https://cpw.cvlcollections.org/files/original/bca0978931a1dc9eab810fca096c6fcc.pdf d5994a87784a46504c6214c1adae59b3 PDF Text Text Colorado Division of Parks and Wildlife July 2013-June 2014 WILDLIFE RESEARCH REPORT State of: Cost Center: Work Package: Task No.: Colorado 3420 1680 N/A Federal Aid Project No. N/A : : : : Division of Parks and Wildlife Avian Research Bird Conservation Avian response to plague management on Colorado prairie dog colonies Period Covered: September 1, 2013 – August 31, 2014 Author: R. Yale Conrey Personnel: D. Tripp, J. Gammonley, CPW; A. Panjabi, E. Youngberg, Rocky Mountain Bird Observatory. All information in this report is preliminary and subject to further evaluation. Information MAY NOT BE PUBLISHED OR QUOTED without permission of the author. Manipulation of these data beyond that contained in this report is discouraged. ABSTRACT Range-wide declines in prairie dog (Cynomys sp.) populations have occurred, and the largest limiting factor in recent decades appears to be the high mortality and colony extirpation associated with plague (Antolin et al. 2002), caused by the bacterium Yersinia pestis. Prairie dog colonies support a diverse community of associated species, many of which are not susceptible to plague but may be indirectly affected. In order to conserve prairie dogs and species associated with their colonies, principally the black-footed ferret (Mustela nigripes), a plague vaccination program is being developed, which may also benefit a suite of species listed in the Conservation Plan for Grassland Species in Colorado (Colorado Division of Wildlife 2003) and the Colorado Sagebrush Conservation Assessment and Strategy (Boyle and Reeder 2005). In Colorado, CPW researchers led by Dan Tripp are surveying colonies before and after bait distribution and conducting a mark-recapture study of prairie dogs and associated small mammal species. As an extension to this project, we initiated research in 2013 on the effects of plague management on avian species associated with prairie dog colonies, with particular focus on species of concern. The 2014 season was the second of three study seasons. We collected the first year of posttreatment data and continued investigating whether avian species associations exist for colonies of Gunnison’s prairie dogs (Cynomys gunnisoni: GUPD); most evidence for associated species comes from black-tailed prairie dogs (C. ludovicianus: BTPD). Since fall 2013, plague epizootics have occurred on one GUPD colony and across half the BTPD study area at four treatment sites and five additional colonies. In September and October 2014, black-footed ferrets were released in three BTPD study colonies, one of which was only 0.5 km from the nearest baited site, which experienced a plague epizootic in fall 2013. In 2014, we detected 95 bird species, with 113 total bird species detected over two seasons, half of which were unique to on- or off-colony sites. We documented 175 plant species in two years, half of which were unique to either BTPD or GUPD sites. Colonies contained a much higher bare ground component with lower vegetation height (heights were similar on- and off-colony in 2013) than off- 1 �colony sites, with shortgrasses dominant at BTPD sites and a more even distribution of plant types at GUPD sites. We detected 13 raptor species during counts. Burrowing owls, northern harriers, ferruginous hawks, prairie falcons, and rough-legged hawks have been detected only on prairie dog colonies (the latter three, only on BTPD colonies, where no off-colony counts were performed). It is unclear whether golden eagles and American kestrels have a higher level of use on- versus off-GUPD colonies, as a strong relationship existed in one year but not the other. Apparent nest success was 47% in 2014 compared to 53% in 2013, with most of the decrease attributable to hail storms and flooding at BTPD sites. An increase in nest numbers was partly due to increased effort and partly to a huge influx of lark buntings during a wet year. We monitored 68 nests of 11 species in 2013 and 225 nests of 15 species in 2014. Our remote camera photos have documented use of vaccine project areas by coyote, badger, fox, and several other carnivores. Overall 4-week carnivore detection rates for May 2013 – April 2014 (naïve occupancy estimates) ranged from 7% for swift fox on BTPD colonies to 54% for coyotes on GUPD colonies. This was the first year of post-treatment data collection on the project, and it will likely take additional years of monitoring to detect potential changes in the avian community caused by plague management, as treated colonies no longer experience extinction events. Thus far, data from vegetation surveys have identified differences between on- and off-colony areas for GUPD sites, but bird data will require further analysis and a third year of data before we draw conclusions regarding the uniqueness of the avian community on GUPD colonies. Work will continue in 2015. 2 �COLORADO PARKS AND WILDLIFE RESEARCH REPORT AVIAN RESPONSE TO PLAGUE MANAGEMENT ON COLORADO PRAIRIE DOG COLONIES REESA C. YALE CONREY PROJECT OBJECTIVES The main long-term objective is to determine whether areas treated to control plague differ from untreated areas in their avian communities. Over time, avian communities on vaccinated prairie dog colonies may differ from those on dusted colonies or those with continued exposure to plague. Shorterterm objectives are to: 1. Determine whether avian species associations exist for Gunnison’s prairie dog colonies. 2. Determine whether insecticidal dusting influences bird density or nest survival. 3. Evaluate response of avian community to other land management that may occur on study areas, such as cattle grazing or flood irrigation. SEGMENT OBJECTIVES This was the second year of data collection and the first year of post-treatment data collection. Specific objectives for 2014 included: 1) Improve protocols. 2) Conduct avian point counts and vegetation surveys at points on and off colonies. 3) Sample vegetation on transects on and off colonies. 4) Quantify use of on- and off-colony sites by breeding and wintering raptors. 5) Find and monitor success of nests for passerines, mountain plover, burrowing owls, and other raptors; characterize vegetation at nests on colonies. 6) Sample carnivores using remote cameras on colonies. INTRODUCTION Wildlife diseases are important to conservation and population dynamics of susceptible species and may also have large indirect effects on non-susceptible species (Antolin et al. 2002). Introduced pathogens have the potential for far-reaching effects on native ecosystems that go beyond the mortality of infected individuals, particularly when a keystone species (Paine 1969) or ecosystem engineer (Jones et al. 1994) is infected. Range-wide declines in prairie dog (Cynomys sp.) populations have occurred, and the largest limiting factor in recent decades appears to be the high mortality and colony extirpation associated with introduced plague (Antolin et al. 2002), caused by the bacterium Yersinia pestis. Plague epidemics were first reported in the western United States in 1899 (Dicke 1926) and in northern Colorado in 1948 (Ecke and Johnson 1952). Instead of living in extensive colonies as they once did, prairie dogs exist in metapopulations of smaller colonies that periodically go extinct and are recolonized (Antolin et al. 2002, Stapp et al. 2004). Prairie dog colonies support a diverse community of associated species (Lomolino and Smith 2004, Smith and Lomolino 2004, Hardwicke 2006, Stapp et al. 2008), many of which are not susceptible to plague but may be indirectly effected. In order to conserve prairie dogs and species associated with their colonies, principally the blackfooted ferret (Mustela nigripes), a plague vaccination program is being tested. Additional species that may benefit from this program include those listed in the Conservation Plan for Grassland Species in Colorado (Colorado Division of Wildlife 2003): burrowing owl (Athene cunicularia: BUOW), mountain plover (Charadrius montanus: MOPL), ferruginous hawk (Buteo regalis: FEHA), and swift fox (Vulpes velox) and in the Colorado Sagebrush Conservation Assessment and Strategy (Boyle and Reeder 2005): Brewer’s sparrow (Spizella breweri: BRSP), green-tailed towhee (Pipilo chlorurus: GTTO), sage sparrow (Artemisiospiza belli: SAGS), sage thrasher (Oreoscoptes montanus: SATH), and vesper sparrow 3 �(Pooecetes gramineus: VESP), as well as BUOW. BUOW and MOPL are known to decline or disappear on colonies that are not reoccupied by prairie dogs after plague epizootics (Butts and Lewis 1982; Sidle et al. 2001; Augustine et al. 2008; Tipton et al. 2008; Conrey 2010), and horned lark, McCown’s longspur, golden eagle, and prairie falcon may benefit from active colonies. From 2013‒2015, researchers in several western states are field-testing the uptake and efficacy (SPV Subcommittee 2011) of a new sylvatic plague vaccine (SPV) for prairie dogs (Rocke et al. 2010). In Colorado, CPW researchers led by Dan Tripp are surveying colonies before and after bait distribution and conducting a mark-recapture study of prairie dogs and associated small mammal species (Tripp and Rocke 2012). As an extension to this project, we are researching the effects of plague management on avian species associated with prairie dog colonies, with particular focus on species of concern. We have collected one year each of pre- and post-treatment data on birds, and additionally intend to demonstrate whether avian species associations exist for colonies of Gunnison’s prairie dogs (Cynomys gunnisoni: GUPD); most evidence for associated species comes from black-tailed prairie dogs (C. ludovicianus: BTPD; Lomolino and Smith 2004, Smith and Lomolino 2004). A vaccine study was initially proposed for white-tailed prairie dogs as well, but work in Colorado will remain limited to BTPD and GUPD. During the initial years of the vaccination project, avian monitoring is also contributing information on responses to climate, grazing, and insecticidal dusting. This project will aid in the development of a standardized protocol for monitoring species associated with prairie dog colonies that could be used state-wide, as called for in the Conservation Plan for Grassland Species in Colorado (Colorado Division of Wildlife 2003). This project involves cooperators from Rocky Mountain Bird Observatory (RMBO), City of Fort Collins, Bureau of Land Management (Gunnison office), National Park Service Florissant Fossil Beds National Monument, and CPW wildlife managers and biologists from Areas 4, 14, and 16. METHODS Study Area Study areas included BTPD colonies in north-central Colorado and GUPD colonies in central Colorado. Baited sites received either vaccine or placebo baits in a blind procedure. Project areas that were selected for the prairie dog vaccine study had adequate numbers of prairie dogs and good access. BTPD (Larimer and Weld Co.) – Study colonies were located in Larimer and Weld Co. adjacent to the Wyoming border at Soapstone Prairie Natural Area (SOAP), managed by City of Fort Collins Natural Areas Program and Meadow Springs Ranch (MSR), managed by City of Fort Collins Utilities Department. These sites are characterized by shortgrass and mixed-grass prairie dominated by grasses (blue grama Bouteloua gracilis and buffalograss B. dactyloides) with smaller amounts of native (scarlet globemallow Sphaeralcea coccinea) and non-native forbs, shrubs, and cactus. Sites were sometimes grazed by cattle at low densities, and some non-baited sites were dusted with deltamethrin to control fleas (and plague). There was much more cattle grazing in 2014 than in 2013, because it was a wetter year with more forage. Both properties were closed to recreational shooting. Mark-resight estimates of BTPD density on shortgrass prairie in Colorado average approximately 10 prairie dogs/acre (Magle et al. 2007). Bird and vegetation surveys were conducted on five SOAP colonies and 20 MSR colonies. There were nine vaccine project areas where raptors, predators, and passerine nests were surveyed: three prairie dog complexes each received vaccine, placebo, and dusting treatments (3 treatments*3 complexes = 9 project areas). 1) The Jack Springs (Jac) colony spanning the SPNA/MSR border contained 100 vaccine acres, 100 control acres, and ~280 dusted acres separated by 200 - 400 m buffer zones. 2) The Barton complex in MSR contained ~130 vaccine acres and ~180 control acres, encompassing much of the Barton south and west colonies (BarS and BarW). Raptor, predator, and passerine nest surveys were also done in the 140 acre Barton east colony (BarE), although the prairie dog crew instead pairs a dusted portion of the 4 �Ferret Center colony (Fer) with the Barton complex. 3) The Ferret Center complex in MSR contained 40 vaccine acres, 40 control acres, and ~478 dusted acres, encompassing the entire North Benson south colony (NBenS) and half of the Ferret Center (Fer) colony, with a 400 m buffer zone separating those treatments. GUPD (Gunnison Basin and Woodland Park) – Study colonies were located in the Gunnison Basin (Gunnison, Saguache, and eastern Montrose Co.) and in the Woodland Park area (Teller Co.). Gunnison Basin (GUNN) sites were managed by the Bureau of Land Management (BLM), Colorado Parks and Wildlife (Miller Ranch State Wildlife Area and Van Tuyl/Cabin Creek SWA), and the U.S. Forest Service Rio Grande National Forest. Woodland Park area (WOOD) sites were managed by the National Park Service Florissant Fossil Beds National Monument (FFB) and Colorado Parks and Wildlife (Dome Rock SWA). These sites are characterized by a mixture of big sagebrush (Artemisia tridentata), rabbitbrush (Chrysothamnus viscidiflorus), prairie grasses and forbs (fringed sagebrush A. frigida) in a matrix of pasture, pine and spruce-fir forests. Sites were sometimes grazed by cattle or sheep at low densities, and all non-baited sites were dusted with deltamethrin to control fleas (and plague). All properties except FFB were open to recreational shooting, but shooting was not prevalent and signage discouraged shooting in vaccine project areas. The study area is within the known range of plague with plague epizootics occurring near the study colonies in 2010 (Tripp et al. unpublished data). Visual counts of Gunnison’s prairie dogs on colonies in Colorado averaged 6.1 prairie dogs/acre in 2010. Bird and vegetation surveys were conducted on 13 GUNN colonies and four WOOD colonies. There were nine vaccine project areas receiving vaccine, placebo, and dusting treatments where raptors, predators, and passerine nests (GUNN only) were surveyed. Because of their small size, entire colonies were treated. 1) The 33 acre Miller Ranch (MR) and 26 acre Kenny Moore (KM) colonies north of Gunnison received vaccine and control treatments, and the 62 acre Power Line (PL) colony 16 km to the south was designated as the paired dusted treatment. 2) The 46 acre Cabin Creek (CC), 37 acre BLM-15 (B15), and 27 acre BLM-5 (B5) colonies southeast of Gunnison received vaccine and control treatments, and the 69 acre BLM-18 (B18) colony was designated as the paired dusted treatment. B5 has been baited in the past due to concerns that GUPD sample size would be too low in B15. All these colonies are within the same complex. 3) Two ~20 acre Florissant Fossil Beds (FFB) and one 24 acre Dome Rock (DR) colony southwest of Woodland Park (150 km east of Gunnison Basin) received vaccine and control treatments. There was no available colony to use as the paired dusted treatment. The original FFB colony (FFBc) sampled in 2013 was discarded as a treatment area due to issues with adjacent private land, and this colony was replaced with two nearby FFB colonies (FFBa and FFBb) in 2014. At the GUPD study sites, we have an additional objective of determining whether avian species associations exist; therefore, we selected off-colony sites for comparison of data from avian point counts, vegetation surveys, and raptor counts. Time and financial constraints precluded nest searching or camera use off colonies. We extended the 250 m point grid off colonies and created a doughnut-shaped region that extended 500 – 1500 m from 2012 colony boundaries. Within these doughnut regions, each year we randomly chose new grids of nine points (3 x 3) to serve as off-colony study areas on public lands. Some grids had fewer than nine points due to land ownership boundaries. For each colony, we surveyed at least two off-colony sites. Some off-colony sites were located in sagebrush, but others were located in forested areas, especially in areas where forest was the dominant cover type (FFB, DR, and USFS property in the Gunnison Basin). Avian Point Counts Each point was surveyed once in May – June 2014. At BTPD study sites, a 250 m grid of points had already been established and surveyed by RMBO from 2006−2013. At GUPD study sites, we created a 250 m grid of points. Within each off-colony grid, we randomly chose a minimum of four points to survey. Two off-colony grids were randomly chosen to pair with each colony, so that we surveyed a 5 �minimum of eight off-colony points per paired colony. For larger colonies containing more than eight survey points, we completed as many point counts off colony as we did on colony. Points were considered to be “on colony” if located within 100 m of the boundary and with good views of the colony. Most point counts were conducted between dawn and 10:00 and were never conducted in rain, hot temperatures (above 30°C), or high winds that made it difficult to hear birds. Regardless of time or weather, we did not conduct counts if we noticed that bird activity (especially singing and calling) was dropping off. We conducted 6-min. point counts, recording each bird’s species, horizontal (radial) distance, sex (if known), use of the prairie dog colony (yes or no), minute of detection (1 – 6), and how it was detected (visual, singing, calling, drumming, fly-over, or other). Membership in a cluster was noted, typically for male-female pairs. After completing the bird count, we recorded weather and site characteristics at each point, including time, temperature, wind speed, cloud cover, management type (typically cattle grazing, sometimes dusting) and whether it was current, from this season or last season, and the presence of excessive noise, roads (primary or secondary), and cliffs or rock outcroppings within 100 m. Within a 50 m radius, we recorded characteristics of tall nesting and perching substrate, including percent cover, height, and dominant species for overstory plants ≥ 3 m and shrubs > 30 cm but < 3 m. Within a 5 m radius, we recorded characteristics of ground cover, including percent cover of grasses (including sedges and rushes), forbs, bare ground, cactus, rock, scat, shrubs, other cover such as lichen, and exotic species. We also recorded the mean height of forbs, dominant exotic species, and the mean height and species of the dominant two grasses. We used the point count protocol designed for Integrated Monitoring of Bird Conservation Regions (IMBCR: Hanni et al. 2012), except that we conducted bird surveys prior to vegetation surveys. This helped to ensure that birds displaced by the observer, including those located at the point itself, were recorded. We also altered IMBCR vegetation survey protocols slightly to make the protocol specific to low stature prairie dog colonies, shortgrass prairie, and sagebrush systems. This was designed to be a quick, visual assessment; a more involved protocol using a Daubenmire frame and robel pole was used on transects and at nests. Vegetation Transects In addition to a visual assessment of vegetation at points, we sampled vegetation on transects and at nests. We completed two transects on vaccine project colonies and paired off-colony sites and at least one transect on non-project sites (both on and off colonies). To locate each transect, we randomly chose a start and an end point from those used in avian point counts. From the start point, we walked along the bearing toward the end point for 240 m, stopping every 20 m to collect vegetation data for a total of 13 points per transect, except on very small colonies. Transect data were collected during the growing season. At each stop point, we recorded the presence of active or inactive prairie dog burrows within 10 m, ground cover, dominant species, and visual obstruction. Percent ground cover was measured within a 50 cm square Daubenmire frame. We recorded the percent bare ground, rock, litter, scat, grass (including sedges and rushes), forb, shrub, cactus, exotic, and other cover. We also recorded the dominant species for each plant category present in the frame. Visual obstruction data were recorded by holding a robel pole at the stop point on the transect and making observations from a distance of 4 m in each cardinal direction with eye level at 1 m. The observer then noted which portions of the 122 cm (4 ft) pole were obstructed by vegetation, identified the plant species obstructing the pole, and noted whether the pole was substantially obstructed or was covered by just a wisp of vegetation (typically a blade of grass). We estimated the height of any structures taller than the pole (a few trees at GUPD off-colony sites). Raptor Counts 6 �Raptors were sometimes sighted during avian point counts, but point counts are not an ideal method for detecting raptors or other uncommon species. Therefore, we chose 1 – 3 locations per vaccine project area (on- and off-colony), positioning observers so that the entire treatment area could be viewed simultaneously. We conducted 30-min. raptor counts, recording each bird’s species, horizontal (radial) distance, sex (if known), time of entry and exit, and behavior (high soar, low soar, directed flight, hover, dive, call, perch, or nest). Membership in a cluster was noted, typically for male-female pairs. This produces a time metric for assessing raptor use of treatment areas and colonies. At the start and end of the count, we recorded weather characteristics, including time, temperature, wind speed, and cloud cover. Raptor counts were conducted after 9:30 from January to March (wintering) and April to August (breeding) and were never conducted in rain. As a supplement to the formal raptor counts, we recorded incidental raptor observations. Nest Searching and Monitoring We searched for MOPL and BUOW nests throughout the study area through visual observation of adult birds, typically in the morning and not in rainy conditions or high winds. MOPL nest in scrapes on the ground in areas with a relatively high bare ground component. BUOW nest in prairie dog burrows, often near colony edges and in burrows with low to moderately-sized mounds. Because these species react more to humans on foot than to vehicles, we conducted surveys from a vehicle whenever possible. When MOPL were detected, we observed the bird, sometimes backing away from the site, and waited for the bird to sit down on a nest. When BUOW were detected, we searched for nests in the vicinity of their perching location; typically males perched conspicuously near the nest burrow during the day. We searched for passerine nests on colonies in vaccine project areas only, because the rope dragging technique (Yackel Adams 2000) that works best for secretive birds and camouflaged nests is time-consuming. Most prairie passerines nest in woven cups on the ground, while shrubland passerines typically place their nests on branches or under shrubs. Each search area was surveyed at least twice. At GUPD sites, each entire colony was searched via rope dragging. At BTPD sites, we searched 40 acre plots, which was the size of the smallest treatment area and an area that could be searched by two people in a half day. For those sites, we nest searched at two plots. We located one plot non-randomly, centered on the area with greatest density of 2013 nest locations. The second plot was randomly located with one corner on a point from the larger grid. Passerine nest searching was typically done after 9:30, because grassland birds are more likely to be in attendance at their nests during the heat of the day, and not in rainy conditions or high winds. At BTPD sites, we dragged a 100 ft (30 m), ½ inch gauge rope with two people at each end, watching for flushing birds in the area ahead of and under the rope. Because the GUPD sites contained a much higher shrub component, it was not possible to drag a heavy rope without continuously getting snagged on vegetation; therefore, we used two different ¼ inch gauge ropes, depending on the shrub component at the site. One was 10 m and the other was 23 m long, held above the vegetation, with heavy hex nuts suspended from smaller ropes to disturb vegetation slightly and flush birds. Additional nests were found during point counts and nest monitoring. When we were unable to find a nest during the initial search, we marked the GPS location and returned at a later date. We likely found the majority of BUOW nests on the landscape using this method (Conrey 2010), but a smaller and unknown proportion of MOPL and passerine nests were found. MOPL and passerine nests were defined as structures containing at least one egg. Because BUOW nests are underground, we defined their nests as burrows with shredded manure present at the entrance (Garcia and Conway 2009), with feathers, regurgitated pellets, and prey remains providing additional evidence of a nest attempt. At the time of nest discovery, we recorded the same weather information that we recorded at points. We also did a rapid visual assessment of vegetation, with more detailed data collected at nest completion when overheating of eggs and nest abandonment was no longer 7 �a concern. We described the nest structure and vegetation immediately around the nest and estimated vegetation height, percent bare ground within a 1 m radius, and whether all, ≥ 50%, < 50%, or none of the nest could be seen from vantage points 5 m to the north and south. BUOW nests were marked with brightly painted wooden stakes placed 10 m north of the nest burrow. MOPL and passerine nests were marked with two small unpainted wooden stakes placed 5 m north and south of the burrow. We collected any pellets that we observed near BUOW nests for possible future dietary analysis. MOPL and BUOW nests, with relatively long incubation and nestling periods, respectively, were monitored at least once per week. Passerine nests were monitored every 2 – 3 days. Starting with the first visit when the nest was discovered, we recorded the time, any management activities (such as cattle grazing), age of eggs and juveniles, and number of eggs, juveniles, and adults present. MOPL and passerine eggs were aged by floating: eggs closer to hatch float higher in the water column. Juveniles were aged according to keys for BUOW (Priest 1997) and LARB (Yackel Adams Unpub. data), and we created our own photographic keys for all other species based on our 2013 observations. Passerine nests were considered successful if at least one fully-feathered juvenile left the nest. Evidence of success included juveniles outside the nest cup, mutes at the edge of the nest, and/or displaying and calling adults, coupled with an intact nest and appropriate timing based on nest age. MOPL nests were considered successful if at least one egg hatched, because their chicks are precocial and can leave the nest area within hours of hatch. Evidence of MOPL success included pip chips, coupled with an intact nest and appropriate timing based on nest age. BUOW nests were considered successful when at least one fledgling aged ≥ 35 days was observed (Thomsen 1971, Davies and Restani 2006, Conrey 2010), because they leave the nest burrow (but may return to it many times) at 10 – 14 days and well before flight or independence are attained. Failed nests were destroyed, contained broken eggs, and/or had eggs or nestlings that disappeared before their expected hatch (MOPL) or fledge (BUOW and passerines) date. For analysis purposes, nests with unknown fate will have their histories truncated back to the last date when the nest was active and will be coded as successful at that time. At nest completion, we recorded the same vegetation data that were collected at points along vegetation transects: presence of prairie dog burrows within 10 m, percent ground cover, and visual obstruction. We placed the Daubenmire frame at 1 m in each cardinal direction and observed the robel pole (placed at the nest) from 4 m in each cardinal direction, producing four readings of each metric. For ground and shrub nests, we also recorded the height of the nest cup above (or below) ground and the plant species or structure type (such as cow paddy) in which (or adjacent to which) the nest was located. Camera Traps Remote cameras (Reconyx Hyperfire Covert IR model PC800) were placed in each vaccine project area to document use by mammalian predators and other wildlife. We increased the number of remote cameras on BTPD colonies in 2014 to better sample their larger size relative to GUPD colonies. We used one camera per small site and 2 – 3 cameras at larger sites. These cameras take photos when triggered by motion from an object that is warmer than ambient temperature. Camera locations were selected to maximize the potential for detections of mammalian predators without the use of baits or lures, which might have acted as attractants and altered the sampling region beyond treatment areas and prairie dog colonies. Cameras were positioned along game trails, aimed at coyote height, and tested before they were armed. Cameras targeted water sources, fence lines, and other landscape features. We set the cameras to take three photos when triggered, with no quiet period between photos. Cameras were deployed in April – May 2014, and several more were added in mid-June. The original nine cameras deployed in May – June of 2013 on BTPD sites were left in place and operated year-round. We checked batteries and SD cards 2 – 6 times per year. Cameras were removed from GUPD sites in September, prior to the advent of winter weather and GUPD hibernation. As a supplement to the 8 �camera data, we recorded incidental observations of predators such as coyotes, foxes, badgers, and rattlesnakes. Databases We designed a database for this project using Microsoft SQL Server 2012 with the data entry interface in Microsoft Access 2007. The database was designed by R. Conrey and D. Conrey, a professional database developer who volunteered his time, to run on the Fort Collins CPW research server. This allows multiple users to simultaneously access the database, while providing for daily backups and improved data security. Users can access a master list of codes for vegetation species, bird species, management types, observers, sites, colonies, and points that if changed, will update throughout the database. Users also access data entry forms for each data type described above, except for photo data. The Reconyx photo database (Newkirk 2014) catalogs photos and stores the metadata associated with them, displaying (but not storing) the photos themselves within Access forms. Users identify species in photos using stand-alone modules which are then loaded into the database. Each photo is examined by at least two observers, and a referee (R. Conrey or field crew leader) resolves any conflicts in IDs. Identification is ongoing, with 2013 photo processing nearly complete. Data Analysis Thus far, all data have been entered and summarized, but data proofing is ongoing and statistical analyses have not yet been completed. We have completed bird and vegetation species lists and summarized ground cover and visual obstruction data collected at transects. For raptor counts, we have calculated a proportional use index, dividing the usage minutes by the total survey minutes for each species and site. Apparent nest success has been calculated as the proportion of nests fledging (BUOW and passerines) or hatching (MOPL) at least one chick. Data will eventually be analyzed using Program DISTANCE to estimate density from point counts, Program MARK to estimate nest survival and occupancy from point counts and camera data, and R, or a similar statistical package, for other data types. RESULTS During the 2014 season, we collected the first year of post-treatment data, following initial bait drop in late summer and fall of 2013. Since fall 2013, plague epizootics have occurred on one GUPD colony and across half the BTPD study area at four treatment sites and five additional colonies. At two of the three pairs of BTPD treatment sites, plague occurred at both the baited sites, meaning that both the control and vaccine areas experienced outbreaks; however, these epizootics started in fall 2013 shortly after vaccine drop and plague was likely present in the system prior to vaccination. All of these areas had small numbers of prairie dogs during May – August 2014, with increasing abundance in the regions that first experienced plague epizootics during 2013. In September 2014, black-footed ferrets were released in two BTPD study colonies (the Roman and Brannigan colonies in the northwest part of Soapstone Prairie Natural Area) that were 3.9 km from the nearest treatment site and 9.2 km from the nearest plague epizootic. In October 2014, additional ferrets were released within the dusted part of a treatment colony (the Ferret Center colony in the southeastern part of Meadow Springs Ranch), which is 0.5 km from the nearest paired baited site and plague epizootic. 