Feral Horse Populations Research Articles

Suppression of reproductive behaviour and gonadal function in female horses-An update.

The behaviour of mares is often detrimental to their performance resulting in frequent demand for methods to suppress gonadal function. In addition, prevention of unintended reproduction especially in feral horse populations may require methods for suppression of gonadal function. Surgical ovariectomy is a safe method but not an acceptable approach in feral mares and undesired in mares where future breeding is considered. There are different approaches for artificial prolongation of the luteal phase resulting in transient inhibition of oestrus and ovulation. Among those, treatment with natural or synthetic progestogens is considered the most common and successful method. Whereas application of intrauterine devices may result in prolongation of luteal function in non-pregnant mares, intrauterine insertion of glass balls is no longer recommended because of complications in individual mares. There are several safer alternatives that may be of interest, especially for population control in free-roaming horses. Treatment with long-acting deslorelin implants inhibited ovulation and oestrus behaviour in mares for limited and variable time intervals in a dose-dependent manner. The effect of GnRH vaccines varies considerably among individual mares, is age dependent, and oestrus-like behaviour may still occur. Contraception via immunization against native porcine or recombinant zona pellucida antigen is successful, but immunocontraception is as much a result of ovarian inactivity as an antibody-based block to sperm-oocyte binding. In conclusion, several treatments for suppression of gonadal function in mares are available, but there are advantages and disadvantages associated that have to be considered. The treatment of choice will thus differ with regard to the demands.

HIGH PREGNANCY RATES IN TWO-YEAR-OLD WILD HORSES

In November 2018, we screened 315 feral Horse mares via veterinary rectal palpation and trans-rectal ultrasound, approximately 1 mo after their capture and removal from the Warm Springs herd management area in Oregon. We documented a pregnancy rate (58.3%) in 2-y-old mares that was nearly 3 times higher than previously published results. For 1-y-old mares (10.8% pregnant), and for mares pooled over all ages (64.4% pregnant), pregnancy rates were within the range of pregnancy rates reported from other feral Horse populations. High pregnancy rates in 2-y-old mares could contribute to higher than expected herd growth rates in feral Horses.

Social determinants of affiliation and cohesion in a population of feral horses

Investigating affiliative behaviors (e.g. proximity, grooming, cooperative behaviors) is essential to understand group cohesion and stability in animal societies, as they may foster, or be fostered by long-term social bonds and ultimately determine an individual’s lifetime reproductive success. Despite growing interest in affiliative behaviors across a wide range of mammalian taxa, little focus has been given to feral horse populations. In this study, we examined the influence of dyadic social factors (sex, rank difference, familiarity and genetic relatedness) on affiliative behaviors and group cohesion. We collected data on 16 feral horse groups in Northern Portugal who were subjected to both predation pressures and anthropogenic interference. A combination of all occurrence sampling methods and drone technology was used to record the horses’ social behavior and spatial data. Same-sex dyads showed a preference to remain in close proximity, however they did not groom as frequently as different-sex dyads. Similar rank individuals were observed in proximity more often than non-similar rank individuals, but only in newly formed groups. Familiar individuals spent more time in proximity and groomed more often than unfamiliar (new) dyads. However, affiliation between conspecifics was not predicted by genetic relatedness. These results suggest that social bonds and group cohesion are mostly sustained by familiar individuals in both stable and newly formed groups. While maintaining close proximity may function to sustain bonds between same-sex individuals, grooming may reinforce both intra-sexual and inter-sexual bonds. Examining the variation of affiliation patterns on horse populations may shed light on their behavioral plasticity and have crucial implications for the welfare and conservation of horse breeds.

