in block five, reviewers of the study have noted that the blocks with higher sampling intensity could result in increased chances of finding pockets of modest hare abundance. The fact that neither every point in the original random sample nor every point in the reduced sample was not surveyed resulted in a nonrandom sample of actual data. Biases associated with the selection of the plots sampled (i.e. closest to roads), makes the use of the data for an unbiased index to snowshoe hare abundance questionable. The tables below show the percent of original plots and the percent of the subset of plots that were surveyed in each block and were entered into the dataset called .newform.xls (obtained from G. Byrne on December 28, 1999). Percentage of Krebs' plots sampled from original random sample. Block: Numberof Original Percentof Number plots samplesize original visited sample 1 2 3 4 5 55 32 34 64 121 200 100 100 200 200 27.5 32.0 34.0 32.0 60.5 Percentage of .Krebs' plots sampled from the reduced sample(every fourth plot from the original sample). Block: Numberof Reduced Percentof plots samplesize reduced Number visited sample 1 2 3 4 5 18 20 22 20 31 50 25 25 50 50 36.0 80.0 88.0 40.0 62.0 • Time frame of surveys The DOW conducted the snowshoe hare pellet surveys throughout the summer of 1998. One reviewer of the study contends that plots that were sampled later in the season had the potential to have more pellets, because there was more time for pellets to be dropped onto the plots and there are more hares due to the addition of hares to the population through parturition. From Figure 1, it appears that each block had an even distribution of plots visited throughout the study period. If there was a time affect influencing the number of pellets on a plot, it appears it would have approximately the same influence in every block. ° • Definition of suitable habitat The DOW protocol details the placement of the Krebs' plots once the randomly selected point is located. Plots were moved from the starting point to the nearest suitable �113 habitat (forest, willow, or mountain shrub habitat). A random number table was used to locate directions and distances to move the point to suitable habitat. If the first randomly selected direction or distance did not locate suitable habitat, another direction and distance was chosen. .' . This protocol effectively reduces the area surveyed during the study to the "suitable habitat" in the study area. Field personnel made subjective determinations of suitable versus nonsuitable habitat, because a rigorous definition of suitable habitat is not easy to follow in the field. The actual size of the area that had a chance of being sampled is unknown and difficult to define. The size of this area could potentially be determined by the delineation of suitable habitat on the GIS vegetation coverage if definitions were consistent throughout the study. Figure 1. Date of Division of Wildlife sampling of plots by block. ..... ~ • ••• ~ 0 0 • 3 m ••• ••• OIi.-3Ir.JaOi/.'r.Ja , . ~ - - - - -••• -- •• - •.... • olino/sa ••• • • ••• - •• • •• • ••••••••• '6/3or.Ja • ••• .... ••• ••• ... -- ••• ••• • •• • .. .. . • ••••••••••••••••••• 07/10/sa 17/z:or.Ja 07n,/sa lB/osr.Ja 0II/1s•••• 'a 1B•••• es•••• sa Date CONCLUSIONS The failure of the DOW snowshoe hare study to validate or follow Krebs' protocol for the estimation of snowshoe hare density prohibits the use of the data in Krebs' estimation formulas. We do not feel this criticism discounts the use of the DOW data for an index of snowshoe hare abundance in the study area. The criticisms surrounding the time component of the study also do not seem to pose a critical problem when producing estimates of hare abundance by block. And if an estimate of suitable habitat can be made using the GIS coverage of the vegetation, the index of abundance can incorporate the amount of area surveyed. �114 Our review of the data revealed problems with the quality of the data. There were twelve missing dates in the dataset, and two dates that were 9 and 49 years before the bulk of the other observations. There were other simple errors in the plot identification variable that indicates the dataset was not verified. We recommend the data be checked for quality control-quality assurance before any analyses are conducted. Ifa truly random sample of Krebs' plots could be identified, we would recommend an analysis of the pellet data using only these plots. But neither all the points in the entire original sample were visited, nor were all the plots identified in the DOW biologists subset sample visited. We feel it is impossible to identify a sample of plots that were selected and visited that do not contain biases associated with the time and logistical constraints of locating and sampling the pre-selected Krebs' plots. Therefore, we do not think this data is capable of producing an index of snowshoe hare abundance that is unbiased. The fieldwork associated with the collection of this data has provided information that can be used to design a study of snowshoe hare abundance that will be adequate to produce estimates by block. The study provides an estimate of the plot to plot variance in the number of pellets in the study area, as well as an idea of the number of randomly placed Krebs' plots that can be surveyed in a season. LITERATURE CITED Krebs, C.J., B.S. Gilbert, S. Boutin and R. Boonstra. 1987. Estimation of snowshoe hare population density from turd transects. Canadian Journal of Zoology. 65: 565567. �115 Colorado Division of Wildlife Wildlife Research Report July 2000 JOB PROGRESS REPORT State of --.!::C~o,!!,lo~rad=o'__ _ Project No. W....:....:._--"'1=53~--"'R"'--..:..:13:::.__ _ Work Package No. __ .:::..0=88""0=<-- -'-- __ Task No. __!.1'-- Mammals Research · ·_-'--' _ __!.B~l~a~ck~· -~foo=ted~F::..;e~rr~e""t_"R""'eco=v.:..:e"""ry..L__ _ _ Monitoring and Managing Disease in Black-footed Ferrets Period Covered: July. 1, 1999 - June 30, 2000. Authors: M. A. Wild and K. T. Castle Personnel: E. Wheeler, E. Schmal, and S. Kasven. ) ABSTRACT \ ) The black-footed ferret is a federally listed endangered species in the United States. Black-footed ferrets have been extirpated from Colorado, but were scheduled to be reintroduced to Moffat County, Colorado in 1999. However, due to high plague activity and low prairie dog densities at the Little Snake Management Area (LSMA), black-footed ferret reintroduction was postponed indefinitely at the site. The secondary site at Coyote Basin, Utah was readied and reintroduction of ferrets was performed there in 1999. The Wolf Creek site in Moffat County, Colorado is also being readied for reintroduction of ferrets, likely in 2001. Our work in support of black-footed ferret reintroduction can be sub-divided into three broad sections: disease monitoring in the proposed release areas in Colorado, care of black-footed ferrets, and flea control as a tool to manage sylvatic plague in prairie dogs and black-footed ferrets. Disease monitoring was performed using collection of coyotes from LSMA in July 1999 and from the Wolf Creek site in February 2000. Two potentially devastating diseases, canine distemper and plague, are present at LSMA. Although prevalence of positive titers to canine distemper virus (CDV) in coyotes have been relatively low over the last 3 years ts33%), the prevalence of positive titers to plague (Yersina pestis) have been high (up to 89% positive). Titers were present in both adult and juvenile animals, suggesting ongoing plague activity in at least some sections of the management area. Samples collected from coyotes at the Wolf Creek site indicated substantially lower disease activity than at LSMA. Prevalence of positive titers to plague and to CDV was 7% of samples collected in February 2000. In general, captive black-footed ferrets maintained at the LSMA breeding and preconditioning pens remained healthy. One ferret died and another was presumed dead in the burrow system after a severe hailstorm. Of 13 kits born at the pens in spring 1999, 12 survived to weaning in fall 1999. These 12 kits in addition to 50 other black-footed ferrets were released at the Coyote Basin site in fall 1999. Twenty ferrets were maintained in the pens overwinter and produced 11 kits in spring 2000. We summarized research results and presented a paper titled "Dose titration and safety oflufenuron fed to captive white-tailed prairie dogs (Cynomys leucurus)" at the 2000 meeting of the �116 Wildlife Disease Association, Jackson, Wyoming. We also performed a trial totes~the efficacy of lufenuron in controlling fleas on captive prairie dogs. One week post-dosing, fleas that fed on prairie dog treated with 500 mglkg lufenuron produced a lower proportion (p < 0.05) of viable rggs than those fed on control prairie dogs (mean 0.16 vs. 0.39, respectively). However, the proportion of viable eggs produced during week 2 post-dosing was low for each group (0.24 treatment vs. 0.17 control). By week 4 postdosing, egg production had fallen to nearly zero in each group; however, the three eggs produced by fleas fed on treated prairie dogs were all viable. Although further, similarly controlled investigation is warranted, preliminary results suggest that given current techniques, the likelihood of lufenuron limiting fleas populations and breaking the plague cycle is not extremely promising. ( �117 MONITORING AND MANAGING DISEASE IN BLACK-FOOTED FERRETS Margaret A. Wild and Kevin T. Castle P. N. OBJECTIVES 1. Monitor disease activity threatening survival of black-footed ferrets reintroduced into the Little Snake Management Area (LSMA). 