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Jason I. Ransom

Publications and source records attributed to Jason I. Ransom.

At least 19 recordsLinked to original sources

Predicted habitat use for reintroduced grizzly bears in the transboundary North Cascades ecosystem

Grizzly bears ( Ursus arctos ) were once numerous in the North Cascades transboundary region of Washington State (United States) and British Columbia (Canada); however, few remain today. To support ongoing reintroduction evaluations, we used simulations based on movement models developed in the Northern Rocky Mountains to predict habitat use by a small founding group of grizzly bears in the North Cascades during the early stages of reintroduction. These simulations represent movements across a spatially explicit landscape based on individual-specific selection and movement parameters. We first evaluated predictive performance in three nearby populations of grizzly bears in the Squamish-Lillooet, McGillvary Mountains, and North Stein-Nahatlatch regions of the Coast Mountains, British Columbia. After applying an elevation-based calibration, predicted habitat use showed strong agreement with GPS location data from 73 collared bears. Across population–sex subgroups, Spearman rank correlations were ≥0.95, with 69.2%–83.3% of locations occurring within the top five habitat classes (covering 50% of mapped habitat) and 17.1%–36.2% in the top class (representing 10%). Overall, 24.5% and 77.9% of locations fell within the top class and top five classes, respectively. Seasonal validation showed strongest predictive performance from May to late summer or fall. Model predictions for the North Cascades indicate that habitat use during early reintroduction is likely to be concentrated in the central and northern mountainous portions of the ecosystem along the US–Canada border. These maps can guide recovery planning when no local bear data are yet available in the North Cascades transboundary region.

British Columbia, Washington

Freezing in a warming climate: Marked declines of a subnivean hibernator after a snow drought

Recent snow droughts associated with unusually warm winters are predicted to increase in frequency and affect species dependent upon snowpack for winter survival. Changes in populations of some cold‐adapted species have been attributed to heat stress or indirect effects on habitat from unusually warm summers, but little is known about the importance of winter weather to population dynamics and how responses to snow drought vary among sympatric species. We evaluated changes in abundance of hoary marmots ( Marmota caligata ) over a period that included a year of record‐low snowpack to identify mechanisms associated with weather and snowpack. To consider interspecies comparisons, our analysis used the same a priori model set as a concurrent study that evaluated responses of American pikas ( Ochotona princeps ) to weather and snowpack in the same study area of North Cascades National Park, Washington, USA. We hypothesized that marmot abundance reflected mechanisms related to heat stress, cold stress, cold exposure without an insulating snowpack, snowpack duration, atmospheric moisture, growing‐season precipitation, or select combinations of these mechanisms. Changes in marmot abundances included a 74% decline from 2007 to 2016 and were best explained by an interaction of chronic dryness with exposure to acute cold without snowpack in winter. Physiological stress during hibernation from exposure to cold, dry air appeared to be the most likely mechanism of change in marmot abundance. Alternative mechanisms associated with changes to winter weather, including early emergence from hibernation or altered vegetation dynamics, had less support. A post hoc assessment of vegetative phenology and productivity did not support vegetation dynamics as a primary driver of marmot abundance across years. Although marmot and pika abundances were explained by strikingly similar models over periods of many years, details of the mechanisms involved likely differ between species because pika abundances increased in areas where marmots declined. Such differences may lead to diverging geographic distributions of these species as global change continues.