Sage‐Grouse Population Dynamics are Adversely Affected by Overabundant Feral Horses

ABSTRACTIn recent decades, feral horse (Equus caballus; horse) populations increased in sagebrush (Artimesia spp.) ecosystems, especially within the Great Basin, to the point of exceeding maximum appropriate management levels (AMLmax), which were set by land administrators to balance resource use by feral horses, livestock, and wildlife. Concomitantly, greater sage‐grouse (Centrocercus urophasianus; sage‐grouse) are sagebrush obligates that have experienced population declines within these same arid environments as a result of steady and continued loss of seasonal habitats. Although a strong body of research indicates that overabundant populations of horses degrade sagebrush ecosystems, empirical evidence linking horse abundance to sage‐grouse population dynamics is missing. Within a Bayesian framework, we employed state‐space models to estimate population rate of change (λ) using 15 years (2005–2019) of count surveys of male sage‐grouse at traditional breeding grounds (i.e., leks) as a function of horse abundance relative to AMLmax and other environmental covariates (e.g., wildfire, precipitation, % sagebrush cover). Additionally, we employed a post hoc impact‐control design to validate existing AMLmax values as related to sage‐grouse population responses, and to help control for environmental stochasticity and broad‐scale oscillations in sage‐grouse abundance. On average, for every 50% increase in horse abundance over AMLmax, our model predicted an annual decline in sage‐grouse abundance by 2.6%. Horse abundance at or below AMLmax coincided with sage‐grouse λ estimates that were consistent with trends at non‐horse areas elsewhere in the study region. Thus, AMLmax, as a whole, appeared to be set adequately in preventing adverse effects to sage‐grouse populations. Results indicated 76%, 97%, and >99% probability of sage‐grouse population decline relative to controls when horse numbers are 2, 2.5, and ≥3 times over AMLmax, respectively. As of 2019, horse herds exceeded AMLmax in Nevada, USA, by >4 times on average across all horse management areas. If feral horse populations continue to grow at current rates unabated, model projections indicate sage‐grouse populations will be reduced within horse‐occupied areas by >70.0% by 2034 (15‐year projection), on average compared to 21.2% estimated for control sites. A monitoring framework that improves on estimating horse abundance and identifying responses of sage‐grouse and other key indicator species (plant and animal) would be beneficial to guide management decisions that promote co‐occurrence of horses with sensitive wildlife and livestock within landscapes subjected to multiple uses. Published 2021. This article is a U.S. Government work and is in the public domain in the USA. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.

Prey Specialization by Cougars on Feral Horses in a Desert Environment

ABSTRACTNatural controls on the distribution, abundance, or growth rates of exotic species are a desirable mode of intervention because of lower costs compared to anthropogenic controls and greater social acceptance. In the Great Basin, cougars (Puma concolor) are the most widely distributed carnivore capable of killing large ungulate prey. Populations of feral horses (Equus ferus) are widely distributed throughout the Great Basin and can grow at rates up to 20%/year. Although cougars exhibit distributional overlap with horses, it has been assumed that predation is minimal because of differences in habitat use and body‐size limitations. To evaluate this hypothesis, we monitored the diets of 21 global positioning system (GPS)‐collared cougars in the western Great Basin (5 males, 8 females) and eastern Sierra Nevada (2 males, 6 females) from 2009–2012. We investigated 1,310 potential kill sites and located prey remains of 820 predation events. We compared prey composition and kill rates of cougars inhabiting the Sierra Nevada and Great Basin, and among male and female cougars across seasons. We used generalized linear mixed models (GLMMs) to examine the effects of prey availability and habitat characteristics on the probability of predation on horses by cougars. Mule deer (Odocoileus hemionus) comprised 91% of prey items killed on the Sierra Nevada reference site but only comprised 29% of prey items in the Great Basin study area. Average annual kill rates for deer differed between the Sierra Nevada ( = 0.85 deer/week, range = 0.44–1.3) and Great Basin ( = 0.21 deer/week, range = 0.00–0.43). Diets of cougars in the Great Basin were composed predominantly of horses (59.6%, n = 460 prey items; 13 individuals). Ten cougars regularly consumed horses, and horses were the most abundant prey in the diet of 8 additional individuals in the Great Basin. Cougars on average killed 0.38 horses/week in the Great Basin (range=0.00–0.94 horses/week). Differences in predation on horses between the sexes of cougars were striking; Great Basin females incorporated more horses across all age classes year‐round, whereas male cougars tended to exploit neonatal young during spring and summer before switching to deer during winter. Within GLMM models, the probability of predation on horses compared to other prey species increased with elevation, horse density, and decreasing density of mule deer on the landscape, and was more likely to occur in sagebrush (Artemesia spp.) than in pinyon (Pinus monophylla)–juniper (Juniperus osteosperma) forests. Behavior of individual cougars accounted for more than a third of the variation explained by our top models predicting predation on horses in the Great Basin. At landscape scales, cougar predation is unlikely to limit the growth of feral horse populations. In the Great Basin ecosystem, however, cougars of both sexes successfully preyed on horses of all age classes. Moreover, some reproductive, female cougars were almost entirely dependent on feral horses year‐round. Taken together, our data suggest that cougars may be an effective predator of feral horses, and that some of our previous assumptions about this relationship should be reevaluated and integrated into management and planning. © 2021 The Wildlife Society.

