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Using fecal DNA and closed-capture models to estimate feral horse population size

Accurate population estimates provide the foundation for managing feral horses ( Equus caballus ferus ) across the western United States. Certain feral horse populations are protected by the Wild and Free-Roaming Horses and Burros Act of 1971 and managed by the Bureau of Land Management (BLM) or the United States Forest Service on designated herd management areas (HMAs) or wild horse territories, respectively. Horses are managed to achieve an appropriate management level (AML), which represents the number of horses determined by BLM to contribute to a thriving natural ecological balance and avoid deterioration of the range. To achieve AML for each HMA, BLM resource managers need accurate and precise population estimates. We tested the use of non-invasive fecal samples in a genetic capture-recapture framework to estimate population size in a closed horse population at the Little Book Cliffs HMA, Colorado, USA, with a known size of 153 individuals. We collected 1,957 samples over 3 independent sampling periods in 2014 and amplified them at 8 microsatellite loci. We applied mark-recapture models to determine population size using 954 samples that amplified at all 8 loci. We subsampled and reanalyzed our dataset to simulate different data collection protocols and evaluated effects on accuracy and precision of estimates using N-mixture modeling, full likelihood closed-capture modeling, and capwire single-occasion modeling that used data from all 3 sampling periods. Our model results were accurate and precise for analyses that used data from all 3 occasions; however, capwire single-occasion modeling was not accurate when we analyzed each sampling period separately. For all subsampling analysis scenarios, reducing sample size decreased precision, whether by reducing number of field staff, field days, or geographic areas surveyed on each period. Reducing spatial coverage of the survey area did not result in accurate population estimates and only marginally lowered the number of samples that would need to be collected to maintain accuracy. Because laboratory analysis contributes the greatest expense for this method ($80 U.S./sample), reducing fecal sample size is advantageous. Our results demonstrate that non-invasive sampling combined with good survey design and careful genetic and capture-recapture analyses can provide an alternative method to estimate the number of feral horses in a closed population. This method may be especially appropriate in situations where aerial inventories are not practical or accurate because of low sighting conditions. But the higher costs associated with laboratory sample analyses may reduce the method's feasibility compared to helicopter surveys.

Colorado↗

Habitat prioritization across large landscapes, multiple seasons, and novel areas: an example using greater sage-grouse in Wyoming

Animal habitat selection is an important and expansive area of research in ecology. In particular, the study of habitat selection is critical in habitat prioritization efforts for species of conservation concern. Landscape planning for species is happening at ever-increasing extents because of the appreciation for the role of landscape-scale patterns in species persistence coupled to improved datasets for species and habitats, and the expanding and intensifying footprint of human land uses on the landscape. We present a large-scale collaborative effort to develop habitat selection models across large landscapes and multiple seasons for prioritizing habitat for a species of conservation concern. Greater sage-grouse ( Centrocercus urophasianus , hereafter sage-grouse) occur in western semi-arid landscapes in North America. Range-wide population declines of this species have been documented, and it is currently considered as “warranted but precluded” from listing under the United States Endangered Species Act. Wyoming is predicted to remain a stronghold for sage-grouse populations and contains approximately 37% of remaining birds. We compiled location data from 14 unique radiotelemetry studies (data collected 1994–2010) and habitat data from high-quality, biologically relevant, geographic information system (GIS) layers across Wyoming. We developed habitat selection models for greater sage-grouse across Wyoming for 3 distinct life stages: 1) nesting, 2) summer, and 3) winter. We developed patch and landscape models across 4 extents, producing statewide and regional (southwest, central, northeast) models for Wyoming. Habitat selection varied among regions and seasons, yet preferred habitat attributes generally matched the extensive literature on sage-grouse seasonal habitat requirements. Across seasons and regions, birds preferred areas with greater percentage sagebrush cover and avoided paved roads, agriculture, and forested areas. Birds consistently preferred areas with higher precipitation in the summer and avoided rugged terrain in the winter. Selection for sagebrush cover varied regionally with stronger selection in the Northeast region, likely because of limited availability, whereas avoidance of paved roads was fairly consistent across regions. We chose resource selection function (RSF) thresholds for each model set (seasonal × regional combination) that delineated important seasonal habitats for sage-grouse. Each model set showed good validation and discriminatory capabilities within study-site boundaries. We applied the nesting-season models to a novel area not included in model development. The percentage of independent nest locations that fell directly within identified important habitat was not overly impressive in the novel area (49%); however, including a 500-m buffer around important habitat captured 98% of independent nest locations within the novel area. We also used leks and associated peak male counts as a proxy for nesting habitat outside of the study sites used to develop the models. A 1.5-km buffer around the important nesting habitat boundaries included 77% of males counted at leks in Wyoming outside of the study sites. Data were not available to quantitatively test the performance of the summer and winter models outside our study sites. The collection of models presented here represents large-scale resource-management planning tools that are a significant advancement to previous tools in terms of spatial and temporal resolution.