Washington

Ecological consequences of anomalies in atmospheric moisture and snowpack

Although increased frequency of extreme‐weather events is one of the most secure predictions associated with contemporary climate change, effects of such events on distribution and abundance of climate‐sensitive species remain poorly understood. Montane ecosystems may be especially sensitive to extreme weather because of complex abiotic and biotic interactions that propagate from climate‐driven reductions in snowpack. Snowpack not only protects subnivean biotas from extreme cold, but also influences forage availability through timing of melt‐off and water availability. We related relative abundances of an alpine mammal, the American pika ( Ochotona princeps ), to measures of weather and snowpack dynamics over an 8‐yr period that included before and after a year of record‐low snowpack in Washington, USA. We sought to (1) quantify any change in pika abundance associated with the snowpack anomaly and (2) identify aspects of weather and snowpack that influenced abundance of pikas. Pikas showed a 1‐yr lag response to the snowpack anomaly and exhibited marked declines in abundance at elevations below 1,400 m simultaneous with increased abundances at higher elevations. Atmospheric moisture, indexed by vapor pressure deficit (VPD), was especially important, evidenced by strong support for the top‐ranked model that included the interaction of VPD with snowpack duration. Notably, our novel application of VPD from gridded climate data for analyses of animal abundances shows strong potential for improving species distribution models because VPD represents an important aspect of weather that influences the physiology and habitat of biota. Pikas were apparently affected by cold stress without snowpack at mid elevations, whereas changes to forage associated with snowpack and VPD were influential at high and low elevations. Our results reveal context dependency in pika responses to weather and illustrate how snow drought can lead to rapid change in the abundance of subnivean animals.

Washington

Trends in landbird density at two national parks in fragmented, mixed-use landscapes of the Pacific Northwest

National parks play a key role in conserving species by providing landscapes where threats from anthropogenic disturbance are reduced. In a recent study of 3 large wilderness parks in the Pacific Northwest, nearly all landbird species were found to be stable or increasing. Nonetheless, contemporary results from the Breeding Bird Survey and mark-recapture studies fuel concerns that some landbirds in the Pacific Northwest are trending in sync with many North America species in widespread decline. Although landbird populations might be thriving in large parks with extensive old-growth forest, those in smaller parks with less intact wilderness and higher ratios of edge-to-interior habitat might reflect the stressors inherent in more human-dominated landscapes. We conducted landbird point-count surveys from 2005 to 2016 in 2 national historical parks situated in the more human-dominated landscapes of this region, San Juan Island National Historical Park and Lewis and Clark National Historical Park. Established primarily to protect cultural resources, these parks lack old-growth forest and consist of relatively small parcels embedded in fragmented, multi-use landscapes. Here, we apply recent developments in point-count analysis to estimate trends in population density for 50 landbird species commonly detected in these small parks, including lagged effects of precipitation and temperature on the annual density of each species, and effects of survey conditions on species detection. All but 3 species exhibited stable densities in both parks, and more than half of the populations analyzed clearly increased in density over the study period. Notable exceptions were single-park declines in Olive-sided Flycatcher ( Contopus cooperi ), Northern Flicker ( Colaptes anratus ), and Hutton's Vireo ( Vireo huttoni ). Annual variation in population density was often related to climate, with generally positive responses to a recent deficit in annual precipitation-as-snow, and more variable responses to higher mean spring temperature. These results reinforce trends estimated for 3 large national parks in the Pacific Northwest, suggesting recent stability of landbird populations in parks of this region, independent of park size or setting.

Northwestern Naturalist

Trophic implications of a phenological paradigm shift: Bald eagles and salmon in a changing climate

Climate change influences apex predators in complex ways, due to their important trophic position, capacity for resource plasticity, and sensitivity to numerous anthropogenic stressors. Bald eagles, an ecologically and culturally significant apex predator, congregate seasonally in high densities on salmon spawning rivers across the Pacific Northwest. One of the largest eagle concentrations is in the Skagit River watershed, which connects the montane wilderness of North Cascades National Park to the Puget Sound. Using multiple long‐term datasets, we evaluated local bald eagle abundance in relation to chum and coho salmon availability; salmon phenology; and the number and timing of flood events in the Skagit. We analysed changes over time as a reflection of climate change impacts, as well as differences between managed and unmanaged portions of the river. We found that peaks in chum salmon and bald eagle presence have advanced at remarkably similar rates ( c . 0.45 days/year), suggesting synchronous phenological responses within this trophic relationship. Yet the temporal relationship between chum salmon spawning and flood events, which remove salmon carcasses from the system, has not remained constant. This has resulted in a paradigm shift whereby the peak of chum spawning now occurs before the first flood event of the season rather than after. The interval between peak chum and first flood event was a significant predictor of bald eagle presence: as this interval grew over time (by nearly one day per year), bald eagle counts declined, with a steady decrease in bald eagle observations since 2002. River section was also an important factor, with fewer flood events, and more eagle observations occurring in the river section experiencing direct hydroelectric flow management. Synthesis and applications . The effects of climate change and hydroelectric management contribute to a complex human footprint in the North Cascades National Park, an otherwise largely natural ecosystem. By accounting for the differential phenological impacts of climate change on bald eagles, salmon, and flood events, Park managers and the operators of the hydroelectric system can more effectively ensure the resilience of the eagle–salmon relationship along the Skagit River.