Using Fecal DNA and Closed‐Capture Models to Estimate Feral Horse Population Size

ABSTRACTAccurate 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. © 2021 The Wildlife Society. This article has been contributed to by US Government employees and their work is in the public domain in the USA.

Long-term storage of feces at -80 °C versus -20 °C is negligible for 16S rRNA amplicon profiling of the equine bacterial microbiome.

The development of next-generation sequencing technologies has spurred a surge of research on bacterial microbiome diversity and function. But despite the rapid growth of the field, many uncertainties remain regarding the impact of differing methodologies on downstream results. Sample storage temperature is conventionally thought to be among the most important factors for ensuring reproducibility across marker gene studies, but to date much of the research on this topic has focused on short-term storage in the context of clinical applications. Consequently, it has remained unclear if storage at −80 °C, widely viewed as the gold standard for long-term archival of feces, is truly required for maintaining sample integrity in amplicon-based studies. A better understanding of the impacts of long-term storage conditions is important given the substantial cost and limited availability of ultra-low temperature freezers. To this end, we compared bacterial microbiome profiles inferred from 16S V3–V4 amplicon sequencing for paired fecal samples obtained from a feral horse population from Sable Island, Nova Scotia, Canada, stored at either −80 °C or −20 °C for 4 years. We found that storage temperature did not significantly affect alpha diversity measures, including amplicon sequence variant (ASV) richness and evenness, and abundance of rare sequence variants, nor presence/absence, relative abundances and phylogenetic diversity weighted measures of beta diversity. These results indicate that storage of equine feces at −20 °C for periods ranging from a few months to a few years is equivalent to storage at −80 °C for amplicon-based microbiome studies, adding to accumulating evidence indicating that standard domestic freezers are both economical and effective for microbiome research.

Feral Horse Space Use and Genetic Characteristics from Fecal DNA

ABSTRACTFeral horses (Equus ferus caballus) in the western United States are managed by the Bureau of Land Management (BLM) and United States Forest Service in designated areas on public lands with a goal of maintaining populations in balance with multiple uses of the landscape. Small, isolated populations can be at risk of extirpation from stochastic events and deleterious genetic effects resulting from inbreeding and reduced heterozygosity. The genetic diversity of feral horse herds is periodically monitored using blood or hair samples collected during management gathers (i.e., occasions when the herd is rounded up). We conducted a study to examine genetic characteristics of the feral horse population at the BLM Little Book Cliffs Herd Management Area (HMA) in Colorado, USA, using non‐invasively collected fecal samples. Additionally, we explored whether genotypes could be used to document space use and potential sub‐population development. We used a random sampling scheme, walking transects in sampling areas covering most of the HMA to find and collect fecal samples of all ages, except those that were deteriorating. We collected >1,800 fecal samples from across the study area in May, August, and October 2014. We then identified unique individuals using a suite of microsatellite loci. Our estimates of genetic diversity from fecal samples were higher than those reported from blood and hair samples taken during recent horse gathers, likely because our sample size and spatial distribution was larger. Genotypes revealed that some individuals were found only in certain parts of the study area and at a higher proportion than random; thus, they could be considered residents in those sampling areas. Using discriminant function analyses, we detected 5 genetic groups in the sample population, but these did not correspond to individuals in specific parts of the study area. Our results support the use of fecal DNA to augment direct observations of horse presence and could be used to detect habitat use and areas of high density. Non‐invasive techniques such as fecal DNA sampling can help managers decide whether new individuals need to be translocated to a closed population to maintain genetic diversity without the human safety and animal welfare concerns associated with gathers and invasive techniques. © 2021 The Wildlife Society.