2014 was also a very wet year at the BTPD site. Normally dry playas and streambeds were full, grasses were tall and formed seedheads in June, and a different suite of species was observed, including a large flush of growth in species normally absent or a minor component of the prairie, such as six weeks fescue and wooly plantain. In response, we saw a huge influx of lark buntings; we had two nests in 2013 and 69 nests in 2014, making lark buntings the most abundant breeders at our sites. McCown’s longspurs declined, while the number of burrowing owl nests doubled. 9 �We conducted 718 avian point counts in 2014 and detected 95 bird species during the breeding season (Table 1), for a total of 113 species detected over two years (one was detected only during raptor counts: App. 1). The most common birds detected were lark bunting, horned lark, western meadowlark, and McCown’s longspur at BTPD sites and Brewer’s sparrow, vesper sparrow, horned lark, sage thrasher, raven, and western meadowlark at GUPD sites, in that order. We have detected 65 species on BTPD colonies, 71 species on GUPD colonies, and 73 species off GUPD colonies. Of the species at GUPD sites, 55 were found both on and off colonies, while 16 were unique to colonies and 17 were unique to offcolony sites. Occupancy and density analyses will not be run until after the 2015 season. We characterized vegetation at 718 point count locations, 225 nests, and ~1400 stop points along 111 transects (Table 1) in 2014. Over two years, we have documented 175 plant species (Table 1, App. 2) or groups (moss, lichen, etc.), half of which were unique to either BTPD or GUPD sites. Most of the plants at GUPD sites occurred at least once both on and off colonies, but 20% of the plant species detected on GUPD colonies have thus far not been detected off colonies. Vegetation transect data (Table 2) suggested that BTPD colonies were dominated by grass (mainly blue grama) and bare ground, with triple the grass coverage of GUPD sites and ~20% more grass than in 2013 when precipitation was closer to the average (Conrey 2013). We had less litter and double the forb coverage at BTPD sites in 2014 as in 2013. GUPD sites had a more even distribution among cover types. Bare ground was the most common cover type at the GUNN sites, both on and off colonies, and GUPD colonies had 14% more bare ground than off-colony areas. At the GUPD sites, shrub cover (mainly rabbitbrush and big sagebrush) was much higher in GUNN while litter and grass cover was much higher in WOOD. On- and off-colony GUPD locations showed larger differences than they did in 2013, especially for bare ground. Exotic cover was 3 – 6% higher in 2014 than in 2013 at GUPD sites but decreased at BTPD sites. Dominant plant species of each type are listed in Table 3, with a complete plant species list in App. 2. Visual obstruction by vegetation was higher at GUPD sites (7 – 16% of the robel pole), which were dominated by rabbitbrush, sagebrush, and taller grasses and forbs, than at BTPD sites (5% of the pole), which were dominated by shortgrasses, but the difference in vegetation heights was much less than in 2013. Approximately 6 cm were obstructed on BTPD colonies, with grasses responsible for most of the obstruction. Differences between on- and off-colony locations at GUPD sites were larger in 2014 than in 2013, with taller vegetation (by 6 cm) off colonies. There tended to be gaps in the vegetation at GUPD sites caused by different structural types: grasses, forbs and litter close to the ground, and branches of shrubs higher up. We conducted raptor counts at 35 locations for a total of 6420 minutes (Table 1) and detected 13 species (Table 4) in 2014. Burrowing owls, northern harriers, ferruginous hawks, prairie falcons, and rough-legged hawks have been detected only on prairie dog colonies (the latter three, only on BTPD colonies, where no off-colony counts were performed). During raptor counts, BUOW were detected only on BTPD colonies, but one individual was again detected during nest searches on a GUPD colony (he did not nest in either year). Swainson’s hawks and turkey vultures were more common on BTPD colonies that at GUPD sites. In contrast, ravens, golden eagles, and red-tailed hawks were more common at GUPD sites. It is unclear whether golden eagles and American kestrels have a higher level of use on- versus offGUPD colonies, as a strong relationship existed in one year but not the other: golden eagles showed the strongest preference for prairie dog colonies during our surveys in 2013 (Table 4). During winter counts on BTPD colonies, Swainson’s hawks and turkey vultures were replaced by rough-legged hawks, and ferruginous hawks became more common than they were during summer. We monitored 225 nests of 15 bird species in 2014, a large increase over the 68 nests of 11 bird species found in 2013 (Conrey 2013; Tables 1, 5). The largest portion of the increase was explained by lark bunting nests on BTPD colonies and Brewer’s sparrow nests at GUPD colonies. Following plague epizootics, burrowing owl nest numbers increased in those colonies. Nest success was average for most 10 �species, with localized episodes of nest destruction on BTPD colonies due to hail and flooding: overall apparent nest success was 47% in 2014 compared to 53% in 2013 (Table 5). We deployed 18 cameras on BTPD colonies and 10 cameras on GUPD colonies (Table 1). BTPD cameras are deployed year-round, because BTPDs do not hibernate, while GUPD cameras were deployed from May into September. We now have 433,272 photos, and most of the 2013 photos have been processed. As expected, many photos recorded prairie dogs, cows, pronghorn, and rabbits. We have documented coyote (Canis latrans), badger (Taxidea taxus), swift fox (Vulpes velox), red fox (V. vulpes), striped skunk (Mephitis mephitis), bobcat (Lynx rufus), and raccoon (Procyon lotor) use of vaccine project areas (listed by number of occurrences in our photos). Overall 4-week detection rates per site (naïve occupancy estimates) for the three most common carnivores from May 2013 – April 2014 ranged from 7% for swift fox on BTPD colonies to 54% for coyotes on GUPD colonies (Table 6). DISCUSSION This was the second year of this study but only the first year of post-treatment data collection on the project, as baits were first dropped in late summer and fall 2013. It will likely take additional years of monitoring to detect potential changes in the avian community caused by plague management, as treated colonies no longer experience extinction events. Since fall 2013, plague epizootics have occurred on one GUPD colony and across half the BTPD study area, with prairie dogs beginning to increase again in all colonies except for one BTPD colony where the epizootic began only recently. At two of the three pairs of BTPD treatment sites, plague occurred at both the baited sites, meaning that both the control and vaccine areas experienced outbreaks; however, these epizootics started in fall 2013 shortly after vaccine drop and plague was likely present in the system prior to vaccination. Prairie dog researchers are interested in whether prairie dogs persisted in treated areas during the outbreak, and whether prairie dog numbers recover more quickly in treated areas. The presence of plague in our study system will contribute to our ability to detect changes in avian communities caused by plague management. This is particularly true for BTPD colonies; additional epizootics beyond the single outbreak at Dome Rock State Wildlife Area will likely be required for us to detect differences on GUPD colonies. In September and October 2014, black-footed ferrets were released in three BTPD study colonies, one of which was only 0.5 km from the nearest baited site, which experienced a plague epizootic in fall 2013. The two release sites (not included in the prairie dog vaccine study) in Soapstone Prairie Natural Area were both dusted and baited with vaccine. Released ferrets have been vaccinated against plague and distemper, and are expected to move quickly into baited sites within the study area. They will likely predate on adult birds and nests, but as 90% of their prey base is typically comprised of prairie dogs (Clark 1986), the overall effect should be minimal. Two of the releases were in regions ~ 4 km from current plague outbreaks, and eight of the 30 released individuals were detected during spotlighting surveys 30 days after the release. It seems likely that at least some of these individuals will survive into the 2015 season, but the fate of the ferrets released into a dusted region 0.5 km from a recent outbreak is less certain. It is unknown whether ferrets will breed in 2015, and young ferrets must be captured to be vaccinated, as they are not expected to consume vaccine baits designed for prairie dogs. 2014 was a much wetter year than 2013 at the BTPD site. Normally dry playas and streambeds were full, grasses were tall and formed seedheads in June, and a different suite of species was observed, including a large flush of growth in species normally absent or a minor component of the prairie. In response, we saw a huge influx of lark buntings; they were the most common species detected in point counts in 2014, and nest numbers increased from two nests in 2013 to 69 nests in 2014, making lark buntings the most abundant breeders at our sites. In contrast, McCown’s longspur numbers declined, while the number of burrowing owl nests doubled. These changes may also be related to concurrent declines in prairie dogs associated with plague epizootics. Vegetation was no longer being clipped, and 11 �extinct areas had much more lush vegetation than usual. Burrowing owl nest numbers increased only in areas that experienced plague epizootics and had persisting (or recovering) prairie dog populations. Burrowing owls also increased in response to plague events (and prairie dog recolonization) in a previous study in northern Colorado (Conrey 2010). Thus far, data from vegetation surveys have identified differences between on- and off-colony areas for GUPD sites, but bird data will require further analysis and a third year of data before we draw conclusions regarding the uniqueness of the avian community on GUPD colonies. Over two seasons, 16 bird species have been detected only on GUPD colonies, not in off-colony areas, but we have not yet modeled occupancy or density of these or other species. Raptor count data are ambiguous; the additional sample size provided by a third study season and new randomly located off-colony sites will hopefully provide enough power to make comparisons. BTPD increase bare ground and alter plant species composition and nutrient cycling rates (Whicker and Detling 1988; Johnson-Nistler et al. 2004); effects of GUPD on vegetation are not well-studied. GUPD do provide refugia and nests for BUOW, kit fox, and small mammals (Miller et al. 1994, Meaney et al. 2006). However, their impact appears less dramatic than that of BTPD (Grant-Hoffman and Detling 2006), perhaps because of differences in habitat, less above-ground activity, little clipping of vegetation, and lower burrow densities (Seglund and Schnurr 2010). We made several improvements to protocols in 2014. First, we began winter raptor counts on BTPD colonies in order to sample wintering species such as rough-legged hawks; FEHA associations with BTPD colonies have also mainly been documented in fall and winter (Smith and Lomolino 2004). We also deployed our cameras on BTPD colonies year-round to sample mammalian carnivores during winter; we did not do any sampling on GUPD colonies after September, as we had no personnel there and GUPD hibernate. Second, we refined nest searching protocols, which improved our success in finding nests. Third, by doing only one round of point counts and hiring two additional temporary staff in 2014, we were able to begin nest searching and raptor counts earlier in the field season and increase sample size. Fourth, we added cameras to more adequately sample large BTPD colonies and hopefully improve occupancy estimation for mammalian carnivores. Another protocol improvement planned for next year should help us to find more MOPL nests; we have gotten permission to use ATVs to search for MOPL nests. We will use the same rope-dragging technique used successfully by Rocky Mountain Bird Observatory to find MOPL nests. However, our sample sizes may still be limited by the apparent decline in MOPL across the region that has been observed in the nearby Pawnee National Grasslands. 2015 will be the third and final study season for the plague vaccine project. After that, the intensity of research and monitoring of bird communities on prairie dog colonies will depend on the presence of plague in the system, presence of black-footed ferrets, available resources, and scale of vaccine use. If the vaccine study suggests that this is a successful approach to plague management, then the vaccine may be more broadly used as a management tool. This would allow us to change the spatial scale of our research, perhaps including larger, more suitable areas for focal species such as BUOW and MOPL. If more MOPL can be located within treated areas, we plan to collaborate with other MOPL researchers across multiple sites to study changes in MOPL dynamics as a response to longer term plague management. Our three years of research will also identify other appropriate focal species, determine whether or not we should sample on GUPD colonies, and lead to a general monitoring protocol for species associated with prairie dog colonies. LITERATURE CITED Antolin, M. F., P. Gober, B. Luce, D. E. Biggins, W. E. V. Pelt, D. B. Seery, M. Lockhart, and M. Ball. 2002. The influence of sylvatic plague on North American wildlife at the landscape level, with 12 �special emphasis on black-footed ferret and prairie dog conservation. Transactions of the 67th North American Wildlife and Natural Resources Conference 67: 104–127. Augustine, D. J., S. J. 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Ph.D. Dissertation, Colorado State University, Fort Collins, Colorado. Conrey, R. Y. 2013. Avian response to plague management on Colorado prairie dog colonies. Wildlife Research Report. Colorado Parks and Wildlife, Fort Collins, Colorado. 25 pg. Davies, J. M. and M. Restani. 2006. Survival and movements of juvenile burrowing owls during the postfledging period. Condor 108:282–291. Dicke, W. M. 1926. Plague in California 1900 – 1925. Proceedings of the 41st Annual Meeting and Conference of State Provincial Health Authority of North America, Atlantic City, New Jersey. Ecke, D. H. and C. W. Johnson. 1952. Plague in Colorado and Texas. Part I. Plague in Colorado. Public Health Monograph No. 6. U. S. Government Printing Office, Washington D.C. Garcia, V. and C. J. Conway. 2009. What constitutes a nesting attempt? Variation in criteria causes bias and hinders comparisons across studies. Auk 126:31–40. Grant-Hoffman, M. N. and J. K. Detling. 2006. 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Smith. 2004. Terrestrial vertebrate communities of black-tailed prairie dog (Cynomys ludovicianus) towns. Biological Conservation 115:89–100. Magle, S. B., B. T. McClintock, D. W. Tripp, G. C. White, M. F. Antolin, and K. R. Crooks. 2007. Mark– resight methodology for estimating population densities for prairie dogs. Journal of Wildlife Management 71: 2067–2073. Meaney, C. A., M. Reed-Eckert, and G. P. Beauvais. 2006. Kit Fox (Vulpes macrotis): a technical conservation assessment. USDA Forest Service, Rocky Mountain Region. http://www.fs.fed.us/r2/projects/scp/assessments/kitfox.pdf [Accessed 2013]. Miller, R. F., T. J. Svejcar, and N. E. West. 1994. Implications of livestock grazing in the Intermountain sagebrush region: plant composition. Pages 101–146 in M. Vavra, W. A. Laycock, and R. D. Pieper, editors. Ecological implications of livestock herbivory in the west. Society for Range Management, Denver, Colorado. 13 �Newkirk, E. S. 2014. CPW Photo Database. Colorado Parks and Wildlife, Fort Collins, Colorado, USA. http://wildlife.state.co.us/Research/Mammal/Pages/MammalsWildlifeResearch.aspx Paine, R. T. 1969. A note on trophic complexity and community stability. American Naturalist 103:91– 93. Priest, J. E. 1997. Age identification of nestling burrowing owls. Journal of Raptor Research Report 9:125–127. Rocke, T. E., N. Pussini, S. R. Smith, J. Williamson, B. Powell, and J. E. Osorio. 2010. Consumption of baits containing raccoon pox-based plague vaccines protects black-tailed prairie dogs (Cynomys ludovicianus). Vector-Borne and Zoonotic Diseases 10: 53–58. Seglund, A. E. and P. M Schnurr. 2010. Colorado Gunnison’s and white-tailed prairie dog conservation strategy. Colorado Division of Wildlife, Denver, Colorado. Sidle, J. G., M. Ball, T. Byer, J. J. Chynoweth, G. Foli, R. Hodorff, G. Moravek, R. Peterson, and D. N. Svingen. 2001. Occurrence of burrowing owls in black-tailed prairie dog colonies on Great Plains National Grasslands. 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Journal of Wildlife Management 72:1001–1006. Tripp, D. W. and T. E. Rocke. 2012. Study plan: Preliminary field trials to evaluate the safety and efficacy of an sylvatic plague vaccine for prairie dogs. Whicker, A. D. and J. K. Detling. 1988. Ecological consequences of prairie dog disturbances: prairie dogs alter grassland patch structure, nutrient cycling, and feeding-site selection by other herbivores. BioScience 38:778–785. Yackel Adams, A. A. 2000. Training notes for grassland bird project. Unpublished report. U.S. Geological Survey Fort Collins Science Center, Fort Collins, Colorado. 8 pp. Yackel Adams, A. A. Unpub data. Lark bunting nestling aging guide. U.S. Geological Survey Fort Collins Science Center, Fort Collins, Colorado. 2 pg. 14 �TABLES Point counts Point count bird species Vegetation transects Vegetation species Raptor count locations Raptor count minutes Nest searching area (acres) Nests Remote cameras Remote camera photos BTPD on 426 56 52 61 9 2760 680 167 18 325063 GUPD on 108 45 30 120 10 1800 300 58 10 44849 GUPD off 184 60 29 123 16 1860 N/A N/A N/A N/A TOTAL 718 95 111 175 35 6420 980 225 28 369912 Table 1. 2014 sample sizes for BTPD and GUPD sites, on and off prairie dog colonies. We did one point count, three winter raptor counts, and 5 – 9 summer raptor counts per location. We surveyed 1 – 2 vegetation transects per site. Nest search plots were surveyed at least twice. Photos listed above were taken between October 2013 and September 2014, and vegetation species include 2013 and 2014 detections. GUPD cameras were deployed during summer, and BTPD cameras were deployed year-round. BTPD % Cover Grass Bare Litter Forb Rock Scat Cactus Shrub Other on 55.9 19.4 8.7 8.3 3.2 1.5 1.4 0.6 1.1 GUNN on 10.5 46.5 9.7 7.8 6.8 2.0 0.1 13.5 3.1 GUPD GUNN WOOD WOOD off on off 16.7 25.7 21.3 32.4 22.3 8.8 12.8 18.3 41.3 6.3 14.9 8.5 10.8 9.3 13.4 1.8 2.8 2.3 0.2 0.0 0.0 15.1 1.3 2.3 4.0 5.6 2.2 Table 2. Ground cover percentages from vegetation transects conducted on BTPD and GUPD sites, on and off prairie dog colonies from June – August 2014. 15 �BTPD Type BTPD on Grass blue grama Forb scarlet globemallow Shrub winterfat Cactus plains pricklypear Other lichen Exotic russian thistle GUNN on junegrass fringed sagebrush rabbitbrush plains pricklypear lichen pinnate tansy mustard GUPD GUNN off WOOD on junegrass blue grama fringed sagebrush fringed sagebrush big sagebrush rabbitbrush plains pricklypear N/A lichen lichen timothy grass kentucky bluegrass WOOD off Poa sp. fringed sagebrush common juniper N/A lichen kentucky bluegrass Table 3. Dominant plant species detected on vegetation transects at BTPD and GUPD sites, on and off prairie dog colonies in June – August 2014. Close seconds included wooly plantain (forb at BTPD on) and ryegrass (exotic at GUNN on). Italics indicate dominant species that differs from 2013. 16 �Species American crow American kestrel burrowing owl common raven Cooper’s hawk ferruginous hawk golden eagle northern harrier prairie falcon red-tailed hawk rough-legged hawk sharp-shinned hawk Swainson's hawk turkey vulture TOTAL minutes 2013 Use Rate (%) BTPD GUPD GUPD on on off 0 0 0.32 2.37 1.25 2.54 14.10 0 0 6.47 24.33 44.60 0 0 0 0.19 0 0 2.12 4.83 0.16 0.13 0 0 0 0 0 0.83 3.17 0 0 0 0 0 0 0 8.91 0 3.33 19.49 0.50 0.32 1560 1200 630 2014 Use Rate (%) BTPD GUPD GUPD on on off 0 0 0 4.94 4.67 2.19 6.94 0 0 3.50 31.89 16.27 0 0 0.11 0.77 0 0 0.46 2.39 4.60 0.46 1.22 0 0.07 0 0 1.79 5.83 9.21 0.84 0 0 0 0.72 0.91 1.82 0.67 1.02 9.74 2.67 5.57 2760 1800 1860 Table 4. Raptor use of vaccine project areas at BTPD and GUPD sites, on and off prairie dog colonies in 2013 and 2014. Use was quantified as time spent in project areas, and use rate = use minutes/total minutes on BTPD, on GUPD, and off GUPD colonies. Data from 2014 include winter counts (Jan – March). Species Brewer's sparrow burrowing owl common raven ferruginous hawk green-tailed towhee horned lark lark bunting McCown's longspur mountain plover sage thrasher Swainson's hawk vesper sparrow western kingbird western meadowlark TOTAL BTPD GUPD 15 33 13 0 1 0 1 0 0 3 36 1 69 0 26 0 2 0 0 13 1 0 0 8 1 0 2 0 167 58 # Nests TOTAL Known Fate 48 44 13 11 1 1 1 1 3 3 37 35 69 67 26 26 2 2 13 13 1 1 8 8 1 1 2 2 225 215 Successful 26 8 1 0 3 14 22 10 1 9 0 7 0 1 102 % Success 0.59 0.73 1.00 0 1.00 0.40 0.33 0.38 0.50 0.69 0 0.88 0 0.50 0.47 Table 5. Nest numbers and fate in vaccine project areas on BTPD and GUPD colonies in 2014. 17 �Species coyote badger swift fox BTPD GUPD 0.485 0.538 0.154 0.115 0.066 0 Table 6. Naïve (minimum) occupancy estimates (do not account for probability of detection) for carnivores based on remote camera photos taken on BTPD and GUPD colonies in 2013. Occupancy was calculated per site over 4-week intervals. 18 �APPENDIX 1: BIRD SPECIES LIST Code AMCO AMCR AMGO AMGP AMKE AMRO AWPE BANS BARS BBMA BCCH BGGN BHCO BHGR BLGR BRBL BRSP BTAH BUOR BUOW CAFI CANG CASP CCLO CHSP CLNU CLSW COFL COGR COHA CONI CORA DCCO DEJU DOWO DUFL EAME EUST EVGR Common Name American coot American crow American goldfinch American golden-plover American kestrel American robin American white pelican bank swallow barn swallow black-billed magpie black-capped chickadee blue-gray gnatcatcher brown-headed cowbird black-headed grosbeak blue grosbeak Brewer's blackbird Brewer's sparrow broad-tailed hummingbird Bullock's oriole burrowing owl Cassin's finch Canada goose Cassin's sparrow chestnut-collared longspur chipping sparrow Clark's nutcracker cliff swallow Cordilleran flycatcher common grackle Cooper's hawk common nighthawk common raven double-crested cormorant dark-eyed junco downy woodpecker dusky flycatcher eastern meadowlark European starling evening grosbeak BTPD on x x x x x x x x GUPD on x x x GUPD off x x x x x x x x x x x x x x x x x x x x x x x x x x x x⁺ x x x x x x x x x x x x x x x x x x* x x x x x x x x x x 19 x x x x x x �FEHA GADW GBHE GOEA GRSP GTGR GTTO GUSG HAWO HETH HOLA HOSP HOWR KILL LARB LASP LBCU LEGO LISP LOSH MALL MCLO MGWA MOBL MOCH MODO MOPL NOFL NOHA NOMO NRWS OSFL PISI PRFA PYNU RBGU RBNU RCKI RNDU RNSA ROPI ROWR RTHA ferruginous hawk gadwall great blue heron golden eagle grasshopper sparrow great-tailed grackle green-tailed towhee Gunnison sage-grouse hairy woodpecker hermit thrush horned lark house sparrow house wren killdeer lark bunting lark sparrow long-billed curlew lesser goldfinch Lincoln's sparrow loggerhead shrike mallard McCown's longspur MacGillivray's warbler mountain bluebird mountain chickadee mourning dove mountain plover northern flicker northern harrier northern mockingbird northern rough-winged swallow olive-sided flycatcher pine siskin prairie falcon pygmy nuthatch ring-billed gull red-breasted nuthatch ruby-crowned kinglet ring-necked duck red-naped sapsucker rock pigeon rock wren red-tailed hawk x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x* x x x x x x x x x x x 20 x x x x x x x x x x x x x x x x x x x x x x x x x �RWBL SACR SAGS SAPH SATH SAVS SOSP SPTO SSHA STJA SWHA TOSO TRSW TUVU VESP VGSW WAVI WBNU WCSP WEBL WEKI WEME WESJ WETA WEWP WIPH WISA WISN WITU YEWA YRWA TOTAL red-winged blackbird sandhill crane sage sparrow Say's phoebe sage thrasher savannah sparrow song sparrow spotted towhee sharp-shinned hawk Steller's jay Swainson's hawk Townsend's solitaire tree swallow turkey vulture vesper sparrow violet-green swallow warbling vireo white-breasted nuthatch white-crowned sparrow western bluebird western kingbird western meadowlark western scrub-jay western tanager western wood-pewee Wilson’s phalarope Williamson's sapsucker Wilson's snipe wild turkey yellow warbler yellow-rumped warbler x x x x x x x x x x x x x x x x x x x x x x x x 65 x x* x x* x* x x x x* x x x x x x x x x x x x x x x x x x x x x x x x 71 x x x x x 73 Table A1. Bird species list for BTPD and GUPD sites during summer 2013−2014. These species were detected during avian point counts with several exceptions. ⁺ = detected while nest searching. * = detected during raptor counts. Rough-legged hawks (RLHA not listed) were detected only during winter raptor counts. 