Washington

Landbird population trends in mountain and historical parks of the North Coast and Cascades Network: 2005–2016 synthesis

Long-term monitoring of landbird populations within the National Park Service (NPS) North Coast and Cascades Inventory and Monitoring Network (NCCN) began in 2005, with the goal of detecting trends to inform the conservation and management of landbirds and their habitats. Here we use 2005–2016 data from over 3500 point-count stations to report landbird occurrence and trends in each of five NCCN parks, including three national parks in mountain wilderness areas (Mount Rainier National Park, North Cascades National Park Complex and Olympic National Park) and two historical parks (Lewis and Clark National Historical Park and San Juan Island National Historical Park). Recent advances in point-count modeling were applied to characterize population trends for 68 landbird species, including up to 41 species in each park. Fitted models suggest that almost all species exhibited stable or increasing trends over the study period. Notable exceptions were a decline in the Olive-sided Flycatcher in two parks and single-park declines in the Norther Flicker, Hutton’s Vireo, Clark’s Nutcracker, Mountain Chickadee, Wilson’s Warbler and Dark-eyed Junco. Negative effects of precipitation-as-snow were supported in over one-third of our population models. Lower precipitation-as-snow in the mountain parks might have contributed to rising landbird densities during the study period. Population density also varied with elevation in mountain parks, but temporal trends were similar among elevational strata for each species analyzed, suggesting no evidence of elevational range-shifts during this study. These results reinforce recent analyses of the first 10 years of point-count data from the three mountain parks (Ray et al. 2017 a). In the current analysis, models were extended to explore effects of covariates on species detection probability. Negative effects of ambient noise level on detection were supported in several cases, but adding covariates of detection generally did not lead to substantial improvements in model fit. In some cases, model fit was improved by reducing the scope of inference to a portion of the focal region, suggesting important effects of habitat heterogeneity.

Natural Resource Report

Recent stability of resident and migratory landbird populations in National Parks of the Pacific Northwest

Monitoring species in National Parks facilitates inference regarding effects of climate change on population dynamics because parks are relatively unaffected by other forms of anthropogenic disturbance. Even at early points in a monitoring program, identifying climate covariates of population density can suggest vulnerabilities to future change. Monitoring landbird populations in parks during the breeding season brings the added benefit of allowing a comparative approach to inference across a large suite of species with diverse requirements. For example, comparing resident and migratory species that vary in exposure to non-park habitats can reveal the relative importance of park effects, such as those related to local climate. We monitored landbirds using breeding-season point-count data collected during 2005–2014 in three wilderness areas of the Pacific Northwest (Mount Rainier, North Cascades, and Olympic National Parks). For 39 species, we estimated recent trends in population density while accounting for individual detection probability using Bayesian hierarchical N -mixture models. Our analyses integrated several recent developments in N -mixture modeling, incorporating interval and distance sampling to estimate distinct components of detection probability while also accommodating count intervals of varying duration, annual variation in the length and number of point-count transects, spatial autocorrelation, random effects, and covariates of detection and density. As covariates of density, we considered metrics of precipitation and temperature hypothesized to affect breeding success. We also considered effects of park and elevational stratum on trend. Regardless of model structure, we estimated stable or increasing densities during 2005–2014 for most populations. Mean trends across species were positive for migrants in every park and for residents in one park. A recent snowfall deficit in this region might have contributed to the positive trend, because population density varied inversely with precipitation-as-snow for both migrants and residents. Densities varied directly but much more weakly with mean spring temperature. Our approach exemplifies an analytical framework for estimating trends from point-count data, and for assessing the role of climatic and other spatiotemporal variables in driving those trends. Understanding population trends and the factors that drive them is critical for adaptive management and resource stewardship in the context of climate change.