2. Develop techniques to manage plague in the LSMA using insect growth regulators applied orally to prairie dogs. SEGMENT OBJECTIVES 1. Provide veterinary care to captive and reintroduced black-footed ferrets. 2. Monitor and manage plague activity in LSMA. METHODS AND MATERIALS Carnivore Disease Survey Infectious diseases can severely impact the success of black-footed ferret (Mustela nigripesy reintroduction efforts. As part of the black-footed ferret reintroduction protocol, we monitored disease activity in carnivores at proposed ferret reintroduction sites: in July 1999 at the Little Snake Management Area (LSMA), Colorado and in February 2000 at the WolfCreek Management Area (WCMA), Colorado. Coyotes (Canis latrans) were collected for post-mortem examination and samples collected as described in the Program Narrative (Wild and Castle 1998). Black-footed Ferret Reintroduction and Veterinary Care I assisted in preparation of the black-footed ferret allocation request submitted to the US Fish and Wildlife Service by the Colorado-Utah black-footed ferret recovery working team in 2000. I provided veterinary care and consultation on health matters for captive black-footed ferrets and black-footed ferret releases. Animal care was performed in accordance with the established protocol (Wild and Castle 1999). Flea Control In Prairie Dogs We completed the experiment "Bioavailability oflufenuron administered orally to captive white-tailed prairie dogs (Cynomys leucurus)" and performed a follow-up experiment "Efficacy of lufenuron for the control of fleas in white-tailed prairie dogs (Cynomy leucurus)". These studies were outlined in Wild and Castle 1998. The detailed study plan for the lufenuron bioavailability study and efficacy trial are reported in Wild and Castle 1999 and in Attachment 1, respectively. �118 RESUL TS AND DISCUSSION Carnivore Disease Survey With the assistance of USDA Wildlife Services and the Bureau of Land Management (BLM) we collected 16 coyotes from the LSMA between 26-30 July 1999 and 14 coyotes from the WolfCreek site on 8-9 February 2000. Coyotes were collected using a combination of calling and aerial gunning. Further collections were not possible due to weather constraints and aircraft availability. No lesions indicative of active disease were noted on gross examination of carcasses. Of the coyotes collected from the LSMA, 31 % (5/16) had positive titers to plague (Fig. 1) using the standard HAIHI test while one additional coyote was positive using the ELISA test. Interestingly, all juveniles sampled this summer (n = 9)' were negative to plague. Because the juvenile coyotes have conswned (sampled) prairie dogs from this summer only, lack of exposure may indicate that the prevalence of plague in prairie dogs in the area is declining. If this is the case, titers in adults would likely be from exposure in previous years. Alternatively, the prey base of the coyotes may have shifted to species that are less commonly infected with plague (e.g., rabbits) if the density' of prairie dogs has been greatly reduced by plague. Forty-three percent of coyotes sampled (6/14 usable samples) had positive titers to tularemia. In contrast to plague, all positive titers to tularemia were observed in juvenile coyotes. As previously observed, tularemia appears to elicit a serologic response of short duration in coyotes. The impact of tularemia on black-footed ferrets and prairie dogs is unknown and warrants investigation. A titer to canine distemper virus (CDV) was found in only one of 14 serum samples tested (Fig. 2). Samples collected from coyotes at the Wolf Creek site indicate substantially lower disease activity than at LSMA. A 9-year-old male coyote had positive titers to plague and to CDV, but all other coyotes (n = 13) were negative to plague and CDV (7% positive; Fig. 3). The adult male and one additional juvenile coyote also had titers to tularemia (14% positive). Black-footed Ferret Reintroduction and Veterinary Care In general, captive black-footed ferrets maintained at LSMA remained healthy. One adult female was treated for an apparent mite infection and secondary bacterial pyoderma. Unfortunately, samples to confirm this diagnosis could not be collected prior to treatment; however, the ferret responded quickly and completely to therapy with ivermectin and amoxicillin. Additional cases of crusty skin have been successfully treated with ivermectin but confirmation of the etiology has not yet been made. A I-yr-old male was found dead and a 4-yr-old female was missing and preswned dead after a severe hailstorm. Necropsy results revealed death from blunt trauma. The death(s) occurred in the new pen complex where shallow burrow systems may have been flooded forcing ferrets to the surface in the severe weather. To avoid this problem in the future, additional above ground shelter will be provided. Of 13 kits born at the pens in spring 1999, 12 survived to weaning in fall 1999. One kit disappeared and is preswned dead in the burrow system. Based on results of carnivore disease monitoring over the past 3 years and prairie dog inventories performed in summer 1999, LSMA was determined to be currently unsuitable habitat for the release of black-footed ferrets. Prairie dog inventories performed by Utah Division of Wildlife showed insufficient densities of prairie dogs to support black-footed ferret reintroduction. As a result, ferrets were not released into LSMA but instead were released into Coyote Basin, Utah. Continued monitoring will determine when (if) LSMA can support reintroduction of black-footed ferrets. A secondary site in Colorado, (WolfCreek) is also being readied for reintroduction of black-footed ferrets in fall 2000 or 2001. I ,_./ �119 The 12 kits produced onsite, in addition to three l-yr-old males, and five 3-yr-old females from the LSMA pens were released at Coyote Basin in November 1999. Additionally, 19 kits and five 3-yr-old females from other captive breeding sites were pre-conditioned at the LSMA prior to release at Coyote Basin. The reintroduction was further supplemented with 28 ferrets released immediately upon arrival from other captive breeding sites without pre-conditioning at LSMA. Prior to release, ferrets were trapped for routine examination, treatment, and identification (Wild and Castle 1999) and a health certificate was issued for each individual. All appeared healthy except for the presence of ectoparasites (ticks, mites, fleas). Individual ferrets were treated with ivermectin and pen dusting was advised. In an attempt to meet our ideal age and sex structure of captive black-footed ferrets (Wild and Castle 1999), we retained seven ferrets and supplemented the population with 13 additional ferrets from other captive breeding facilities. Seven black-footed ferrets (five 1-yr-old females and two males) were retained in captivity at the LSMA pens. In November 1999, six adult females, 'three female kits, and four male kits were added to this breeding group bringing the total number of ferrets at the LSMA pens to 20. Ferrets were maintained under the standard care protocol (Wild and Castle 1999). Females were paired with males in spring 2000, and four litters resulted. One litter was apparently consumed by the female when about 1 day of age. The other three litters yielded 11 kits. Flea Control In Prairie Dogs We presented results of the lufenuron dose titration study at the 2000 meeting of the Wildlife Disease Association, Jackson, Wyoming. The abstract of that presentation read: DOSE-TITRATION ANDSAFETYOFLUFENURONFEDTOCAPTIVEWHITE-TAILED PRAIRIEDOGS(CYNOMYS