21 �APPENDIX 2: PLANT SPECIES LIST Code Family Grasses, Sedges, and Rushes ACHY Poaceae AGCR Poaceae ARPU Poaceae BODA Poaceae BOGR Poaceae BRIN Poaceae BRTE Poaceae CAIN Cyperaceae CALO Poaceae CARE Cyperaceae ELEL Poaceae ELGL Poaceae ELRE Poaceae ELTR Poaceae FEID Poaceae FEST Poaceae HECO Poaceae HOJU Poaceae JUBA Juncaceae KOMA Poaceae LOPE Poaceae MUHL Poaceae MUMO Poaceae NAVI Poaceae PASM Poaceae PHPR Poaceae Scientific Name Common Name Achnatherum hymenoides Agropyron cristatum Aristida purpurea Bouteloua dactyloides Bouteloua gracillis Bromus inermis Bromus tectorum Carex inops ssp. heliophila Calamovilfa longifolia Carex Spp. Elymus elymoides Elymus glaucus Elymus repens Elymus trachycaulus Festuca idahoensis Festuca Spp. Hesperostipa comata Hordeum jubatum Juncus balticus Koeleria macrantha Lolium perenne Muhlenbergia Spp. Muhlenbergia montana Nassella viridula Pascopyrum smithii Phleum pratense Indian Ricegrass Crested Wheatgrass Purple Threeawn Buffalograss Blue Grama Smooth Brome Cheatgrass Sun Sedge Prairie Sandreed Sedge Spp. (Unidentified) Squirreltail Blue Wildrye Quackgrass Slender Wheatgrass Idaho Fescue Unknown Fescue Needle & Thread Grass Foxtail Barley Baltic Rush Junegrass Annual Ryegrass Muhlenbergia Spp. Mountain Muhly Green Needlegrass Western Wheatgrass Timothy Grass 22 Exotic x x x BTPD on x x x x x x x x x x x x x x x x x x x x x x x x x x x x x GUPD GUPD on off x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x �PIMI POFE POPR RUSH SCSC SOBI SPCR VUOC Forbs ACMI ANAR ANMI ANSE ARAB ARAN ARFR ARPO ARUV ASBI ASDR BASC BEPL CALI CAMI CHBE CHEN CHGL CHLE CHWA CIAR CICA Poaceae Poaceae Poaceae Juncaceae Poaceae Poaceae Poaceae Poaceae Piptatherum micranthum Poa fendleriana Poa pratensis Juncus Spp. Schizachyrium scoparium Sorghum bicolor Sporobolus cryptandrus Vulpia octoflora Littleseed Ricegrass Muttongrass Kentucky Bluegrass Rush Spp. Little Bluestem Sorghum Sand Dropseed Six Weeks Fescue Asteraceae Apiaceae Asteraceae Primulaceae Asteraceae Rosaceae Asteraceae Papavaraceae Ericaceae Fabaceae Fabaceae Chenopodiaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Chenopodiaceae Chenopodiaceae Euphorbiaceae Chenopodiaceae Chenopodiaceae Asteraceae Asteraceae Achillea millefolium Angelica arguta Antennaria microphylla Androsace septentrionalis Artemisia absinthium Argentina anserina Artemisia frigida Argemone polyanthemos Arctostaphylos uva-ursi Astragalus bisulcatus Astragalus drummondii Bassia scoparia Besseya plantaginea Castilleja linariifolia Castilleja miniata Chenopodium berlandieri Chenopodium Spp. Chamaesyce glyptosperma Chenopodium leptophyllum Chenopodium watsonii Cirsium arvense Cirsium canescens Common Yarrow White Angelica Littleleaf Pussytoes Pygmy-flower Rock-Jasmine Absinth Wormwood Silver Weed Fringed Sagebrush Crested/ Annual Pricklepoppy Bearberry (Kinnikinnick) Two-grooved Milkvetch Drummond's Milkvetch Kochia/ Mexican Fireweed White River Coral Drops Narrowleaf Paintbrush Scarlet Paintbrush Netseed Lambsquarter Goosefoot Spp. (Unidentified) Small Ribseed Sandmat Narrowleaf Goosefoot Watson's Goosefoot Canada Thistle Prairie/ Plains/ Creamy Thistle 23 x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x �CIUN CRTH DESO ERCA ERCE ERCO ERFL ERLO EROV ERRA ERSP ERST ERUMA ERUMM FRVE FRSP GABO GECA GETR GRSQ GUSA HAFL HEPA HEUC HEVI IRMI LAOC LEDE LEVI LEVU LILE Asteraceae Boraginaceae Brassicaceae Asteraceae Polygonaceae Asteraceae Asteraceae Polygonaceae Polygonaceae Polygonaceae Asteraceae Asteraceae Polygonaceae Polygonaceae Rosaceae Gentianaceae Rubiaceae Geraniaceae Rosaceae Asteraceae Asteraceae Boraginaceae Saxifragaceae Saxifragaceae Asteraceae Iridaceae Boraginaceae Brassicaceae Brassicaceae Asteraceae Linaceae Cirsium undulatum Cryptantha thyrsiflora Descurainia sophia Erigeron canus Eriogonum cernuum Erigeron compositus Erigeron flagellaris Eriogonum lonchophyllum Eriogonum ovalifolium Eriogonum racemosum Erigeron speciosus Erigeron strigosus Eriogonum umbellatum v. aureum Eriogonum umbellatum v. majus Fragaria vesca Frasera speciosa Galium boreale Geranium caespitosum Geum triflorum Grindelia squarrosa Gutierrezia sarothrae Hackelia floribunda Heuchera parvifolia Heuchera Spp. Heterotheca villosa Iris missouriensis Lappula occidentalis Lepidium densiflorum Lepidium virginicum Leucanthemum vulgare Linum lewisii Wavyleaf Thistle Calcareous Cryptantha Pinnate Tansy Mustard Hoary Fleabane Nodding Buckwheat Cutleaf Daisy Trailing Fleabane Spearleaf Buckwheat Cushion Buckwheat Redroot Buckwheat Showy Fleabane Prairie Fleabane Sulphur Buckwheat Creamy Buckwheat Woodland Strawberry Monument Plant Bedstraw Pineywoods Geranium Old Man's Whiskers Curlycup Gumweed Broom Snakeweed Many-Flowered Stickseed Littleleaf Alumroot Alumroot Spp. (Unidentified) Hairy False Golden Aster Rocky Mountain Iris Western Sticktight/ Stickweed Common Pepperweed Virginia Pepperweed Ox Eye Daisy Blue Flax 24 x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x �LIPU LUAR LUWY LYJU MAPI MARA MARE MATA MEAR MEHU MELA OECO ORLU OXLA OXSE PEBA PECR PHHO PHLO PIOP PLMA PLPA POGR POOL POPE PSLA RHRO SATR SAXI SEIN SELA Asteraceae Fabaceae Fabaceae Asteraceae Asteraceae Liliaceae Berberidaceae Asteraceae Lamiaceae Boraginaceae Boraginaceae Onagraceae Scrophulariaceae Fabaceae Fabaceae Scrophulariaceae Scrophulariaceae Polemoniaceae Polemoniaceae Asteraceae Plantaginaceae Plantaginaceae Rosaceae Portulacaceae Polygonaceae Fabaceae Crassulaceae Chenopodiaceae Saxifragaceae Asteraceae Crassulaceae Liatris punctata Lupinus argenteus Lupinus wyethii Lygodesmia juncea Machaeranthera pinnatifida Maianthemum racemosum Mahonia repens Machaeranthera tanacetifolia Mentha arvensis Mertensia humilis Mertensia lanceolata Oenothera coronopifolia Orthocarpus luteus Oxytropis lambertii Oxytropis sericea Penstemon barbatus Penstemon crandallii Phlox hoodii Phlox longifolia Picradeniopsis oppositifolia Plantago major Plantago patagonica Potentilla gracilis Portulaca oleracea Polygonum persicaria Psoralidium lanceolatum Rhodiola rosea Salsola tragus Saxifraga Spp. Senecio integerrimus Sedum lanceolatum Gayfeather/ Dotted Blazing Star Silvery Lupine Wyeth's Lupine Skeletonweed/ Rush Skeleton Plant Lacy Tansyaster Feathery False Solomon's Seal Oregon Grape Tanseyleaf Tansyaster Wild Mint Mountain Bluebells Prairie Bluebells Crownleaf Evening Primrose Yellow Owl's Clover Lambert Crazyweed White Locoweed Beardlip Beardtongue Crandall's Beardtongue Spiny Phlox Longleaf Phlox Opposite Leaf Bahia Common Plantain Woolly Plantain Slender Cinquefoil Common Purslane/ Little Hogweed Spotted Ladysthumb Lemon Scurfpea King's Crown Russian Thistle / Tumbleweed Saxifrage Spp. (Unidentified) Lambstongue Ragwort Spearleaf Stonecrop 25 x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x �SENE SOMI SOTR SPCO SPFA TAOF TAVU THAR THRH TRHY TRPR URDI VIAM VICI WYAM Shrubs ACGL AMAL ARTR ATCA CEMO CHVI DAFR EREF ERNA JUCO JUSC KRLA PRAM PRVI PUTR Asteraceae Asteraceae Solanaceae Malvaceae Euphorbiaceae Asteraceae Asteraceae Brassicaceae Fabaceae Fabaceae Fabaceae Urticaceae Fabaceae Fabaceae Asteraceae Senecio Spp. Solidago missouriensis Solanum triflorum Sphaeralcea coccinea Taraxacum officinale Tanacetum vulgare Thlaspi arvense Thermopsis rhombifolia Trifolium hybridum Trifolium pratense Urtica gracilis Vicia americana Vicia Spp. Wyethia amplexicaulis Groundsel Spp. (Unidentified) Missouri Goldenrod Cutleaf Nightshade Scarlet Globemallow Spurge (Unidentified) Dandelion Common Tansy Field Pennycress Golden Pea Alsike clover Red Clover Stinging Nettle American Vetch Vetch Spp. (Unidentified) Mule-ears Aceraceae Rosaceae Asteraceae Chenopodiaceae Rosaceae Asteraceae Rosaceae Polygonaceae Asteraceae Cupressaceae Cupressaceae Chenopodiaceae Rosaceae Rosaceae Rosaceae Acer glabrum Amelanchier alnifolia Artemisia tridentata Atriplex canescens Cercocarpus montanus Chrysothamnus viscidiflorus Dasiphora fruticosa Eriogonum effusum Ericameria nauseosa Juniperus communis ssp. alpina Juniperus scopulorum Krascheninnikovia lanata Prunus americana Prunus virginiana Purshia tridentata Mountain Maple Serviceberry Big Sagebrush Fourwing Saltbush Mountain Mahogany Douglas Rabbitbrush Shrubby Cinquefoil Spreading Buckwheat Rubber /Grey Rabbitbrush Common Juniper Rocky Mountain Juniper Winter Fat American Plum Western Chokecherry Bitter Brush 26 x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x �RHTR Anacardiaceae RIAU Grossulariaceae RICE Grossulariaceae ROAR Rosaceae ROWO Rosaceae RUID Rosaceae SALI Salicaceae SYOC Caprifoliaceae TECA Asteraceae XANT Asteraceae YUGL Agavaceae Trees ABLA Pinaceae PIEN Pinaceae PIFL Pinaceae PIPO Pinaceae PIPU Pinaceae POAN Salicaceae PODE Salicaceae POTR Salicaceae PSME Pinaceae Cacti and Other Plants ESVI Cactaceae FUNG LICH MOSS OPFR Cactaceae OPPO Cactaceae PESI Cactaceae Total Rhus trilobata Ribes aureum Ribes cereum Rosa arkansana Rosa woodsii Rubus idaeus Salix Spp. Symphoricarpos occidentalis Tetradymia canescens Xanthium spp. Yucca glauca Skunkbrush Sumac/ Squawbrush Golden Currant Wax Currant Prairie/ Wild/ Porter Prairie Rose Woods' Rose Wild Red Raspberry Willow Spp. (Unidentified) Western Snowberry Spineless Horsebrush Cocklebur sp. Great Plains Yucca/ Soapweed Yucca Abies lasiocarpa Picea engelmannii Pinus flexilis Pinus ponderosa Picea pungens Populus angustifolia Populus deltoides ssp. wislizenii Populus tremuloides Pseudotsuga menziesii Subalpine Fir Engelmann Spruce Limber Pine Ponderosa Pine Blue Spruce Narrow-leaved Cottonwood Rio Grande Cottonwood Quaking Aspen Douglas Fir Escobaria vivipara Pincushion Cactus Fungi/ Mushroom/ Basidiomycota Lichen Moss Brittle Pricklypear Plains Pricklypear Mountain Ball Cactus Opuntia fragilis Opuntia polyacantha Pediocactus simpsonii 175 species x x x x x x x x x x x x x x x x x x 61 x x x x x x x x 22 27 x x x x x x x x x x 120 x x x x x x x x x x x x x x x 123 �Table A2. Plant species list for BTPD and GUPD sites for 2013−2014. 28 � https://cpw.cvlcollections.org/files/original/bf86e9d3437fa730175e8b43166a8759.pdf afef4eb559c99f50493194e94dd8b7a9 PDF Text Text Colorado Division of Parks and Wildlife July 2014-June 2015 WILDLIFE RESEARCH REPORT State of: Cost Center: Work Package: Task No.: Colorado 3420 1680 N/A Federal Aid Project No. N/A : : : : Division of Parks and Wildlife Avian Research Bird Conservation Avian response to plague management on Colorado prairie dog colonies Period Covered: September 1, 2014 – August 31, 2015 Author: R. Yale Conrey Principle Investigators: R. Yale Conrey, D. Tripp, J. Gammonley, CPW; A. Panjabi, E. Youngberg, Bird Conservancy of the Rockies (formerly Rocky Mountain Bird Observatory). Collaborators: Miranda Middleton (CPW), Bird Conservancy of the Rockies (formerly Rocky Mountain Bird Observatory), City of Fort Collins Natural Areas and Utilities Programs, Bureau of Land Management (Gunnison office), National Park Service Florissant Fossil Beds National Monument, and CPW wildlife managers and biologists from Areas 4, 14, and 16. All information in this report is preliminary and subject to further evaluation. Information MAY NOT BE PUBLISHED OR QUOTED without permission of the author. Manipulation of these data beyond that contained in this report is discouraged. EXTENDED ABSTRACT Range-wide declines in prairie dog (Cynomys sp.) populations have occurred, and the largest limiting factor in recent decades appears to be the high mortality and colony extirpation associated with plague (Antolin et al. 2002), caused by the bacterium Yersinia pestis. Prairie dog colonies support a diverse community of associated species, many of which are not susceptible to plague but may be indirectly affected. In order to conserve prairie dogs and species associated with their colonies, principally the black-footed ferret (Mustela nigripes), a plague vaccination program is being developed (Fig. 1a), which may also benefit a suite of species (Fig. 1b) listed in the Conservation Plan for Grassland Species in Colorado (Colorado Division of Wildlife 2003) and the Colorado Sagebrush Conservation Assessment and Strategy (Boyle and Reeder 2005). CPW is involved in a multi-state, multi-agency study of prairie dogs and associated small mammal species; the objective is to determine whether survival is enhanced by the experimental vaccine compared to use of placebo or insecticide to control fleas, an important vector of plague. They are also interested in how patterns of prairie dog abundance and occupancy change when plague epizootics occur in plots receiving different treatments. As an extension to this project, we initiated research in 2013 on the effects of plague management on avian species associated with prairie dog colonies, with particular focus on species of concern. Our main long-term objective is to determine whether areas treated to control plague differ from untreated areas in their avian communities. Shorterterm objectives are to 1) Determine how plague affects avian species and their predators associated with 1 �prairie dog colonies; 2) Determine whether avian species associations exist for colonies of Gunnison’s prairie dogs (C. gunnisoni: GUPD); most evidence for associated species comes from black-tailed prairie dogs (C. ludovicianus: BTPD); 3) Determine whether insecticidal dusting influences bird density or nest survival; 4) Evaluate the importance of covariates such as weather and cattle grazing. Study areas included BTPD colonies in north-central Colorado and GUPD colonies in westcentral Colorado. BTPD study colonies were dominated by short and mid-grasses and located in Larimer and Weld Co. adjacent to the Wyoming border, managed by the City of Fort Collins. GUPD study colonies were dominated by sagebrush mixed with other shrubs and grasses and located in the Gunnison Basin (Gunnison, Saguache, and eastern Montrose Co.) and Woodland Park area (Teller Co.), managed by the Bureau of Land Management, U.S. Forest Service, National Park Service, and CPW. The 2015 season was the third of three study seasons associated with phase 1 of this avian research project, which coincided with Wildlife Health’s 3-year efficacy trials for the plague vaccine (Fig. 1a). In Colorado, CPW Wildlife Health Program staff led by Dan Tripp surveyed colonies before and after bait distribution and conducted a mark-recapture study of prairie dogs and associated small mammal species. Treated areas were arranged in triplets with one vaccine, placebo, and dusted site per group; baited sites were assigned vaccine or placebo baits in a blind procedure. We have collected 1 year of pre-treatment avian data and 2 years of post-treatment data. We created a 250 m point grid to sample all treated and untreated prairie dog colonies on public land within the study region, and on GUPD colonies, we created a doughnut-shaped region that extended 500 – 1500 m from colony boundaries and randomly chose grids of nine points (3 x 3) to serve as off-colony study areas. Although breeding season data collection for phase 1 is complete, analyses are ongoing. Bird occupancy, density, and species composition will be estimated from point count data. Summer and winter counts of diurnal raptors and early season passive and call-playback surveys of mountain plover (Charadrius montanus) and burrowing owls (Athene cunicularia) were used to sample species that are rarely detected during point counts. On-colony nest survival rates will be estimated for passerines (Fig. 1d, e) and burrowing owls. Remote camera data will be used to estimate summer and winter on-colony occupancy rates for mammalian carnivores, including coyotes, badgers, and swift fox. Finally, we have quantified percent ground cover, visual obstruction (Fig. 1c), and species composition of vegetation at points, nests, and along randomly located transects. Since fall 2013, plague epizootics have occurred on one GUPD colony and across ~70% of the BTPD study area. In September and October 2014 and 2015, black-footed ferrets were released in three BTPD study colonies. Precipitation (Fig. 2) has varied greatly over the three years of this study, particularly on BTPD sites, from slightly dry to very wet, compared to the 30-year average. At this point, data analyses are all preliminary, with detailed analysis to follow during 2015 – 2016. From 2013 – 2015, we detected 130 bird species during the breeding season. During BTPD colony surveys, at least three bird species (Brewer’s blackbird Euphagus cyanocephalus, Brewer’s sparrow Spizella breweri, and vesper sparrow Pooecetes gramineus) appeared to have higher detection rates on active prairie dog colonies, while two species (grasshopper sparrow Ammodramus savannarum and lark bunting Calamospiza melanocorys) appeared to have higher detection rates on colonies with extinct or severely reduced prairie dog populations following plague outbreaks (Table 2). During surveys on and off GUPD colonies, seven species (Brewer’s sparrow, common raven Corvus corax, horned lark Eremophila alpestris, red-winged blackbird Agelaius phoeniceus, sage thrasher Oreoscoptes montanus, vesper sparrow, and western meadowlark Sturnella neglecta) appeared to show associations with colonies, while four species (darkeyed junco Junco hyemalis, green-tailed towhee Pipilo chlorurus, mountain chickadee Poecile gambeli, and western wood-pewee Contopus sordidulus) had higher detection rates off colonies, and many others showed no pattern (Table 3). We documented 217 plant species over three years. Colonies contained a higher bare ground component with lower vegetation heights than off-colony sites, with shortgrasses dominant at BTPD sites and a more even distribution of grasses, forbs, and shrubs at GUPD sites. Vegetation species composition was highly variable at BTPD sites over time, with increasing grasses and forbs and decreasing bare ground during an El Niño event associated with high rainfall during the 2 �growing season (Table 5, Fig. 2). We detected 17 raptor species during on- and off-colony counts. Burrowing owls, northern harriers Circus cyaneus, ferruginous hawks Buteo regalis (Fig. 1b), and roughlegged hawks Buteo lagopus were detected only on prairie dog colonies (ferruginous hawks, only on BTPD colonies). Apparent nest success varied between 50 and 57%, except that it was 40% on BTPD colonies and 69% at GUPD colonies in 2014. The decrease at BTPD colonies was likely attributable to hail storms and flooding during the peak nesting season in 2014, but prior to doing a thorough nest survival analysis, there was no obvious explanation for increased survival at GUPD colonies that year. The increase in nest numbers after 2013 was partly due to increased effort and partly to a huge influx of lark buntings during El Niño and following widespread plague events. In > 1 million remote camera photos, we have documented decreased coyote activity and increased swift fox activity over three years. Swift fox occurred only on BTPD colonies and badgers were more commonly detected there, while coyotes were equally common across all sites on BTPD and GUPD colonies. This was the third year of data collection on this project, and it will likely take additional years of monitoring to detect potential changes in the avian community caused by different types of plague management, as treated colonies no longer experience extinction events. Regardless of the efficacy of plague vaccine versus insecticide in reducing plague impacts, the vaccine will continue to be an important tool due to cost/benefit of its use and increasing evidence that fleas are evolving resistance to deltamethrin. Preliminary data suggest that bird densities do vary according to the status of prairie dogs on a colony, with differences between active colonies and those with extirpated or severely reduced prairie dog populations following plague outbreaks. These data also suggest several species may be associated with GUPD, including Brewer’s sparrow, vesper sparrow, and sage thrasher. Vegetation surveys have also identified differences between on- and off-colony areas. Raptor and camera data collection will continue through winter 2015/2016. Afterward, the field-based portion of phase 1 of this avian research project will be complete. We anticipate that phase 2 of this project will have a larger spatial scale, with the plague vaccine used more broadly as a management tool, but will focus on fewer data collection methods, species, and/or sites. Our study areas and the focus of data collection may shift, depending on availability of vaccine baits, management priorities, and results of avian data analysis from phase 1. 3 �COLORADO PARKS AND WILDLIFE RESEARCH REPORT AVIAN RESPONSE TO PLAGUE MANAGEMENT ON COLORADO PRAIRIE DOG COLONIES REESA C. YALE CONREY INTRODUCTION Wildlife diseases are important to conservation and population dynamics of susceptible species and may also have large indirect effects on non-susceptible species (Antolin et al. 2002). Introduced pathogens have the potential for far-reaching effects on native ecosystems that go beyond the mortality of infected individuals, particularly when a keystone species (Paine 1969) or ecosystem engineer (Jones et al. 1994) is infected. Range-wide declines in prairie dog (Cynomys sp.) populations have occurred, and the largest limiting factor in recent decades appears to be the high mortality and colony extirpation associated with introduced plague (Antolin et al. 2002), caused by the bacterium Yersinia pestis. Plague epidemics were first reported in the western United States in 1899 (Dicke 1926) and in northern Colorado in 1948 (Ecke and Johnson 1952). Instead of living in extensive colonies as they once did, prairie dogs exist in metapopulations of smaller colonies that periodically go extinct and are recolonized (Antolin et al. 2002, Stapp et al. 2004). Prairie dog colonies support a diverse community of associated species (Lomolino and Smith 2004, Smith and Lomolino 2004, Hardwicke 2006, Stapp et al. 2008), many of which are not susceptible to plague but may be indirectly effected. In order to conserve prairie dogs and species associated with their colonies, principally the blackfooted ferret (Mustela nigripes), a plague vaccination program is being tested (Fig. 1a). Additional species that may benefit from this program include those listed in the Conservation Plan for Grassland Species in Colorado (Colorado Division of Wildlife 2003): burrowing owl (Athene cunicularia: BUOW), mountain plover (Charadrius montanus: MOPL), ferruginous hawk (Buteo regalis: FEHA, Fig. 1b), and swift fox (Vulpes velox) and in the Colorado Sagebrush Conservation Assessment and Strategy (Boyle and Reeder 2005): Brewer’s sparrow (Spizella breweri: BRSP), green-tailed towhee (Pipilo chlorurus: GTTO), sage sparrow (Artemisiospiza belli: SAGS), sage thrasher (Oreoscoptes montanus: SATH), and vesper sparrow (Pooecetes gramineus: VESP), as well as BUOW. BUOW and MOPL are known to decline or disappear on colonies that are not reoccupied by prairie dogs after plague epizootics (Butts and Lewis 1982; Sidle et al. 2001; Augustine et al. 2008; Tipton et al. 2008; Conrey 2010), and horned lark (Eremophila alpestris), McCown’s longspur (Rhynchophanes mccownii), golden eagle (Aquila chrysaetos), prairie falcon (Falco mexicanus), and other birds may benefit from active colonies. From 2013‒2015, researchers in several western states are field-testing the uptake and efficacy (SPV Subcommittee 2011) of a new sylvatic plague vaccine (SPV) for prairie dogs (Rocke et al. 2010); the objective is to determine whether survival of prairie dogs and other small mammals is enhanced by the experimental vaccine compared to use of placebo or insecticide to control fleas, an important vector of plague. In Colorado, CPW researchers led by Dan Tripp are surveying colonies before and after bait distribution and conducting a mark-recapture study of prairie dogs and associated small mammal species (Tripp and Rocke 2012). As an extension to this project, we initiated research in 2013 on the effects of plague management on avian species associated with prairie dog colonies, with particular focus on species of concern. Our main long-term objective is to determine whether areas treated to control plague differ from untreated areas in their avian communities. Shorter-term objectives are to 1) Determine how plague affects avian species and their predators associated with prairie dog colonies; 2) Determine whether avian species associations exist for colonies of Gunnison’s prairie dogs (C. gunnisoni: GUPD); most evidence for associated species comes from black-tailed prairie dogs (C. ludovicianus: BTPD; Lomolino and Smith 2004, Smith and Lomolino 2004); 3) Determine whether insecticidal dusting 4 �influences bird density or nest survival; 4) Evaluate the importance of covariates such as weather and cattle grazing. This was the third and final year of phase 1 data collection for this project. We have collected one year of pre- and two years of post-treatment data on birds. During the initial years of the vaccination project, avian monitoring will also contribute information on responses to plague, insecticidal dusting, weather, and cattle grazing. This project will aid in the development of a standardized protocol for monitoring species associated with prairie dog colonies that could be used state-wide, as called for in the Conservation Plan for Grassland Species in Colorado (Colorado Division of Wildlife 2003). METHODS Study Area Study areas included BTPD colonies in north-central Colorado and GUPD colonies in westcentral Colorado. Baited sites received either vaccine or placebo baits in a blind procedure. Project areas that were selected for the prairie dog vaccine study had adequate numbers of prairie dogs and good access. BTPD (Larimer and Weld Co.) – Study colonies were located in Larimer and Weld Co. adjacent to the Wyoming border at Soapstone Prairie Natural Area (SOAP), managed by City of Fort Collins Natural Areas Program and Meadow Springs Ranch (MSR), managed by City of Fort Collins Utilities Department. These sites are characterized by shortgrass and mixed-grass prairie dominated by grasses (blue grama Bouteloua gracilis and buffalograss B. dactyloides) with smaller amounts of native (scarlet globemallow Sphaeralcea coccinea) and non-native forbs, shrubs, and cactus. Sites were sometimes grazed by cattle at low densities, and some non-baited sites were dusted with deltamethrin to control fleas (and plague). There was much more cattle grazing in 2014 − 2015 than in 2013, because these were wetter years (Fig. 2) with more forage. Both properties were closed to recreational shooting. Mark-resight estimates of BTPD density on shortgrass prairie in Colorado average approximately 10 prairie dogs/acre (Magle et al. 2007). Bird and vegetation surveys were conducted on nine SOAP colonies and 23 MSR colonies. There were nine vaccine project areas where raptors, predators, and passerine nests were surveyed: three prairie dog complexes each received vaccine, placebo, and dusting treatments (3 treatments*3 complexes = 9 project areas). 1) The Jack Springs (Jac) colony spanning the SPNA/MSR border contained 100 vaccine acres, 100 control acres, and ~280 dusted acres separated by 200 - 400 m buffer zones. 2) The Barton complex in MSR contained ~130 vaccine acres and ~180 control acres, encompassing much of the Barton south and west colonies (BarS and BarW). Raptor, predator, and passerine nest surveys were also done in the 140 acre Barton east colony (BarE), although the prairie dog crew instead paired a dusted portion of the Ferret Center colony (Fer) with the Barton complex. 3) The Ferret Center complex in MSR contained 40 vaccine acres, 40 control acres, and ~478 dusted acres, encompassing the entire North Bulger south colony and half of the Ferret Center (Fer) colony, with a 400 m buffer zone separating those treatments. GUPD (Gunnison Basin and Woodland Park) – Study colonies were located in the Gunnison Basin (Gunnison, Saguache, and eastern Montrose Co.) and in the Woodland Park area (Teller Co.). Gunnison Basin (GUNN) sites were managed by the Bureau of Land Management (BLM), Colorado Parks and Wildlife (Miller Ranch State Wildlife Area and Van Tuyl/Cabin Creek SWA), and the U.S. Forest Service Rio Grande National Forest. Woodland Park area (WOOD) sites were managed by the National Park Service Florissant Fossil Beds National Monument (FFB) and Colorado Parks and Wildlife (Dome Rock SWA). These sites are characterized by a mixture of big sagebrush (Artemisia tridentata), rabbitbrush (Chrysothamnus viscidiflorus), prairie grasses and forbs (fringed sagebrush A. frigida) in a matrix of pasture, pine and spruce-fir forests. Sites were sometimes grazed by cattle or sheep at low densities, and all non-baited sites were dusted with deltamethrin to control fleas (and plague). All 5 �properties except FFB were open to recreational shooting, but shooting was not prevalent and signage discouraged shooting in vaccine project areas. The study area is within the known range of plague with plague epizootics occurring near the study colonies in 2010 (Tripp et al. unpublished data). Visual counts of Gunnison’s prairie dogs on colonies in Colorado averaged 6.1 prairie dogs/acre in 2010. Bird and vegetation surveys were conducted on 14 GUNN colonies and six WOOD colonies. There were nine vaccine project areas receiving vaccine, placebo, and dusting treatments where raptors, predators, and passerine nests (GUNN only) were surveyed. Because of their small size, entire colonies were treated. 1) The 33 acre Miller Ranch (MR) and 26 acre Kenny Moore (KM) colonies north of Gunnison received vaccine and control treatments, and the 62 acre Power Line (PL) colony 16 km to the south was designated as the paired dusted treatment. 2) The 46 acre Cabin Creek (CC), 37 acre BLM-15 (B15), and 27 acre BLM-5 (B5) colonies southeast of Gunnison received vaccine and control treatments, and the 69 acre BLM-18 (B18) colony was designated as the paired dusted treatment. B5 has been baited in the past due to concerns that GUPD sample size would be too low in B15. All these colonies are within the same complex. 