Ecological Interactions Involving Feral Horses and Predators: Review with Implications for Biodiversity Conservation

ABSTRACTFor many ecosystems, feral horses are increasingly becoming an important if not dominant component of ungulate biomass and hence influence on community dynamics. Yet we still know little of how horses contribute to key ecological interactions including predator‐prey and indirect competitive relationships at a community level. Notably, feral species like horses can exhibit life‐history traits that differ from that of native (mainly artiodactyl) herbivore competitors. Artificial selection for traits like increased, early, or extended reproduction that have yet to be reversed by natural selection, coupled with naturally selected differences in anatomy and behavior, in addition to unique management objectives for horses compared to other species, means that the dynamics of feral horse populations are not likely to align with what might be expected of other large herbivores. Unexpected population dynamics and inherent biological asymmetries between native ungulates and feral horses may therefore influence the former via direct competition for shared resources and through enemy‐mediated interactions like apparent competition. In several localities feral horses now co‐exist with multiple native prey species, some of which are in decline or are species at risk. Compounding risks to native species from direct or indirect competitive exclusion by horses is the unique nature and socio‐political context of feral horse management, which tends towards allowing horse populations to be limited largely by natural, density‐dependent factors. We summarize the inherent asymmetries between feral horse biology and that of other ungulate prey species with consequences for conservation, focusing on predator‐prey and emerging indirect interactions in multi‐prey systems, and highlight future directions to address key knowledge gaps in our understanding of how feral horses may now be contributing to the (re)structuring of food webs. Observations of patterns of rapid growth and decline, and associated skews in sex ratios of feral horse populations, indicate a heightened potential for indirect interactions among large ungulate prey species, where there is a prevalence of feral horses as preferred prey, particularly where native prey are declining. In places like western North America, we expect predator‐prey interactions involving feral horses to become an increasingly important factor in the conservation of wildlife. This applies not only to economically or culturally important game species but also at‐risk species, both predators (e.g., wolves [Canis lupus], grizzly bears [Ursus arctos]) and prey (e.g., woodland caribou [Rangifer tarandus caribou]), necessitating an ecological understanding of the role of horses in natural environments that goes beyond that of population control. © 2021 The Wildlife Society.

A repeatable and quantitative DNA metabarcoding assay to characterize mixed strongyle infections in horses

Horses are ubiquitously infected by a diversity of gastro-intestinal parasitic helminths. Of particular importance are nematodes of the family Strongylidae, which can significantly impact horse health and performance. However, knowledge about equine strongyles remains limited due to our inability to identify most species non-invasively using traditional morphological techniques. We developed a new internal transcribed spacer 2 (ITS2) DNA metabarcoding ‘nemabiome’ assay to characterise mixed strongyle infections in horses and assessed its performance by applying it to pools of infective larvae from fecal samples from an experimental herd in Kentucky, USA and two feral horse populations from Sable Island and Alberta, Canada. In addition to reporting the detection of 33 different species with high confidence, we illustrate the assay’s repeatability by comparing results generated from aliquots from the same fecal samples and from individual horses sampled repeatedly over multiple days or months. We also validate the quantitative potential of the assay by demonstrating that the proportion of amplicon reads assigned to different species scales linearly with the number of larvae present. This new tool significantly improves equine strongyle diagnostics, presenting opportunities for research on species-specific anthelmintic resistance and the causes and consequences of variation in mixed infections.

Does size matter? Examining the possible mechanisms of multi-stallion groups in horse societies

In some feral horse populations, adult females are either associated with a single male or multiple males (stallions). However, little is known on why such groups with more than one male exist, considering that stallions fight to monopolize females. Body size is often an important determinant of male fighting ability and/or dominance rank and, consequently, reproductive success. Stallions may, therefore, vary in “quality” which could be a differentiating factor between single-stallion and multi-stallion males. We observed feral Garrano horses in Northern Portugal and examined the relationship between body size, sex, group type and number of females. Using a non-invasive laser distance meter, we determined individual body length and height by overlapping photos of the horses and a measuring tape. We tested the accuracy of remote measurements by comparison with manually measuring domestic horses and found no statistical differences between the two methods. Thus, we assume the values obtained were reliable and the methodology used was a useful tool for assessing morphological characteristics. Results showed no significant differences in body length and height for all parameters assessed, suggesting that size is not a determinant factor for feral horse society. Thus, other factors may be more important to explain the existence of single and multi-stallion groups.