LEUCURUS) KEVINT. CASTLEColorado Division of Wildlife, 317 W. Prospect St. Fort Collins, CO, 80521, MARGARETA. WILD, Colorado Division of Wildlife, 317 W. Prospect St. Fort Collins, CO, 80521, andS. CRAIGPARKS,Novartis Animal Health, P.O. Box 26402, Greensboro, NC 27404. Plague is a zoonotic disease that impacts populations of prairie dogs (Cynomys spp.) and other species, such as black-footed ferrets, that rely on them for food and shelter. Yersinia pestis, the etiological agent of plague, is transmitted primarily by the bite of an infective flea. Recently developed compounds used to control fleas in pet animals offer a promising alternative to insecticide dusts for the control of fleas in wild rodents. Lufenuron is a lipid-soluble insect growth regulator with ovicidal and larvicidal activity. Lufenuron is efficacious for controlling fleas in cats and dogs at blood concentrations above 50-100 parts per billion (Ppb). A single oral dose oflufenuron has been shown to be effective in controlling the cat flea (Ctenocephalides fobs fobs) for at least 30 days in treated cats and dogs. To date, there have been no studies conducted to detennine the duration of lufenuron blood concentrations in any prairie dog species. We compared lufenuron blood concentrations in white-tailed prairie dogs (c. leucurus) during periods of activity (nontorpid group) and hibernation (torpid group) during January-March 1999. We hypothesized that ifhigh serum concentrations of lufenuron could be maintained over winter during hibernation or for> 1 mo in active prairie dogs, the compound may be effective for use in breaking the plague cycle. Thirty captive WTPD were fed 300 mg/kg lufenuron; half the animals were allowed to become torpid, while the other half were kept awake. All animals remained healthy throughout the 9 week study period. Prairie dogs in the active group gained weight, while those in the torpid group lost weight over the 9 weeks. Blood was drawn from each animal prior to dosing, one week after dosing, then every other week until week 9 post-dosing. Serum was harvested and tested by HPLC for lufenuron concentration. Blood lufenuron concentration did not differ between the groups one week post-dosing. Concentration in both groups decreased over time, but the �120 concentration in torpid animals declined at a more gradual rate; after weeks 3, 5, and 7, lufenuron levels in torpid WTPD were significantly higher than levels in nontorpid WTPD. After nine weeks, blood levels were again similar, and had approached the limit of detection (10 ppb). Blood levels in nontorpid WTPD declined to <50 ppb after 3 weeks, while levels in torpid WTPD declined to <50 ppb after 7 weeks. Future studies will be required to determine efficacy of lufenuron in controlling fleas on WTPD. If effective blood concentrations are similar to dogs and cats, however, frequent dosing would be required to control flea numbers on prairie dogs and thus break the plague cycle. Results from this experiment indicated that blood concentrations of lufenuron did not reach initial levels as high as anticipated, nor were the concentrations maintained above 50 ppb for as long as anticipated (Fig. 4). The most likely cause of these low concentrations was poor absorption of the drug by prairie dogs. This may be due to the difference in gut morphology between rodents and carnivores. Alternatively, other aspects of pharmacokenetics may have been responsible for the low serum levels in prairie dogs despite dosing at rates 10-30 times higher than those recommended for cats and dogs, respectively. We followed up the dose titration experiment with an experiment to test the efficacy of orallufenuron to control fleas on captive prairie dogs. Based on data from the bioavailability study, we increased the lufenuron dose to 500 mg/kg. Serum samples from the study have been submitted for lufenuron assay. Results are pending. Interpretation of efficacy results will rely on these serum lufenuron levels; however, we were able to make some preliminary comparisons between performance of fleas fed on treatment and control prairie dogs. One week post-dosing, fleas that fed on treated prairie dog produced a lower proportion (p < 0.05) of viable eggs than those fed on control prairie dogs (mean 0.16 vs. 039, respectively). Unfortunately, the proportion of viable eggs produced during week 2 post-dosing was low for each group (0.24 treatment vs. 0.17 control). By week 4 post-dosing, egg production had fallen to nearly zero in each group; however, the three eggs produced by fleas fed on treated prairie dogs were all viable. We are uncertain of the cause of the dramatic reduction in egg production and viability observed during the course of the experiment. Anecdotal reports suggest that flea production may be influenced seasonally (or at least cyclically) despite attempts to maintain controlled environmental conditions in the insectary (Metzger, Pers. Comm.). Regardless, it is unfortunate that serum levels oflufenuron decreased more rapidly than we had expected and that data did not support the hypothesis that lufenuron would be an effective means to significantly reduce flea production over the summer. Although further, similarly controlled investigation is warranted, preliminary results suggest that given current techniques, the likelihood of lufenuron limiting fleas populations and breaking the plague cycle is not extremely promising. Therefore, experiments into efficacy of controlling flea infestations in simulated burrow environments and in the field (described in Wild and Castle 1998) will not be performed. LITERATURE CITED Wild, M. A. and K. T.' Castle. 1998. Monitoring Div. Wildl. Res. Rep., 0880-1, Jul1997 - Jun Wild, M. A. and K. T. Castle. 1999. Monitoring Div. Wildl. Res. Rep., 0880-1, Jul1998 - Jun and managing disease in black-footed ferrets. Colorado 1998, Fort Collins. and managing disease in black-footed ferrets. Colorado 1999, Fort Collins. �121 Attachment I Kevin T. Castle, Margaret A. Wild, and S. Craig Parks Colorado Division of Wildlife Foothills Wildlife Research Facility (KTC and MAW) Novartis Animal Health, Greensboro, NC (SCP) Introduction Black-footed ferret (Mustela nigripes) recovery plans call for the reintroduction of ferrets to sites characterized by the presence of viable populations of prairie dogs (Cynomys spp.), which provide food and shelter for ferrets. Unfortunately, prairie dogs inhabiting many potential reintroduction sites carry fleas that can serve as vectors ofYersinia pestis, the causative agent of plague (Ubico et al, 1988). Prairie dogs and ferrets are both highly susceptible to plague (Barnes, 1993; Williams et al., 1994) so the chances of successful ferret reintroduction will be enhanced if the numbers of fleas infesting a prairie dog colony can be significantly reduced. Lufenuron is a benzoylphenylurea derivative which inhibits formation of chitin in the exoskeleton of insects (Cohen, 1987). A single oral dose oflufenuron has been shown effective in controlling the cat flea (Ctenocephalides felis felis) for at least 30 days in treated cats (Blagburn et al., 1994) and dogs (Hink et al., 1994; Blagburn et al., 1995). No studies on the efficacy oflufenuron have been conducted with prairie dogs; however, Davis (1997) reported a significant reduction in fleas on free-ranging ground squirrels (Spermophilus beecheyi) that had been treated with lufenuron. We are performing a series of experiments to test the applicability of lufenuron to control fleas in captive, and ultimately wild, white-tailed prairie dogs (Cynomys leucurus). Thus far in pilot studies we have determined standard husbandry, maintenance, and handling protocols for captive prairie dogs and determined a test dose oflufenuron based upon a bioavailability study. We have also attempted to establish an insectary colony of a flea species (Oropsylla tuberculata) that naturally infests prairie dogs and their burrows. Oropsylla fleas are among the most common prairie dog fleas, and have been implicated in the transmission of plague in prairie dogs (Ubico et al., 1988). Fleas of this genus are nest fleas that infest the host only to obtain a blood meal. At other times the fleas select microenvironments within in the burrow system that are conducive to successful reproduction and survival. In pilot studies we were unable to develop methods to successfully maintain and produce self-sustaining populations of 0. tuberculata in an insectary. However, a related species, 0. montana, which naturally infests ground squirrels (e.g. Spermophilus beechyi) has been successfully maintained