3) Two ~20 acre Florissant Fossil Beds (FFB) and one 24 acre Dome Rock (DR) colony southwest of Woodland Park (150 km east of Gunnison Basin) received vaccine and control treatments. There was no available colony to use as the paired dusted treatment. At the GUPD study sites, we have an additional objective of determining whether avian species associations exist; therefore, we selected off-colony sites for comparison of data from avian point counts, vegetation surveys, and raptor counts. Time and financial constraints precluded nest searching or camera use off colonies. We extended the 250 m point grid off colonies and created a doughnut-shaped region that extended 500 – 1500 m from colony boundaries. Within these doughnut regions, each year we randomly chose new grids of nine points (3 x 3) to serve as off-colony study areas on public lands. Some grids had fewer than nine points due to land ownership boundaries. For each colony, we surveyed at least two off-colony sites. Some off-colony sites were located in sagebrush, but others were located in forested areas, especially in areas where forest was the dominant cover type (FFB, DR, and USFS property in the Gunnison Basin). Avian Point Counts Each point was surveyed once in May – June 2013 – 2015. At BTPD study sites, a 250 m grid of points had already been established and surveyed by RMBO from 2006−2013. At GUPD study sites, we created a 250 m grid of points. Within each off-colony grid, we randomly chose a minimum of four points to survey. Two off-colony grids were randomly chosen to pair with each colony, so that we surveyed a minimum of eight off-colony points per paired colony. For larger colonies containing more than eight survey points, we completed as many point counts per off-colony colony as we did on colony. Points were considered to be “on colony” if located within 100 m of the boundary and with good views of the colony. Most point counts were conducted between dawn and 10:00 and were never conducted in rain, hot temperatures (above 30°C), or high winds that made it difficult to hear birds. Regardless of time or weather, we did not conduct counts if we noticed that bird activity (especially singing and calling) was dropping off. In addition to these breeding season counts, we conducted early counts for mountain plover and burrowing owls, which arrive earlier than passerines and are poorly detected in regular point counts, in late April – early May 2015. At all grid locations where these species were likely to be detected, we first did 6-min. passive point count surveys. Passive surveys were followed by 9-min.call-playback surveys at a selection of grid locations spaced at least 700 m to avoid calling in birds and double-counting them. The call-playback sequence was 3 min. of silence, 3 min. of plover calls (30-sec. each of display, chirp, and chick calls, each separated by 30-sec. of silence), and 3 min. of owl calls (two 30-sec. segments of coo call and 30-sec. of alarm call, each separated by 30-sec. of silence). Surveys were conducted from ATVs, 6 �because plover and owls are much less likely to be disturbed by observers in vehicles than by observers on foot. We conducted 6-min.breeding season point counts, recording each bird’s species, horizontal (radial) distance, sex (if known), use of the prairie dog colony (yes or no), minute of detection (1 – 6), and how it was detected (visual, singing, calling, drumming, fly-over, or other). Membership in a cluster was noted, typically for male-female pairs. After completing the bird count, we recorded weather and site characteristics at each point, including time, temperature, wind speed, cloud cover, management type (typically cattle grazing, sometimes dusting) and whether it was current, from this season or last season, and the presence of excessive noise, roads (primary or secondary), and cliffs or rock outcroppings within 100 m. Within a 50 m radius, we recorded characteristics of tall nesting and perching substrate, including percent cover, height, and dominant species for overstory plants ≥ 3 m and shrubs > 30 cm but < 3 m. Within a 5 m radius, we recorded characteristics of ground cover, including percent cover of grasses (including sedges and rushes), forbs, shrubs, cactus, litter, bare ground, rock, scat, other cover such as lichen, and exotic species. We also recorded the dominant exotic species, and the mean height and species of the dominant grass. We used the point count protocol designed for Integrated Monitoring of Bird Conservation Regions (IMBCR: Hanni et al. 2012), except that we conducted bird surveys prior to vegetation surveys. This helped to ensure that birds displaced by the observer, including those located at the point itself, were recorded. We also altered IMBCR vegetation survey protocols slightly to make the protocol specific to low stature prairie dog colonies, shortgrass prairie, and sagebrush systems. This was designed to be a quick, visual assessment; a more involved protocol using a Daubenmire frame and robel pole was used on transects and at nests. Vegetation Transects In addition to a visual assessment of vegetation at points, we sampled vegetation on transects and at nests. We completed two transects on vaccine project colonies and paired off-colony sites and at least one transect on non-project sites (both on and off colonies). To locate each transect, we randomly chose a start and an end point from those used in avian point counts. From the start point, we walked along the bearing toward the end point for 240 m, stopping every 20 m to collect vegetation data for a total of 13 points per transect, except on very small colonies. Transect data were collected during the growing season, most during July and August. At each stop point, we recorded the presence of active or inactive prairie dog burrows within 10 m, ground cover, dominant species, and visual obstruction (Fig. 1c). Percent ground cover was measured within a 50 cm square Daubenmire frame. We recorded the percent bare ground, rock, litter, scat, grass (including sedges and rushes), forb, shrub, cactus, exotic, and other cover. We also recorded the dominant species for each plant category present in the frame. Visual obstruction data were recorded by holding a robel pole at the stop point on the transect and making observations from a distance of 4 m in each cardinal direction with eye level at 1 m. The observer then noted which portions of the 122 cm (4 ft) pole were obstructed by vegetation, identified the plant species obstructing the pole, and noted whether the pole was substantially obstructed or was covered by just a wisp of vegetation (typically a blade of grass). We estimated the height of any structures taller than the pole (a few trees at GUPD off-colony sites). Raptor Counts Raptors were sometimes sighted during avian point counts, but point counts are not an ideal method for detecting raptors or other uncommon species. Therefore, we chose 1 – 3 locations per vaccine project area (on- and off-colony), positioning observers so that the entire treatment area could be viewed simultaneously. We conducted 30-min. raptor counts, recording each bird’s species, horizontal (radial) distance, sex (if known), time of entry and exit, and behavior (high soar, low soar, directed flight, hover, 7 �dive, call, perch, or nest). Membership in a cluster was noted, typically for male-female pairs. This produces a time metric for assessing raptor use of treatment areas and colonies. At the start and end of the count, we recorded weather characteristics, including time, temperature, wind speed, and cloud cover. Raptor counts were conducted after 9:30 from November to early March (wintering) and April to August (breeding) and were never conducted in rain. As a supplement to the formal raptor counts, we recorded incidental raptor observations. Nest Searching and Monitoring We searched for mountain plover and burrowing owl nests throughout the study area through visual observation of adult birds, typically in the morning and not in rainy conditions or high winds. Target search regions included areas where MOPL and BUOW were detected during call-playback surveys, areas with nests in previous years, and other areas with appropriate habitat. MOPL nest in scrapes on the ground in areas with a relatively high bare ground component. BUOW nest in prairie dog burrows, often near colony edges and in burrows with low to moderately-sized mounds. Because these species react more to humans on foot than to vehicles, we conducted surveys from a vehicle whenever possible. Where ATV access was possible, we nest searched for MOPL using two ATVs with a 30 m rope suspended between them. Bungies on each end of the rope allowed observers to keep the rope taut and watch for running birds. When MOPL were detected, we observed the bird, sometimes backing away from the site, and waited for the bird to sit down on a nest. When BUOW were detected, we searched for nests in the vicinity of their perching location; typically males perched conspicuously near the nest burrow during the day. We searched for passerine nests (Fig. 1d) on colonies in vaccine project areas only, because the rope dragging technique (Yackel Adams 2000) that works best for secretive birds and camouflaged nests is time-consuming. Most prairie passerines nest in woven cups on the ground, while shrubland passerines typically place their nests on branches or under shrubs. Each search area was surveyed at least twice. At GUPD sites, each entire colony was searched via rope dragging. At BTPD sites, we searched 40 acre plots, which was the size of the smallest treatment area and an area that could be searched by two people in a half day. For those sites, we nest searched at two plots. We located one plot non-randomly, centered on the area with greatest density of the previous year’s nest locations. The second plot was randomly located with one corner on a point from the larger 250 m grid. Passerine nest searching was typically done after 9:30, because grassland birds are more likely to be in attendance at their nests during the heat of the day, and not in rainy conditions or high winds. At BTPD sites, we dragged a 100 ft (30 m), ½ inch gauge rope with two people at each end, watching for flushing birds in the area ahead of and under the rope. Because the GUPD sites contained a much higher shrub component, it was not possible to drag a heavy rope without continuously getting snagged on vegetation; therefore, we used two different ¼ inch gauge ropes, depending on the shrub component at the site. One was 10 m and the other was 23 m long, held above the vegetation, with heavy hex nuts suspended from smaller ropes to disturb vegetation slightly and flush birds. A few locations with very dense shrub cover were searched by disturbing vegetation with 4 foot wooden dowels in each outstretched hand. Additional nests were found during point counts and nest monitoring. When we were unable to find a nest during the initial search, we marked the GPS location and returned at a later date. We likely found the majority of BUOW nests on the landscape using this method (Conrey 2010), but a smaller and unknown proportion of MOPL and passerine nests were found. MOPL and passerine nests were defined as structures containing at least one egg. Because BUOW nests are underground, we defined their nests as burrows with shredded manure present at the entrance (Garcia and Conway 2009), with feathers, regurgitated pellets, and prey remains providing additional evidence of a nest attempt. Some BUOW burrows were probed with an electronic camera on the end of a structure similar to a plumbing snake (Peeper System 2.3, Sandpiper Technologies). At the 8 �time of nest discovery, we recorded the same weather information that we recorded at points. We also did a rapid visual assessment of vegetation, with more detailed data collected at nest completion when overheating of eggs and nest abandonment was no longer a concern. We described the nest structure and vegetation immediately around the nest and estimated vegetation height, percent bare ground within a 1 m radius, and whether all, ≥ 50%, < 50%, or none of the nest could be seen from vantage points 5 m to the north and south. BUOW nests were marked with brightly painted wooden stakes placed 10 m north of the nest burrow. MOPL and passerine nests were marked with two small unpainted wooden stakes placed 5 m north and south of the burrow. We collected any pellets that we observed near BUOW nests for possible future dietary analysis. MOPL and BUOW nests, with relatively long incubation and nestling periods, respectively, were monitored at least once per week. Passerine nests were monitored every 2 – 3 days. Starting with the first visit when the nest was discovered, we recorded the time, any management activities (such as cattle grazing), age of eggs and juveniles, and number of eggs, juveniles, and adults present. MOPL and passerine eggs were aged by floating (Fig. 1e): eggs closer to hatch float higher in the water column. Juveniles were aged according to keys for BUOW (Priest 1997) and LARB (Yackel Adams Unpub. data), and we created our own photographic keys for all other species based on our 2013 observations. Passerine nests were considered successful if at least one fully-feathered juvenile left the nest. Evidence of success included juveniles outside the nest cup, mutes at the edge of the nest, and/or displaying and calling adults, coupled with an intact nest and appropriate timing based on nest age. MOPL nests were considered successful if at least one egg hatched, because their chicks are precocial and can leave the nest area within hours of hatch. Evidence of MOPL success included pip chips, coupled with an intact nest and appropriate timing based on nest age. BUOW nests were considered successful when at least one fledgling aged ≥ 35 days was observed (Thomsen 1971, Davies and Restani 2006, Conrey 2010), because they leave the nest burrow (but may return to it many times) at 10 – 14 days and well before flight or independence are attained. Failed nests were destroyed, contained broken eggs, and/or had eggs or nestlings that disappeared before their expected hatch (MOPL) or fledge (BUOW and passerines) date. For analysis purposes, nests with unknown fate will have their histories truncated back to the last date when the nest was active and will be coded as successful at that time. At nest completion, we recorded the same vegetation data that were collected at points along vegetation transects: presence of prairie dog burrows within 10 m, percent ground cover, and visual obstruction. We placed the Daubenmire frame at 1 m in each cardinal direction and observed the robel pole (placed at the nest) from 4 m in each cardinal direction, producing four readings of each metric. For ground and shrub nests, we also recorded the height of the nest cup above (or below) ground and the plant species or structure type (such as cow paddy) in which (or adjacent to which) the nest was located. Remote Cameras Remote cameras (Reconyx Hyperfire Covert IR model PC800) were placed in each vaccine project area to document use by mammalian predators and other wildlife. In 2013, we had just one camera per treatment, but we increased the number of remote cameras on BTPD colonies in 2014 – 2015 to better sample their larger size relative to GUPD colonies. In 2015, we used two cameras at small BTPD sites and 3 – 6 cameras at larger sites. These cameras take photos when triggered by motion from an object that is warmer than ambient temperature. Camera locations were selected to maximize the potential for detections of mammalian predators without the use of baits or lures, which might have acted as attractants and altered the sampling region beyond treatment areas and prairie dog colonies. Cameras were positioned along game trails, aimed at coyote height, and tested before they were armed. Cameras targeted water sources, fence lines, and other landscape features and were positioned in space such that the entire treatment was sampled as thoroughly as possible. We set the cameras to take three photos when triggered, with no quiet period between photos. 9 �Most cameras were deployed in April – early May of each year. Most of the cameras at BTPD sites were left in place and operated year-round. We checked batteries and SD cards at least 3 times during summer and once during winter, but most camera checks were more frequent (once or twice per month from May - September), particularly when cows were present on a site, as they sometimes disrupted camera operation. Cameras were removed from GUPD sites in September, prior to the advent of winter weather and GUPD hibernation. As a supplement to the camera data, we recorded incidental observations of predators such as coyotes, foxes, and badgers. Databases We designed a database for this project using Microsoft SQL Server 2012 with the data entry interface in Microsoft Access 2007. The database was designed by R. Conrey and D. Conrey, a professional database developer who volunteered his time, to run on the Fort Collins CPW research server. This allows multiple users to simultaneously access the database, while providing for daily backups and improved data security. Users can access a master list of codes for vegetation species, bird species, management types, observers, sites, colonies, and points that if changed, will update throughout the database. Users also access data entry forms for each data type described above, except for photo data. The Reconyx photo database (Newkirk 2014) catalogs photos and stores the metadata associated with them, displaying (but not storing) the photos themselves within Access forms. Users identify species in photos using stand-alone modules which are then loaded into the database. Each photo is examined by at least two observers, and a referee (R. Conrey or field crew leader) resolves any conflicts in IDs. Identification is ongoing. Data Analysis Thus far, all data have been entered and summarized, but data proofing is ongoing and statistical analyses have not yet been completed. For point counts, we calculated use rate for each species by dividing the number of detections by the number of points per site*year. We completed bird and vegetation species lists and summarized ground cover data collected at transects. For raptor counts, we calculated a proportional use index, dividing the usage minutes by the total survey minutes for each site*year. Apparent nest success was calculated as the proportion of nests fledging (raptors and passerines) or hatching (MOPL) at least one chick. Naïve (minimum) occupancy estimates were calculated for carnivores based on remote camera photos taken during 2-week intervals starting 1 April of each year, separated into breeding and non-breeding periods. Data will eventually be analyzed using Program DISTANCE to estimate density from point counts, Program MARK to estimate nest survival and occupancy from point counts and camera data, and R for other data types. RESULTS We have collected one year of pre-treatment data and two years of post-treatment data since initial bait drop in late summer and fall of 2013. Since fall 2013, plague epizootics have occurred on one GUPD colony and across ~70% of the BTPD study area at 15 colonies. During each year of our study, one of the three pairs of BTPD treatment sites experienced a plague outbreak, spanning the entire treatment area by fall 2015, aside from a dusted region around the USFWS black-footed ferret breeding center. Plague occurred at both the baited sites in each pair, meaning that both the control and vaccine areas experienced outbreaks. This epizootic started in fall 2013 shortly after vaccine drop, so plague was present in the system prior to vaccination. Colonies that experienced epizootics in 2013 experienced total or near extirpation of prairie dogs, but beginning in 2014, treated areas experienced losses of ~50 – 95% (two of these were treated with placebo baits). All of these areas had small numbers of prairie dogs with increasing abundance within one year of the outbreak, but populations were severely reduced compared pre-plague levels. In September 2014 and 2015, black-footed ferrets were released in two BTPD study 10 �colonies (the Roman and Brannigan colonies in the northwest part of Soapstone Prairie Natural Area). In October 2014 and 2015, additional ferrets were released within the dusted part of a treatment colony (the Ferret Center colony in the southeastern part of Meadow Springs Ranch), which is 0.5 km from the nearest paired baited site and plague epizootic. In 2014, 42 total ferrets were released on BTPD study colonies, with a similar number planned for release in 2015. 2014 and 2015 were very wet summers at the BTPD site compared to the 30-year average, whereas precipitation at GUPD sites was dry to normal except for a wet May in 2015 (Fig. 2). Normally dry playas and streambeds were full, grasses were tall and formed seedheads in June, and a different suite of species was observed, including a large flush of growth in species normally absent or a minor component of the prairie, such as six weeks fescue (Vulpia octoflora) and wooly plantain (Plantago patagonica). We saw a huge influx of lark buntings (Calamospiza melanocorys), which typically nest in or near taller vegetation than is typically found on these BTPD colonies; we had two nests in 2013, 69 nests in 2014, and 39 nests in 2015, making lark buntings the most abundant breeders at our sites, along with McCown’s longspurs. We conducted 2,284 avian point counts in 2013 − 2015 and detected 130 bird species during the breeding season (Table 1, App. 1). The most common birds detected were horned lark, lark bunting, western meadowlark, and McCown’s longspur at BTPD sites and Brewer’s sparrow, vesper sparrow, green-tailed towhee, sage thrasher, horned lark, western meadowlark (Sturnella neglecta), and common raven (Corvus corax) at GUPD sites, in that order. We detected 83 species on BTPD colonies, 81 species on GUPD colonies, and 80 species off GUPD colonies. Occupancy and density analyses have not yet been completed, but we have compared use rates (number of detections per point) across BTPD colonies with differing prairie dog activity status and across GUPD sites, on and off colonies. During BTPD colony surveys, at least three bird species (Brewer’s blackbird Euphagus cyanocephalus, Brewer’s sparrow, and vesper sparrow) appeared to have higher detection rates on active prairie dog colonies, while two species (grasshopper sparrow Ammodramus savannarum and lark bunting) appeared to have higher detection rates on colonies with extinct or severely reduced prairie dog populations following plague outbreaks (Table 2). During surveys on and off GUPD colonies, seven species (Brewer’s sparrow, common raven, horned lark, red-winged blackbird Agelaius phoeniceus, sage thrasher, vesper sparrow, and western meadowlark) appeared to show associations with colonies, while four species (dark-eyed junco Junco hyemalis, green-tailed towhee Pipilo chlorurus, mountain chickadee Poecile gambeli, and western wood-pewee Contopus sordidulus) had higher detection rates off colonies, and many others showed no pattern (Table 3). In future density models, we will test the importance of these factors along with other covariates such as weather, vegetation characteristics, cattle grazing, and predator occupancy rates. We characterized vegetation at 2,284 point count locations, 543 nests, and ~4000 stop points along 321 transects in 2013 – 2015 and documented 217 species (Table 1, App. 2). Vegetation transect data (Table 4) suggested that BTPD colonies were dominated by grass (mainly blue grama) and bare ground, with triple the grass coverage of GUPD sites and ~20% more grass during the El Niño event of 2014 – 2015 than in 2013 when precipitation was closer to the average (Table 5, Fig. 2). We observed half the litter, a 10% drop in bare ground, and double the forb coverage at BTPD sites over the same time period (Table 5). GUPD sites had a more even distribution among cover types (Table 4): bare ground was the most common cover type at the GUNN sites, both on and off colonies, and GUPD colonies had 14% more bare ground and shorter vegetation heights than off-colony areas. At the GUPD sites, shrub cover (mainly big sagebrush) was much higher in GUNN while grass (mainly blue grama), litter, and forb (mainly fringed sagebrush) cover were much higher in WOOD. Exotic cover was low: 1% at BTPD, 1 – 2% at WOOD, and 4 – 5% at GUNN sites (Table 4). The most common exotics were grasses (Table 6) that were likely purposefully planted for forage in the past. Visual obstruction by vegetation (Fig. 1c) was lowest on BTPD colonies (8.8 cm), moderate at WOOD sites (13.0 cm on colonies, 15.5 cm off colonies), 11 �and highest at GUNN sites (18.0 cm on colonies, 22.9 cm off colonies). Vegetation heights increased each year of our study on BTPD colonies, with grasses responsible for most of the obstruction. Forbs had the highest species richness, followed by grasses and shrubs. Dominant plant species of each type are listed in Table 6, with a complete plant species list in App. 2. We conducted raptor counts at 75 locations for a total of 20,250 minutes (Table 1) and detected 17 species in 2013 – 2015 (Table 7). Burrowing owls, northern harriers (Circus cyaneus), ferruginous hawks (Fig. 1b), and rough-legged hawks (Buteo lagopus) were detected only on prairie dog colonies (ferruginous hawks, only on BTPD colonies). Burrowing owls, Swainson’s hawks (Buteo swainsoni) and turkey vultures (Cathartes aura) were more common on BTPD colonies, while ravens and red-tailed hawks (Buteo jamaicensis) were more common at GUPD sites. During winter counts on BTPD colonies, burrowing owls, Swainson’s hawks, and turkey vultures were replaced by rough-legged hawks, and ferruginous hawks and northern harriers became more common than they were during summer. Relationships between raptor use and plague, GUPD colony associations, and other factors have not yet been explored. We monitored 543 nests (Fig. 1d, e) of 21 bird species, with apparent nest success (fate) of 52% in 2013 – 2015 (Tables 1, 8). Apparent nest success varied between 50 and 57%, except that it was 69% at GUPD colonies and 40% on BTPD colonies in 2014, a wet year with frequent hail storms at BTPD sites (Fig. 2). The increase in nest numbers after 2013 was partly due to increased effort and partly to a huge influx of lark buntings during El Niño and following widespread plague events. Following plague epizootics, burrowing owl nest numbers increased in those colonies. Comparing nesting data to point count data, it appears that the species that preferred to sing and display on active BTPD colonies (versus reduced or extinct colonies) do not often nest on those colonies (Tables 2, 8), although Brewer’s and vesper sparrows did commonly sing and nest on GUPD colonies (Tables 3, 8). Daily nest survival will be estimated using known fate models in Program MARK, and the importance of covariates such as plague, weather, vegetation characteristics, cattle grazing, and predator occupancy rates will be explored. Between 2013 and 2015, we increased remote camera numbers from 18 to 43, with 32 cameras on BTPD colonies and 11 cameras on GUPD colonies in 2015 (Table 1). We have analyzed > 1.13 million photos (Table 1), and more photos will be taken this winter at BTPD colonies. BTPD cameras are deployed year-round, because BTPDs do not hibernate, while GUPD cameras were deployed from May − September. As expected, many photos recorded prairie dogs, cows, pronghorn (Antilocapra americana), and rabbits. We have documented coyote (Canis latrans), swift fox (Vulpes velox), badger (Taxidea taxus), striped skunk (Mephitis mephitis), red fox (V. vulpes), bobcat (Lynx rufus), black-footed ferret, and raccoon (Procyon lotor) use of vaccine project areas (listed by number of occurrences in our photos). We observed decreased coyote activity and increased swift fox activity from 2013 – 2015 (Table 9). Swift fox and skunks occurred only on BTPD colonies and badgers were more commonly detected there, while coyotes were equally common across all sites on BTPD and GUPD colonies. Overall 2-week detection rates per site (naïve occupancy estimates) for the three most common carnivores ranged from 3% for badgers on both BTPD colonies (April – July 2014) and GUPD colonies (August – Sept 2014) to 51% for coyotes on BTPD colonies (August 2014 – March 2015; Table 9). DISCUSSION The 2015 season was the third of three study seasons associated with phase 1 of this avian research project, which coincided with Wildlife Health’s 3-year efficacy trials for the plague vaccine. We have collected one year of pre-treatment data and two years of post-treatment data. It will likely take additional years of monitoring to detect potential changes in the avian community caused by plague management, as treated colonies no longer experience extinction events. Therefore, data analyses will 12 �focus on shorter-term objectives, including evaluating the importance of plague on BTPD colonies and colony associations for birds and GUPD. Since fall 2013, plague epizootics have occurred on one GUPD colony and across ~70% of the BTPD study area. All of these areas had small numbers of prairie dogs with increasing abundance within one year of the outbreak, but populations were severely reduced compared to pre-plague levels. Plague outbreaks have occurred across all treated BTPD sites, but plague was likely present in the system prior to vaccination, and immunity takes time to develop after vaccination (D. Tripp, pers. comm.). Prairie dog researchers are interested in whether prairie dogs persist in treated areas during outbreaks, recover more quickly in treated areas, and have higher survival during enzootic periods (without obvious outbreaks). At least three bird species (Brewer’s blackbird, Brewer’s sparrow, and vesper sparrow) appeared to have higher detection rates on active prairie dog colonies, while two species (grasshopper sparrow and lark bunting) appeared to have higher detection rates on colonies with extinct or severely reduced prairie dog populations following plague outbreaks. Higher numbers of grasshopper sparrows and lark buntings coincided with high May – June precipitation, which likely contributed to high flea numbers and conditions favorable to a plague epizootic. Conditions across much of the BTPD study area in 2014 – 2015 began to support grassland birds that favor taller herbaceous vegetation over those that favor shorter vegetation, such as horned larks and McCown’s longspurs. Interestingly, these two species did not show an association with active BTPD colonies in our initial exploration of the data, while Brewer’s and vesper sparrows, which nest in shrubs and dense herbaceous vegetation, respectively, did have higher detection rates on active colonies. Further analyses of species density and nest survival that include other covariates, such as weather and predator occupancy, will elucidate these relationships. Thus far, data from avian point counts and vegetation surveys have identified differences between on- and off-colony areas for GUPD sites, but more analyses are required before we draw conclusions regarding the uniqueness of the avian community on GUPD colonies. Point count data suggest several species may be associated with GUPD, including Brewer’s sparrow, vesper sparrow, and sage thrasher. GUPD colonies contained a higher bare ground component with lower vegetation heights than off-colony sites. BTPD increase bare ground and alter plant species composition and nutrient cycling rates (Whicker and Detling 1988; Johnson-Nistler et al. 2004); effects of GUPD on vegetation are not well-studied. GUPD do provide refugia and nests for BUOW, kit fox, and small mammals (Miller et al. 1994, Meaney et al. 2006). However, their impact appears less dramatic than that of BTPD (Grant-Hoffman and Detling 2006), perhaps because of differences in habitat, less above-ground activity, little clipping of vegetation, and lower burrow densities (Seglund and Schnurr 2010). Precipitation has varied greatly over the three years of this study, particularly on BTPD sites, from slightly dry to very wet, compared to the 30-year average. In summer 2014 and 2015, normally dry playas and streambeds were full, grasses were tall and formed seedheads in June, and a different suite of species was observed, including a large flush of growth in species normally absent or a minor component of the prairie. In response, we saw a huge influx of lark buntings to BTPD sites; they were the most common species detected in point counts and nest searches in 2014, and nest numbers continued to be high in 2015. In contrast, McCown’s longspur numbers declined in 2014, then rebounded in 2015, while the number of burrowing owl nests doubled each year. These changes may also be related to concurrent declines in prairie dogs associated with plague epizootics, which accelerate during El Niño events when cooler, wetter springs and summers result in high numbers of fleas, a major vector of plague. Vegetation was no longer being clipped, and extinct areas had much more lush vegetation than usual. Burrowing owl nest numbers increased only in areas that experienced plague epizootics and had persisting (or recovering) prairie dog populations. Burrowing owls also increased in response to plague events (and prairie dog recolonization) in a previous study in northern Colorado (Conrey 2010). High nest numbers did not correspond with high nest success; apparent nest success declined from 54% in 2013 to 40% in 2015 at BTPD sites, and then recovered to 57% in 2015. In contrast, precipitation at GUPD sites remained 13 �average to low throughout 2013 – 2015, except for high moisture in May 2015. Apparent nest success on GUPD colonies increased from 50% in 2013 to 69% in 2014, then returned to 51% in 2015. The decrease in nest success at BTPD colonies was likely attributable to hail storms and flooding during the peak nesting season in 2014 (there was less hail and flooding in 2015), but prior to doing a thorough nest survival analysis, there was no obvious explanation for increased survival at GUPD colonies that year. Further analysis of these patterns will be elucidated by daily nest survival models that include factors such as precipitation, prairie dog status, predator occupancy, and other factors. In September and October 2014, 42 black-footed ferrets were released in three BTPD study colonies, one of which was only 0.5 km from the nearest baited site, which experienced a plague epizootic in fall 2013. Additional ferrets were released in 2015, and spotlight surveys in 2015 documented a lactating female. The two release sites (not included in the prairie dog vaccine study) in Soapstone Prairie Natural Area were both dusted and baited with vaccine, and the release site near the Ferret Center is a dusted treatment within the vaccine study. Released ferrets have been vaccinated against plague and distemper, and were expected to disperse into baited sites within the study area. We began documenting ferrets in our remote camera photos in October 2015, but thus far, no ferrets have been seen or photographed outside of the colonies where they were released. At this point, young ferrets must be captured to be vaccinated, as they are not expected to consume vaccine baits designed for prairie dogs, but work on an oral vaccine for ferrets is in progress. Ferrets will likely predate on adult birds and nests, but as 90% of their prey base is typically comprised of prairie dogs (Clark 1986), the overall effect on avian species should be minimal. Raptor and camera data collection will continue through winter 2015/2016. Afterward, the fieldbased portion of phase 1 of this avian research project will be complete. Regardless of the efficacy of plague vaccine versus insecticide in reducing plague impacts, the vaccine will continue to be an important tool due to cost/benefit of its use and increasing evidence that fleas are evolving resistance to deltamethrin. After 2015, scale of use of plague vaccine will depend on the cost and availability of vaccine and analyses of prairie dog survival. The focus and intensity of research and monitoring of bird communities on prairie dog colonies will depend on the presence of plague in the system, presence of black-footed ferrets, available resources, and scale of vaccine use. If the vaccine study suggests that this is a successful approach to plague management, then the vaccine may be more broadly used as a management tool. This would allow us to change the spatial scale of our research during phase 2 of this project, perhaps including larger, more suitable areas for focal species such as BUOW and MOPL. If more MOPL can be located within treated areas, we plan to collaborate with other MOPL researchers across multiple sites to study changes in MOPL dynamics as a response to longer term plague management. Analyses this fall will also identify other appropriate focal species, determine whether or not we should continue sampling on GUPD colonies, and provide guidance for a general monitoring protocol for species associated with prairie dog colonies. LITERATURE CITED Antolin, M. F., P. Gober, B. Luce, D. E. Biggins, W. E. V. Pelt, D. B. Seery, M. Lockhart, and M. Ball. 2002. The influence of sylvatic plague on North American wildlife at the landscape level, with special emphasis on black-footed ferret and prairie dog conservation. Transactions of the 67th North American Wildlife and Natural Resources Conference 67: 104–127. Augustine, D. J., S. J. Dinsmore, M. B. Wunder, V. J. Dreitz, and F. L. Knopf. 