Ecosphere

Development of a robust analytical framework for assessing landbird trends, dynamics and relationships with environmental covariates in the North Coast and Cascades Network

During 2015-2016, we completed development of a new analytical framework for landbird population monitoring data from the National Park Service (NPS) North Coast and Cascades Inventory and Monitoring Network (NCCN). This new tool for analysis combines several recent advances in modeling population status and trends using point-count data and is designed to supersede the approach previously slated for analysis of trends in the NCCN and other networks, including the Sierra Nevada Network (SIEN). Advances supported by the new model-based approach include 1) the use of combined data on distance and time of detection to estimate detection probability without assuming perfect detection at zero distance, 2) seamless accommodation of variation in sampling effort and missing data, and 3) straightforward estimation of the effects of downscaled climate and other local habitat characteristics on spatial and temporal trends in landbird populations. No changes in the current field protocol are necessary to facilitate the new analyses. We applied several versions of the new model to data from each of 39 species recorded in the three mountain parks of the NCCN, estimating trends and climate relationships for each species during 2005-2014. Our methods and results are also reported in a manuscript in revision for the journal Ecosphere (hereafter, Ray et al.). Here, we summarize the methods and results outlined in depth by Ray et al., discuss benefits of the new analytical framework, and provide recommendations for its application to synthetic analyses of long-term data from the NCCN and SIEN. All code necessary for implementing the new analyses is provided within the Appendices to this report, in the form of fully annotated scripts written in the open-access programming languages R and JAGS.

Natural Resource Report

Practical bias correction in aerial surveys of large mammals: Validation of hybrid double-observer with sightability method against known abundance of feral horse (Equus caballus) populations

Reliably estimating wildlife abundance is fundamental to effective management. Aerial surveys are one of the only spatially robust tools for estimating large mammal populations, but statistical sampling methods are required to address detection biases that affect accuracy and precision of the estimates. Although various methods for correcting aerial survey bias are employed on large mammal species around the world, these have rarely been rigorously validated. Several populations of feral horses ( Equus caballus ) in the western United States have been intensively studied, resulting in identification of all unique individuals. This provided a rare opportunity to test aerial survey bias correction on populations of known abundance. We hypothesized that a hybrid method combining simultaneous double-observer and sightability bias correction techniques would accurately estimate abundance. We validated this integrated technique on populations of known size and also on a pair of surveys before and after a known number was removed. Our analysis identified several covariates across the surveys that explained and corrected biases in the estimates. All six tests on known populations produced estimates with deviations from the known value ranging from -8.5% to +13.7% and <0.7 standard errors. Precision varied widely, from 6.1% CV to 25.0% CV. In contrast, the pair of surveys conducted around a known management removal produced an estimated change in population between the surveys that was significantly larger than the known reduction. Although the deviation between was only 9.1%, the precision estimate (CV = 1.6%) may have been artificially low. It was apparent that use of a helicopter in those surveys perturbed the horses, introducing detection error and heterogeneity in a manner that could not be corrected by our statistical models. Our results validate the hybrid method, highlight its potentially broad applicability, identify some limitations, and provide insight and guidance for improving survey designs.