in the insectary using methods of M. Metzger and K. Gage (pers. comm). These fleas will feed on prairie dogs and successfully reproduce after ingesting a blood meal. Because of its similarity to O. tuberculata, we will use O. montana as a model to determine if flea reproduction can be controlled in lufenuron-treated prairie dogs. A chambered-flea system has been used to test on-host viability and fecundity of fleas on cats (Thomas et al. 1996) and laboratory mice (D. Engelthaller, pers. comm.). In this system, fleas are contained in a chamber attached to the host, and can obtain a blood meal through a mesh screen. This system has several advantages over other methods of artificial infestation. First, because a known number of adults can be placed into the chamber, flea mortality is easily noted, and survivorship can be determined. Second, sex ratios can be adjusted to maximize egg production. lbird, eggs can be recovered readily, to determine viability. Fourth, the environment in the chamber will be warm and humid, and therefore conducive to adult and egg survival. Finally, fleas will not be able to leave the prairie dog, and therefore will not be able to infest caretakers or other animals. �122 In this study we will: 1) develop techniques for artificially infesting white-tailed prairie dogs with chambered populations of fleas, 2) test the flea control efficacy oflufenuron fed to white-tailed prairie dogs artificially infested with fleas, and 3) compare efficacy of flea control with serum lufenuron levels. Our working hypothesis is that fleas which feed on lufenuron-dosed prairie dogs will have reduced survival of eggs and larvae compared to fleas which feed on control animals that receive no lufenuron. Methods Prairie Dog Maintenance We will use 31 captive white-tailed prairie dogs maintained at the Foothills Wildlife Research Facility in Ft. Collins in our experiment. Prairie dogs will be housed singly (ifused in experiments) or in pairs (if used for flea colony maintenance) in custom-designed cages (100 em x 500 ern x 600 em; Wild and Castle 1998). Prairie dogs will be observed daily, and will have ad libitum access to Teklad Rodent Blocks and water. Windows will provide a natural photoperiod, and a combination of timed heaters and air conditioners will be used to keep ambient temperature between 10 and 25° C. Twenty prairie dogs (10 males and 10 females) will be blocked by sex and randomly divided equally into a treatment group and a control group for the experiment. The treatment group will receive lufenuron while the control group will not. Because all animals cannot be housed in one building due to space limitations, the groups will be split evenly between two separate but similar buildings, to help control for any interbuilding differences (e.g. temperature, humidity) that may exist. The sample size of20 prairie dogs will allow us to detect a :::::91 % reduction in the number of eggs that successfully hatch in the treated group given alpha = 0.10 and beta = 0.90. An additional 11 captive prairie dogs will be maintained under similar conditions, but will not participate in the experiment. Instead, they will be used in the maintenance of the flea colony (see below). Lufenuron Dosing Prairie dogs in the experimental group will be fasted for 24 h prior to dosing; water will be available during the fast. After the fasting period (day 0), each experimental animal will be weighed, then offered a bolus dose oflufenuron (300 mg/kg body mass). Lufenuron will be mixed thoroughly in approximately 10 g of highly palatable bait (ground rat chow and molasses); control animals will receive bait only. We previously determined that a majority of fasted prairie dogs consume about 90-95% of their dose in less than 12 h. We will observe the progress of bait ingestion in each animal to determine when the dose is ingested. Remaining bait will be removed after 24 h. We will weigh the baitllufenuron mixture before and after the prairie dogs are dosed, in order to calculate the actual dose ingested. After dosing, normal feeding will resume. Flea Infestation An initial stock oflaboratory-reared, disease-free fleas (0. montana) will be obtained from Dr. Kenneth Gage at the Centers for Disease Control and Prevention (CDC) in Ft. Collins. Fleas will be housed in 500 ml glass jars containing larva-rearing media, to allow the population to be self-sustaining. Rearing media consists ofwheast (Red Star Biologicals), dried beef blood (Monfort Biologicals), powdered dog chow, and sand. Fleas will be maintained in an incubator at about 22-23° C and :::::70%relative humidity (M. Metzger, pers. comm.) to ensure optimal reproduction. A natural photoperiod will be approximated within the incubator using 'fluorescent lights and a timer. Adult fleas must ingest a blood meal in order to reproduce. We will use two methods to provide blood meals to fleas held in the insectary for propagation of the colony. The first method will follow an established protocol (Castle and Wild 1998) to provide blood to the adult fleas in each rearing chamber, using neonatal rodents. Briefly, when fleas are in need of a blood meal (1-2 times per week), we will obtain . ~ , �123 neonatal rodents from a private colony. The neonates will be placed into the insectaries with fleas for up to 24 h; previous work has shown that over 80% of the neonates are alive after 24 h. No food or water will be provided for the neonates, as they are strictly dependent on nursing. After feeding by the fleas, neonates will he euthanized by an overdose of inhalant anesthetic. Once per week, for 7 weeks, we will utilize the 11 non-experimental prairie dogs as blood sources, using the chambered flea technique described below. While the use of neonatal rodents is an efficient, approved method of providing blood to fleas, neonatal rodents are not always available from private colonies. Prairie dogs will therefore serve the dual purposes of providing blood meals when neonates are unavailable, and minimizing the number of neonatal rodents sacrificed. One week prior to study initiation, and on study days 2, 7, 14,21,28,35, and 42 we will place flea chambers on experimental prairie dogs. Fleas feeding on treated prairie dogs will potentially be exposed to lufenuron from this blood meal. We will collect 50 adult female and 30 adult male fleas from the insectary and place theminto a chamber (2.5 em diameter). Each chamber will be·attached to a prairie dog so the fleas can obtain a blood meal. To attach the chambers, prairie dogs will be anesthetized using isoflurane delivered by a vaporizer. Respiration and depth of anesthesia will be monitored. A patch of fur on the dorsal thorax caudal to the shoulders will be shaved. A flea chamber will be placed on the skin, and taped into place using Elastikon and Vet-rap. The prairie dog will then be placed in a 30 em x 20 cm x 20 em holding box to recover from anesthesia, and will be monitored while the chamber is attached. Chambers will remain on each prairie dog for 30 min. At the end of the feeding time, the prairie dog will again be anesthetized with isoflurane, and the tape and chamber will be removed. Prairie dogs will be returned to their cages after recovery from anesthesia. Fleas will be observed for evidence of feeding by observation under a IO-2Ox microscope, and blood-filled fleas will be placed in plastic vials and put into our insectary for egg recovery. 