2008. Response of mountain plovers to plague-driven dynamics of black-tailed prairie dog colonies. Landscape Ecology 23:689–697. Boyle, S. A. and D. R. Reeder. 2005. Colorado sagebrush: a conservation assessment and strategy. Colorado Division of Wildlife, Grand Junction, Colorado. Butts, K. O. and J. C. Lewis. 1982. The importance of prairie dog colonies to burrowing owls in 14 �Oklahoma. Proceedings of the Oklahoma Academy of Sciences 62:46–52. Clark, Tim W. 1986. Some guidelines for management of the black-footed ferret. Great Basin Naturalist Memoirs 8:160–168. Colorado Division of Wildlife. 2003. Conservation plan for grassland species in Colorado. Colorado Division of Wildlife, Denver, Colorado. Conrey, R. Y. 2010. Breeding success, prey use, and mark-resight estimation of burrowing owls nesting on black-tailed prairie dog towns: plague affects a non-susceptible raptor. Ph.D. Dissertation, Colorado State University, Fort Collins, Colorado. Conrey, R. Y. 2013. Avian response to plague management on Colorado prairie dog colonies. Wildlife Research Report. Colorado Parks and Wildlife, Fort Collins, Colorado. 25 pg. Davies, J. M. and M. Restani. 2006. Survival and movements of juvenile burrowing owls during the postfledging period. Condor 108:282–291. Dicke, W. M. 1926. Plague in California 1900 – 1925. Proceedings of the 41st Annual Meeting and Conference of State Provincial Health Authority of North America, Atlantic City, New Jersey. Ecke, D. H. and C. W. Johnson. 1952. Plague in Colorado and Texas. Part I. Plague in Colorado. Public Health Monograph No. 6. U. S. Government Printing Office, Washington D.C. Garcia, V. and C. J. Conway. 2009. What constitutes a nesting attempt? Variation in criteria causes bias and hinders comparisons across studies. Auk 126:31–40. Grant-Hoffman, M. N. and J. K. Detling. 2006. Vegetation on Gunnison’s prairie dog colonies in Southwestern Colorado. Rangeland Ecology and Management 59:73–79. Hanni, D. J., C. M. White, R. A. Sparks, J. A. Blakesley, J. J. Birek, N. J. Van Lanen, J. A. Fogg, J. M. Berven, and M. A. McLaren. 2012. Integrated Monitoring of Bird Conservation Regions (IMBCR): Field protocol for spatially-balanced sampling of landbird populations. Unpublished report. Rocky Mountain Bird Observatory, Brighton, Colorado. 39 pp. Hardwicke, K. 2006. Prairie dogs, plants, and pollinators: tri-trophic interactions affect plant-insect floral visitor webs in shortgrass steppe. Ph.D. Dissertation. Colorado State University, Fort Collins, Colorado. Johnson-Nistler, C. M., B. F. Sowell, H. W. Sherwood, and C. L. Wambolt. 2004. Black-tailed prairie dog effects on Montana’s mixed-grass prairie. Journal of Range Management. 57:641–648. Jones, C. G., J. H. Lawton, and M. Shachak. 1994. Organisms as ecosystem engineers. Oikos 69:373– 386. Lomolino, M. V. and G. A. Smith. 2004. Terrestrial vertebrate communities of black-tailed prairie dog (Cynomys ludovicianus) towns. Biological Conservation 115:89–100. Magle, S. B., B. T. McClintock, D. W. Tripp, G. C. White, M. F. Antolin, and K. R. Crooks. 2007. Mark– resight methodology for estimating population densities for prairie dogs. Journal of Wildlife Management 71: 2067–2073. Meaney, C. A., M. Reed-Eckert, and G. P. Beauvais. 2006. Kit Fox (Vulpes macrotis): a technical conservation assessment. USDA Forest Service, Rocky Mountain Region. http://www.fs.fed.us/r2/projects/scp/assessments/kitfox.pdf [Accessed 2013]. Miller, R. F., T. J. Svejcar, and N. E. West. 1994. Implications of livestock grazing in the Intermountain sagebrush region: plant composition. Pages 101–146 in M. Vavra, W. A. Laycock, and R. D. Pieper, editors. Ecological implications of livestock herbivory in the west. Society for Range Management, Denver, Colorado. Newkirk, E. S. 2014. CPW Photo Database. Colorado Parks and Wildlife, Fort Collins, Colorado, USA. http://wildlife.state.co.us/Research/Mammal/Pages/MammalsWildlifeResearch.aspx Paine, R. T. 1969. A note on trophic complexity and community stability. American Naturalist 103:91– 93. Priest, J. E. 1997. Age identification of nestling burrowing owls. Journal of Raptor Research Report 9:125–127. 15 �Rocke, T. E., N. Pussini, S. R. Smith, J. Williamson, B. Powell, and J. E. Osorio. 2010. Consumption of baits containing raccoon pox-based plague vaccines protects black-tailed prairie dogs (Cynomys ludovicianus). Vector-Borne and Zoonotic Diseases 10: 53–58. Seglund, A. E. and P. M Schnurr. 2010. Colorado Gunnison’s and white-tailed prairie dog conservation strategy. Colorado Division of Wildlife, Denver, Colorado. Sidle, J. G., M. Ball, T. Byer, J. J. Chynoweth, G. Foli, R. Hodorff, G. Moravek, R. Peterson, and D. N. Svingen. 2001. Occurrence of burrowing owls in black-tailed prairie dog colonies on Great Plains National Grasslands. Journal of Raptor Research 35:316–321. Smith, G. A. and M. V. Lomolino. 2004. Black-tailed prairie dogs and the structure of avian communities on the shortgrass plains. Oecologia 138:592–602. SPV Subcommittee. 2011. Call for partners: Phase II field studies on oral sylvatic plague vaccine (SPV) for prairie dogs. Stapp, P., M. F. Antolin, and M. Ball. 2004. Patterns of extinction in prairie dog metapopulations: plague outbreaks follow El Nino events. Frontiers in Ecology and the Environment 2:235–240. Stapp, P., B. Van Horne, and M. D. Lindquist. 2008. Ecology of mammals of the shortgrass steppe. Pages 132–180 in W. K. Lauenroth and I. C. Burke, editors. Ecology of the shortgrass steppe: a longterm perspective. Oxford University Press, New York, New York. Thomsen, L. 1971. Behavior and ecology of burrowing owls on the Oakland Municipal Airport. Condor 73:177–192. Tipton, H. C., V. J. Dreitz, and P. F. Doherty, Jr. 2008. Occupancy of mountain plover and burrowing owl in Colorado. Journal of Wildlife Management 72:1001–1006. Tripp, D. W. and T. E. Rocke. 2012. Study plan: Preliminary field trials to evaluate the safety and efficacy of an sylvatic plague vaccine for prairie dogs. Whicker, A. D. and J. K. Detling. 1988. Ecological consequences of prairie dog disturbances: prairie dogs alter grassland patch structure, nutrient cycling, and feeding-site selection by other herbivores. BioScience 38:778–785. Yackel Adams, A. A. 2000. Training notes for grassland bird project. Unpublished report. U.S. Geological Survey Fort Collins Science Center, Fort Collins, Colorado. 8 pp. Yackel Adams, A. A. Unpub data. Lark bunting nestling aging guide. U.S. Geological Survey Fort Collins Science Center, Fort Collins, Colorado. 2 pg. 16 �FIGURE LEGENDS Figure 1. Photos from BTPD and GUPD sites during 2013 – 2015. a) GUPD consuming experimental bait. b) Ferruginous hawk seen during a winter raptor count on a BTPD colony. c) Visual obstruction measurement at a GUPD site. d) Horned lark nest on a BTPD colony. e) Estimating age of a lark bunting nest by floating eggs on a BTPD colony. Figure 2. Monthly precipitation at BTPD (Larimer and Weld Co.) and GUPD (Gunnison Co.) sites from 2013 – 2015, including the 30-year averages. Data were taken from the nearest weather station. BTPD data are shown with solid lines and square symbols. GUPD data are shown with dashed lines and round symbols. 17 �TABLES Point counts Bird species Vegetation transects Vegetation species Raptor count locations Raptor count minutes Nests Remote camera locations Remote camera photos BTPD on 1,274 83 148 91 9 7,890 413 32 985,819 GUPD on 446 81 84 152 13 5,010 115 11 145,992 GUPD off TOTAL 564 2,284 80 130 89 321 155 217 53 75 7,350 20,250 15 543 N/A 43 N/A 1,131,811 Table 1. 2013 – 2015 sample sizes for BTPD and GUPD sites, on and off prairie dog colonies. Off-colony locations were randomly (re-)chosen each year. We did one point count, 3 – 5 winter raptor counts, and 5 – 9 summer raptor counts per location. We surveyed 1 – 2 vegetation transects per site. Nest search plots were surveyed at least twice. GUPD cameras were deployed during summer, and BTPD cameras were deployed year-round. Camera numbers were highest in 2015, when new cameras were added while previous years’ locations were retained. 18 �CODE SPECIES BRBL BRSP VESP BARS BHCO CORA EUST HOLA LASP MCLO RWBL WEME GRSP LARB Brewer's Blackbird Brewer's Sparrow Vesper Sparrow Barn Swallow Brown-headed Cowbird Common Raven European Starling Horned Lark Lark Sparrow McCown's Longspur Red-Winged Blackbird Western Meadowlark Grasshopper Sparrow Lark Bunting 2013 Use Rate A R+E 0.067 0.053 0.116 0.063 0.371 0.116 0.156 0.200 0.049 0.053 0.192 0.084 0.000 0.021 6.656 7.505 0.045 0.084 2.509 1.947 0.138 0.253 1.531 2.337 0.018 0.000 3.817 3.316 2014 Use Rate A R+E 0.115 0.035 0.107 0.018 0.202 0.029 0.074 0.194 0.025 0.459 0.086 0.135 0.206 0.088 3.864 3.253 0.045 0.006 1.099 0.812 0.329 0.312 1.593 1.759 0.012 0.029 2.506 5.471 2015 Use Rate A R+E 0.815 0.045 0.095 0.111 0.481 0.177 0.143 0.075 0.069 0.018 0.079 0.111 0.011 0.192 3.709 3.562 0.116 0.168 0.762 1.453 0.640 0.279 2.587 2.270 0.323 0.486 1.201 2.195 TOTAL COUNT 223 116 298 157 119 147 133 5583 108 1780 402 2496 235 3667 Table 2. Bird use rates for the most common species detected during avian point counts on BTPD prairie dog colonies with varying prairie dog activity status: Active (A), Reduced (R) after a plague event, and Extinct (E). Use rate was calculated by dividing the number of detections by the total number of points per colony per year. The first three species were more common on active colonies, those in the middle showed no preference, and the last two species were more common on reduced or extinct colonies. Data reported here do not yet account for probability of detection or the location of individual birds inside or outside of treatment area boundaries. 19 �CODE SPECIES BRSP CORA HOLA RWBL SATH VESP WEME AMRO BBMA BTLH NOFL ROWR DEJU GTTO MOCH WEWP Brewer's Sparrow Common Raven Horned Lark Red-Winged Blackbird Sage Thrasher Vesper Sparrow Western Meadowlark American Robin Black-billed Magpie Broad-tailed Hummingbird Northern Flicker Rock Wren Dark-eyed Junco Green-tailed Towhee Mountain Chickadee Western Wood-pewee 2013 Use Rate In Out 2.240 0.663 0.700 0.143 0.880 0.367 0.170 0.071 1.350 0.133 3.030 0.888 0.910 0.694 0.370 0.357 0.260 0.102 0.210 0.143 0.240 0.286 0.240 0.214 0.110 0.184 0.640 0.969 0.040 0.133 0.200 0.347 2014 Use Rate In Out 1.095 0.827 0.514 0.189 0.333 0.438 0.133 0.124 0.486 0.249 0.838 0.492 0.371 0.319 0.210 0.195 0.086 0.054 0.124 0.108 0.114 0.059 0.019 0.130 0.019 0.086 0.333 0.341 0.057 0.178 0.057 0.173 2015 Use Rate In Out 1.512 1.242 0.518 0.527 0.619 0.329 0.369 0.177 0.494 0.404 1.048 0.430 0.488 0.332 0.196 0.383 0.107 0.123 0.179 0.343 0.095 0.137 0.125 0.137 0.071 0.162 0.345 0.549 0.012 0.126 0.196 0.141 TOTAL COUNT 1155 406 435 172 440 864 431 269 107 193 129 130 104 467 93 164 Table 3. Bird use rates for the most common species detected during avian point counts on GUPD sites, comparing points located in prairie dog colonies to those 500 – 1500 m outside colony boundaries. Use rate was calculated by dividing the number of detections by the total number of points per colony per year. The first seven species were more common in colonies, those in the middle showed no preference, and the last four species were more common outside colonies. Data reported here do not yet account for probability of detection or the location of individual birds inside or outside of colony or off-colony grid boundaries. 20 �% Cover Grass Bare Litter Forb Rock Scat Cactus Shrub Other Exotic BTPD BTPD on 51.9 17.5 14.8 7.7 3.5 1.7 1.2 0.9 0.9 0.9 GUNN on 12.7 38.5 12.7 7.1 10.3 2.2 0.1 14.1 2.5 3.8 GUPD GUNN WOOD WOOD off on off 15.9 24.0 16.8 24.3 15.1 9.8 14.9 24.6 36.9 8.4 16.9 14.9 13.9 12.8 15.9 2.0 1.7 1.1 0.1 0.0 0.0 16.3 1.8 1.6 4.3 3.1 3.0 5.1 2.0 1.0 Table 4. Ground cover percentages from vegetation transects conducted on BTPD and GUPD sites, on and off prairie dog colonies, averaged from June – August 2013 – 2015. % Cover Grass Litter Bare Forb 2013 36.7 27.2 22.8 3.9 2014 55.9 8.7 19.4 8.3 2015 55.7 14.3 13.3 9 Table 5. Ground cover percentages for dominant vegetation types from 2013 – 2015 for BTPD sites in north central Colorado. 21 �BTPD Type BTPD on Grass blue grama Forb scarlet globemallow Shrub winterfat Cactus plains pricklypear Other lichen Exotic netseed lambsquarter GUNN on western wheatgrass fringed sagebrush big sagebrush plains pricklypear lichen crested wheatgrass GUPD GUNN off WOOD on junegrass blue grama fringed sagebrush fringed sagebrush big sagebrush rabbitbrush plains pricklypear N/A lichen lichen kentucky bluegrass smooth brome WOOD off blue grama fringed sagebrush common juniper N/A lichen smooth brome Table 6. Dominant plant species detected on vegetation transects at BTPD and GUPD sites, on and off prairie dog colonies in June – August 2013 – 2015. 22 �Species American Kestrel Burrowing Owl Common Raven Ferruginous Hawk Golden Eagle Loggerhead Shrike Northern Goshawk Northern Harrier Prairie Falcon Rough-legged Hawk Red-tailed Hawk Sharp-shinned Hawk Swainson’s Hawk Turkey Vulture TOTAL min 2013 Use Rate (%) BTPD GUPD GUPD in in out 1.89 0.76 2.32 13.84 0.00 0.00 6.35 11.43 74.20 0.13 0.00 0.00 2.08 5.81 0.14 0.00 0.00 0.00 0.00 0.00 0.00 0.13 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.63 2.67 0.00 0.00 0.00 0.00 8.55 0.00 2.75 19.12 0.29 0.29 1590 1050 690 2014 Use Rate (%) BTPD GUPD GUPD in in out 4.27 4.67 1.41 6.00 0.00 0.00 3.58 31.89 10.45 1.36 0.00 0.00 0.67 2.39 2.96 0.00 2.89 0.58 0.00 0.00 0.00 0.42 1.22 0.00 0.06 0.00 0.00 0.73 0.00 0.00 2.09 5.83 5.91 0.00 0.72 0.58 1.76 0.67 0.65 8.48 2.67 3.57 3300 1800 2910 2015 Use Rate (%) BTPD GUPD GUPD TOTAL in in out min 1.13 2.55 1.60 469 31.03 1.39 0.00 1379 4.67 10.79 12.53 2572 4.60 0.00 0.00 185 0.97 0.83 0.59 315 0.00 0.00 0.00 69 0.00 0.00 0.80 30 0.63 0.00 0.00 57 0.97 0.00 0.03 32 0.13 0.28 0.00 34 1.07 8.19 1.28 641 0.00 0.00 0.00 30 3.70 0.23 0.08 363 3.10 2.13 1.09 921 3000 2160 3750 20250 Table 7. Raptor use of vaccine project areas at BTPD and GUPD sites, on and off prairie dog colonies in 2013 – 2015. Use was quantified as time spent in project areas, and use rate = use minutes/total minutes in BTPD, in GUPD, and off GUPD colonies. Data include breeding counts (late April – August) at all sites and wintering counts (November – early March) at BTPD sites. Species with small sample sizes are not shown: American crow, Cooper’s hawk, and osprey (2 min each). 23 �Species American Avocet American Kestrel Brewer's Blackbird Brewer's Sparrow Burrowing Owl Common Raven Ferruginous Hawk Grasshopper Sparrow Green-tailed Towhee Horned Lark Killdeer Lark Bunting Lark Sparrow McCown's Longspur Mountain Plover Sage Thrasher Swainson's Hawk Unknown Bird Vesper Sparrow Western Kingbird Western Meadowlark TOTAL 2013 Fate (n) BTPD GUPD (0) (0) (0) (0) (0) (0) 1.00 (3) 1.00 (4) 0.71 (7) (0) (0) (0) 0.00 (1) (0) (0) (0) (0) (0) 0.55 (12) 0.00 (1) 1.00 (1) (0) 1.00 (2) (0) (0) (0) 0.31 (27) (0) 1.00 (3) (0) (0) 0.00 (1) 1.00 (2) (0) 0.00 (1) (0) (0) 0.00 (4) (0) (0) (0) (0) 0.54 (58) 0.50 (10) 2014 Fate (n) BTPD GUPD (0) (0) (0) (0) (0) (0) 0.57 (15) 0.60 (33) 0.73 (13) (0) 1.00 (1) (0) 0.00 (1) (0) (0) (0) (0) 1.00 (3) 0.38 (36) 1.00 (1) (0) (0) 0.33 (69) (0) (0) (0) 0.38 (26) (0) 0.50 (2) (0) (0) 0.69 (13) 0.00 (1) (0) 0.00 (1) (0) (0) 0.88 (8) 0.00 (1) (0) 0.50 (2) (0) 0.40 (169) 0.69 (58) 2015 Fate (n) BTPD GUPD 0.50 (2) (0) 0.67 (3) (0) (0) 0.00 (1) 0.25 (4) 0.47 (37) 0.88 (25) 0.00 (2) 1.00 (1) (0) 1.00 (1) (0) 0.50 (2) (0) (0) 0.00 (1) 0.44 (36) (0) (0) (0) 0.55 (39) (0) 0.50 (2) 1.00 (1) 0.55 (57) (0) 0.50 (3) (0) (0) 0.67 (10) 1.00 (3) (0) 0.00 (1) (0) (0) 0.71 (7) 0.00 (2) (0) 0.00 (1) (0) 0.57 (182) 0.51 (59) Fate (n) TOTAL 0.50 (2) 0.67 (3) 0.00 (1) 0.56 (96) 0.78 (47) 1.00 (2) 0.67 (3) 0.50 (2) 0.75 (4) 0.44 (86) 1.00 (1) 0.42 (110) 0.67 (3) 0.45 (110) 0.71 (8) 0.65 (24) 0.83 (6) 0.00 (3) 0.71 (19) 0.00 (3) 0.33 (3) 0.52 (536) Table 8. Nest fate (n = sample size) in vaccine project areas on BTPD and GUPD colonies during 2013 – 2015. These are naïve apparent nest survival estimates. 24 �Coyote BTPD GUPD 2013: April - July 2013: August - March 2014: April - July 2014: August - March 2015: April - July 2015: August - Oct 0.42 0.44 0.46 0.36 0.51 0.38 0.21 0.48 0.27 0.47 0.26 0.38 Badger Swift Fox BTPD GUPD BTPD GUPD 0 0 0.13 0 0.10 0.11 0.07 0 0.03 0.05 0.14 0 0.18 0.03 0.47 0 0.14 0.08 0.30 0 0.09 0 0.46 0 Table 9. Naïve (minimum) occupancy estimates (do not account for probability of detection) for carnivores based on remote camera photos taken on BTPD and GUPD colonies in 2013 – 2015. Each site had one camera in 2013, and additional cameras were deployed in 2014 – 2015. Occupancy was calculated per site over 2-week intervals. Occupancy rates are shown separately for the breeding season versus the remainder of the year. 25 �FIGURES a b c Sean Streich Sean Streich d e Miranda Middleton Miranda Middleton Figure 1. Photos from BTPD and GUPD sites during 2013 – 2015. a) GUPD consuming experimental bait. b) Ferruginous hawk seen during a winter raptor count on a BTPD colony. c) Visual obstruction measurement at a GUPD site. d) Horned lark nest on a BTPD colony. e) Estimating age of a lark bunting nest by floating eggs on a BTPD colony. 26 �Precipitation 200 180 160 140 Precipitation (mm) BTPD 2013 BTPD 2014 120 BTPD 2015 BTPD average 100 GUPD 2013 GUPD 2014 80 GUPD 2015 GUPD average 60 40 20 0 April May June July Figure 2. Monthly precipitation at BTPD (Larimer and Weld Co.) and GUPD (Gunnison Co.) sites from 2013 – 2015, including the 30-year averages. Data were taken from the nearest weather station. BTPD data are shown with solid lines and square symbols. GUPD data are shown with dashed lines and round symbols. 27 �APPENDIX 1: BIRD SPECIES LIST CODE AMAV AMCO AMCR AMGO AMGP AMKE AMRO AMWI AWPE BAEA BAIS BANS BARS BBMA BCCH BGGN BHCO BHGR BRBL BRSP BTLH BUOW BWTE CAFI CANG CASP CCLO CCSP CHSP CLNU CLSW COFL COGR COHA CONI CORA DCCO DEJU SPECIES American Avocet American Coot American Crow American Goldfinch American Golden-plover American Kestrel American Robin American Wigeon American White Pelican Bald Eagle Baird's Sparrow Bank Swallow Barn Swallow Black-billed Magpie Black-capped Chickadee Blue-gray Gnatcatcher Brown-headed Cowbird Black-headed Grosbeak Brewer's Blackbird Brewer's Sparrow Broad-tailed Hummingbird Burrowing Owl Blue-winged Teal Cassin's Finch Canada Goose Cassin's Sparrow Chestnut-collared Longspur Clay-colored Sparrow Chipping Sparrow Clark's Nutcracker Cliff Swallow Cordilleran Flycatcher Common Grackle Cooper's Hawk Common Nighthawk Common Raven Double-crested Cormorant Dark-eyed Junco BTPD Lar/Weld In x x x x x x x x x x x x Gunn In GUPD Gunn Wood Out In x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x Wood Out x x* x x x x x x x x x x x x x x x x x x x x x x x x x x 28 x x x x x x x* x x x x x x �DOWO DUFL EABL EAKI EAME ECDO EUST EVGR FEHA GADW GBHE GOEA GRAJ GRSP GTGR GTTO GUSG GWTE HAWO HETH HOLA HOSP HOWR KILL LARB LASP LBCU LEGO LISP LOSH MALL MCLO MGWA MOBL MOCH MODO MOPL NOFL NOGO NOHA NOMO NRWS NSHO Downy Woodpecker Dusky Flycatcher Eastern Bluebird Eastern Kingbird Eastern Meadowlark Eurasian Collared-Dove European Starling Evening Grosbeak Ferruginous Hawk Gadwall Great Blue Heron Golden Eagle Gray Jay Grasshopper Sparrow Great-tailed Grackle Green-tailed Towhee Gunnison Sage-grouse Green-winged Teal Hairy Woodpecker Hermit Thrush Horned Lark House Sparrow House Wren Killdeer Lark Bunting Lark Sparrow Long-billed Curlew Lesser Goldfinch Lincoln's Sparrow Loggerhead Shrike Mallard McCown's Longspur MacGillivray's Warbler Mountain Bluebird Mountain Chickadee Mourning Dove Mountain Plover Northern Flicker Northern Goshawk Northern Harrier Northern Mockingbird Northern Rough-winged Swallow Northern Shoveler x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x* x x x 29 x x x x �OSFL OSPR PIGR PISI PRFA PYNU RBGU RBNU RCKI RNDU RNEP RNSA ROPI ROWR RTHA RWBL SACR SAGS SAPH SATH SAVS SNBU SOSP SPTO SSHA STJA SWHA TOSO TRES TUVU VESP VGSW WAVI WBNU WCSP WEBL WEKI WEME WESJ WETA WEWP WHIM WIPH Olive-sided Flycatcher Osprey Pine Grosbeak Pine Siskin Prairie Falcon Pygmy Nuthatch Ring-billed Gull Red-breasted Nuthatch Ruby-crowned Kinglet Ring-necked Duck Ring-necked Pheasant Red-naped Sapsucker Rock Pigeon Rock Wren Red-tailed Hawk Red-winged Blackbird Sandhill Crane Sage Sparrow Say's Phoebe Sage Thrasher Savannah Sparrow Snow Bunting Song Sparrow Spotted Towhee Sharp-shinned Hawk Stellar's Jay Swainson's Hawk Townsend's Solitaire Tree Swallow Turkey Vulture Vesper Sparrow Violet-green Swallow Warbling Vireo White-breasted Nuthatch White-crowned Sparrow Western Bluebird Western Kingbird Western Meadowlark Western Scrub-Jay Western Tanager Western Wood-pewee Whimbrel Wilson's Phalarope x x* x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 30 x x x x x x x x x x x x x* x x x x x x x x x x x x x x x* x x* x x x* x x x x x x x x x x x x x x x x x �WISA WISN WITU YEWA YHBL YRWA TOTAL Williamson's Sapsucker Wilson's Snipe Wild Turkey Yellow Warbler Yellow-headed Blackbird Yellow-rumped Warbler x x x 83 x x x x x x x x x x x x 68 x 69 x 54 50 Table A1. Bird species list for BTPD and GUPD sites during summer 2013−2015. These species were detected during avian point counts with several exceptions. * = detected during raptor counts. Roughlegged hawks (RLHA not listed) were detected only during winter raptor counts. 31 �APPENDIX 2: PLANT SPECIES LIST Code Family Scientific Name Common Name Exotic BTPD BTPD In Gunn In GUPD Gunn Wood Out In x x x x x x x Wood Out Grasses, Sedges, and Rushes ACHY Poaceae Achnatherum hymenoides Indian Ricegrass ACLE Poaceae Achnatherum lettermanii Letterman’s Needlegrass AGCR Poaceae Agropyron cristatum Crested Wheatgrass x x AGST Poaceae Agrostis stolonifera Creeping Bentgrass x x ARPU Poaceae Aristida purpurea Purple Threeawn x x BODA Poaceae Bouteloua dactyloides Buffalograss x x x x BOGR Poaceae Bouteloua gracillis Blue Grama x x x x x BRCI Poaceae Bromus ciliatus Fringed Brome x x x BRHO Poaceae Bromus hordeaceus Soft Brome x BRIN Poaceae Bromus inermis Smooth Brome x BRPO Poaceae Bromus porteri Porter Brome BRTE Poaceae Bromus tectorum Cheatgrass CAIN Cyperaceae Carex inops ssp. heliophila Sun Sedge x CALO Poaceae Calamovilfa longifolia Prairie Sandreed CARE Cyperaceae Carex Spp. Sedge Spp. (Unidentified) ELEL Poaceae Elymus elymoides Squirreltail ELGL Poaceae Elymus glaucus Blue Wildrye ELRE Poaceae Elymus repens Quackgrass ELTR Poaceae Elymus trachycaulus Slender Wheatgrass FEAR Poaceae Festuca arizonica Arizona Fescue FEID Poaceae Festuca idahoensis Idaho Fescue FEST Poaceae Festuca Spp. Unknown Fescue x x HECO Poaceae Hesperostipa comata Needle & Thread Grass x x x HOJU Poaceae Hordeum jubatum Foxtail Barley x x x JUBA Juncaceae Juncus balticus Baltic Rush x x 32 x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x �KOMA Poaceae Koeleria macrantha Junegrass LOPE Poaceae Lolium perenne Annual Ryegrass MUHL Poaceae Muhlenbergia Spp. Muhlenbergia Spp. MUMO Poaceae Muhlenbergia montana Mountain Muhly MUTO Poaceae Muhlenbergia torreyi Ring Muhly NAVI Poaceae Nassella viridula Green Needlegrass PASM Poaceae Pascopyrum smithii Western Wheatgrass PHPR Poaceae Phleum pratense Timothy Grass PIMI Poaceae Piptatherum micranthum POFE Poaceae Poa fendleriana POPR Poaceae Poa pratensis Kentucky Bluegrass RUSH Juncaceae Juncus Spp. Rush Spp. SCPA Poaceae Schedonnardus paniculatus Tumblegrass SCPR Poaceae Schedonorus pratensis Meadow Fescue SCSC Poaceae Schizachyrium scoparium Little Bluestem SOBI Poaceae Sorghum bicolor Sorghum SPCR Poaceae Sporobolus cryptandrus Sand Dropseed x VUOC Poaceae Vulpia octoflora Six Weeks Fescue x ACMI Asteraceae Achillea millefolium Common Yarrow ACRE Asteraceae Acroptilon repens Russian Knapweed x AMBL Amaranthaceae Amaranthus blitoides Prostrate Pigweed x ANAR Apiaceae Angelica arguta White Angelica x ANMI Asteraceae Antennaria microphylla Littleleaf Pussytoes ANMU Ranunculaceae Anemone multifida Cutleaf Anemone ANSE Primulaceae Androsace septentrionalis Pygmy-flower Rock-Jasmine ARAB Asteraceae Artemisia absinthium Absinth Wormwood ARAN Rosaceae Argentina anserina Silver Weed ARCA Asteraceae Artemisia campestris Field Sagewort ARCO Caryophyllaceae Arenaria congesta Ballhead Sandwort ARFE Caryophyllaceae Arenaria fendleri Fendler's Sandwort x x x x x x x x x x x x x x x x x x x x x x x Littleseed Ricegrass x x x x Muttongrass x x x x x x x x x x x x x x x x x x x x x x x Forbs 33 x x x x x x x x x x x x x x x x x x x x x x x x x x �ARFR Asteraceae Artemisia frigida Fringed Sagebrush ARLU Asteraceae Artesmia ludoviciana Prairie Sage x x x ARPO Papavaraceae Argemone polyanthemos Crested/ Annual Pricklepoppy ARUV Ericaceae Arctostaphylos uva-ursi Bearberry (Kinnikinnick) ASBI Fabaceae Astragalus bisulcatus Two-grooved Milkvetch x x x ASDR Fabaceae Astragalus drummondii Drummond's Milkvetch x x x ASMI Fabaceae Astragalus miser Timber Milkvetch x ASPA Fabaceae Astragalus parryii Parry's Milkvetch x ASSH Fabaceae Astralagus shortianus Short's Milkvetch BASC Chenopodiaceae Bassia scoparia Kochia/ Mexican Fireweed BEPL Scrophulariaceae Besseya plantaginea White River Coral Drops CALI Scrophulariaceae Castilleja linariifolia Narrowleaf Paintbrush x CAMI Scrophulariaceae Castilleja miniata Scarlet Paintbrush CHBE Chenopodiaceae Chenopodium berlandieri Netseed Lambsquarter CHEN Chenopodiaceae Chenopodium Spp. CHGL Euphorbiaceae CHLE CHSE x x x x x x x x x x x x x x Goosefoot Spp. (Unidentified) x x Chamaesyce glyptosperma Small Ribseed Sandmat x x Chenopodiaceae Chenopodium leptophyllum Narrowleaf Goosefoot Euphorbiaceae Chamaesyce serpyllifolia Thyme-leaf spurge x CHWA Chenopodiaceae Chenopodium watsonii Watson's Goosefoot x CIAR Asteraceae Cirsium arvense Canada Thistle CICA Asteraceae Cirsium canescens Prairie/ Plains/ Creamy Thistle CIUN Asteraceae Cirsium undulatum Wavyleaf Thistle COAR Convolvulaceae Convolvulus arvensis Field Bindweed CRTH Boraginaceae Cryptantha thyrsiflora Calcareous Cryptantha DESO Brassicaceae Descurainia sophia Pinnate Tansy Mustard EQAR Equisetaceae Equisetum arvense Field Horsetail ERAS Brassicaceae Erysimum asperum Western Wallflower ERCA Asteraceae Erigeron canus Hoary Fleabane x ERCE Polygonaceae Eriogonum cernuum Nodding Buckwheat x ERCO Asteraceae Erigeron compositus Cutleaf Daisy ERFL Asteraceae Erigeron flagellaris Trailing Fleabane 34 x x x x x x x x x x x x x x x x x x x x x x x x x x x �ERLO Polygonaceae Eriogonum lonchophyllum Spearleaf Buckwheat x EROV Polygonaceae Eriogonum ovalifolium Cushion Buckwheat x ERRA Polygonaceae Eriogonum racemosum Redroot Buckwheat x x ERSP Asteraceae Erigeron speciosus Showy Fleabane x x ERST Asteraceae Erigeron strigosus Prairie Fleabane x x ERUMA Polygonaceae Eriogonum umbellatum v. aureum Sulphur Buckwheat x x x ERUMM Polygonaceae Eriogonum umbellatum v. majus Creamy Buckwheat x x x FRVE Rosaceae Fragaria vesca Woodland Strawberry x x FRSP Gentianaceae Frasera speciosa Monument Plant x GABO Rubiaceae Galium boreale Bedstraw x GECA Geraniaceae Geranium caespitosum Pineywoods Geranium GETR Rosaceae Geum triflorum Old Man's Whiskers GEVI Geraniaceae Geranium viscosissimum Sticky Purple Geranium GRSQ Asteraceae Grindelia squarrosa Curlycup Gumweed x x GUSA Asteraceae Gutierrezia sarothrae Broom Snakeweed x x x HAFL