Wyoming↗

Seasonal resource selection and movement ecology of free-ranging horses in the western United States

Understanding factors driving resource selection and habitat use of different species is an important component of management and conservation. Feral horses ( Equus caballus ) are free ranging across various vegetation types in the western United States, yet few studies have quantified their resource selection and seasonal use. We conducted a study to determine effects of vegetation community, distance to water, and topographic variables on seasonal resource selection in 2 feral horse populations in Great Basin sagebrush ( Artemisia spp.) ecosystems of west-central Utah, USA: Conger Herd Management Area (HMA) and Frisco HMA. We deployed global positioning system (GPS) radio-collars on 38 female horses and GPS-transmitters braided and glued into the tail hair of 14 males, collecting locations every 2 hours for 1–4 years between 2016 and 2020. We calculated home range size and core use area of social groups (harems) and bachelor males using auto-correlated kernel density estimators for each biologically defined season (breeding, fall, and winter) per study year. We examined seasonal home range size and overlap of harem groups and bachelor males and compared movement speed of bachelors and harems among seasons. We determined seasonal resource selection in a use-availability framework using resource selection functions. We hypothesized that horses would select for areas of high herbaceous vegetation, that water would be a key variable in resource selection models like other equids, and home range size in winter would be largest because horses can eat snow for hydration and could therefore roam farther from surface water. Mean annual home range size was 103.12 ± 37.38 km 2 (SD) for Conger harems and 117.47 ± 32.75 km 2 for Frisco harems. At Conger there was no difference in home range size between harem groups and bachelor males, but home range size was smaller in winter than other seasons, whereas winter home range size at Frisco was larger than other seasons. Bachelor males moved at higher speeds than harems during all seasons, and harem groups from both populations had lower movement speeds in winter. Harem groups had distinct winter ranges with little overlap on breeding season ranges. In both populations, all horses selected for herbaceous vegetation types and avoided forest relative to shrubland throughout the year. Harems at Frisco were consistently located closer to water sources, whereas selection for water sources by Conger harems varied seasonally, with winter having the lowest selection. Harem groups at Conger had an average of 10.6% of their home ranges outside the HMA boundary and Frisco harems had up to 66.8% outside, likely because of the horseshoe shape of Frisco HMA in which shrub meadows (foraging areas) comprise the horseshoe center, which is outside the HMA. Our results highlight the importance of water sources, which were a key predictor of horse movement patterns in our study. We emphasize the utility of telemetry devices to understand resource selection of feral horses at a fine scale, enabling management to be more targeted and facilitate planning.

Utah↗

Comparing methods to estimate feral burro abundance

Obtaining precise and unbiased estimates of feral burro ( Equus asinus ) abundance in the western United States is challenging due to their cryptic pelage and the rugged terrain they inhabit. Management agencies employ helicopter-based, simultaneous double-observer sightability surveys (hereafter denoted as DOS) to estimate abundance of burros; but the DOS method routinely produces negatively biased estimates due to residual heterogeneity in detection probability. Consequently, testing alternative methods to improve upon current procedures is warranted. Residual heterogeneity in DOS surveys can be minimized by including radio-collared individuals in the population. Alternatively, if distance measurements are recorded, residual heterogeneity can also be reduced via a mark-recapture distance sampling (MRDS) approach. Aerial infrared (IR) surveys offer a safer alternative than helicopter-based surveys because they can be flown at a higher altitude and require fewer observers in the aircraft. Further, IR surveys using a distance sampling approach have been shown to generate accurate and precise estimates of feral horse ( E. caballus ) populations. Accordingly, we compared results of surveys using aerial IR distance sampling, the standard DOS survey, a DOS survey incorporating detections of radio-collared individuals, and an MRDS analysis of a feral burro population with a known minimum population size in central Utah, winter 2015–2016 and spring 2016. The minimum number of burros known alive during the winter and spring surveys were 236 and 136, respectively. The average detection probability of IR surveys was P = 0.88 (SE = 0.16) and distance models produced estimates of 127 burros (95% CIs = 99–175) for the winter survey, and 94 burros (CIs = 72–134) for the spring survey. Mean detection probability of the standard DOS surveys was P = 0.78 (SE = 0.09), and model-generated abundance estimates were 155 burros (CIs = 133–227) in winter, and 92 burros (CIs = 79–139) in spring. Incorporating detections of radio-collared individuals in the DOS survey resulted in a decreased detection probability ( P = 0.46; SE = 0.06) and increased abundance estimates to 267 (CIs = 169–571) and 155 (CIs = 128–263) for winter and spring, respectively. Mark-recapture distance sampling produced a mean detection probability of P = 0.48 (SE = 0.12) and resulted in estimates of 282 (CIs = 178–385) and 169 (CIs = 73–310) burros in winter and spring, respectively. Our study demonstrated that aerial IR surveys conducted using standard distance sampling can produce precise estimates of burro population sizes; however, estimates were negatively biased relative to the known population size. Small sample size limits generalization of our results, but the IR-based distance approach did not improve upon DOS surveys. Accounting for residual heterogeneity through use of radio-collars and mark-recapture distance sampling eliminated the negative bias from the standard DOS survey but decreased survey precision. Managers will need to decide whether unbiased but less precise abundance estimates are preferable compared to a more precise, but biased, estimate.