0. montana fleas typically lay eggs within 2-3 days of a blood meal (M. Metzger, pers. comm.), so adult dishes will be monitored every day for 7 days to monitor egg production. Eggs will be removed from the adult dish and placed in new vials inside the insectary; they will be observed every day for larval emergence. The total number of eggs produced by fleas from each prairie dog will be recorded, as will the number oflarva that emerge eachday. Unfed adult fleas will be returned to the insectary (prior to lufenuron dosing) or preserved in alcohol (after lufenuron dosing). Blood Collection Concentration oflufenuron in the blood may be an indicator of efficacy of flea control. To determine the relationship between blood lufenuron concentration and egg viability and larval development, we will collect 3 ml of blood from each anesthetized prairie dog prior to chamber attachment on each sampling day. Blood will be collected by jugular venipuncture and placed into a glass vacutainer blood tube without anticoagulant. Alternatively, ifwe are unable to collect an adequate blood sample peripherally, we will collect blood from the vena cava or directly from the heart. Although cardiac puncture is considered to be a safe procedure for collection oflarge volumes of blood from laboratory rodents (CCAC, 1984), due to the increased risks involved with cardiac puncture, we will use this technique only to obtain critical samples. Serum will be harvested and frozen within 4 h of collection. Serum lufenuron concentration will be determined by HPLC at Novartis Laboratories. Based on our pilot studies and other literature (Blagburn et al. 1994, 1995; Thomas et al. 1996) we do not anticipate any prairie dog mortality due to the flea infestations, lufenuron dosing regime or blood collection, but if a severe allergic reaction or other health problem associated with our procedures occurs, the prairie dog will be removed from the study and provided veterinary care or euthanized with an overdose of inhalant anesthetic or barbiturate. A complete post-mortem examination will be performed on any animal that may die during the experimental period. �124 Data Analysis Efficacy of lufenuron will be determined by comparing developmental success of eggs produced by fleas exposed to treated prairie dogs vs. eggs produced by fleas exposed to control group prairie dogs. These formulas will be used in efficacy determination: Developmental success = number of larvae hatched x 100 number of eggs collected Percentage efficacy=mean developmental success (control) - mean developmental success (treated) x 100 mean developmental success (control) Mean developmental success values for eggs collected from prairie dogs at each sampling period will be analyzed using analysis of covariance, using blood lufenuron concentration at each time step as the covariate. Group responses will be considered significantly different if p < 0.10. Literature Cited Barnes, A. M. 1993. A review of plague and its relevance to prairie dog populations and the black-footed ferret. Proceedings of the symposium on the management of prairie dog complexes for the reintroduction of the black-footed ferret. J. L. Oldemeyer, D. E. Biggins, B. J. Miller, and R Crete, eds. 96pp. Blagburn, B. L., J. L. Vaughan, D. S. Lindsay, and G. L. Tebbitt. 1994. Efficacy dosage titration of lufenuron against developmental stages of fleas (Ctenocephalides felis felis) in cats. American Journal of Veterinary Research 55: 98-101. Blagburn, B. L., C. M. Hendrix, J. L. Vaughan, D. S. Lindsay, and S. H. Barnett. 1995. Efficacy of lufenuron against developmental stages of fleas (Ctenocephalides felis felis) in dogs housed in simulated home environments. American Journal of Veterinary Research 56: 464-467. Castle, K. T. and M. A. Wild. 1998. Protocol for the use of neonatal rodents as a blood source for insectary-reared fleas. Colorado Division of Wildlife Animal Care and Use Committee Study Plan. Canadian Council on Animal Care (CCAC). 1984. GUide to the care and use of experimental animals, Vol. 2. Ottawa, Ont., Canada. Cohen, E. 1987. Interference with chitin biosynthesis in insects. In Chitin and benzoylphenyl ureas, series entomologica, vol. 38, J. E. Wright and A. Retnakaran (eds), Dr. W. Junk, Publishers, Boston, pp.3342. Davis, R M. 1997. Use of an orally administered insect development inhibitor (lufenuron) as a flea control agent in the California ground squirrel, Spermophilus beecheyi. Fourth International Symposium on Ectoparasites of Pets, pp. 31-35. Hilton, D. F. 1. 1971. A method for rearing fleas of ground squirrels. Transactions of the Royal Society of Tropical Medicine and Hygiene. 66: 188-189. Hink, W. F., M. Zakson, and S. Barnett. 1994. Evaluation ofa single oral dose oflufenuron to control flea infestations in dogs. American Journal of Veterinary Research 55: 822-824. Thomas, R E., L. Wallenfels, and I. Popeil. 1996. On-host viability and fecundity of Ctenocephalides felis (Siphonaptera: Pulicidae), using a novel chambered flea technique. Journal of Medical Entomology 33: 250-256. Ubico, S. R, G. O. Maupin, K. A. Fagerstone, and R G. McLean. 1988. A plague epizootic in the whitetailed prairie dogs (Cynomys leucurus) of Meeteetse, Wyoming. Journal of Wildlife Diseases 24: 399-406. Williams, E. S., K. Mills, D. R. Kwiatkowski, E. T. Thome, and A. Boerger-Fields. 1994. Plague in a black-footed ferret (Mustela nigripes). Journal of Wildlife Diseases 30:581-585. Wild, M. A. and K. T. Castle 1998. Monitoring and managing disease in black-footed ferrets. Colorado Division of Wildlife Program Narrative, Project # W-153-R �125 • Plague-Neg 20 o Plague-Pas 15 •... Q) .0 E 10 :J Z 5 1997-W 1997-S 1998-W 1998-S 1999-W 1999-S Fig. la. Prevalence of exposure to plague in juvenile coyotes from the Little Snake Management Area, Colorado, from winter 1997 through summer 1999. 20 ue-Pos 15 Q) .0 E 10 :J Z 5 1997-W 1997-8 1998-W 1998-8 1999-W 1999-8 Fig. lb. Prevalence of exposure to plague in adult coyotes from the Little Snake Management Area, Colorado, from winter 1997 through summer 1999. Data from winter 1997 (age class unknown) provided by M Albee. �126 25 -,.--_' 0 CDV positive II1II CDV negative 20 1997-W 1997-S 1998-W 1998-S 1999-W 1999-S Fig. 2. Prevalence of exposure to canine distemper virus (CDVY in coyotes from the Little Snake Management Area, Colorado, from winter i997·through summer 1999. All positive coyotes were adults with the exception of one juvenile in summer 1998. Data from winter 1997 (age class unknown) provided by M. Albee. 20 15 .. .se 10 Z=' 5 0 Plague CDV Fig. 3. Prevalence of exposure to plague and canine distemper virus in adult coyotes from the Wolf Creek site, Colorado in winter 2000. 200 § .~ 175 •..• 150 1:l 8 ,-)25 ~.D 8500 § •... § ~ j 75 • 50 • 25 • • • O+---.---~--~--~----.---.---~---.---. 2 3 4 5 6 7 8 9 10 Week Post-Dosing Fig. 4. Mean serum lufenuron concentrations of active (+) and torpid (.) prairie dogs orally dosed with 300 mg/kg lufenuron. � https://cpw.cvlcollections.org/files/original/a3816a28b63f15aded34fbf4785f7874.pdf bc5ab3c68360a122ed82e894fbbf3217 PDF Text Text 127 Colorado Division of Wildlife Wildlife Research Report July 2000 JOB PROGRESS State of REPORT Colorado Project No. Cost Center 3430 W-153-R-13 Work Package No. _---'3:;...;:0'-"'0..:...1 Task No. 1 Period Covered: Mammals Program Deer Conservation _ Investigating Factors Contributing to Declining Mule Deer Numbers July 1 1999 June 30, 2000 Author: T. M. Pojar Personnel: _) .-- / T. Baker, T. Beck, C. Bishop, G. Bock, D. Coven, L. Dehart, B. Diamond, B. Dreher, J. Ellenberger, V. Graham, J. Griggs, D. Gustine, B. Hoffner, B. Lamont, M. King, K. Larsen, M. Mclain, H. McNally, K. Miller, M. W. Miller, E. Myers, J. Olterman, D . Schweitzer, T. Spraker, B. Watkins, S. Znamenacek. ABSTRACT -, ), /i: d Long-term data indicate that fawn:doe ratios have shown a significant and consistent decline over the past 20 years in most deer management units in Colorado as well as in most of the western states. This has resulted in a general decline in mule deer (Odocoileus hemionus) density and diminished recreational potential from this major resource. It is important for the Division of Wildlife to understand the factors contributing to the reduced fawn:doe ratios and the declining deer populations. Neonatal survival (birth to 6 months of age) is an important component of population recruitment. This is the second year of monitoring cause-specific neonatal fawn mortality on the Uncompahgre Plateau. In 1999, a sample of 50 fawns was radioed and during the 2000 fawning season 88 fawns were captured. Fawns were captured as . near birth as possible and are monitored to about 6 months of age. A new design radio collar that weighed 1109 was used on the 2000 season fawns. The collar was expandable and used latex tubing to facilitate drop-off in about 6 months. Neonatal fawn survival for 1999 was 38%. Of the 50 fawns collared, 30% died of sickness/starvation, 26% from predators (12% coyote, 8% feline, and 6% bear), 4% from unknown causes, and 2% from poaching. Thirty-one of 50 fawns died with 58% dying from causes other than predation. Monitoring is in progress for the 2000 season fawns. Early season 2000 mortality is less than it was in 1999. Deaths from sickness/starvation are half (10% vs. 20%) what they were in 1999 (as of . August 1) and predation is about three-fourths (15% vs. 20% that of 1999 (Figure 1). June 2000 was warm and dry, which may have affected early fawn survival. On the Uncompahgre, December fawn:doe ratios are negatively correlated (r = -0.8183) with the preceding June precipitation (R~.6697, P=0.0038). However, mean temperature for June had a weak positive correlation (r = 0.2756) and was not significant (R2 =.0.0760, P = 0.4409). The peak offawning was between June 14thand June 26th when 93% of the �128 fawns were caught (Figure 2). Year 2000 fawns were slightly larger than the 1999 fawns in terms of body weight and hindfoot length. Neither the difference in weight, 4.4692 (2000)vs. 4.3714 kg (1999), nor the difference in hing foot length, 10.474 (2000) vs. 10.297 (1999) inches, was statistically different, P = 0.5845 and P = 0.1309, respectively. �129 IMPACT OF PREDATION AND VEGETATIVE COVER ON MULE DEER FAWN SURVIVAL Thomas M. Pojar P. N. OBJECTIVES 1. Identify agents of neonatal mule deer fawn mortality from birth to 6 months of age. SEGMENT OBJECTIVES 1. Capture and radio collar 30-35 neonatal fawns each on the Uncompahgre Plateau (D-19), Middle Park (D-9), and Red Feathers (D-4). 