Boraginaceae Hackelia floribunda Many-Flowered Stickseed x x HEPA Saxifragaceae Heuchera parvifolia Littleleaf Alumroot x x HEUC Saxifragaceae Heuchera Spp. Alumroot Spp. (Unidentified) HEVI Asteraceae Heterotheca villosa Hairy False Golden Aster HYFI Asteraceae Hymenopappus filifolius Fineleaf Hymenopappus IRMI Iridaceae Iris missouriensis Rocky Mountain Iris IVAX Asteraceae Iva axillaris Povertyweed x LAOC Boraginaceae Lappula occidentalis Western Sticktight/ Stickweed x LASC Asteraceae Laennecia schiedeana Pineland Horseweed x LEDE Brassicaceae Lepidium densiflorum Common Pepperweed LERA Brassicaceae Lepidium ramosissimum Many-branched Pepperweed LEVI Brassicaceae Lepidium virginicum Virginia Pepperweed LEVU Asteraceae Leucanthemum vulgare Ox Eye Daisy LILE Linaceae Linum lewisii Blue Flax LIPU Asteraceae Liatris punctata Gayfeather/ Blazing Star LUAR Fabaceae Lupinus argenteus Silvery Lupine 35 x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x �LUWY Fabaceae Lupinus wyethii Wyeth's Lupine LYJU Asteraceae Lygodesmia juncea Skeletonweed x MAPI Asteraceae Machaeranthera pinnatifida Lacy Tansyaster x MARA Liliaceae Maianthemum racemosum Feathery False Solomon's Seal MARE Berberidaceae Mahonia repens Oregon Grape MATA Asteraceae Machaeranthera tanacetifolia Tanseyleaf Tansyaster MEAR Lamiaceae Mentha arvensis Wild Mint MEHU Boraginaceae Mertensia humilis Mountain Bluebells MELA Boraginaceae Mertensia lanceolata Prairie Bluebells MEOF Fabaceae Melilotus officinalis Yellow Sweetclover x x MESA Fabaceae Medicago sativa Alfalfa x x MILI Nyctaginaceae Mirabilis linearis Narrowleaf Four o'clock OECA Onagraceae Oenothera caespitosa Tufted Evening Primrose OECO Onagraceae Oenothera coronopifolia Crownleaf Evening Primrose x OESU Onagraceae Oenothera suffrutescens Scarlet Beeblossom x ORLU Orobanchaceae Orobanche ludoviciana Louisiana Broomrape ORLU2 Scrophulariaceae Orthocarpus luteus Yellow Owl's Clover x OXLA Fabaceae Oxytropis lambertii Lambert Crazyweed x OXSE Fabaceae Oxytropis sericea White Locoweed PEBA Scrophulariaceae Penstemon barbatus Beardlip Beardtongue PECR Scrophulariaceae Penstemon crandallii Crandall's Beardtongue PHBE Brassicaceae Physaria bellii Front Range Twinpod x PHHO Polemoniaceae Phlox hoodii Spiny Phlox x PHLO Polemoniaceae Phlox longifolia Longleaf Phlox PIOP Asteraceae Picradeniopsis oppositifolia Opposite Leaf Bahia PLMA Plantaginaceae Plantago major Common Plantain PLPA Plantaginaceae Plantago patagonica Woolly Plantain POGR Rosaceae Potentilla gracilis Slender Cinquefoil POHI Rosaceae Potentilla hippiana Woolly Cinquefoil POOL Portulacaceae Portulaca oleracea Common Purslane/ Hogweed x POPE Polygonaceae Polygonum persicaria Spotted Ladysthumb x 36 x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x �PSLA Fabaceae Psoralidium lanceolatum Lemon Scurfpea x RACO Asteraceae Ratibida columnifera Upright Prairie Coneflower x RHRO Crassulaceae Rhodiola rosea King's Crown x SATR Chenopodiaceae Salsola tragus Russian Thistle / Tumbleweed x SAXI Saxifragaceae Saxifraga Spp. Saxifrage Spp. (Unidentified) x x SEIN Asteraceae Senecio integerrimus Lambstongue Ragwort x x SELA Crassulaceae Sedum lanceolatum Spearleaf Stonecrop x x SENE Asteraceae Senecio Spp. Groundsel Spp. (Unidentified) SOMI Asteraceae Solidago missouriensis Missouri Goldenrod SONU Fabaceae Sophora nuttalliana Silky Sophora x SOTR Solanaceae Solanum triflorum Cutleaf Nightshade x x SPCO Malvaceae Sphaeralcea coccinea Scarlet Globemallow x x SPFA Euphorbiaceae Spurge (Unidentified) x SYLA Asteraceae Symphyotrichum lanceolatum White Panicle Aster TAOF Asteraceae Taraxacum officinale Dandelion TAVU Asteraceae Tanacetum vulgare Common Tansy THAR Brassicaceae Thlaspi arvense Field Pennycress THRH Fabaceae Thermopsis rhombifolia Golden Pea TRDU Asteraceae Tragopogon dubius Yellow Salsify x TRHY Fabaceae Trifolium hybridum Alsike Clover x TRPR Fabaceae Trifolium pratense Red Clover x URDI Urticaceae Urtica gracilis Stinging Nettle x VIAM Fabaceae Vicia americana American Vetch x VICI Fabaceae Vicia Spp. Vetch Spp. (Unidentified) WYAM Asteraceae Wyethia amplexicaulis Mule-ears ACGL Aceraceae Acer glabrum Mountain Maple AMAL Rosaceae Amelanchier alnifolia Serviceberry ARCA Asteraceae Artemisia cana Silver Sagebrush ARTR Asteraceae Artemisia tridentata Big Sagebrush x ATCA Chenopodiaceae Atriplex canescens Fourwing Saltbush x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x Shrubs 37 x x x x x x x x x �CEMO Rosaceae Cercocarpus montanus Mountain Mahogany x x x x x CHVI Asteraceae Chrysothamnus viscidiflorus Douglas Rabbitbrush x x x x x DAFR Rosaceae Dasiphora fruticosa Shrubby Cinquefoil x x x x EREF Polygonaceae Eriogonum effusum Spreading Buckwheat x ERNA Asteraceae Ericameria nauseosa Rubber /Grey Rabbitbrush x x x x HODU Rosaceae Holodiscus dumosus Rockspirea JUCO Cupressaceae Juniperus communis ssp. alpina Common Juniper x x JUSC Cupressaceae Juniperus scopulorum Rocky Mountain Juniper x x KRLA Chenopodiaceae Krascheninnikovia lanata Winter Fat x x PRAM Rosaceae Prunus americana American Plum x x PRVI Rosaceae Prunus virginiana Western Chokecherry x x PUTR Rosaceae Purshia tridentata Bitter Brush x x RHTR Anacardiaceae Rhus trilobata Skunkbrush Sumac x x RIAU Grossulariaceae Ribes aureum Golden Currant x x x RICE Grossulariaceae Ribes cereum Wax Currant x ROAR Rosaceae Rosa arkansana Prairie/ Wild Rose x ROWO Rosaceae Rosa woodsii Woods' Rose RUID Rosaceae Rubus idaeus Wild Red Raspberry SALI Salicaceae Salix Spp. Willow Spp. (Unidentified) SYOC Caprifoliaceae Symphoricarpos occidentalis Western Snowberry TECA Asteraceae Tetradymia canescens Spineless Horsebrush x XANT Asteraceae Xanthium spp. Cocklebur sp. x YUGL Agavaceae Yucca glauca Great Plains Yucca x ABLA Pinaceae Abies lasiocarpa Subalpine Fir PIEN Pinaceae Picea engelmannii Engelmann Spruce PIFL Pinaceae Pinus flexilis Limber Pine PIPO Pinaceae Pinus ponderosa Ponderosa Pine PIPU Pinaceae Picea pungens Blue Spruce POAN Salicaceae Populus angustifolia Narrow-leaved Cottonwood PODE Salicaceae Populus deltoides ssp. wislizenii Rio Grande Cottonwood x x x x x x x x x x x x x x x x x x x x x x x x x Trees 38 x x x x x x x x x x x x x x x x x x x �POTR Salicaceae Populus tremuloides Quaking Aspen x x PSME Pinaceae Pseudotsuga menziesii Douglas Fir x x x x Escobaria vivipara Pincushion Cactus x FUNG Fungi x x x LICH Lichen x x x x x MOSS Moss x x x x x 74 75 x Cacti and Other Plants ESVI Cactaceae OPFR Cactaceae Opuntia fragilis Brittle Pricklypear OPPO Cactaceae Opuntia polyacantha Plains Pricklypear PESI Cactaceae Pediocactus simpsonii Mountain Ball Cactus Total x x x x Table A2. Plant species list for BTPD and GUPD sites on and off prairie dog colonies for 2013−2015. 39 x x 33 217 species x 91 127 127 � https://cpw.cvlcollections.org/files/original/1171a1cd2436b2def8131c52577a67c8.pdf 2698fc33204a411287eb495acddbc297 PDF Text Text Colorado Division of Parks and Wildlife July 2015-June 2016 WILDLIFE RESEARCH REPORT State of: Cost Center: Work Package: Task No.: Colorado 3420 1680 N/A Federal Aid Project No. N/A : : : : Division of Parks and Wildlife Avian Research Bird Conservation Avian response to plague management on Colorado prairie dog colonies Period Covered: September 1, 2015 – August 31, 2016 Author: R. Yale Conrey Principle Investigators: R. Yale Conrey, D. Tripp, J. Gammonley, CPW; A. Panjabi, E. Youngberg, Bird Conservancy of the Rockies (formerly Rocky Mountain Bird Observatory). Collaborators: Miranda Middleton (CPW), Bird Conservancy of the Rockies (formerly Rocky Mountain Bird Observatory), City of Fort Collins Natural Areas and Utilities Programs, Michael Wunder and Allison Pierce (University of Colorado Denver), Bureau of Land Management (Cañon City office), and CPW wildlife managers, biologists, park rangers, and property technicians from Areas 1 and 4. All information in this report is preliminary and subject to further evaluation. Information MAY NOT BE PUBLISHED OR QUOTED without permission of the author. Manipulation of these data beyond that contained in this report is discouraged. EXTENDED ABSTRACT Range-wide declines in prairie dog (Cynomys sp.) populations have occurred, and the largest limiting factor in recent decades appears to be the high mortality and colony extirpation associated with plague (Antolin et al. 2002), caused by the bacterium Yersinia pestis. Prairie dog colonies support a diverse community of associated species, many of which are not susceptible to plague but may be indirectly affected. In order to conserve prairie dogs and species associated with their colonies, principally the black-footed ferret (Mustela nigripes), a plague vaccination program is being developed (Fig. 1a), which may also benefit a suite of species (Fig. 1b, d) listed in the Conservation Plan for Grassland Species in Colorado (Colorado Division of Wildlife 2003) and the Colorado Sagebrush Conservation Assessment and Strategy (Boyle and Reeder 2005). CPW is involved in a multi-state, multi-agency study of prairie dogs and associated small mammal species; the objective is to determine whether survival is enhanced by the experimental vaccine compared to use of placebo or insecticide to control fleas, an important vector of plague. They are also interested in how patterns of prairie dog abundance and occupancy change when plague epizootics occur in plots receiving different treatments. As an extension to this project, we initiated research in 2013 on the effects of plague management on avian species associated with prairie dog colonies, with particular focus on species of concern. Our main long-term objective is to determine whether areas treated to control plague differ from untreated areas in their avian communities. Shorter-term objectives are to 1) Determine how plague affects avian species and their 1 �predators associated with prairie dog colonies; 2) Determine whether avian species associations exist for colonies of Gunnison’s prairie dogs (C. gunnisoni: GUPD); most evidence for associated species comes from black-tailed prairie dogs (C. ludovicianus: BTPD); 3) Determine whether insecticidal dusting influences bird density or nest survival; 4) Evaluate the importance of covariates such as weather and cattle grazing. Study areas in 2013–2015 included BTPD colonies in north-central Colorado and GUPD colonies in west-central Colorado, but in 2016, we replaced the western Colorado location with South Park, a high elevation site in central Colorado. Study sites were characterized by short and mid-grasses, dominated by blue grama (Bouteloua gracilis). Our first three years of research (2013−2015) coincided with Wildlife Health’s 3-year efficacy trials for an oral plague vaccine for prairie dogs (Fig. 1a). In Colorado, CPW Wildlife Health Program staff led by Dan Tripp surveyed colonies before and after bait distribution and conducted a mark-recapture study of prairie dogs and associated small mammal species. Treated areas were arranged in triplets with one vaccine, placebo, and dusted site per group; baited sites were assigned vaccine or placebo baits in a blind procedure. In 2016, CPW Wildlife Health did further research on bait distribution and design, but the vaccine was in the initial stages of becoming a management tool, more broadly used in ferret reintroduction sites and in some high priority GUPD colonies. We continued work at our BTPD site in 2016, and with our collaborators from Bird Conservancy of the Rockies, we now have 11 years of point count data spanning two plague cycles. In 2015, we completed three years of work comparing on- and off-colony study areas at GUPD sites in western Colorado, and in 2016 shifted our focus to South Park, an area with locally high densities of mountain plover (Charadrius montanus) and a possible reintroduction site for GUPD, which have been extirpated from most formerly occupied areas. South Park contains several small active GUPD colonies, and large areas of potentially suitable habitat with no prairie dogs. We continued to conduct avian point counts in 2016, and bird occupancy, density, and species composition will be estimated from these data. Summer and winter counts of diurnal raptors and early season passive and call-playback surveys of mountain plover (Charadrius montanus) and burrowing owls (Athene cunicularia: Fig. 1d) were used to sample high priority species that are rarely detected during point counts. We searched for nests of these species as well, but did not repeat the intensive passerine nest monitoring program of previous years. We continued to collect remote camera data, which will be used to estimate summer and winter on-colony occupancy rates for mammalian carnivores, including coyotes, badgers, and swift fox (Fig. 1e). Finally, we again quantified percent ground cover, visual obstruction (Fig. 1c), and species composition of vegetation at points, nests, and along randomly located transects. Since fall 2013, plague epizootics have impacted ~80% of the BTPD study area. In September and October 2014 and 2015, black-footed ferrets were released in three BTPD study colonies. Precipitation (Fig. 2) has varied greatly during this study, particularly on BTPD sites, from slightly dry to very wet, compared to the 30-year average. At this point, all data analyses are preliminary. From 2013 – 2016, we detected 137 bird species during the breeding season. During BTPD colony surveys, at least three bird species (Brewer’s blackbird Euphagus cyanocephalus, horned lark Eremophila alpestris, and vesper sparrow Pooecetes gramineus) appeared to have higher detection rates on active prairie dog colonies, while three species (European starling Sturnus vulgaris, grasshopper sparrow Ammodramus savannarum, and lark bunting Calamospiza melanocorys) appeared to have higher detection rates on colonies with extinct or severely reduced prairie dog populations following plague outbreaks (Table 2). Trend analysis of point count data from 2006−2016, which included two plague events, is ongoing with our collaborators. We documented 230 plant species over four years. Colonies contained a higher bare ground component with lower vegetation heights than off-colony sites, with shortgrasses dominant at BTPD sites, short and mid-grasses at GUPD sites in South Park, and a more even distribution of grasses, forbs, and shrubs at GUPD sites in western Colorado. Vegetation species composition was highly variable at BTPD sites over time, with increasing grass cover and decreasing bare ground following an El Niño event associated with high rainfall during the growing season (Table 4, Fig. 2). We detected 18 raptor 2 �species during on- and off-colony counts. Burrowing owls, northern harriers Circus cyaneus, ferruginous hawks Buteo regalis (Fig. 1b), and rough-legged hawks Buteo lagopus were detected only on prairie dog colonies, and half the raptor species we observed used on-colony areas more than off-colony areas during our surveys, with the remainder appearing to show no preference or having low sample size (Table 6). Patterns in the raptor data on active colonies vs. plagued colonies have not yet been examined. Despite three rounds of nest searching for mountain plover in South Park, nests were found only in a known area of high density in James Mark Jones State Wildlife Area, which was monitored by our collaborators at University of Colorado − Denver. However, the number of burrowing owl nests at our BTPD site has doubled during each year of our study following an El Niño event and widespread plague, which may have created conditions with high prey and burrow availability in a landscape that continues to contain many active (if small) prairie dog colonies. Finally, in > 10.8 million remote camera photos, we have documented use of colonies by 8 species of mammalian carnivores, with increasing swift fox activity over the course of this study. Swift fox occurred only on BTPD colonies and badgers were more commonly detected there, while coyotes were equally common across all sites on BTPD and GUPD colonies. These species are known nest predators with large home ranges and carry fleas that could potentially move plague across a broad area. This was the fourth year of data collection on this project, and it will likely take additional years of monitoring to detect potential changes in the avian community caused by different types of plague management, as treated colonies no longer experience extinction events. Regardless of the efficacy of plague vaccine versus insecticide in reducing plague impacts, the vaccine will continue to be an important tool due to cost/benefit of its use and increasing evidence that fleas are evolving resistance to deltamethrin. Preliminary data suggest that bird densities do vary according to the status of prairie dogs on a colony, with differences between active colonies and those with extirpated or severely reduced prairie dog populations following plague outbreaks. Vegetation surveys have also identified differences between on- and off-colony areas. Raptor and camera data collection will continue through winter 2016/2017. During the 2017 breeding season, we hope to continue collecting point count, raptor count, and vegetation data on BTPD colonies with different activity levels, following the extensive 2013−2015 plague outbreak. Beyond that, there are several possibilities for continued research. The GUPD reintroduction planned for South Park is on hold until new funding is obtained; our avian work on GUPD sites may similarly go on hiatus. We may work more in ferret reintroduction sites, on private lands, or in a new location with little or no plague management (such as Pawnee National Grassland) for comparison with our current BTPD site north of Fort Collins. 3 �COLORADO PARKS AND WILDLIFE RESEARCH REPORT AVIAN RESPONSE TO PLAGUE MANAGEMENT ON COLORADO PRAIRIE DOG COLONIES REESA C. YALE CONREY INTRODUCTION Wildlife diseases are important to conservation and population dynamics of susceptible species and may also have large indirect effects on non-susceptible species (Antolin et al. 2002). Introduced pathogens have the potential for far-reaching effects on native ecosystems that go beyond the mortality of infected individuals, particularly when a keystone species (Paine 1969) or ecosystem engineer (Jones et al. 1994) is infected. Range-wide declines in prairie dog (Cynomys sp.) populations have occurred, and the largest limiting factor in recent decades appears to be the high mortality and colony extirpation associated with introduced plague (Antolin et al. 2002), caused by the bacterium Yersinia pestis. Plague epidemics were first reported in the western United States in 1899 (Dicke 1926) and in northern Colorado in 1948 (Ecke and Johnson 1952). Instead of living in extensive colonies as they once did, prairie dogs exist in metapopulations of smaller colonies that periodically go extinct and are recolonized (Antolin et al. 2002, Stapp et al. 2004). Prairie dog colonies support a diverse community of associated species (Lomolino and Smith 2004, Smith and Lomolino 2004, Hardwicke 2006, Stapp et al. 2008), many of which are not susceptible to plague but may be indirectly affected. In order to conserve prairie dogs and species associated with their colonies, principally the blackfooted ferret (Mustela nigripes), a plague vaccination program is being tested (Fig. 1a). Additional species that may benefit from this program include those listed in the Conservation Plan for Grassland Species in Colorado (Colorado Division of Wildlife 2003): burrowing owl (Athene cunicularia: BUOW, Fig. 1d), mountain plover (Charadrius montanus: MOPL), ferruginous hawk (Buteo regalis: FEHA, Fig. 1b), and swift fox (Vulpes velox) and in the Colorado Sagebrush Conservation Assessment and Strategy (Boyle and Reeder 2005): Brewer’s sparrow (Spizella breweri: BRSP), green-tailed towhee (Pipilo chlorurus: GTTO), sage sparrow (Artemisiospiza belli: SAGS), sage thrasher (Oreoscoptes montanus: SATH), and vesper sparrow (Pooecetes gramineus: VESP), as well as BUOW. BUOW and MOPL are known to decline or disappear on colonies that are not reoccupied by prairie dogs after plague epizootics (Butts and Lewis 1982; Sidle et al. 2001; Augustine et al. 2008; Tipton et al. 2008; Conrey 2010), and horned lark (Eremophila alpestris), McCown’s longspur (Rhynchophanes mccownii), golden eagle (Aquila chrysaetos), prairie falcon (Falco mexicanus), and other birds may benefit from active colonies. From 2013‒2015, researchers in several western states field-tested the uptake and efficacy (SPV Subcommittee 2011) of a new sylvatic plague vaccine (SPV) for prairie dogs (Rocke et al. 2010); the objective is to determine whether survival of prairie dogs and other small mammals is enhanced by the experimental vaccine compared to use of placebo or insecticide to control fleas, an important vector of plague. In Colorado, CPW researchers led by Dan Tripp surveyed colonies before and after bait distribution and conducted a mark-recapture study of prairie dogs and associated small mammal species (Tripp and Rocke 2012). Data analyses are ongoing. As an extension to this project, we initiated research in 2013 on the effects of plague management on avian species associated with prairie dog colonies, with particular focus on species of concern. Our main long-term objective is to determine whether areas treated to control plague differ from untreated areas in their avian communities. Shorter-term objectives are to 1) Determine how plague affects avian species and their predators associated with prairie dog colonies; 2) Determine whether avian species associations exist for colonies of Gunnison’s prairie dogs (C. gunnisoni: GUPD); most evidence for associated species comes from black-tailed prairie dogs (C. ludovicianus: BTPD; Lomolino and Smith 2004, Smith and Lomolino 2004); 3) Determine whether insecticidal dusting 4 �influences bird density or nest survival; 4) Evaluate the importance of covariates such as weather and cattle grazing. We continued work at our BTPD site in 2016 and have now collected one year of pre- and three years of post-treatment data on birds. Our collaborators at Bird Conservancy of the Rockies have a point count dataset at this site extending back to 2006. This location north of Fort Collins is also a ferret reintroduction site, and CPW Wildlife Health has continued to research bait design and distribution there. In 2015, we completed three years of work comparing on- and off-colony study areas at GUPD sites in western Colorado, and in 2016 shifted our focus to South Park, an area with locally high densities of mountain plover and a possible reintroduction site for GUPD, which have been extirpated from most formerly occupied areas. METHODS Study Area Study areas included BTPD colonies in north-central Colorado and GUPD colonies and a possible reintroduction site in central Colorado. Baited sites received either vaccine or placebo baits in a blind procedure. Project areas that were selected for the prairie dog vaccine study had adequate numbers of prairie dogs and good access. BTPD (Larimer and Weld Co.) – Study colonies were located in Larimer and Weld Co. adjacent to the Wyoming border at Soapstone Prairie Natural Area (SOAP), managed by City of Fort Collins Natural Areas Program and Meadow Springs Ranch (MSR), managed by City of Fort Collins Utilities Department. These sites are characterized by shortgrass and mixed-grass prairie dominated by grasses (blue grama Bouteloua gracilis and buffalograss B. dactyloides) with smaller amounts of native (scarlet globemallow Sphaeralcea coccinea) and non-native forbs, shrubs, and cactus. Sites were sometimes grazed by cattle at low densities, and some non-baited sites were dusted with deltamethrin to control fleas (and plague). There was much more cattle grazing in 2014 − 2016 than in 2013, because 2014 – 2015 were wet years (Fig. 2), and the high levels of forage persisted into 2016. Both properties were closed to recreational shooting. Mark-resight estimates of BTPD density on shortgrass prairie in Colorado average approximately 10 prairie dogs/acre (Magle et al. 2007). Bird, vegetation, and camera surveys were conducted on 10 SOAP colonies and 22 MSR colonies. There were nine vaccine project areas where raptors, predators, and passerine nests were surveyed: three prairie dog complexes each received vaccine, placebo, and dusting treatments from 2013 − 2015 (3 treatments*3 complexes = 9 project areas). In 2016, colonies near ferret reintroduction sites were dusted and vaccinated, the other MSR treatment areas received only vaccine, and the remaining colonies within the study area received no treatment. The following colony areas were mapped prior to the 2013 season, but active acreage was significantly less by 2016, following plague epizootics: 1) The Jack Springs (Jac) colony spanning the SPNA/MSR border contained 100 vaccine acres, 100 control acres, and ~280 dusted acres separated by 200 - 400 m buffer zones. 2) The Barton complex in MSR contained ~130 vaccine acres and ~180 control acres, encompassing much of the Barton south and west colonies (BarS and BarW). Raptor, predator, and passerine nest surveys were also done in the 140 acre Barton east colony (BarE), which received no treatment. 3) The Ferret Center complex in MSR contained 40 vaccine acres, 40 control acres, and ~478 dusted acres, encompassing the entire North Bulger south colony and half of the Ferret Center (Fer) colony, with a 400 m buffer zone separating those treatments. GUPD (South Park) – Study sites were located in South Park (Park Co.), which contains several small active GUPD colonies and large areas of potentially suitable habitat with no prairie dogs. South Park sites were managed by the Bureau of Land Management (BLM) and Colorado Parks and Wildlife (James Mark Jones State Wildlife Area, Tomahawk SWA, Charlie Meyers SWA, Spinney Mountain SWA, Spinney Mountain State Park, and Eleven Mile SP). These high elevation sites (~3000 m) are 5 �characterized by shortgrass and mixed-grass prairie dominated by grasses (blue grama and plains bluegrass Poa arida) with a relatively high amount of bare ground and forb cover (fringed sage Artemisia frigida). This is a large open mountain “park” that is isolated by surrounding forested areas and higher mountain peaks. Sites were sometimes grazed by cattle at low densities, and all sites received vaccine baits and were dusted with deltamethrin to control fleas (and plague) in 2016. Shooting was uncommon at our sites. The study area is within the known range of plague (Tripp et al. unpublished data). Bird, vegetation, and camera surveys were conducted on five colonies and six plots within the proposed reintroduction area. Because of their small size and conservation importance, entire colonies were treated with both vaccine and insecticide. Colonies were located within 2 km of Spinney Mountain and Elevenmile Canyon Reservoirs: Spinney South (92 acres), Spinney Dam (28 acres), Charlie Meyers (94 acres), Cross Creek (29 acres), and 11 Mile (84 acres). James Mark Jones and Tomahawk SWAs are 17,429 and 1,655 acres, respectively, so we sampled six plots within the SWAs. Three 1 km2 plots were randomly located within the area where Allison Pierce (UC-Denver) found the majority of her plover nests in 2015. Three additional plots (~ 1.5 km2) were non-randomly assigned to the remaining portions of JMJ and Tomahawk that we mapped (with the assistance of Michelle Flenner in CPW’s GIS group) as potentially suitable for mountain plover and GUPD, based on terrain and vegetation cover. Avian Point Counts Each point was surveyed once in May – June. At BTPD study sites, a 250 m grid of points had already been established and surveyed by BCR from 2006−2013. At GUPD study sites, we created a 250 m grid of points and surveyed those within 100 m of colonies and within the selected plots on JMJ and Tomahawk SWAs. Points were considered to be “on colony” if located within 100 m of the boundary and with good views of the colony. Most point counts were conducted between dawn and 10:00 and were never conducted in rain, hot temperatures (above 30°C), or high winds that made it difficult to hear birds. Regardless of time or weather, we did not conduct counts if we noticed that bird activity (especially singing and calling) was dropping off. In addition to these point counts, we conducted separate counts for burrowing owls, which are poorly detected in regular point counts. We used a similar protocol for MOPL and BUOW in 2015, but this did not yield many MOPL nests, and following three years of very limited MOPL detections, we did not repeat that protocol for MOPL in 2016. We did a 3 min. passive survey followed by 3 min. of callplayback at grid locations spaced at least 700 m to avoid calling in birds and double-counting them, so that the whole colony was visible within 500 m of a survey point. The call-playback sequence was 3 min. of silence and 3 min. of owl calls (two 30-sec. segments of coo call and 30-sec. of alarm call, each separated by 30-sec. of silence). Surveys were conducted from trucks or ATVs where possible, because owls are much less likely to be disturbed by observers in vehicles than by observers on foot. Where not possible, surveys on foot were completed from colony edges or other locations prior to moving through them and disturbing owls. BUOW counts were conducted three times within a 2-week period by at least two different observers, in order to facilitate analysis using a removal method, and they were immediately followed by nest searching where owls were detected. We conducted 6-min.breeding season point counts, recording each bird’s species, horizontal (radial) distance, sex (if known), use of the prairie dog colony (yes or no), minute of detection (1 – 6), and how it was detected (visual, singing, calling, drumming, fly-over, or other). Membership in a cluster was noted, typically for male-female pairs. After completing the bird count, we recorded weather and site characteristics at each point, including time, temperature, wind speed, cloud cover, management type (typically cattle grazing, sometimes dusting) and whether it was current, from this season or last season, and the presence of excessive noise, roads (primary or secondary), and cliffs or rock outcroppings within 100 m. Within a 50 m radius, we recorded characteristics of tall nesting and perching substrate, including percent cover, height, and dominant species for overstory plants ≥ 3 m and shrubs > 30 cm but < 3 m. 6 �Within a 5 m radius, we recorded characteristics of ground cover, including percent cover of grasses (including sedges and rushes), forbs, shrubs, cactus, litter, bare ground, rock, scat, other cover such as lichen, and exotic species. We also recorded the dominant exotic