Utah↗

Ecology of band-tailed pigeons in Oregon

The ecology of band-tailed pigeons ( Columba fasciata ) was investigated in western Oregon to assess the roles of survival and recruitment to population status and the relation of food, minerals, and diet to reproduction strategy. Band-tailed pigeons congregated at mineral deposits to consume minerals from mid-June to mid-September. Males generally arrived and departed between daylight and 1000 h and females between 0930 and 1200 h, the inverse of the nest attentive schedules for the sexes. The pigeons used one or several adjacent mineral sites throughout the summer and most returned to the same mineral site in subsequent years. Band-tailed pigeons were resident from April through September; migration apparently did not begin before late September. Three indirect sources of evidence indicated that fledging began in June, reached a peak in mid- August, and continued until late September. Second year (SY) band-tailed pigeons apparently began nesting later than and were about one-third as productive as adults. In Oregon, most of the adult population could fledge as many as two young over the 100-110-day nesting period, resulting in a maximum potential recruitment of 47-50% juveniles. We estimated that the fall population contained about 23% juveniles, 12% yearlings, and 65% adults. Mean annual survival of adults was 63.7%, but was year-specific. To maintain a stable population with the estimated survival rates required production near the biotic potential (40.8%). Conversely, a stable population could be attained with the estimated rate of recruitment (23%) and high but realistic rates of survival (adults, 83%; juveniles, 68%). Counts of band-tailed pigeons at mineral sites in Oregon indicated that the population had undergone two periods of modest increase (2.4-7.1% per year) and two periods of sharp decline (10.4-11.1% per year). In 1988 the population index was only 34% of the 1950-88 average. Harvest of band-tailed pigeons in the three Pacific Coast states averaged 414,000 from 1957 to 1983; about 55% of the harvest took place in California, 23% in Washington, and 22% in Oregon. In Oregon, about 65% of the harvest took place in the first 10 days of September and 40% was at mineral sites. Fewer juveniles were shot at mineral sites (13%) than at feeding areas (25%). Hunting at mineral sites was directed at experienced breeders and may be particularly detrimental to the population. However, the overall effect of hunting on abundance was not determined. Band-tailed pigeons fed extensively on elder ( Sambucus spp.) and cascara buckthorn ( Rhamnus purshiana ) berries while nesting. The emergence of Pacific red elder (S. callicarpa) berries in June provided the food resources necessary for initiation of reproduction in Oregon, and probably throughout the Northwest. Elder berries contain little calcium (0.06-0.12%), therefore, the pigeons in the Northwest may require a mineral supplement in their diet. Mineral sites may be the scarcest resource required for reproduction by band-tailed pigeons in the Northwest.