2. Measure vegetative density and height at fawn bed sites. 3. Measure distance between successive bed sites for individual fawns. 4. Compare fawn survival and causes of mortality among 3 vegetational and ecologically different study areas. 5. Correlate height and density of vegetation at fawn birth sites and bed sites with percent of fawns killed by predators. 6. Compare height and density of vegetation at fawn bed sites and random sites. INTRODUCTION Major changes in the original Program Narrative include: 1. Limit the fawn collaring effort to the Uncompahgre Plateau (D-19). Because of the size of the area and diversity of vegetative communities on the Uncompahgre and because of intense political interest in this area, all resources were concentrated on the Uncompahgre during this segment. The target sampling intensity was increased to 80 fawns with emphasis placed on getting the sample widely distributed over the entire 2,300 square mile area. 2. Abandon the idea of measuring vegetation at bed sites because of evidence that persistent disturbance of bed sites reduces survival of young ungulates (Philips 1998). 3. The logistics of placing an adequate crew in remote fawning areas and the unknown of whether or not fawns could be captured in sufficient numbers given deer density, terrain, and vegetative cover encountered negated many of the objectives in the PN. In essence, this investigation was reduced to placing emphasis on capturing fawns on the Uncompahgre and monitoring their survival and causes of mortality. Summer fawn survival and causes of mortality are 2 important factors impacting deer population performance. STUDY AREA The study area is described in Pojar and Andelt (1999). METHODS Radio collared does were used to locate fawning areas although no special effort was made to capture the fawns of radioed does. The strategy for capturing fawns involved observing does for behavioral and physical signs, such as udder development, of having fawned. If it was determined that these indicators suggested that the doe had fawned, the area was searched for the fawn(s). Once located, �130 the fawn was approached from behind (out of direct eyesight) and restrained for weighing, measuring, and putting on the radio collar. Processing usually took less than 5 minutes. The collars used were made of 1.5-inch elastic with an expansion ratio of 1.5:1 and equipped with surgical tubing to allow the collar to drop off after about 6 months. Each collar weighed 110 g. The circumference of the collar was 10 inches, 6 of that was elastic. The 6 inches of elastic has the ability to expand to 9 inches. A l-inch loop was sewn in the elastic reducing the collar circumference to 9 inches. The loop was sewn with 4 stitches of cotton thread with the intent that these would rip out within 2-3 weeks after the collar was put on a fawn. When the collar was put on a fawn it was 9 inches in circumference and fit quite loosely. When the loop was released the circumference increased to 10 inches and as the elastic relaxed the collar could expand another 3 inches giving a totally expanded circumference of 13 inches. Collars were hung outside in the shade for several days before use to help dissipate any fabric and human scents associated with manufacture. When taken to the field in workers backpacks, the collars were sealed in zip loc bags. If a collar was to be recycled, i.e. removed from a dead fawn and reused, it was washed in tap water (no soap), hung out to dry, then placed in aplastic bag with native forage. RESULTS By recycling some of the collars, 88 fawns were radioed. The sample was roughly distributed according to deer density on the summer range. The sample of radioed fawns were distributed across the Uncompahgre Plateau as follows: North (Cold Springs) 29, central (25 Mesa) 16, south (Celesca) 33, and far south (10). Thus far, the collars are expanding as expected. From recovered collars, it appears the cotton stitching rips out in about 4 weeks or less. There was only one instance of suspected slippage of a collar and it is possible the fawn was killed or scavenged and the collar carried away from the carcass. Of the fawns recovered, there was no sign of the collar causing abrasions on the skin, however, there was slight hair loss in some cases from the collar being loose enough to rotate or move on the neck. The fawns captured during 2000 had a mean weight of 4.47 kg compared to 4.37 kg in 1999; this difference was not significant (P=0.5845). The hind foot length of 10.48 inches in 2000 was not different from that of 1999, 10.30 inches (P=0.1309). There was a noticeable difference in the weather, both precipitation and temperature, during the fawning period between 1999 and 2000 with the most recent year being warmer and dryer. Early sickness/starvation deaths (through August 1) in 2000 were half (10% vs. 20%) what they were in 1999 (Figure 1). It was theorized that the warmer, dryer June weather was a key factor in neonate fawn survival. Weather data for the Montrose area was tabulated for the past 10 years. I tested the hypothesis that June temperature and precipitation are related to the subsequent early winter (December) fawn:doe ratios collected during helicopter classification surveys done by management personnel. The correlation with precipitation was highly significant and negatively correlated (r = -0.8183) with subsequent fawn:doe ratios, R2 = 0.67, P = 0.0038; temperature had a slight positive correlation (r = 0.2756) with fawn:doe ratios but it was not significant, R2 = 0.0760, P = 0.4432. Although the search for newborn fawns began in early June, the first fawns were not caught until June 14thin 2000. In the 13-day period between June 14thand June 26th, 93% of the fawns were caught. The last fawn was caught on July 9th (Figure 2). In 1999, the 17-day period from June 13 to June 30,96% of the fawns were captured; the first fawns were captured on June 9th. From this data there is no evidence that a double fawning peak is occurring on the Uncompahgre Plateau. Data on captured fawns is in Table 1. Although the information for 2000 regarding mortalities only goes through August Pt, Figure 3 shows a comparison of total cause-specific mortality for 1999 (total) and 2000 (through August 1~. �131 LITERATURE CITED Philips, G. E. 1998. Effects of human-induced disturbance during calving season on reproductive success of elk in the upper Eagle River valley. Ph. D. Dissertation, Colorado State University, Ft. Collins. Pojar, T. M. and W. F. Andelt. 1999. Investigation offactors contributing to declining mule deer numbers. Wildlife Research Report, Mammals Research, Federal Aid Projects, Job Progress Report, Project W-153-R-12, Colorado Division of Wildlife, Fort Collins, Colorado, USA. Table l. Neonatal mule deer fawns captured and radio collared on the Uncompahgre Plateau, Colorado 2000. The mortality data is very preliminary and represents only deaths through August 1,2000. Probable Cause Radio I.D. Date of Capture location UTM Hindfoot Date Sex Wt Recovered of Death Capture Coordinates (kg) (inches) East North 150.023 6-15 703980 4268055 F 10.75 4.4 150.033 6-18 700954 4278742 F 10.25 4.7 150.044 6-16 702309 4280011 F 11.125 5.3 150.054 6-24 703446 4267697 F 11.25 5.1 150.064 6-18 700353 10.625 4274667 M 5.0 150.074 6-24 703050 4267256 M 11.00 4.4 150.084 6-24 703442 4267726 M 6.1 11.00 150.094 6-18 704622 4274176 M 11.00 5.0 150.104 6-22 700255 10.625 4282798 F 5.5 150.114 6-19 697815 5.3 11.375 4283738 M 150.134 6-20 6+ 11.875 703390 4279744 M Sick/starve 150.146 6-19 7-9 701207 4279027 M 9.75 3.9 150.155 6-20 702179 10.125 4275357 F 4.1 150.163 6-18 700357 10.50 4283325 M 4.2 7-19 Sick/starve 150.184 6-22 703468 10.00 4279642 M 4.0 150.192 6-20 702293 4.5 10.125 4281859 M 150.204 6-22 '4.7 700428 4283374 M 10.25 150.214 6-26 704463 10.50 4272215 F 4.7 Coyote 150.223 6-21 751358 6-23 4236752 M 3.6 10.50 150.234 6-20 727652 10.125 4252092 F 3.5 723136 150.244 6-26 4257872 F 10.625 5.7 150.254 6-22 730938 4264027 M 3.9 9.625 150.263 6-19 750827 4230756 M 4.0 10.25 150.272 6-25 700728 10.25 4284815 M 3.75 150.283 6-25 703475 10.50 4279355 F 5.0 10.00 150.294 6-26 241593 4235614 M 4.2 Sick/starve 150.303 6-21 760162 11.25 7-31 4237065 M 5.2 7-3 Bear 150.314 6-23 732743 4251998 M 11.125 5.0 150.324 6-26 241806 9.50 4236255 M 2.9 721362 10.00 8-1 Coyote 150.334 6-22 4264368 M 4.6 150.344 6-20 720095 4265861 F 5.2 11.00 Coyote 150.353 6-24 730048 9.75 7-27 4263042 F 3.4 150.362 6-20 748878 4231358 M 10.625 4.6 150.374 6-21 756526 10.75 4248692 M 5.1 150.383 6-26 728565 4254798 M 3.4 9.875 Coyote 150.393 6-21 10.125 7-5 743115 4255001 F 3.8 150.423 6-25 10.25 730831 4263910 M 4.0 150.434 6-18 757228 4232141 F 5.3 11.25 7-17 _~LcJcl~L ___ 6-20 4.