species, and the mean height and species of the dominant grass. We used the point count protocol designed for Integrated Monitoring of Bird Conservation Regions (IMBCR: Hanni et al. 2012), except that we conducted bird surveys prior to vegetation surveys. This helped to ensure that birds displaced by the observer, including those located at the point itself, were recorded. We also altered IMBCR vegetation survey protocols slightly to make the protocol specific to low stature prairie dog colonies, shortgrass prairie, and sagebrush systems. This was designed to be a quick, visual assessment; a more involved protocol using a Daubenmire frame and robel pole was used on transects and at nests. Vegetation Transects In addition to a visual assessment of vegetation at points, we sampled vegetation on transects and at nests. We completed two transects on all colonies and South Park plots, except that only one transect could be located on the smallest colonies. To locate each transect, we randomly chose a start and an end point from those used in avian point counts. From the start point, we walked along the bearing toward the end point for 240 m, stopping every 20 m to collect vegetation data for a total of 13 points per transect, except on very small colonies. Transect data were collected during the growing season, mainly from midJune through August. At each stop point, we recorded the presence of active or inactive prairie dog burrows within 10 m, ground cover, dominant species, and visual obstruction (Fig. 1c). Percent ground cover was measured within a 50 cm square Daubenmire frame. We recorded the percent bare ground, rock, litter, scat, grass (including sedges and rushes), forb, shrub, cactus, exotic, and other cover. We also recorded the dominant species for each plant category present in the frame. Visual obstruction data were recorded by holding a robel pole at the stop point on the transect and making observations from a distance of 4 m in each cardinal direction with eye level at 1 m. The observer then noted which portions of the 122 cm (4 ft) pole were obstructed by vegetation, identified the plant species obstructing the pole, and noted whether the pole was substantially obstructed or was covered by just a wisp of vegetation (typically a blade of grass). We estimated the height of any structures taller than the pole. Raptor Counts Raptors were sometimes sighted during avian point counts, but point counts are not an ideal method for detecting raptors or other uncommon species. Therefore, we chose 1 – 2 locations per vaccine project area and plot, positioning observers so that the entire treatment area could be viewed simultaneously. We conducted 30-min. raptor counts, recording each bird’s species, horizontal (radial) distance, sex (if known), time of entry and exit, and behavior (high soar, low soar, directed flight, hover, dive, call, perch, or nest). Membership in a cluster was noted, typically for male-female pairs. This produces a time metric for assessing raptor use of treatment areas and colonies. At the start and end of the count, we recorded weather characteristics, including time, temperature, wind speed, and cloud cover. Raptor counts were conducted after 9:30 from November to early March (wintering) and April to August (breeding) and were never conducted in rain. As a supplement to the formal raptor counts, we recorded incidental raptor observations. Nest Searching and Monitoring We searched for mountain plover and burrowing owl nests throughout the study area through visual observation of adult birds, typically in the morning and not in rainy conditions or high winds. Target search regions included areas where MOPL and BUOW were detected during call-playback surveys, areas with nests in previous years, and other areas with appropriate habitat. MOPL nest in 7 �scrapes on the ground in areas with a relatively high bare ground component. BUOW nest in prairie dog burrows, often near colony edges and in burrows with low to moderately-sized mounds. Because these species react more to humans on foot than to vehicles, we conducted surveys from a vehicle whenever possible. Where ATV access was possible, we nest searched for MOPL using two ATVs with a 30 m rope suspended between them. Bungies on each end of the rope allowed observers to keep the rope taut and watch for running birds. When MOPL were detected, we observed the bird, sometimes backing away from the site, and waited for the bird to sit down on a nest. When BUOW were detected, we searched for nests in the vicinity of their perching location; typically males perched conspicuously near the nest burrow during the day. Additional nests were found during point counts and nest monitoring. When we were unable to find a nest during the initial search, we marked the GPS location and returned at a later date. We likely found the majority of BUOW nests on the landscape using this method (Conrey 2010). We also monitored all other raptor nests that we found. We searched for passerine nests in 2013 – 2015 but did not repeat this intensive protocol in 2016. MOPL nests were defined as structures containing at least one egg. Because BUOW nests are underground, we defined their nests as burrows with shredded manure present at the entrance (Garcia and Conway 2009), with feathers, regurgitated pellets, and prey remains providing additional evidence of a nest attempt. Some BUOW burrows were probed with an electronic camera on the end of a structure similar to a plumbing snake (Peeper System 2.3, Sandpiper Technologies). At the time of nest discovery, we recorded the same weather information that we recorded at points. We also did a rapid visual assessment of vegetation, with more detailed data collected at nest completion when overheating of eggs and nest abandonment was no longer a concern. We described the nest structure and vegetation immediately around the nest and estimated vegetation height, percent bare ground within a 1 m radius, and whether all, ≥ 50%, < 50%, or none of the nest could be seen from vantage points 5 m to the north and south. BUOW nests were marked with brightly painted wooden stakes placed 10 m north of the nest burrow. Other raptor nests were unmarked, but they were very easy to locate. We collected any pellets that we observed near BUOW nests for possible future dietary analysis. Nests were monitored at least once per week. Starting with the first visit when the nest was discovered, we recorded the time, any management activities (such as cattle grazing), age of juveniles, and number of eggs (where they could be seen), juveniles, and adults present. Juveniles were aged according to keys for BUOW (Priest 1997) and by comparison with the presumed hatch date for other raptors, based on our observations. Non-BUOW raptor nests were considered successful if at least one fully-feathered juvenile left the nest. Evidence of success included juveniles outside the nest, and/or displaying and calling adults, coupled with an intact nest and appropriate timing based on nest age. BUOW nests were considered successful when at least one fledgling aged ≥ 35 days was observed (Thomsen 1971, Davies and Restani 2006, Conrey 2010), because they leave the nest burrow (but may return to it many times) at 10 – 14 days and well before flight or independence are attained. Failed nests were destroyed, contained broken eggs, and/or had eggs or nestlings that disappeared before their expected fledge date. For analysis purposes, nests with unknown fate will have their histories truncated back to the last date when the nest was active and will be coded as successful at that time. At nest completion, we recorded the same vegetation data that were collected at points along vegetation transects: presence of prairie dog burrows within 10 m, percent ground cover, and visual obstruction. We placed the Daubenmire frame at 1 m in each cardinal direction and observed the robel pole (placed at the nest) from 4 m in each cardinal direction, producing four readings of each metric. 8 �Remote Cameras Remote cameras (Reconyx Hyperfire Covert IR model PC800) were placed in each vaccine project area to document use by mammalian predators and other wildlife. In 2013, we had just one camera per treatment, but we increased the number of remote cameras on BTPD colonies in 2014 – 2016 to better sample their larger size relative to GUPD colonies. We also deployed two cameras in the larger South Park colonies and plots. These cameras take photos when triggered by motion from an object that is warmer than ambient temperature. Camera locations were selected to maximize the potential for detections of mammalian predators without the use of baits or lures, which might have acted as attractants and altered the sampling region beyond treatment areas and prairie dog colonies. Cameras were positioned along game trails, aimed at coyote height, and tested before they were armed. Cameras targeted water sources, fence lines, and other landscape features and were positioned in space such that the entire treatment was sampled as thoroughly as possible. We set the cameras to take three photos when triggered, with no quiet period between photos. Most cameras were deployed in April – May, and most of the cameras at BTPD sites were left in place and operated year-round. We checked batteries and SD cards at least 3 times during summer and once during winter, but most camera checks were more frequent (once or twice per month from May September), particularly when cows were present on a site, as they sometimes disrupted camera operation. Cameras were removed from GUPD sites in September, prior to the advent of winter weather and GUPD hibernation. As a supplement to the camera data, we recorded incidental observations of predators such as coyotes, foxes, and badgers. Databases We designed a database for this project using Microsoft SQL Server 2012 with the data entry interface in Microsoft Access 2007. The database was designed by R. Conrey and D. Conrey, a professional database developer who volunteered his time, to run on the Fort Collins CPW research server. This allows multiple users to simultaneously access the database, while providing for daily backups and improved data security. Users can access a master list of codes for vegetation species, bird species, management types, observers, sites, colonies, and points that if changed, will update throughout the database. Users also access data entry forms for each data type described above, except for photo data. The Reconyx photo database (Newkirk 2014) catalogs photos and stores the metadata associated with them, displaying (but not storing) the photos themselves within Access forms. Users identify species in photos using stand-alone modules which are then loaded into the database. Each photo is examined by at least two observers, and a referee (R. Conrey or field crew leader) resolves any conflicts in IDs. Identification is ongoing. Data Analysis Thus far, all data have been entered and summarized, but data proofing is ongoing and statistical analyses have not yet been completed. For point counts, we calculated use rate for each species by dividing the number of detections by the number of points per site. We completed bird and vegetation species lists and summarized ground cover data collected at transects. For raptor counts, we calculated a proportional use index, dividing the usage minutes by the total survey minutes for each site. Apparent nest success was calculated as the proportion of nests fledging at least one chick. Naïve (minimum) occupancy estimates were calculated for carnivores based on remote camera photos taken during 2-week intervals starting 1 April of each year, separated into breeding and non-breeding periods. Data will eventually be analyzed using Program DISTANCE to estimate density from point counts, Program MARK to estimate nest survival and occupancy from point counts and camera data, and R for other data types. 9 �RESULTS We have collected one year of pre-treatment data and three years of post-treatment data at BTPD sites. For GUPD sites, we completed 3 years of work at Gunnison Basin and the Woodland Park area in 2015 and began work in South Park in 2016, prior to a planned GUPD reintroduction that is now on hold until funding can be obtained. Since fall 2013, plague epizootics have occurred across ~80% of the BTPD study area at 21 colonies. During each of the first three years our study, one of the three pairs of BTPD treatment sites experienced a plague outbreak, spanning the entire treatment area by fall 2015, aside from a dusted region around the USFWS black-footed ferret breeding center. Plague occurred at both the baited sites in each pair, meaning that both the control and vaccine areas experienced outbreaks. This epizootic started in fall 2013 shortly after vaccine drop, so plague was present in the system prior to vaccination. Colonies that experienced epizootics in 2013 experienced total or near extirpation of prairie dogs, but beginning in 2014, treated areas experienced losses of ~50 – 95% (two of these were treated with placebo baits). All of these areas had small numbers of prairie dogs with increasing abundance within 1 - 2 years of the outbreak, but populations were severely reduced compared pre-plague levels. Each year in September 2014 − 2016, black-footed ferrets were released in two BTPD study colonies (the Roman and Brannigan colonies in the northwest part of Soapstone Prairie Natural Area). In October 2014 and 2016, additional ferrets were released within the dusted part of a treatment colony (the Ferret Center colony in the southeastern part of Meadow Springs Ranch), which is 0.5 km from the nearest paired baited site and plague epizootic. 68 total ferrets were released on these properties, and a small amount of reproduction has been documented. Although 2013 and 2016 were fairly normal to slightly dry, 2014 and 2015 were very wet summers at the BTPD site compared to the 30-year average (Fig. 2). Normally dry playas and streambeds were full, grasses were tall and formed seedheads in June, and a different suite of species was observed, including a large flush of growth in species normally absent or a minor component of the prairie, such as six weeks fescue (Vulpia octoflora) and wooly plantain (Plantago patagonica). In 2016, these species were more common than blue grama grass or the forbs more commonly seen in previous years. We also saw a huge influx of lark buntings (Calamospiza melanocorys), which typically nest in or near taller vegetation than is typically found on these BTPD colonies; we had two nests in 2013, 69 nests in 2014, and 39 nests in 2015, making lark buntings the most abundant breeders at our sites, along with McCown’s longspurs. We conducted 1,734 avian point counts in 2013 − 2016 (more than 2,000, if repeat counts are included) and detected 137 bird species during the breeding season (Table 1, App. 1). The most common birds detected were horned lark, western meadowlark (Sturnella neglecta), lark bunting, and McCown’s longspur at BTPD sites and Brewer’s sparrow, vesper sparrow, horned lark, western meadowlark, greentailed towhee, common raven (Corvus corax), and sage thrasher at GUPD sites, in that order. We detected 85 species on BTPD colonies, 84 species on GUPD colonies, and 82 species off GUPD colonies. Baird’s sparrows (Ammodramus bairdii) were first detected at our BTPD sites in 2015 and appear to be becoming more widely distributed. Occupancy and density analyses have not yet been completed, but we have compared use rates (number of detections per point) across BTPD colonies with differing prairie dog activity status. During BTPD colony surveys, at least three bird species (Brewer’s blackbird Euphagus cyanocephalus, horned lark Eremophila alpestris, and vesper sparrow Pooecetes gramineus) appeared to have higher detection rates on active prairie dog colonies, while three species (European starling Sturnus vulgaris, grasshopper sparrow Ammodramus savannarum, and lark bunting Calamospiza melanocorys) appeared to have higher detection rates on colonies with extinct or severely reduced prairie dog populations following plague outbreaks (Table 2). In future density models, we will test the importance of these factors along with other covariates such as weather, vegetation characteristics, cattle grazing, and predator occupancy rates. 10 �We characterized vegetation at 1,734 point count locations, 607 nests, and ~5100 stop points along 404 transects in 2013 – 2016 and documented 230 species (Table 1, App. 2). Vegetation transect data (Table 3) suggested that BTPD colonies were dominated by grasses (mainly blue grama until 2016, when six weeks fescue was more common), litter, and bare ground, with triple the grass coverage of GUPD sites and rapidly increasing grass cover (doubled by 2016) during and after the El Niño event of 2014 – 2015 (Table 4, Fig. 2). We observed half the litter, a third of the bare ground, and a temporary increase in forb coverage at BTPD sites over the same time period (Table 4). GUPD sites in South Park were also dominated by grasses but had more forb cover and bare ground (Table 3): GUPD colonies had more bare ground and less forb cover than off-colony areas. Exotic cover was < 1% at all sites (Table 3). Visual obstruction by vegetation (Fig. 1c) was ≤ 9 cm at all sites in 2016. Vegetation heights increased each year of our study on BTPD colonies, with grasses responsible for most of the obstruction. Forbs had the highest species richness, followed by grasses and shrubs. Dominant plant species of each type are listed in Table 5, with a complete plant species list in App. 2. We conducted raptor counts at 86 locations for a total of 26,480 minutes (Table 1) and detected 18 species from 2013 – 2016 (Table 6). Burrowing owls, northern harriers Circus cyaneus, ferruginous hawks Buteo regalis (Fig. 1b), and rough-legged hawks Buteo lagopus were detected only on prairie dog colonies, and half the raptor species we observed used on-colony areas more than off-colony areas during our surveys, with the remainder appearing to show no preference or having low sample size (Table 6). During winter counts on BTPD colonies, burrowing owls, Swainson’s hawks, and turkey vultures were replaced by rough-legged hawks, and ferruginous hawks and northern harriers became more common than they were during summer. Relationships between raptor use and plague have not yet been analyzed. We monitored 64 raptor nests in 2016 (Table 1), mainly burrowing owl nests, which had an apparent nest success rate of 76% (Table 7). Despite three rounds of nest searching for mountain plover in South Park, nests were found only in a known area of high density in James Mark Jones State Wildlife Area, which was monitored by our collaborators at University of Colorado − Denver. The number of burrowing owl nests at our BTPD site has doubled during each year of our study (Table 7) following an El Niño event and widespread plague. Daily nest survival will be estimated using known fate models in Program MARK, and the importance of covariates such as plague, weather, vegetation characteristics, cattle grazing, and predator occupancy rates will be explored. We deployed 38 remote cameras in 2016 that took 339,006 photos from April to August, with > 10.8 million photos analyzed since 2013 (Table 1). BTPD cameras are deployed year-round, because BTPDs do not hibernate, while GUPD cameras were deployed from May − September. As expected, many photos recorded prairie dogs, cows, pronghorn (Antilocapra americana), and rabbits. We have documented coyote (Canis latrans), swift fox (Vulpes velox), badger (Taxidea taxus), striped skunk (Mephitis mephitis), red fox (V. vulpes), bobcat (Lynx rufus), black-footed ferret, and raccoon (Procyon lotor) use of vaccine project areas (listed by number of occurrences in our photos), and one black bear (Ursus americanus) using a proposed GUPD reintroduction site in South Park. We observed numerous instances of cooperative hunting behavior between coyotes and badgers in our photos (Fig. 1e). Swift fox, which appear to have increased steadily from 2013 – 2016, and skunks occurred only on BTPD colonies and badgers were more commonly detected there, while coyotes were equally common across all sites on BTPD and GUPD colonies (Table 8). Overall 2-week detection rates per site (naïve occupancy estimates) for the three most common carnivores ranged from 3% for badgers on both BTPD colonies (April – July 2014) and GUPD colonies (August – Sept 2014) to 64% for swift fox on BTPD colonies (August 2015 – March 2016; Table 8). 11 �DISCUSSION The 2016 season was our fourth year of research on BTPD sites and the first year of research at South Park, which contains several small GUPD colonies and large areas where prairie dogs have been extirpated. In 2013 – 2015, CPW Wildlife Health conducted efficacy trials for the experimental oral plague vaccine for prairie dogs. Although the vaccine was still considered experimental in 2016, its use was at a broader scale as a management tool to combat plague. We have collected one year of pretreatment data and three years of post-treatment data at our BTPD site, which has experienced a major plague event beginning in 2013. It will likely take additional years of monitoring to detect potential changes in the avian community caused by plague management, as treated colonies no longer experience extinction events. Therefore, initial data analyses will focus on shorter-term objectives, including evaluating the importance of plague on BTPD colonies and differences between on and off-colony areas at GUPD sites. Since fall 2013, plague epizootics have occurred across ~80% of the BTPD study area. All of these areas had small numbers of prairie dogs with increasing abundance within 1 - 2 years of the outbreak, but populations were severely reduced compared to pre-plague levels. Plague outbreaks have occurred across all treated BTPD sites, but plague was likely present in the system prior to vaccination, and immunity takes months to develop after vaccination (D. Tripp, pers. comm.). Prairie dog researchers are interested in whether prairie dogs persist in treated areas during outbreaks, recover more quickly in treated areas, and have higher survival during enzootic periods (without obvious outbreaks). At least three bird species (Brewer’s blackbird, horned lark, and vesper sparrow) appeared to have higher detection rates on active prairie dog colonies, while three species (European starling, grasshopper sparrow, and lark bunting) appeared to have higher detection rates on colonies with extinct or severely reduced prairie dog populations following plague outbreaks. Higher numbers of grasshopper sparrows and lark buntings coincided with high May – June precipitation, which likely contributed to high flea numbers and conditions favorable to a plague epizootic. Conditions across much of the BTPD study area in 2014 – 2015 began to support grassland birds that favor taller herbaceous vegetation over those that favor shorter vegetation, such as horned larks and McCown’s longspurs. Further analyses of species density and nest survival that include other covariates, such as weather and predator occupancy, will elucidate these relationships. Precipitation has varied greatly during this study (Fig. 2), particularly on BTPD sites, from slightly dry to very wet, compared to the 30-year average. In summer 2014 and 2015, normally dry playas and streambeds were full, grasses were tall and formed seedheads in June, and a different suite of species was observed, including a large flush of growth in species normally absent or a minor component of the prairie such as six weeks fescue. In response, we saw a huge influx of lark buntings to BTPD sites; they were the most common species detected in point counts and nest searches in 2014, and nest numbers continued to be high in 2015. In contrast, McCown’s longspur numbers declined in 2014, then rebounded in 2015, while the number of burrowing owl nests doubled each year. These changes may also be related to concurrent declines in prairie dogs associated with plague epizootics, which accelerate during El Niño events when cooler, wetter springs and summers result in high numbers of fleas, a major vector of plague. Vegetation was no longer being clipped, and extinct areas had much more lush vegetation than usual. Burrowing owl nest numbers increased only in areas that experienced plague epizootics and had persisting (or recovering) prairie dog populations. This may have created conditions with high prey and burrow availability in a landscape that continues to contain many active (if small) prairie dog colonies. Burrowing owls also increased in response to plague events (and prairie dog recolonization) in a previous study in northern Colorado (Conrey 2010). In September and October 2014, 42 black-footed ferrets were released in three BTPD study colonies, one of which was only 0.5 km from the nearest baited site, which experienced a plague epizootic 12 �in fall 2013. Twenty-six additional ferrets were released in 2015 − 2016, and surveys documented a lactating female and a kit. The two release sites (not included in the prairie dog vaccine study) in Soapstone Prairie Natural Area were both dusted and baited with vaccine, and the release site near the Ferret Center was a dusted treatment within the vaccine study through 2015 and was both dusted and vaccinated in 2016. Released ferrets have been vaccinated against plague and distemper, and were expected to disperse into adjacent colonies; however, most of the nearby colonies have experienced plague outbreaks. We began documenting ferrets in our remote camera photos in October 2015, but thus far, no ferrets have been seen or photographed outside of the colonies where they were released. At this point, young ferrets must be captured to be vaccinated, as they are not expected to consume vaccine baits designed for prairie dogs, but work on an oral vaccine for ferrets is in progress. Ferrets will likely predate on adult birds and nests, but as 90% of their prey base is typically comprised of prairie dogs (Clark 1986), the overall effect on avian species should be minimal. This was the fourth year of data collection on this project, and it will likely take additional years of monitoring to detect potential changes in the avian community caused by different types of plague management, as treated colonies no longer experience extinction events. Regardless of the efficacy of plague vaccine versus insecticide in reducing plague impacts, the vaccine will continue to be an important tool due to cost/benefit of its use and increasing evidence that fleas are evolving resistance to deltamethrin. Preliminary data suggest that bird densities do vary according to the status of prairie dogs on a colony, with differences between active colonies and those with extirpated or severely reduced prairie dog populations following plague outbreaks. Vegetation surveys have also identified differences between on- and off-colony areas. Raptor and camera data collection will continue through winter 2016/2017. During the 2017 breeding season, we hope to continue collecting point count, raptor count, and vegetation data on BTPD colonies with different activity levels, following the extensive 2013−2015 plague outbreak. Beyond that, there are several possibilities for continued research. The GUPD reintroduction planned for South Park is on hold until new funding is obtained; our avian work on GUPD sites may similarly go on hiatus. We may work more in ferret reintroduction sites, on private lands, or in a new location with little or no plague management (such as Pawnee National Grassland) for comparison with our current BTPD site north of Fort Collins. The first publications planned from this research include an analysis of point count data at our BTPD site from 2006 – present (with collaborators from BCR), including two complete plague cycles, and a comparison of on- and off-colony GUPD sites regarding their avian and vegetation communities. Future publications will compare passerine nest survival, raptor use, and mammalian carnivore occupancy on active colonies versus those with extirpated or severely reduced prairie dog populations following plague outbreaks. LITERATURE CITED Antolin, M. F., P. Gober, B. Luce, D. E. Biggins, W. E. V. Pelt, D. B. Seery, M. Lockhart, and M. Ball. 2002. The influence of sylvatic plague on North American wildlife at the landscape level, with special emphasis on black-footed ferret and prairie dog conservation. Transactions of the 67th North American Wildlife and Natural Resources Conference 67: 104–127. Augustine, D. J., S. J. Dinsmore, M. B. Wunder, V. J. Dreitz, and F. L. Knopf. 2008. Response of mountain plovers to plague-driven dynamics of black-tailed prairie dog colonies. Landscape Ecology 23:689–697. Boyle, S. A. and D. R. Reeder. 2005. Colorado sagebrush: a conservation assessment and strategy. Colorado Division of Wildlife, Grand Junction, Colorado. Butts, K. O. and J. C. Lewis. 1982. The importance of prairie dog colonies to burrowing owls in Oklahoma. Proceedings of the Oklahoma Academy of Sciences 62:46–52. Clark, Tim W. 1986. Some guidelines for management of the black-footed ferret. Great Basin Naturalist Memoirs 8:160–168. 13 �Colorado Division of Wildlife. 2003. Conservation plan for grassland species in Colorado. Colorado Division of Wildlife, Denver, Colorado. Conrey, R. Y. 2010. Breeding success, prey use, and mark-resight estimation of burrowing owls nesting on black-tailed prairie dog towns: plague affects a non-susceptible raptor. Ph.D. Dissertation, Colorado State University, Fort Collins, Colorado. Conrey, R. Y. 2013. Avian response to plague management on Colorado prairie dog colonies. Wildlife Research Report. Colorado Parks and Wildlife, Fort Collins, Colorado. 25 pg. Davies, J. M. and M. Restani. 2006. Survival and movements of juvenile burrowing owls during the postfledging period. Condor 108:282–291. Dicke, W. M. 1926. Plague in California 1900 – 1925. Proceedings of the 41st Annual Meeting and Conference of State Provincial Health Authority of North America, Atlantic City, New Jersey. Ecke, D. H. and C. W. Johnson. 1952. Plague in Colorado and Texas. Part I. Plague in Colorado. Public Health Monograph No. 6. U. S. Government Printing Office, Washington D.C. Garcia, V. and C. J. Conway. 2009. What constitutes a nesting attempt? Variation in criteria causes bias and hinders comparisons across studies. Auk 126:31–40. Grant-Hoffman, M. N. and J. K. Detling. 2006. Vegetation on Gunnison’s prairie dog colonies in Southwestern Colorado. Rangeland Ecology and Management 59:73–79. Hanni, D. J., C. M. White, R. A. Sparks, J. A. Blakesley, J. J. Birek, N. J. Van Lanen, J. A. Fogg, J. M. Berven, and M. A. McLaren. 