Colorado, Nevada, Utah, Wyoming

Behavior of feral horses in response to culling and GnRH immunocontraception

Wildlife management actions can alter fundamental behaviors of individuals and groups,which may directly impact their life history parameters in unforeseen ways. This is especially true for highly social animals because changes in one individual&rsquo;s behavior can cascade throughout its social network. When resources to support populations of social animals are limited and populations become locally overabundant, managers are faced with the daunting challenge of decreasing population size without disrupting core behavioral processes. Increasingly, managers are turning to fertility control technologies to supplement culling in efforts to suppress population growth, but little is quantitatively known about how either of these management tools affects behavior. Gonadotropin releasing hormone (GnRH) is a small neuropeptide that performs an obligatory role in mammalian reproduction and has been formulated into the immunocontraceptive GonaCon-BTM. We investigated the influences of this vaccine on behavior of feral horses ( Equus caballus ) at Theodore Roosevelt National Park, North Dakota, USA, for a year preceding and a year following nonlethal culling and GnRH-vaccine treatment. We observed horses during the breeding season and found only minimal differences in time budget behaviors of free-ranging female feral horses treated with GnRH and those treated with saline. The differences observed were consistent with the metabolic demands of pregnancy and lactation. We observed similar social behaviors between treatment groups, reflecting limited reproductive behavior among control females due to high rates of pregnancy and suppressed reproductive behavior among treated females due to GnRH-inhibited ovarian activity. In the treatment year, band stallion age was the only supported factor influencing herding behavior (P < 0.001), harem-tending behavior (P < 0.001), and agonistic behavior (P = 0.02). There was no difference between the mean body condition of control females (4.9 (95% CI = 4.7&ndash;5.1)) and treated females(4.8 (95% CI = 4.7&ndash;4.9)). Band fidelity among all females increased 25.7% in the year fol-lowing vaccination and culling, despite the social perturbation associated with removal of conspecifics. Herding behavior by stallions decreased 50.7% following treatment and culling (P < 0.001), while harem-tending behavior increased 195.0% (P < 0.001). The amount of available forage influenced harem-tending, reproductive, and agonistic behavior in the year following culling and treatment (P < 0.04). These changes reflected the expected nexus between a species with polygynous social structure and strong group fidelity and the large instantaneous change in population density and demography coincident with culling.Behavioral responses to such perturbation may be synergistic in reducing grazing pressure by decreasing energetically expensive competitive behaviors, but further investigation is needed to explicitly test this hypothesis.

Applied Animal Behaviour Science

Influence of nonnative and native ungulate biomass and seasonal precipitation on vegetation production in a Great Basin ecosystem

The negative effects of equid grazers in semiarid ecosystems of the American West have been considered disproportionate to the influence of native ungulates in these systems because of equids' large body size, hoof shape, and short history on the landscape relative to native ungulates. Tools that can analyze the degree of influence of various ungulate herbivores in an ecosystem and separate effects of ungulates from effects of other variables (climate, anthropomorphic disturbances) can be useful to managers in determining the location of nonnative herbivore impacts and assessing the effect of management actions targeted at different ungulate populations. We used remotely sensed data to determine the influence of native and nonnative ungulates and climate on vegetation productivity at wildlife refuges in Oregon and Nevada. Our findings indicate that ungulate biomass density, particularly equid biomass density, and precipitation in winter and spring had the greatest influence on normalized difference vegetation index (NDVI) values. Our results concur with those of other researchers, who found that drought exacerbated the impacts of ungulate herbivores in arid systems.

Nevada, Oregon

Ecological feedbacks can reduce population-level efficacy of wildlife fertility control

1. Anthropogenic stress on natural systems, particularly the fragmentation of landscapes and the extirpation of predators from food webs, has intensified the need to regulate abundance of wildlife populations with management. Controlling population growth using fertility control has been considered for almost four decades, but nearly all research has focused on understanding effects of fertility control agents on individual animals. Questions about the efficacy of fertility control as a way to control populations remain largely unanswered. 2. Collateral consequences of contraception can produce unexpected changes in birth rates, survival, immigration and emigration that may reduce the effectiveness of regulating animal abundance. The magnitude and frequency of such effects vary with species-specific social and reproductive systems, as well as connectivity of populations. Developing models that incorporate static demographic parameters from populations not controlled by contraception may bias predictions of fertility control efficacy. 3. Many population-level studies demonstrate that changes in survival and immigration induced by fertility control can compensate for the reduction in births caused by contraception. The most successful cases of regulating populations using fertility control come from applications of contraceptives to small, closed populations of gregarious and easily accessed species. 4. Fertility control can result in artificial selection pressures on the population and may lead to long-term unintentional genetic consequences. The magnitude of such selection is dependent on individual heritability and behavioural traits, as well as environmental variation. 5. Synthesis and applications . Understanding species' life-history strategies, biology, behavioural ecology and ecological context is critical to developing realistic expectations of regulating populations using fertility control. Before time, effort and funding are invested in wildlife contraception, managers may need to consider the possibility that many species and populations can compensate for reduction in fecundity, and this could minimize any reduction in population growth rate.