Feral Horse Population Model and Body Condition: Useful Management Tools in Tornquist Park, Argentina?

ABSTRACTFeral invasive mammals challenge management strategies worldwide including the feral horse (Equus caballus) population in Tornquist Park, Argentina. In this study, I constructed population matrix models using demography data collected between 1995 and 2002. I validated the models with independent counts from 2008 to 2016. I individually identified feral horses and recorded them as I walked a fixed path that covered 20 km2. I performed an elasticity analysis to estimate the influence of the demographic parameters on population growth rate. In addition, I explored whether body condition score could be a proxy for carrying capacity. Model projections with the highest values of fecundity and survival exhibited by females of all ages during 1995–2002 fit the counts from 2008 to 2016, when growth rate was moderate (λ = 1.10), very well. Elasticity of adult female survival (0.54) was 4 times higher than for fecundity and survival of all other ages. Body condition of adult females at the end of winter, 2012–2013, after reductions in density, was improved compared with 2001–2002 when the population approached carrying capacity. The validated population model together with body condition score could allow managers to take more informed decisions when planning the control of feral horses in Tornquist Park. This approach could also be applied to other small, closed, feral horse populations and other large‐mammal populations. © 2020 The Wildlife Society.

Causes and consequences of an unusually male-biased adult sex ratio in an unmanaged feral horse population.

The adult sex ratio (ASR) is important within ecology due to its predicted effects on behaviour, demography and evolution, but research examining the causes and consequences of ASR bias have lagged behind the studies of sex ratios at earlier life stages. Although ungulate ASR is relatively well-studied, exceptions to the usual female-biased ASR challenge our understanding of the underlying drivers of biased ASR and provide an opportunity to better understand its consequences. Some feral ungulate populations, including multiple horse populations, exhibit unusually male-biased ASR. For example, research suggests that the feral horse Equus ferus caballus population on Sable Island, Nova Scotia, Canada may exhibit a male-biased ASR. Such exceptions to the rule provide a valuable opportunity to reveal the contributions of environmental context and trait differences to ASR bias. We aimed to test for bias in Sable Island horse ASR, identify the demographic drivers of bias, and explore its demographic and social consequences. To do this, we used life history, movement and group membership information for hundreds of horses followed through a long-term individual-based study between 2007 and 2018. Sable Island horse ASR is male biased and this skew has increased over time, reaching 62% male in 2018. Our life table response experiment suggested that ASR skew was driven predominantly by male-biased adult survival. Further analyses pointed to sex-biased survival being driven by reduced female survival post-reproduction. Male-biased ASR was associated with reduced harem sizes, an increase in the number of social groups on the island, and reduced reproduction in young females. Our results support the idea that male-biased ASR in feral ungulate populations may be caused by a combination of high population density and high reproductive output. We suggest that female-biased mortality may be caused by females continuing to reproduce at high density, and thus being more susceptible to resource shortages. Thus, our results highlight the strong context dependence of ASR. Furthermore, our work indicates the potential for ASR to substantially alter a population's social organisation. Such changes in social structure could have knock-on consequences for demography by altering the formation/stability of social relationships, or competition for matings.

Seasonality of feral horse grazing and invasion of Pinus halepensis in grasslands of the Austral Pampean Mountains (Argentina): management considerations