2012. Integrated Monitoring of Bird Conservation Regions (IMBCR): Field protocol for spatially-balanced sampling of landbird populations. Unpublished report. Rocky Mountain Bird Observatory, Brighton, Colorado. 39 pp. Hardwicke, K. 2006. Prairie dogs, plants, and pollinators: tri-trophic interactions affect plant-insect floral visitor webs in shortgrass steppe. Ph.D. Dissertation. Colorado State University, Fort Collins, Colorado. Johnson-Nistler, C. M., B. F. Sowell, H. W. Sherwood, and C. L. Wambolt. 2004. Black-tailed prairie dog effects on Montana’s mixed-grass prairie. Journal of Range Management. 57:641–648. Jones, C. G., J. H. Lawton, and M. Shachak. 1994. Organisms as ecosystem engineers. Oikos 69:373– 386. Lomolino, M. V. and G. A. Smith. 2004. Terrestrial vertebrate communities of black-tailed prairie dog (Cynomys ludovicianus) towns. Biological Conservation 115:89–100. Magle, S. B., B. T. McClintock, D. W. Tripp, G. C. White, M. F. Antolin, and K. R. Crooks. 2007. Mark– resight methodology for estimating population densities for prairie dogs. Journal of Wildlife Management 71: 2067–2073. Meaney, C. A., M. Reed-Eckert, and G. P. Beauvais. 2006. Kit Fox (Vulpes macrotis): a technical conservation assessment. USDA Forest Service, Rocky Mountain Region. http://www.fs.fed.us/r2/projects/scp/assessments/kitfox.pdf [Accessed 2013]. Miller, R. F., T. J. Svejcar, and N. E. West. 1994. Implications of livestock grazing in the Intermountain sagebrush region: plant composition. Pages 101–146 in M. Vavra, W. A. Laycock, and R. D. Pieper, editors. Ecological implications of livestock herbivory in the west. Society for Range Management, Denver, Colorado. Newkirk, E. S. 2014. CPW Photo Database. Colorado Parks and Wildlife, Fort Collins, Colorado, USA. http://wildlife.state.co.us/Research/Mammal/Pages/MammalsWildlifeResearch.aspx Paine, R. T. 1969. A note on trophic complexity and community stability. American Naturalist 103:91– 93. Priest, J. E. 1997. Age identification of nestling burrowing owls. Journal of Raptor Research Report 9:125–127. Rocke, T. E., N. Pussini, S. R. Smith, J. Williamson, B. Powell, and J. E. Osorio. 2010. Consumption of baits containing raccoon pox-based plague vaccines protects black-tailed prairie dogs (Cynomys ludovicianus). Vector-Borne and Zoonotic Diseases 10: 53–58. Seglund, A. E. and P. M Schnurr. 2010. Colorado Gunnison’s and white-tailed prairie dog conservation strategy. Colorado Division of Wildlife, Denver, Colorado. 14 �Sidle, J. G., M. Ball, T. Byer, J. J. Chynoweth, G. Foli, R. Hodorff, G. Moravek, R. Peterson, and D. N. Svingen. 2001. Occurrence of burrowing owls in black-tailed prairie dog colonies on Great Plains National Grasslands. Journal of Raptor Research 35:316–321. Smith, G. A. and M. V. Lomolino. 2004. Black-tailed prairie dogs and the structure of avian communities on the shortgrass plains. Oecologia 138:592–602. SPV Subcommittee. 2011. Call for partners: Phase II field studies on oral sylvatic plague vaccine (SPV) for prairie dogs. Stapp, P., M. F. Antolin, and M. Ball. 2004. Patterns of extinction in prairie dog metapopulations: plague outbreaks follow El Nino events. Frontiers in Ecology and the Environment 2:235–240. Stapp, P., B. Van Horne, and M. D. Lindquist. 2008. Ecology of mammals of the shortgrass steppe. Pages 132–180 in W. K. Lauenroth and I. C. Burke, editors. Ecology of the shortgrass steppe: a longterm perspective. Oxford University Press, New York, New York. Thomsen, L. 1971. Behavior and ecology of burrowing owls on the Oakland Municipal Airport. Condor 73:177–192. Tipton, H. C., V. J. Dreitz, and P. F. Doherty, Jr. 2008. Occupancy of mountain plover and burrowing owl in Colorado. Journal of Wildlife Management 72:1001–1006. Tripp, D. W. and T. E. Rocke. 2012. Study plan: Preliminary field trials to evaluate the safety and efficacy of an sylvatic plague vaccine for prairie dogs. Whicker, A. D. and J. K. Detling. 1988. Ecological consequences of prairie dog disturbances: prairie dogs alter grassland patch structure, nutrient cycling, and feeding-site selection by other herbivores. BioScience 38:778–785. Yackel Adams, A. A. 2000. Training notes for grassland bird project. Unpublished report. U.S. Geological Survey Fort Collins Science Center, Fort Collins, Colorado. 8 pp. Yackel Adams, A. A. Unpub data. Lark bunting nestling aging guide. U.S. Geological Survey Fort Collins Science Center, Fort Collins, Colorado. 2 pg. 15 �FIGURE LEGENDS Figure 1. Photos from BTPD and GUPD sites during 2013 – 2016. a) GUPD consuming experimental bait. b) Ferruginous hawk seen during a winter raptor count. c) Visual obstruction measurement. d) Burrowing owl on BTPD site. e) Coyote and badger photographed by remote camera. Figure 2. Monthly precipitation at BTPD (Larimer and Weld Co.) sites from 2013 – 2016, including the 30-year average. Data were taken from the nearest weather station. 16 �TABLES Point counts Bird species Vegetation transects Vegetation species Raptor count locations Raptor count minutes Nests Remote camera locations Remote camera photos BTPD on GUPD on GUPD off 301 59 66 53 9 3,440 64 24 312,272 32 38 7 18 5 1,710 0 7 8,716 119 53 12 22 6 570 0 7 18,018 TOTAL TOTAL 2016 2013−16 452 1,734 81 137 85 404 75 230 20 86 5,720 26,480 64 607 38 58 339,006 10,842,301 Table 1. Sample sizes for BTPD and GUPD sites, on and off prairie dog colonies. We did one point count, 5 winter raptor counts, and 5 – 9 summer raptor counts per location. We surveyed 1 – 2 vegetation transects per plot each year. Photos from 2016 were taken in April – August. GUPD cameras were deployed during summer, and BTPD cameras were deployed year-round. SPECIES Brewer's Blackbird Horned Lark Vesper Sparrow Barn Swallow Brewer's Sparrow Common Raven McCown's Longspur Red-Winged Blackbird Western Meadowlark European Starling Grasshopper Sparrow Lark Bunting Use Rate A R+E 0.069 0.249 3.899 4.642 0.135 0.324 0.122 0.122 0.101 0.089 0.120 0.114 1.379 1.544 0.395 0.330 2.566 2.812 0.070 0.123 0.173 0.290 2.231 2.797 TOTAL COUNT 257 6764 369 192 150 185 2301 574 4235 151 361 3948 Table 2. Bird use rates for the most common species detected during avian point counts on BTPD colonies (Larimer and Weld Co.) with varying prairie dog activity status: Active (A), Reduced (R) after a plague event, and Extinct (E). Use rate was calculated by dividing the number of detections by the total number of points surveyed. The first three species were more common on active colonies, those in the middle showed no preference, and the last three species were more common on reduced or extinct colonies. Data reported here do not yet account for probability of detection or the location of individual birds inside or outside of treatment area boundaries. 17 �% Cover Grass Litter Bare Forb Rock Cactus Scat Shrub Other Exotic BTPD on 71.0 11.9 7.5 4.8 1.7 1.1 1.1 0.6 0.4 0.4 GUPD on 44.4 10.7 27.2 9.9 3.5 0.0 0.6 0.2 3.5 0.3 GUPD off 41.2 4.7 16.5 23.5 2.6 0.0 0.9 0.6 9.9 0.0 Table 3. Ground cover percentages from vegetation transects conducted on BTPD (Larimer and Weld Co.) and GUPD (Park Co.) sites, on and off prairie dog colonies, from June – August 2016. % Cover Grass Litter Bare Forb 2013 36.7 27.2 22.8 3.9 2014 55.9 8.7 19.4 8.3 2015 55.7 14.3 13.3 9 2016 71.0 11.9 7.5 4.8 Table 4. Ground cover percentages for dominant vegetation types from 2013 – 2016 for BTPD sites (Larimer and Weld Co.) in north central Colorado. Type Grass Forb Shrub Cactus Other Exotic BTPD on six weeks fescue woolly plantain rubber rabbitbrush plains pricklypear lichen field pennycress GUPD on blue grama fringed sagebrush winterfat N/A lichen field pennycress GUPD off plains bluegrass fringed sagebrush rabbitbrush sp. N/A lichen N/A Table 5. Dominant plant species detected on vegetation transects at BTPD (Larimer and Weld Co.) and GUPD (Park Co.) sites, on and off prairie dog colonies in June – August 2016. 18 �Species American Kestrel Burrowing Owl Common Raven Ferruginous Hawk Golden Eagle Loggerhead Shrike Northern Goshawk Northern Harrier Prairie Falcon Rough-legged Hawk Red-tailed Hawk Sharp-shinned Hawk Swainson’s Hawk Turkey Vulture TOTAL min BTPD in 2.26 23.02 3.23 2.21 1.09 0.00 0.00 0.46 0.52 0.24 1.34 0.00 4.80 6.07 11840 GUPD in GUPD out 2.30 1.47 0.45 0.00 17.50 16.70 0.03 0.00 1.79 2.08 0.78 0.21 0.00 0.38 0.48 0.00 0.33 0.04 0.09 0.00 5.46 3.04 0.19 0.21 0.45 0.52 2.66 1.85 6690 7950 TOTAL min 539 2756 2881 264 414 69 30 86 86 35 766 30 639 1044 26480 Table 6. Raptor use of vaccine project areas at BTPD and GUPD sites, on and off prairie dog colonies in 2013 – 2016. Use was quantified as time spent in project areas, and use rate = 100*(use minutes/total minutes) in BTPD, in GUPD, and off GUPD colonies. Data include breeding counts (late April – August) at all sites and wintering counts (November – early March) at BTPD sites. Species with small sample sizes are not shown: American crow, bald eagle, Cooper’s hawk, and osprey (2 min each). Species American Kestrel Burrowing Owl Common Raven Ferruginous Hawk Great Horned Owl Swainson's Hawk 2013 Fate (n) (0) 0.71 (7) (0) 0.00 (1) (0) 1.00 (2) 2014 Fate (n) (0) 0.73 (13) 1.00 (1) 0.00 (1) (0) 0.00 (1) 2015 Fate (n) 0.67 (3) 0.88 (25) 1.00 (1) 1.00 (1) (0) 1.00 (3) 2016 Fate (n) 1.00 (5) 0.76 (54) 1.00 (1) 1.00 (1) 0.00 (1) 0.50 (2) TOTAL 0.88 (8) 0.78 (99) 1.00 (3) 0.50 (4) 0.00 (1) 0.75 (8) Table 7. Nest fate (n = sample size) for raptors in vaccine project areas on BTPD colonies (Larimer and Weld Co.) during 2013 – 2016. These are naïve apparent nest survival estimates. 19 �Coyote BTPD GUPD 2013: April - July 2013: August - March 2014: April - July 2014: August - March 2015: April - July 2015: August - March 2016: April - July 2016: August 0.42 0.44 0.46 0.36 0.51 0.38 0.54 0.61 0.52 0.48 0.27 0.47 0.26 0.38 0.31 0.33 Badger Swift Fox BTPD GUPD BTPD GUPD 0 0 0.13 0 0.10 0.11 0.07 0 0.03 0.05 0.14 0 0.18 0.03 0.47 0 0.13 0.06 0.29 0 0.11 0.02 0.64 0 0.14 0.28 0.51 0 0.26 0.05 0.63 0 Table 8. Naïve (minimum) occupancy estimates (do not account for probability of detection) for carnivores based on remote camera photos taken on BTPD and GUPD colonies in 2013 – 2016. BTPD cameras were deployed year-round, while GUPD cameras were deployed during summer only, so GUPD colony estimates apply only to May – July and August – early September. GUPD cameras were deployed in Gunnison Basin and Woodland Park in 2013−2015 and in South Park in 2016. Seven cameras deployed in 2016 in areas where GUPD prairie dogs have been extirpated yielded estimates of 0.26 (April – July) and 0.19 (August) for coyotes, with no badgers or swift fox. Occupancy was calculated per site over 2week intervals. 20 �FIGURES a b c Sean Streich Sean Streich e d Walter Wehtje Figure 1. Photos from BTPD and GUPD sites during 2013 – 2016. a) GUPD consuming experimental bait. b) Ferruginous hawk seen during a winter raptor count. c) Visual obstruction measurement. d) Burrowing owl on BTPD site. e) Coyote and badger photographed by remote camera. 21 �BTPD Precipitation 200 180 160 Precipitation (mm) 140 120 2013 2014 100 2015 2016 80 average 60 40 20 0 April May June July Figure 2. Monthly precipitation at BTPD (Larimer and Weld Co.) sites from 2013 – 2016, including the 30-year average. Data were taken from the nearest weather station. 22 �APPENDIX 1: BIRD SPECIES LIST CODE AMAV AMCO AMCR AMGO AMGP AMKE AMRO AMWI AWPE BAEA BAIS BANS BARS BBMA BCCH BGGN BHCO BHGR BRBL BRSP BTLH BTPI BUOW BWTE CAFI CAGU SPECIES American Avocet American Coot American Crow American Goldfinch American Golden-plover American Kestrel American Robin American Wigeon American White Pelican Bald Eagle Baird's Sparrow Bank Swallow Barn Swallow Black-billed Magpie Black-capped Chickadee Blue-gray Gnatcatcher Brown-headed Cowbird Black-headed Grosbeak Brewer's Blackbird Brewer's Sparrow Broad-tailed Hummingbird Band-tailed Pigeon Burrowing Owl Blue-winged Teal Cassin's Finch California Gull BTPD BTPD In x x x x x x x x x x x x Gunn In Gunn Out x x x South In South Out x x x x x x x x x x x x x x x x x x x x x x x x x x x x GUPD Wood Wood In Out x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 23 �CANG CASP CCLO CCSP CHSP CLNU CLSW COFL COGR COHA CONI CORA DCCO DEJU DOWO DUFL EABL EAKI EAME ECDO EUST EVGR FEHA GADW GBHE GOEA GRAJ GRSP GTGR GTTO GUSG Canada Goose Cassin's Sparrow Chestnut-collared Longspur Clay-colored Sparrow Chipping Sparrow Clark's Nutcracker Cliff Swallow Cordilleran Flycatcher Common Grackle Cooper's Hawk Common Nighthawk Common Raven Double-crested Cormorant Dark-eyed Junco Downy Woodpecker Dusky Flycatcher Eastern Bluebird Eastern Kingbird Eastern Meadowlark Eurasian Collared-Dove European Starling Evening Grosbeak Ferruginous Hawk Gadwall Great Blue Heron Golden Eagle Gray Jay Grasshopper Sparrow Great-tailed Grackle Green-tailed Towhee Gunnison Sage-grouse x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 24 x x �GWTE HAWO HETH HOLA HOME HOSP HOWR KILL LARB LASP LBCU LEGO LISP LOSH MALL MCLO MGWA MOBL MOCH MODO MOPL NOFL NOGO NOHA NOMO NRWS NSHO OSFL OSPR PIGR PISI Green-winged Teal Hairy Woodpecker Hermit Thrush Horned Lark Hooded Merganser House Sparrow House Wren Killdeer Lark Bunting Lark Sparrow Long-billed Curlew Lesser Goldfinch Lincoln's Sparrow Loggerhead Shrike Mallard McCown's Longspur MacGillivray's Warbler Mountain Bluebird Mountain Chickadee Mourning Dove Mountain Plover Northern Flicker Northern Goshawk Northern Harrier Northern Mockingbird Northern Rough-winged Swallow Northern Shoveler Olive-sided Flycatcher Osprey Pine Grosbeak Pine Siskin x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 25 x x �PRFA PYNU RBGU RBNU RCKI RECR RNDU RNEP RNSA ROOS ROPI ROWR RTHA RWBL SACR SAGS SAPH SATH SAVS SNBU SOSP SPSA SPTO SSHA STJA SWHA TOSO TRES TUVU VESP VGSW Prairie Falcon Pygmy Nuthatch Ring-billed Gull Red-breasted Nuthatch Ruby-crowned Kinglet Red Crossbill Ring-necked Duck Ring-necked Pheasant Red-naped Sapsucker Rooster Rock Pigeon Rock Wren Red-tailed Hawk Red-winged Blackbird Sandhill Crane Sage Sparrow Say's Phoebe Sage Thrasher Savannah Sparrow Snow Bunting Song Sparrow Spotted Sandpiper Spotted Towhee Sharp-shinned Hawk Stellar's Jay Swainson's Hawk Townsend's Solitaire Tree Swallow Turkey Vulture Vesper Sparrow Violet-green Swallow x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 26 x x x x x x x x x x x x x x x �WAVI WBNU WCSP WEBL WEGR WEKI WEME WESJ WETA WEWP WHIM WIPH WISA WISN WITU YEWA YHBL YRWA TOTAL Warbling Vireo White-breasted Nuthatch White-crowned Sparrow Western Bluebird Western Grebe Western Kingbird Western Meadowlark Western Scrub-Jay Western Tanager Western Wood-pewee Whimbrel Wilson's Phalarope Williamson's Sapsucker Wilson's Snipe Wild Turkey Yellow Warbler Yellow-headed Blackbird Yellow-rumped Warbler 137 species x x x x x x x x x x x x x x x x x x x x x x x 89 x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 68 x 68 x 54 x x 50 33 46 Table A1. Bird species list for BTPD and GUPD sites on and off prairie dog colonies in Colorado during summer 2013−2016. These species were detected during avian point counts conducted during the breeding season. Gunn and Wood sites were sampled 2013−2015, while South was sampled only in 2016. 27 �APPENDIX 2: PLANT SPECIES LIST Code Family Scientific Name Common Name Exotic BTPD BTPD In Gunn In Gunn Out x x x x x x x GUPD Wood Wood In Out South In South Out x x x x x x Grasses, Sedges, and Rushes ACHY Poaceae Achnatherum hymenoides Indian Ricegrass ACLE Poaceae Achnatherum lettermanii Letterman’s Needlegrass AGCR Poaceae Agropyron cristatum Crested Wheatgrass x x AGST Poaceae Agrostis stolonifera Creeping Bentgrass x x ARPU Poaceae Aristida purpurea Purple Threeawn x x BODA Poaceae Bouteloua dactyloides Buffalograss x x x x BOGR Poaceae Bouteloua gracilis Blue Grama x x x x x BRCI Poaceae Bromus ciliatus Fringed Brome x x x BRHO Poaceae Bromus hordeaceus Soft Brome x BRIN Poaceae Bromus inermis Smooth Brome x BRPO Poaceae Bromus porteri Porter Brome BRTE Poaceae Bromus tectorum Cheatgrass CAIN Cyperaceae Carex inops ssp. heliophila Sun Sedge CALO Poaceae Calamovilfa longifolia Prairie Sandreed CARE Cyperaceae Carex Spp. Sedge Spp. (Unident.) x ELEL Poaceae Elymus elymoides Squirreltail x ELGL Poaceae Elymus glaucus Blue Wildrye ELRE Poaceae Elymus repens Quackgrass ELTR Poaceae Elymus trachycaulus Slender Wheatgrass FEAR Poaceae Festuca arizonica Arizona Fescue FEID Poaceae Festuca idahoensis Idaho Fescue FEST Poaceae Festuca Spp. Unknown Fescue x x HECO Poaceae Hesperostipa comata Needle & Thread Grass x x x HOJU Poaceae Hordeum jubatum Foxtail Barley x x x JUBA Juncaceae Juncus balticus Baltic Rush x x x KOMA Poaceae Koeleria macrantha Junegrass x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 28 x x x x x x x �LOPE Poaceae Lolium perenne Annual Ryegrass MUFI Poaceae Muhlenbergia filiformis Pullup Muhly x x MUHL Poaceae Muhlenbergia Spp. Muhlenbergia Spp. MUMO Poaceae Muhlenbergia montana Mountain Muhly MUTO Poaceae Muhlenbergia torreyi Ring Muhly NAVI Poaceae Nassella viridula Green Needlegrass PASM Poaceae Pascopyrum smithii Western Wheatgrass PHPR Poaceae Phleum pratense Timothy Grass PIMI Poaceae Piptatherum micranthum Littleseed Ricegrass POAR Poaceae Poa arida Plains Bluegrass POFE Poaceae Poa fendleriana Muttongrass POLE Poaceae Poa leptocoma Blue Marshgrass POPR Poaceae Poa pratensis Kentucky Bluegrass x x RUSH Juncaceae Juncus Spp. Rush Spp. x x SCPA Poaceae Schedonnardus paniculatus Tumblegrass x SCPR Poaceae Schedonorus pratensis Meadow Fescue SCSC Poaceae Schizachyrium scoparium Little Bluestem SOBI Poaceae Sorghum bicolor Sorghum SPCR Poaceae Sporobolus cryptandrus Sand Dropseed x VUOC Poaceae Vulpia octoflora Six Weeks Fescue x ACMI Asteraceae Achillea millefolium Common Yarrow ACRE Asteraceae Acroptilon repens Russian Knapweed ALCE Liliaceae Allium cernuum Nodding Onion AMBL Amaranthaceae Amaranthus blitoides Prostrate Pigweed AMBR Asteraceae Ambrosia Sp. Ambrosia sp. ANAR Apiaceae Angelica arguta White Angelica ANMI Asteraceae Antennaria microphylla Littleleaf Pussytoes ANMU Ranunculaceae Anemone multifida Cutleaf Anemone x ANOC Primulaceae Androsace occidentalis Western Rockjasmine x ANPA Asteraceae Antennaria parvifolia Small-leaf Pussytoes x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x Forbs x x x x x x x x x x 29 x x x x x x x x x x �ANSE Primulaceae Androsace septentrionalis Pygmy-flower Rockjasmine ARAB Asteraceae Artemisia absinthium Absinth Wormwood ARAN Rosaceae Argentina anserina Silver Weed ARCA Asteraceae Artemisia campestris Field Sagewort ARCO Caryophyllaceae Arenaria congesta Ballhead Sandwort ARFE Caryophyllaceae Arenaria fendleri Fendler's Sandwort x ARFR Asteraceae Artemisia frigida Fringed Sagebrush x ARLU Asteraceae Artesmia ludoviciana Prairie Sage ARPO Papavaraceae Argemone polyanthemos Crested Pricklypoppy ARUV Ericaceae Arctostaphylos uva-ursi Bearberry (Kinnikinnick) ASBI Fabaceae Astragalus bisulcatus Two-grooved Milkvetch x x x ASDR Fabaceae Astragalus drummondii Drummond's Milkvetch x x x ASMI Fabaceae Astragalus miser Timber Milkvetch x ASPA Fabaceae Astragalus parryii Parry's Milkvetch x ASSH Fabaceae Astralagus shortianus Short's Milkvetch BASC Chenopodiaceae Bassia scoparia Kochia/ Mexican Fireweed BEPL Scrophulariaceae Besseya plantaginea White River Coral Drops CALI Scrophulariaceae Castilleja linariifolia Narrowleaf Paintbrush x x CAMI Scrophulariaceae Castilleja miniata Scarlet Paintbrush x x CHBE Chenopodiaceae Chenopodium berlandieri Netseed Lambsquarter x x CHEN Chenopodiaceae Chenopodium Spp. Goosefoot Spp. (Unident.) x x CHGL Euphorbiaceae Chamaesyce glyptosperma Small Ribseed Sandmat x x CHHI Chenopodiaceae Chenopodium hians Hians Goosefoot CHLE Chenopodiaceae Chenopodium leptophyllum Narrowleaf Goosefoot CHSE Euphorbiaceae Chamaesyce serpyllifolia Thyme-leaf spurge x CHWA Chenopodiaceae Chenopodium watsonii Watson's Goosefoot x CIAR Asteraceae Cirsium arvense Canada Thistle CICA Asteraceae Cirsium canescens Prairie/Creamy Thistle CIUN Asteraceae Cirsium undulatum Wavyleaf Thistle COAR Convolvulaceae Convolvulus arvensis Field Bindweed CRTH Boraginaceae Cryptantha thyrsiflora Calcareous Cryptantha x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 30 x x x x x x x x x �DESO Brassicaceae Descurainia sophia Pinnate Tansy Mustard DRAB Brassicaceae Draba sp. Unknown Mustard (Draba) x x EQAR Equisetaceae Equisetum arvense Field Horsetail ERAS Brassicaceae Erysimum asperum Western Wallflower ERCA Asteraceae Erigeron canus Hoary Fleabane x ERCE Polygonaceae Eriogonum cernuum Nodding Buckwheat x ERCO Asteraceae Erigeron compositus Cutleaf Daisy ERFL Asteraceae Erigeron flagellaris Trailing Fleabane ERLO Polygonaceae Eriogonum lonchophyllum Spearleaf Buckwheat x EROV Polygonaceae Eriogonum ovalifolium Cushion Buckwheat x ERRA Polygonaceae Eriogonum racemosum Redroot Buckwheat x x ERSP Asteraceae Erigeron speciosus Showy Fleabane x x ERST Asteraceae Erigeron strigosus Prairie Fleabane x x ERUMA Polygonaceae Eriogonum umbellatum v. aureum Sulphur Buckwheat x x x ERUMM Polygonaceae Eriogonum umbellatum v. majus Creamy Buckwheat x x x FRVE Rosaceae Fragaria vesca Woodland Strawberry x x FRSP Gentianaceae Frasera speciosa Monument Plant x GABO Rubiaceae Galium boreale Bedstraw x GECA Geraniaceae Geranium caespitosum Pineywoods Geranium GETR Rosaceae Geum triflorum Old Man's Whiskers GEVI Geraniaceae Geranium viscosissimum Sticky Purple Geranium GRSQ Asteraceae Grindelia squarrosa Curlycup Gumweed x x GUSA Asteraceae Gutierrezia sarothrae Broom Snakeweed x x x HAFL Boraginaceae Hackelia floribunda Many-Flowered Stickseed x x HEPA Saxifragaceae Heuchera parvifolia Littleleaf Alumroot x x HEUC Saxifragaceae Heuchera Spp. Alumroot Spp. (Unident.) HEVI Asteraceae Heterotheca villosa Hairy False Golden Aster HYFI Asteraceae Hymenopappus filifolius Fineleaf Hymenopappus HYRI Asteraceae Hymenoxys richardsonii Pingue Rubberweed IRMI Iridaceae Iris missouriensis Rocky Mountain Iris IVAX Asteraceae Iva axillaris Povertyweed x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x 31 x �LALA Fabaceae Lathyrus latifolius Perennial Pea LAOC Boraginaceae Lappula occidentalis Flatspine Stickseed x x x LASC Asteraceae Laennecia schiedeana Pineland Horseweed x LEDE Brassicaceae Lepidium densiflorum Common Pepperweed LERA Brassicaceae Lepidium ramosissimum Many-branched Pepperweed LEVI Brassicaceae Lepidium virginicum Virginia Pepperweed LEVU Asteraceae Leucanthemum vulgare Ox Eye Daisy LILE Linaceae Linum lewisii Blue Flax LIPU Asteraceae Liatris punctata Gayfeather/ Blazing Star x x LUAR Fabaceae Lupinus argenteus Silvery Lupine x x LUWY Fabaceae Lupinus wyethii Wyeth's Lupine x x LYJU Asteraceae Lygodesmia juncea Skeletonweed x MAPI Asteraceae Machaeranthera pinnatifida Lacy Tansyaster x MARA Liliaceae Maianthemum racemosum Feathery False Lily of Valley MARE Berberidaceae Mahonia repens Oregon Grape MATA Asteraceae Machaeranthera tanacetifolia Tanseyleaf Tansyaster x MAVE Marsileaceae Marsilea vestita Hairy Waterclover x MEAR Lamiaceae Mentha arvensis Wild Mint MEHU Boraginaceae Mertensia humilis Mountain Bluebells MELA Boraginaceae Mertensia lanceolata Prairie Bluebells MEOF Fabaceae Melilotus officinalis Yellow Sweetclover x x MESA Fabaceae Medicago sativa Alfalfa x x MILI Nyctaginaceae Mirabilis linearis Narrowleaf Four o'clock OECA Onagraceae Oenothera caespitosa Tufted Evening Primrose OECO Onagraceae Oenothera coronopifolia Crownleaf Evening Primrose x OESU Onagraceae Oenothera suffrutescens Scarlet Beeblossom x ORLU Orobanchaceae Orobanche ludoviciana Louisiana Broomrape ORLU2 Scrophulariaceae Orthocarpus luteus Yellow Owl's Clover OXLA Fabaceae Oxytropis lambertii Lambert Crazyweed OXSE Fabaceae Oxytropis sericea White Locoweed PEBA Scrophulariaceae Penstemon barbatus Beardlip Beardtongue x x x x x x x x x x x x x x x x x x x x x 32 x x x x x x x x x x x x x x x x x x x x x x �PECR Scrophulariaceae Penstemon crandallii Crandall's Beardtongue PHBE Brassicaceae Physaria bellii Front Range Twinpod x PHHO Polemoniaceae Phlox hoodii Spiny Phlox x PHLO Polemoniaceae Phlox longifolia Longleaf Phlox PIOP Asteraceae Picradeniopsis oppositifolia Opposite Leaf Bahia PLMA Plantaginaceae Plantago major Common Plantain PLPA Plantaginaceae Plantago patagonica Woolly Plantain POGR Rosaceae Potentilla gracilis Slender Cinquefoil POHI Rosaceae Potentilla hippiana Woolly Cinquefoil POOL Portulacaceae Portulaca oleracea Common Purslane/ Hogweed x POPE Polygonaceae Polygonum persicaria Spotted Ladysthumb x PSLA Fabaceae Psoralidium lanceolatum Lemon Scurfpea RACO Asteraceae Ratibida columnifera Upright Prairie Coneflower RHRO Crassulaceae Rhodiola rosea King's Crown x SATR Chenopodiaceae Salsola tragus Russian Thistle/Tumbleweed x SAXI Saxifragaceae Saxifraga Spp. Saxifrage Spp. (Unident.) SEDE Selaginellaceae Selaginella densa Lesser Spikemoss SEIN Asteraceae Senecio integerrimus SELA Crassulaceae SENE x x x x x x x x x x x x x x x x x x x x x x x x x x x Lambstongue Ragwort x x Sedum lanceolatum Spearleaf Stonecrop x x Asteraceae Senecio Spp. Groundsel Spp. (Unident.) SOMI Asteraceae Solidago missouriensis Missouri Goldenrod SONU Fabaceae Sophora nuttalliana Silky Sophora x SOTR Solanaceae Solanum triflorum Cutleaf Nightshade x x SPCO Malvaceae Sphaeralcea coccinea Scarlet Globemallow x x SPFA Euphorbiaceae Spurge (Unident.) x SYLA Asteraceae Symphyotrichum lanceolatum White Panicle Aster TAOF Asteraceae Taraxacum officinale Dandelion TAVU Asteraceae Tanacetum vulgare Common Tansy THAR Brassicaceae Thlaspi arvense Field Pennycress THRH Fabaceae Thermopsis rhombifolia Golden Pea TRDU Asteraceae Tragopogon dubius Yellow Salsify x x x x x x x x x x x x x x x x x x x 33 x x x �TRHY Fabaceae Trifolium hybridum Alsike Clover x TRPR Fabaceae Trifolium pratense Red Clover x x URDI Urticaceae Urtica gracilis Stinging Nettle x VIAM Fabaceae Vicia americana American Vetch x VICI Fabaceae Vicia Spp. Vetch Spp. (Unident.) WYAM Asteraceae Wyethia amplexicaulis Mule-ears ACGL Aceraceae Acer glabrum Mountain Maple AMAL Rosaceae Amelanchier alnifolia Serviceberry x x ARCA Asteraceae Artemisia cana Silver Sagebrush x x ARTR Asteraceae Artemisia tridentata Big Sagebrush x x x x x ATCA Chenopodiaceae Atriplex canescens Fourwing Saltbush x x CEMO Rosaceae Cercocarpus montanus Mountain Mahogany x x x x x CHVI Asteraceae Chrysothamnus viscidiflorus Douglas Rabbitbrush x x x x x DAFR Rosaceae Dasiphora fruticosa Shrubby Cinquefoil x x x x EREF Polygonaceae Eriogonum effusum Spreading Buckwheat x ERNA Asteraceae Ericameria nauseosa Rubber /Grey Rabbitbrush x HODU Rosaceae Holodiscus dumosus Rockspirea JUCO Cupressaceae Juniperus communis ssp. alpina Common Juniper x x JUSC Cupressaceae Juniperus scopulorum Rocky Mountain Juniper x x KRLA Chenopodiaceae Krascheninnikovia lanata Winterfat x x PRAM Rosaceae Prunus americana American Plum x x PRVI Rosaceae Prunus virginiana Western Chokecherry x x PUTR Rosaceae Purshia tridentata Bitter Brush x x RASP Asteraceae RHTR Anacardiaceae Rhus trilobata Skunkbrush Sumac x x RIAU Grossulariaceae Ribes aureum Golden Currant x x x RICE Grossulariaceae Ribes cereum Wax Currant x ROAR Rosaceae Rosa arkansana Prairie/ Wild Rose x ROWO Rosaceae Rosa woodsii Woods' Rose RUID Rosaceae Rubus idaeus Wild Red Raspberry x x x x x x x Shrubs x x x x x x x x x x x x x x Rabbitbrush (Unident.) 34 x x x x x x x x x x x x �SALI Salicaceae Salix Spp. Willow Spp. (Unident.) SYOC Caprifoliaceae Symphoricarpos occidentalis Western Snowberry x TECA Asteraceae Tetradymia canescens Spineless Horsebrush x XANT Asteraceae Xanthium spp. Cocklebur sp. x YUGL Agavaceae Yucca glauca Great Plains Yucca x ABLA Pinaceae Abies lasiocarpa Subalpine Fir PIEN Pinaceae Picea engelmannii Engelmann Spruce PIFL Pinaceae Pinus flexilis Limber Pine PIPO Pinaceae Pinus ponderosa Ponderosa Pine PIPU Pinaceae Picea pungens Blue Spruce POAN Salicaceae Populus angustifolia Narrow-leaved Cottonwood PODE Salicaceae Populus deltoides ssp. wislizenii POTR Salicaceae Populus tremuloides PSME Pinaceae x x x x x x x x Trees x x x x x x x x x x x x x x x x x x x x x Rio Grande Cottonwood x x Quaking Aspen x x Pseudotsuga menziesii Douglas Fir x x Escobaria vivipara x Cacti and Other Plants ESVI Pincushion Cactus x FUNG Cactaceae Fungi x x x LICH Lichen x x x x x MOSS Moss x x x x x x x 74 75 OPFR Cactaceae Opuntia fragilis Brittle Pricklypear OPPO Cactaceae Opuntia polyacantha Plains Pricklypear PESI Cactaceae Pediocactus simpsonii Mountain Ball Cactus Total x x x 21 24 x x x 34 230 species x 96 127 127 Table A2. Plant species list for BTPD and GUPD sites on and off prairie dog colonies in Colorado for 2013−2016. Gunn and Wood sites were sampled 2013−2015, while South was sampled only in 2016. 35 � Dublin Core The Dublin Core metadata element set is common to all Omeka records, including items, files, and collections. For more information see, http://dublincore.org/documents/dces/. Title A name given to the resource Reports Text A resource consisting primarily of words for reading. Examples include books, letters, dissertations, poems, newspapers, articles, archives of mailing lists. Note that facsimiles or images of texts are still of the genre Text. Dublin Core The Dublin Core metadata element set is common to all Omeka records, including items, files, and collections. For more information see, http://dublincore.org/documents/dces/. Title A name given to the resource Avian response to plague management on Colorado prairie dog colonies Description An account of the resource Range-wide declines in prairie dog (<em>Cynomys</em> sp.) populations have occurred, and the largest limiting factor in recent decades appears to be the high mortality and colony extirpation associated with plague (Antolin et al. 2002), caused by the bacterium <em>Yersinia pestis</em>. Prairie dog colonies support a diverse community of associated species, many of which are not susceptible to plague but may be indirectly affected. Creator An entity primarily responsible for making the resource Yale Conrey, Reesa Subject The topic of the resource Prairie dog <em>Yersinia pestis</em> Black-footed ferret Plague Avian species Extent The size or duration of the resource. various pages Date Created Date of creation of the resource. 2014-2016 Rights Information about rights held in and over the resource <a href="http://rightsstatements.org/vocab/NoC-NC/1.0/">No Copyright - Non-Commercial Use Only</a> Type The nature or genre of the resource Text Format The file format, physical medium, or dimensions of the resource application/pdf Language A language of the resource English Is Part Of A related resource in which the described resource is physically or logically included. Cost Center 3420 Avian Research. Work Package 1680 Bird conservation