Journal of Applied Ecology

Contraception can lead to trophic asynchrony between birth pulse and resources

Abiotic inputs such as photoperiod and temperature can regulate reproductive cyclicity in many species. When humans perturb this process by intervening in reproductive cycles, the ecological consequences may be profound. Trophic mismatches between birth pulse and resources in wildlife species may cascade toward decreased survival and threaten the viability of small populations. We followed feral horses (Equus caballus) in three populations for a longitudinal study of the transient immunocontraceptive porcine zona pellucida (PZP), and found that repeated vaccinations extended the duration of infertility far beyond the targeted period. After the targeted years of infertility, the probability of parturition from post-treated females was 25.6% compared to 64.1% for untreated females, when the data were constrained only to females that had demonstrated fertility prior to the study. Estimated time to parturition increased 411.3 days per year of consecutive historical treatment. Births from untreated females in these temperate latitude populations were observed to peak in the middle of May, indicating peak conception occurred around the previous summer solstice. When the post-treated females did conceive and give birth, parturition was an estimated 31.5 days later than births from untreated females, resulting in asynchrony with peak forage availability. The latest neonate born to a post-treated female arrived 7.5 months after the peak in births from untreated females, indicating conception occurred within 24–31 days of the winter solstice. These results demonstrate surprising physiological plasticity for temperate latitude horses, and indicate that while photoperiod and temperature are powerful inputs driving the biological rhythms of conception and birth in horses, these inputs may not limit their ability to conceive under perturbed conditions. The protracted infertility observed in PZP-treated horses may be of benefit for managing overabundant wildlife, but also suggests caution for use in small refugia or rare species.

PLoS ONE

A collaborative approach for estimating terrestrial wildlife abundance

Accurately estimating abundance of wildlife is critical for establishing effective conservation and management strategies. Aerial methodologies for estimating abundance are common in developed countries, but they are often impractical for remote areas of developing countries where many of the world's endangered and threatened fauna exist. The alternative terrestrial methodologies can be constrained by limitations on access, technology, and human resources, and have rarely been comprehensively conducted for large terrestrial mammals at landscape scales. We attempted to overcome these problems by incorporating local peoples into a simultaneous point count of Asiatic wild ass (Equus hemionus) and goitered gazelle (Gazella subgutturosa) across the Great Gobi B Strictly Protected Area, Mongolia. Paired observers collected abundance and covariate metrics at 50 observation points and we estimated population sizes using distance sampling theory, but also assessed individual observer error to examine potential bias introduced by the large number of minimally trained observers. We estimated 5671 (95% CI = 3611&ndash;8907) wild asses and 5909 (95% CI = 3762&ndash;9279) gazelle inhabited the 11,027 km 2 study area at the time of our survey and found that the methodology developed was robust at absorbing the logistical challenges and wide range of observer abilities. This initiative serves as a functional model for estimating terrestrial wildlife abundance while integrating local people into scientific and conservation projects. This, in turn, creates vested interest in conservation by the people who are most influential in, and most affected by, the outcomes.