~~!~2~_ __I4.~~I~ ___ J':____4.]___ J.9.:~Q_____ -----------__ l_?.9A4.4. -------- �132 Table 1 Continued Radio I.D. Date of Capture 150.455 150.464 150.473 150.484 150.495 150.504 150.514 150.524 150.545 150.553 150.564 150.573 150.584 150.593 150.604 151.114 151.123 151.144 151.153 151.164 151.173 151.195 151.203 151.233 151.245 151.254 151.263 151.274 151.284 151.294 151.314 151.324 151.333 150.013A 150.013B 150. 124A 150. 124B 150.172A 150.172B 150.403A 150.403B 150.415A 150.415B 150.533A 150.533B 151.134A 151.134B 151.183A 151.l83B 6-19 6-16 6-19 6-14 6-17 6-19 6-16 6-19 6-19 6-23 6-17 6-15 6-17 6-14 6-20 6-15 6-16 6-18 6-18 6-18 6-18 6-18 6-26 6-17 6-17 6-14 6-14 6-22 6-22 6-22 6-19 6-22 6-15 6-26 7-6 6-24 6-27 6-20 6-26 6-26 7-9 6-18 6-26 6-21 6-29 6-16 6-30 6-18 7-5 Capture location UTM Coordinates North East 762275 4233303 755233 4247651 760381 4232242 754488 4235165 756570 4233083 762275 4233303 4247828 754392 727428 4251901 749686 4229850 755706 4231637 4246702 757702 752002 4249457 754295 4247567 4249079 752525 754867 4231692 756942 4231730 754848 4331643 751910 4249670 751910 4249670 4249165 752547 752639 4248689 742905 4240385 731972 4251646 4251207 750332 754406 4248275 248077 4233576 248077 4233576 762261 4224531 761180 4220892 761688 4221672 761310 4220980 761688 4221672 761539 4223050 703376 4267562 704293 4269378 705226 4269426 4274463 704284 702297 4280730 730004 4261815 728978 4254395 702133 4280026 730056 4262079 730074 4261926 751358 4236752 707752 4270072 755167 4231731 707752 4270072 746380 4247269 701385 4279114 Sex M M M F M M M F M M F M M M F M F F F F F M F M M F M F F F F F F F F F F M M M M F F M F F F F M Wt (kg) 5.7 3.1 5.3 3.8 4.1 5.2 4.5 3.6 5.1 3.8 3.0 4.7 4.8 3.6 4.8 4.3 3.6 3.8 3.6 5.0 4.1 4.7 5.7 4.7 5.2 4.3 4.6 4.8 6.1 3.4 3.1 3.6 3.5 6.0 5.8 3.4 4.0 4.9 5.0' 4.3 6.7 3.5 3.0 3.7 6.1 4.4 5.0 4.0 6+ Hindfoot (inches) 11.50 9.625 10.50 10.50 10.25 10.75 10.625 10.25 10.25 10.50 9.625 11.00 11.125 10.125 10.25 10.50 9.875 10.25 9.75 10.50 10.625 10.75 11.125 10.25 11.375 10.83 10.25 10.43 12.20 10.16 8.86 10.00 9.76 11.00 11.00 8.00 10.25 11.00 10.875 10.625 11.50 9.875 8.875 10.50 10.625 10.75 11.50 10.125 11.75 Date Recovered Probable Cause of Death 7-8 Sick/starve 6-30 7-31 6-25 Coyote Bear?· Sick/starve 6-23 Sick/starve 7-3 Bear 6-24 . 7-5 6-23 7-31 6-21 6-26 7-6 Sick/starve Bear Coyote Sick/starve Coyote Coyote Coyote �133 YEAR 2000 - Through Number of fawns radioed Number of fawn deaths Cause Predation Sick/starve Total Alive % of morts Number 13 9 66 August 1 88 % of Total Sample 22 Predation 'lb % of total 59% 41% 15% 10% 75% Sick/starve 10% Total Alive 75% YEAR 1999 - Through Number of fawns radioed Number of fawn deaths Cause Predation Sick/starve Total Alive Number 10 10 30 August 1 50 20 % ofTo~1 % of morts % of total 50% 20% 50% 20% 60% Sample Predation 20% Figure 1. Comparison of cause-specific mule deer fawn mortality, Uncompahgre Plateau, Colorado. represents mortality from collaring through August 1st. 14 "'C Cl) 12 •... .aa. 10 C'O () 8 - - 6 I-- o c ~ u. o o Z 4 f- f- 2 o lLU .IL_I_ Jul8 Jun 14 Jun 18 Jun 22 Jun 26 Jun 30 Jul4 Jun 16 Jun 20 Jun 24 Jun 28 Jut 2 Jut6 Figure 2. Capture dates for year 2000 fawns from the Uncompahgre Plateau, Colorado. Data �CAUSE-SPECIFIC FAWN MORTALITY - UNCOMPAHGRE 2000 (Through Aug 1) Number of fawns radloe 88 Num ber of fawn deaths 22 % of mort % of total 36% 9% 18% 5% 5% 1% 41% 10% 75% •....• w .j::o. r----------------------------, Cause Number Coyote 8 Bear 4 Unk Pred 1 Sick/starve 9 Total Alive 66 % '" Number Cause Coyote 6 4 Feline Bear 3 Sick/starve 15 Unknown 2 Poached 1 Total alive 19 Bear 5% Unk Pred 1% Sick/starve 10% or Mortalltl •• ,... CAUSE-SPECIFIC FAWN MORTALITY - UNCOMPAHGRE 1999 (Total for year) Number of fawns radloe 50 ...--'---------------------------, Number of fawn deaths ·31 % of mort % of total 19% 12% 13% 8% 10% 6% 48% 30% Totalilive 6% 4% 38% 3% 2% 38% % % of Total Sample ./. of Total Sample Coyote 12% Bear 6% or Mortalltl.s •..". Unknown 4% Sick/starve 30% '''' Figure 3. Comparison of cause-specific mortality for 1999 and 2000, Uncompahgre Plateau, Colorado. The tabulation of 1999 includes the entire period from fawn capture to approximately 6 months of age. The data for 2000 only goes through August 1, 2000 (the date of this writing). �135 \ Colorado Division of Wildlife Wildlife Research Report July 2000 \ JOB PROGRESS REPORT State of Project No. Work Package No. __ Task No. Cost Center 3430 W-153-R-13 Mammals Research Program -=3,-"0-,,,-0-,,-1 Deer Conservation _ 4 Period Covered: Authors: Colorado Effects of Habitat Enrichment on Mule Deer Recruitment and Survival Rates July 1, 1999 - June 30, 2000 C. J. Bishop and G. C. White L. H. Carpenter, D. Coven, D. J. Freddy, R. B. Gill, R. Harthan, J. Sazma, B. E. Watkins, and B. Welch Personnel: ) ABSTRACT A Program Narrative (Appendix A) was developed to test the effects of habitat enrichment on mule deer recruitment and survival rates on the Uncompahgre Plateau in southwest Colorado. Post hoc analyses were completed to evaluate the effects of buck harvest on age and sex ratios (Appendix B), and to roughly estimate the total number of deer in Colorado (Appendix C). ·C., ,: .• �136 ;: I -, �137 EFFECTS OF HABITAT ENRICHMENT ON MULE DEER RECRUITMENT AND SURVIVAL RATES C. J. Bishop and G. C. White P. N. OBJECTIVES 1. To conduct a one-year pilot study to assess the logistical feasibility of the proposed study and to gather preliminary data to improve the study's efficiency and experimental design. 2. To determine experimentally whether enhancing mule deer nutrition during winter and early spring by supplemental feeding increases December fawn:doe ratios and overwinter fawn survival. 3. To determine experimentally to what extent habitat treatments replicate the effect of enhanced nutrition from supplemental feeding. SEGMENT OBJECTIVES 1. Prepare a Program Narrative. 2. Plan field logistics and purchase equipment and supplies in preparation for initial field work beginning in FY 2000-01. INTRODUCTION Mule deer numbers apparently declined during the 1990's throughout much of the West, and have clearly decreased since the peak population levels documented in the 1940's-60's (Gill et al. 1999, Unsworth et al. 1999). Biologists and sportsmen alike have concerns as to what factors may be responsible for declining population trends. Although previous and current research indicates that multiple interacting factors are responsible, habitat and predation have received the focus of attention. A number of studies have evaluated whether predator control increases deer survival, yet results are highly variable (Connolly 1981, Ballard et al.1999). Together, predator control studies with adequate rigor indicate that predation effects on mule deer are variable as a result of time-specific and site-specific factors. Studies which have demonstrated deer population responses to predator control treatments have failed to determine whether predation is ultimately more limiting than habitat. Numerous research studies have evaluted mule deer habitat quality, but virtually no studies have documented population responses to habitat improvements. In many areas where declining deer numbers are of concern, predation is common yet habitat quality appears to have declined. The question remains as to whether predation or habitat is more limiting to mule deer in these situations, and whether habitat quality can be improved for the benefit of deer. We designed a field experiment to measure deer population responses to habitat enrichment treatments. We will conduct the study on the Uncompahgre Plateau, where several predator species are present in abundant