Oregon↗

Red fox predation on breeding ducks in midcontinent North America

Red fox ( Vulpes vulpes ) predation on nesting ducks was assessed by examining 1,857 adult duck remains found at 1,432 fox rearing dens from 1968 to 1973. Dabbling ducks were much more vulnerable to foxes than diving ducks. Dabbling ducks (1,798) found at dens consisted of 27% blue-winged teals ( Anas discors ), 23% mallards ( A. platyrhynchos ), 20% northern pintails (A. acuta), 9% northern shovelers ( Spatula clypeata ), 8% gadwalls ( A. strepera ), 3% green-winged teals ( A. crecca ), 2% American wigeons ( A. americana ), and 10% unidentified. Relative abundance of individual species and nesting chronology were the most important factors affecting composition of ducks taken by foxes. Seventy-six percent of 1,376 adult dabbling ducks and 40% of 30 adult diving ducks for which sex was determined were hens. In western North Dakota and western South Dakota, 65% of mallard and northern pintail remains found at dens were hens compared with 76% in eastern North Dakota and eastern South Dakota ( P < 0.05). Percentage hens varied among the 5 most common dabbling ducks found at dens. In eastern North Dakota and eastern South Dakota, where predation on ducks was greatest, an average of 64% of gadwall, 73% of northern pintail, 81% of blue-winged teal, 81% of mallard, and 90% of northern shoveler remains found at dens were hens. Percentage hens among duck remains found at dens increased as the duck nesting season progressed. Numbers of adult ducks found at individual dens ranged from 0 to 67. The average number of ducks found in and around den entrances was used as an index of fox predation rates on ducks. Predation rate indices ranged from 0.01 duck/den in Iowa to 1.80 ducks/den in eastern North Dakota. Average annual predation rate indices for dabbling ducks in a 3-county intensive study area in eastern North Dakota were closely correlated with May pond numbers ( r = 0.874, P < 0.10) and duck population size ( r = 0.930, P < 0.05), but all species were not affected in the same manner or to the same degree. Drought had least effect on populations and predation rate indices of mallards and gadwalls and had greatest effect on those of northern pintails and northern shovelers. Hens of early nesting species were more vulnerable to foxes than hens of late nesting species. Predation rate indices were expanded to estimate total numbers of ducks taken by fox families during the denning season. Estimated numbers of dabbling ducks taken annually by individual fox families in 2 physiographic regions comprising the intensive study area ranged from 16.1 to 65.9. Predation was highest during wet years and lowest during dry years and averaged lower, but was more variable, in the region where tillage was greatest and wetland water levels were least stable. Predation in the intensive study area averaged 2.97 adult dabbling ducks/ km 2 /year and represented an estimated average annual loss of 13.5% of hen and 4.5% of drake populations in that area. Of 5,402 individual food items found at dens in the intensive study area, 24% were adult ducks. Ducks made up an estimated maximum average of 16% of the prey biomass required by fox families during the denning season. The average annual take of adult ducks by foxes in the midcontinent area was estimated to be about 900,000. This estimate included both scavenged and fox-killed ducks, as well as ducks taken after the denning season. Fox impact on midcontinent ducks was greatest in eastern North Dakota where both fox and duck densities were relatively high. Predation in that area was likely increased by environmental factors, especially intensive agriculture that concentrated nesting and reduced prey abundance. Predation by red foxes and other predators severely reduces duck production in the midcontinent area. Effective management to increase waterfowl production will necessitate coping with or reducing high levels of predation.

Iowa, Manitoba, Minnesota, Nebraska, North Dakota,↗

Current land bird distribution and trends in population abundance between 1982 and 2012 on Rota, Mariana Islands

The western Pacific island of Rota is the fourth largest human-inhabited island in the Mariana archipelago and designated an Endemic Bird Area. Between 1982 and 2012, 12 point-transect distance-sampling surveys were conducted to assess bird population status. Surveys did not consistently sample the entire island; thus, we used a ratio estimator to estimate bird abundances in strata not sampled during every survey. Trends in population size were reliably estimated for 11 of 13 bird species, and 7 species declined over the 30-y time series, including the island collared-dove Streptopelia bitorquata , white-throated ground-dove Gallicolumba xanthonura , Mariana fruit-dove Ptilinopus roseicapilla , collared kingfisher Todiramphus chloris orii , Micronesian myzomela Myzomela rubratra , black drongo Dicrurus macrocercus , and Mariana crow Corvus kubaryi . The endangered Mariana crow (x̄ = 81 birds, 95% CI 30&ndash;202) declined sharply to fewer than 200 individuals in 2012, down from 1,491 birds in 1982 (95% CI = 815&ndash;3,115). Trends increased for white tern Gygis alba , rufous fantail Rhipidura rufifrons mariae , and Micronesian starling Aplonis opaca . Numbers of the endangered Rota white-eye Zosterops rotensis declined from 1982 to the late 1990s but returned to 1980s levels by 2012, resulting in an overall stable trend. Trends for the yellow bittern Ixobrychus sinensis were inconclusive. Eurasian tree sparrow Passer montanus trends were not assessed; however, their numbers in 1982 and 2012 were similar. Occupancy models of the 2012 survey data revealed general patterns of land cover use and detectability among 12 species that could be reliably modeled. Occupancy was not assessed for the Eurasian tree sparrow because of insufficient detections. Based on the 2012 survey, bird distribution and abundance across Rota revealed three general patterns: 1) range restriction, including Mariana crow, Rota white-eye, and Eurasian tree sparrow; 2) widespread distribution, low abundance, including collared kingfisher, island collared-dove, white-throated ground-dove, Mariana fruit-dove, white tern, yellow bittern, black drongo, and Micronesian myzomela; and 3) widespread distribution, high abundance, including rufous fantail and Micronesian starling. The Mariana crow was dispersed around the periphery of the island in steep forested land-cover types. In contrast, the Rota white-eye was restricted to the high-elevation mesa. Only for the white-throated ground-dove was there a significant difference among cover types, with lower occupancy in open field than in forested areas. Vegetation was included in the best-fit occupancy models for yellow bittern, black drongo, Micronesian myzomela, and Micronesian starling, but vegetation type was not a significant variable nor included in the top models for the remaining five species: white tern, island collared-dove, Mariana fruit-dove, collared kingfisher, and rufous fantail. Given declining population trends, the Rota bird-monitoring program could benefit from establishing threshold and alert limits and identifying alternative research and management actions. Continued monitoring and demographic sampling, in conjunction with ecological studies, are needed to understand why most bird species on Rota are declining, identify the causative agents, and assess effectiveness of conservation actions, especially for the Mariana crow.