Biological Conservation

Detection probability in aerial surveys of feral horses

Observation bias pervades data collected during aerial surveys of large animals, and although some sources can be mitigated with informed planning, others must be addressed using valid sampling techniques that carefully model detection probability. Nonetheless, aerial surveys are frequently employed to count large mammals without applying such methods to account for heterogeneity in visibility of animal groups on the landscape. This often leaves managers and interest groups at odds over decisions that are not adequately informed. I analyzed detection of feral horse (Equus caballus) groups by dual independent observers from 24 fixed-wing and 16 helicopter flights using mixed-effect logistic regression models to investigate potential sources of observation bias. I accounted for observer skill, population location, and aircraft type in the model structure and analyzed the effects of group size, sun effect (position related to observer), vegetation type, topography, cloud cover, percent snow cover, and observer fatigue on detection of horse groups. The most important model-averaged effects for both fixed-wing and helicopter surveys included group size (fixed-wing: odds ratio = 0.891, 95% CI = 0.850–0.935; helicopter: odds ratio = 0.640, 95% CI = 0.587–0.698) and sun effect (fixed-wing: odds ratio = 0.632, 95% CI = 0.350–1.141; helicopter: odds ratio = 0.194, 95% CI = 0.080–0.470). Observer fatigue was also an important effect in the best model for helicopter surveys, with detection probability declining after 3 hr of survey time (odds ratio = 0.278, 95% CI = 0.144–0.537). Biases arising from sun effect and observer fatigue can be mitigated by pre-flight survey design. Other sources of bias, such as those arising from group size, topography, and vegetation can only be addressed by employing valid sampling techniques such as double sampling, mark–resight (batch-marked animals), mark–recapture (uniquely marked and identifiable animals), sightability bias correction models, and line transect distance sampling; however, some of these techniques may still only partially correct for negative observation biases.

Journal of Wildlife Management

Demography of the Pryor Mountain wild horses, 1993-2007

Wild horses (Equus caballus) at Pryor Mountain were studied by direct observation from 1993 through 2007. All horses present were individually identifiable on the basis of coat coloration, head and leg markings, gender, and band associations. Of the 609 horses either present prior to foaling in 1993 or born since, ages were precisely known for 491 (observed as a foal). Ages for 52 horses were estimated through tooth eruption and wear patterns, and for the remaining 66 horses through body size, morphology, and anecdotal evidence concerning when they were present on the range. At varying intensities, never less than 30 days per year, all horses were inventoried and their band associations noted. Foals were paired with dams based on observations of attachment during the early days and weeks of life. Year of death was determined by identification of the carcass where possible. In the absence of finding a carcass, an animal that was not observed for 2 years was considered to have died in the year that it went missing. Animals that were removed from the herd and mares that were part of a contraception study were excluded from calculations of survival and foaling rates, respectively, as appropriate. The average prefoaling population over the 15 years of the study was 148.8 animals (range = 120-187), and the annual foal crop averaged 32.1 (range = 23-40). Large removals (19-60 animals) in four years helped maintain the herd at this level; apparent growth rate (calculated as though removals had not occurred) was 9.6 percent annually (? = 1.096, range = 0.977-1.220). This annual growth rate is relatively low compared to that for many western horse herds, at least in part because of a decline in foal survival. Sex ratio of the foal crop varied widely among years, but pooled across years did not differ from 50:50. Sex ratio in the herd changed mostly as a result of removals. The average age of both males and females in the herd increased during the course of the study. Annual survival of males did not differ from that of females, nor did gender affect annual survival of foals. Pooled across years, ages, and sexes, the annual survival rate was 0.899. Annual foal survival rate was 0.697 and declined through time, with a tendency toward recovery in 2005-2007. Foal survival was higher in larger bands, but did not differ between foals born to primiparous and multiparous mares. A few 2-year-old mares produced foals; foaling rate (excluding contracepted mares and foals they produced) increased through age 10, remained high through age 15, and declined thereafter. Overall foaling rate for mares =3 years of age was 0.576 foals per mare, with no apparent trend during the period of our study. Foaling rate in years following gathers was somewhat lower than in other years. There was a positive relation between foaling rate and band size. Primiparous mares were somewhat less likely to foal in the following year than were multiparous mares. Most stallions that acquired a harem did so at age 5 or 6, and the average age of harem stallions increased during our study. Most harems had 1-3 mares =2 years of age, but harem size varied with age of the stallion, increasing through about age 11 and declining thereafter. About 6 percent of bands had a satellite stallion (=5 years of age), but the mean number of mares did not differ between single- and multistallion bands. Most stallions left their natal band at age 2 or 3, but 17 percent remained with their natal band until age 4 or 5. Foal survival rate was positively related to precipitation, suggesting a possible link to forage production and availability mediated through mare fitness. There also was evidence for density-dependent population regulation, as both population growth rate and survival rate were negatively correlated with population size from the previous year. These and other factors were not sufficient to stabilize the population during our period of study, however, as evidenced by the necess