numbers. Predator numbers will not be manipulated in any way. Habitat enrichment treatments will consist of supplemental feed provided to deer during the winter. If December fawn:doe ratios and overwinter fawn survival improve as a direct result of-the supplemental feed, then we can presume that deer nutrition is ultimately more limiting than predation. The field experiment also incorporates habitat manipulation treatments, which will consist of prescribed fire or mechanical techniques to set back succession of pinyon-juniper habitat in an effort to improve the vigor and quality of winter habitat for mule deer. Deer population responses will be measured in relation to the habitat manipulations in the same manner as the supplemental feed. Thus, the experiment allows us to determine whether nutritional quality �138 of habitat is ultimately more limiting than predation in a late-seral pinyon-juniper/sagebrush if so, whether habitat can be effectively improved for mule deer. landscape, and There have been several challenges to implementing this study. First, using supplemental feed in a field study sends a contradictory message to the public given the Division of Wildlife's policy on winter feeding. Second, the study is very expensive and therefore requires considerable Division support to implement. Third, logistical concerns raise the issue of whether the study can be carried out appropriately, such that response variables are representative of treatment differences and measured without bias yet with adequate precision. Finally, some personnel have questioned the need for such a study, particularly given the expense. In result, we gave a number of presentations explaining the study in detail to generate widespread support throughout the Division. We also decided to conduct a pilot study in the first year to address logistical concerns and to allow more time to generate necessary funding. MATERIALS Program Narrative AND METHODS Development We developed a Program Narrative (Appendix A), which addresses both the l-year pilot study as well as the complete 4-6 year study. The P. N. has not been peer-reviewed at this point, but will be peer-reviewed in early FY 2000-01 prior to the pilot study. The P. N. will be modified as necessary using results from the pilot study, and once again peer-reviewed assuming there are significant changes. We based the P. N. on an initial study proposal submitted by Gary White and Len Carpenter. We conducted a thorough literature review and discussed study methodologies with numerous Division personnel to refine field methods. We spent time in the field with the local Area Biologist, Bruce Watkins, and District Wildlife Manager, Dale Coven, assessing the Uncompahgre Plateau for potential treatment and control areas. We worked closely with the Uncompahgre Ecosystem Restoration Project (UERP) committee to select treatment/control sites in the context of proposed habitat manipulation areas. Uncompahgre Ecosystem Restoration Project The Uncompahgre Ecosystem Restoration Project (UERP) was initiated by the Division of Wildlife after former Director John Mumma allocated $500,000 of capital construction funds to habitat improvements for the benefit of mule deer. UERP is comprised of individuals from DOW, U.S. Forest Service, Bureau of Land Management (BLM), and the Colorado West Public Lands Partnership (PLP). Since the initial $500,000, more funds have been obtained for UERP by the other agencies and PLP. Mike Mclain (AWM) and Bruce Watkins (Area Biologist) have represented DOW's interests in UERP from the onset, while Jim Garner (Habitat Biologist) and I joined the effort this past winter. UERP's mission is to improve ecosystem health on the Uncompahgre Plateau through an integrated effort of the various agencies and groups involved. A primary goal is to increase species diversity, age diversity, and the quality of habitat communities on the Plateau in order to improve the distribution and quality of mule deer winter range. To accomplish this goal, UERP is using a landscape approach to treat habitat using various techniques, primarily prescribed fire and mechanical treatments. All treatments will be re-seeded with shrub-forb-grass mixtures in an attempt to prevent invasive weeds from establishing and to increase forage diversity for mule deer. Deer population responses on one of these habitat treatment areas will be intensively monitored as part of our study's experimental design. The various other habitat treatments will be implemented in a pattern of treatment and control areas such that long-term deer responses can be monitored using the current mule deer population monitoring system on the Uncompahgre Plateau. UERP is currently in the process of hiring a technical coordinator and a policy/education coordinator to facilitate project implementation. The technical coordinator will assist with project design, implementation, and evaluation; provide technical expertise and budget oversight; assist with grant writing and production �139 of project reports; and ensure that proposed projects comply with state and federal regulations. The policy/education coordinator will provide information concerning the project and generate support from government agencies, stakeholders, and the general public. In essence, the policy/education coordinator will serve a classic "information and education" role for UERP. Once the coordinators are hired, we will begin identifying specific treatment sites and proceed with actual habitat treatments. UERP's current .timeline should enable some habitat manipulations to be completed in 2-3 years as needed for our habitat enrichment study. NEPA Requirements Since much of the field work for our habitat enrichment study will be conducted on BLM lands, we are subject to the requirements of the National Environmental Policy Act (NEPA). There are 2 types of experimental treatments in our study: (1) supplemental feeding and (2) habitat manipulations. We submitted our P. N. along with a detailed description of the supplemental feeding protocols to the Uncompahgre Field Office of the BLM. They determined that the feeding portion of our study meets the Categorical Exclusion requirement ofNEP A, thereby granting approval to proceed without further review. As explained above, all habitat manipulations will be performed by UERP, which assumes the responsibility for obtaining NEP A approval and any other regulatory clearances. Presentations We presented our habitat enrichment study to both Division personnel and other groups throughout the year. For Division personnel, our goal was to explain the study and address a variety of concerns that had been raised. It was our intention to gain support for the study from the various leaders in the Wildlife Programs Branch, as well as managers and biologists in the West Region arid particularly those in Area 18. ..·For non-Division groups, we discussed the various issues affecting mule deer populations, and described the habitat enrichment study in light of how the Division of Wildlife is attempting to address mule deer concerns. We gave presentations at the following meetings, workshops, etc.: • DWM Training Session (for incoming D\VMs) - 11512000, Denver • West Section Terrestrial Biologist Staff Meeting - 2/7/2000, Grand Junction • CSU StudentslFaculty, Wildlife Field Studies Class Trip - 3/1112000, Uncompahgre Plateau • Wildlife Programs Branch Meeting - 3/14/2000, Denver • Hunter Education Instructors, Big Game Workshop - 4/15/2000, Montrose • Hunter Education Instructors, Masters Workshop - 5/6/2000, Winter Park • Colorado Elementary and Secondary Education Teachers, Teacher Education Program 6/23/2000, Ridgway State ParklUncompahgre Plateau • Area 18 Staff Meeting -7/27/2000, Lone Cone Cabin The following presentation was also given, but was unrelated to the habitat enrichment study: • Large Mammal Capture Techniques and Radio-Telemetry, CSU Wildlife Management Short Course - 312912000, Fort Collins Other Activities Mule Deer Legislative Report Along with Mammals Research Leader, Bruce Gill, and other members of the research staff, I assisted in the writing, editing, and publication of "Declining Mule Deer Populations in Colorado: Reasons and Responses". The report was written for and submitted to the Colorado Legislature at their request. The report was distributed widely via hard copies and the internet. I was responsible for responding to a number of comments the Division received concerning the report. I presented the information contained in the report in several of the presentations listed above. �140 Mule D