Rota↗

Population ecology of breeding Pacific common eiders on the Yukon-Kuskokwim Delta, Alaska

Populations of Pacific common eiders (Somateria mollissima v-nigrum) on the Yukon-Kuskokwim Delta (YKD) in western Alaska declined by 50–90% from 1957 to 1992 and then stabilized at reduced numbers from the early 1990s to the present. We investigated the underlying processes affecting their population dynamics by collection and analysis of demographic data from Pacific common eiders at 3 sites on the YKD (1991–2004) for 29 site-years. We examined variation in components of reproduction, tested hypotheses about the influence of specific ecological factors on life-history variables, and investigated their relative contributions to local population dynamics. Reproductive output was low and variable, both within and among individuals, whereas apparent survival of adult females was high and relatively invariant (0.89 ± 0.005). All reproductive parameters varied across study sites and years. Clutch initiation dates ranged from 4 May to 28 June, with peak (modal) initiation occurring on 26 May. Females at an island study site consistently initiated clutches 3–5 days earlier in each year than those on 2 mainland sites. Population variance in nest initiation date was negatively related to the peak, suggesting increased synchrony in years of delayed initiation. On average, total clutch size (laid) ranged from 4.8 to 6.6 eggs, and declined with date of nest initiation. After accounting for partial predation and non-viability of eggs, average clutch size at hatch ranged from 2.0 to 5.8 eggs. Within seasons, daily survival probability (DSP) of nests was lowest during egg-laying and late-initiation dates. Estimated nest survival varied considerably across sites and years (mean = 0.55, range: 0.06–0.92), but process variance in nest survival was relatively low (0.02, CI: 0.01–0.05), indicating that most variance was likely attributed to sampling error. We found evidence that observer effects may have reduced overall nest survival by 0.0–0.36 across site-years. Study sites with lower sample sizes and more frequent visitations appeared to experience greater observer effects. In general, Pacific common eiders exhibited high spatio-temporal variance in reproductive components. Larger clutch sizes and high nest survival at early initiation dates suggested directional selection favoring early nesting. However, stochastic environmental effects may have precluded response to this apparent selection pressure. Our results suggest that females breeding early in the season have the greatest reproductive value, as these birds lay the largest clutches and have the highest probability of successfully hatching. We developed stochastic, stage-based, matrix population models that incorporated observed spatio-temporal (process) variance and co-variation in vital rates, and projected the stable stage distribution () and population growth rate (λ). We used perturbation analyses to examine the relative influence of changes in vital rates on λ and variance decomposition to assess the proportion of variation in λ explained by process variation in each vital rate. In addition to matrix-based λ, we estimated λ using capture–recapture approaches, and log-linear regression. We found the stable age distribution for Pacific common eiders was weighted heavily towards experienced adult females (≥4 yr of age), and all calculations of λ indicated that the YKD population was stable to slightly increasing (λmatrix = 1.02, CI: 1.00–1.04); λreverse-capture–recapture = 1.05, CI: 0.99–1.11; λlog-linear = 1.04, CI: 0.98–1.10). Perturbation analyses suggested the population would respond most dramatically to changes in adult female survival (relative influence of adult survival was 1.5 times that of fecundity), whereas retrospective variation in λ was primarily explained by fecundity parameters (60%), particularly duckling survival (42%). Among components of fecundity, sensitivities were highest for duckling survival, suggesti

Alaska↗