Scientific Investigations Report

Injection-Site Reactions in Wild Horses (Equus caballus) Receiving an Immunocontraceptive Vaccine

The U.S. Geological Survey and the Bureau of Land Management are conducting research on the efficacy of the immunocontraceptive agent porcine zona pellucida (PZP) in reducing fertility of wild horses (Equus caballus). As an antigen, PZP stimulates antibody production when injected into many mammalian species. These antibodies bind to the external surface of the ovum, preventing fertilization. By itself, PZP is only weakly immunogenic and is therefore delivered with an adjuvant, most commonly one of the Freund adjuvants, designed to further stimulate antibody production. Freund's complete adjuvant (FCA) in particular is known to be very effective, but may also be associated with undesirable side effects such as formation of abscesses at injection sites. Such reactions may be exacerbated when accompanied by the additional trauma of a remotely delivered dart. Because horses in our three study herds were individually identifiable by color markings and harem association, we were able to monitor mares for injection-site reactions (abscesses, nodules, swelling, and stiffness) following inoculation with PZP. In 100 injections delivered by hand we observed a single nodule, two instances of swelling, and no other reactions. In two herds that received remotely delivered (dart) injections, the frequency of reactions was about 1 and 6 percent for abscesses, 25 percent for nodules (both herds), 11 and 33 percent for swelling, and 1 and 12 percent for stiffness. Abscesses were too infrequent to allow meaningful analysis of the relation to covariates, but for the other types of reactions we used logistic regression to examine the relation of occurrence to the delivery method (rifle or CO2-powered blowgun), adjuvant (FCA, Freund's modified adjuvant, and Freund's incomplete adjuvant), dart trauma (normal or abnormal), and age of mare. Abnormal dart trauma included cases where the dart hit bone or the needle broke off. We found strong evidence (odds ratio = 5.023, P = 0.001) for a higher probability of occurrence of swelling when darts were delivered by blowgun. We found some evidence (odds ratio = 8.729, P = 0.07) that abnormal dart trauma led to a higher frequency of nodule formation. Nodules were the most common reactions observed and often persisted for a year or more, but in our observations they did not appear to change any animal's range of movement or locomotor patterns and in most cases did not appear to differ in magnitude from naturally occurring injuries or scars. We were unable to perform histological examinations of these nodules, but they may be similar to granulomas reported by other investigators following administration of Freund's adjuvant.

Scientific Investigations Report

Quantifying Equid Behavior - A Research Ethogram for Free-Roaming Feral Horses

Feral horses (Equus caballus) are globally distributed in free-roaming populations on all continents except Antarctica and occupy a wide range of habitats including forest, grassland, desert, and montane environments. The largest populations occur in Australia and North America and have been the subject of scientific study for decades, yet guidelines and ethograms for feral horse behavioral research are largely absent in the scientific literature. The U.S. Geological Survey (USGS) Fort Collins Science Center conducted research on the influences of the immunocontraceptive porcine zona pellucida (PZP) on feral horse behavior from 2003-2006 in three discrete populations in the American west. These populations were the Little Book Cliffs Wild Horse Range in Colorado, McCullough Peaks Herd Management Area in Wyoming, and Pryor Mountain Wild Horse Range in Montana; the research effort included over 1,800 hours of behavioral observations of 317 adult free-roaming feral horses. An ethogram was developed during the course of this study to facilitate accurate scientific data collection on feral horse behavior, which is often challenging to quantify. By developing this set of discrete behavioral definitions and a set of strict research protocols, scientists were better able to address both applied questions, such as behavioral changes related to fertility control, and theoretical questions, such as understanding networks and dominance hierarchies within social groups of equids.

Techniques and Methods