The introduction of domestic ungulates has altered the natural grazing regime, promoting the invasion of exotic woody plants being seasonality or stoking season one of the key factors regulating the grazing regime. The Pampean Mountain grasslands of Central Argentina have been intensively transformed by the presence of feral horse populations that have increased the last 40 years, coinciding with an increased abundance of Pinus halepensis dispersed from cultivated stands. In this study, we evaluated the effect of the seasonal variation in perennial grasses defoliation (no defoliation; continuous defoliation; defoliation during autumn-winter and defoliation during spring-summer) on the establishment of P. halepensis seedlings and the response of the dominant perennial grasses (Nasella tenuis and Piptochaetium lejopodum) to defoliation in areas with different grazing histories, following a split-plot design. The establishment of P. halepensis was highest in the grazed areas and in treatments where the defoliation of the perennial grasses was practiced throughout the year or during autumn-winter. Perennial grasses respond differently to defoliation according to the grazing history. In the grazed areas, defoliation negatively affected the perennial grasses during all the study period, showing the scarce resilience after grazing while in ungrazed areas, the perennial grasses were little affected by the seasonality of defoliation. Both, above- and belowground competition affected the establishment of P. halepensis, according to defoliation treatment and grazing history. The information obtained allows the elaboration of management recommendations to avoid the propagation of P. halepensis; therefore, the seasonal management of the grazing is a recommended practice toprevent or reduce the intensity of feral horse grazing during the autumn and winter months.

Rising up to the challenge of their rivals: Mare infidelity intensifies stallion response to playback of aggressive conspecific vocalizations

Management tools like immunocontraception can alter the behavior of target animals, but the extent to which they affect non-target individuals has received less attention. The feral horse (Equus caballus) population on Shackleford Banks, North Carolina is an ideal system with which these questions may be explored, as management of the population with the immunocontraceptive agent porcine zona pellucida (PZP) has resulted in an increased propensity for females to change social groups and thus, decreased social stability. During the study, on average, females made 1.4 group changes per day (range = 0–18.5 group changes per day): females previously treated with PZP made 1.8 group changes per day, while females that had never been treated made 1.2 group changes per day. Between May and August 2017, we used playbacks of aggressive male vocalizations (squeals) and human voices (reciting “hello horse”) to assess changes in stallion responses to male rivals versus socially irrelevant stimuli in the context of female turnover. Over the course of the study, males were observed for 9.4 h on average (range = 2.4–20.5 h). Males spent more time vigilant (estimate = 12.431, P = 0.016, x¯squeal = 30 s, x¯control = 19 s) and were more likely to approach the speaker following squeal playbacks than controls (estimate = 2.325, P = 0.039). Males’ latency to return to normal behaviors varied depending on whether the playback was conducted in the weeks before, during, or after group changes occurred (P = 0.025, x¯before = 26 s, x¯during = 39 s, x¯after = 53 s). Male responses were not affected by the total number of female group changes a male experienced during the observation period (P > 0.4), suggesting the effects are more context-dependent and not long-lasting. These findings suggest mare turnover can impact stallion responsiveness to potential rivals. As previously contracepted mares change groups more often than untreated mares and stallions exhibit prolonged responses to aggressive vocalizations after experiencing a female group change, contraception-induced changes to mare behavior may lead to increased male aggression in response to intruding rivals, which could be associated with greater energy expenditure. Finally, our work demonstrates that playback experiments are a useful tool for studying feral horse behavior in the wild. As the need for population control of different species continues to expand, rigorous investigations of immunocontraception’s effects on non-target animals are critical if agencies are to manage populations most effectively.

Not playing by the rules: Unusual patterns in the epidemiology of parasites in a natural population of feral horses (Equus caballus) on Sable Island, Canada.

Sable Island, Nova Scotia, Canada hosts one of few natural populations of feral horses (Equus caballus) never exposed to anthelmintics. Coproculture revealed cyathostomes, Strongylus equinus, S. edentatus, and S. vulgaris, with S. equinus (unusually) dominating in adult horses and cyathostomes dominating in young horses (<3 years of age). We examined 35 horses found dead in the springs of 2017 and 2018, as well as fecal samples from live horses in spring (n = 45) and summer 2018 (n = 236) using McMaster fecal flotation and Baermann larval sedimentation on fresh samples, and modified Wisconsin flotation and sucrose gradient immunofluorescent assay for Giardia and Cryptosporidium on frozen samples. Mean strongyle fecal egg counts were 666 eggs per gram (EPG) in dead horses, 689 EPG in live horses in spring, and 1105 EPG in summer; domestic horses are usually treated at counts exceeding 200 EPG. Adult horses (unusually) had patent infections with the lungworm Dictyocaulus arnfieldi and ascarids (Parascaris spp.), and in spring, dead horses had 5 times higher odds of having patent ascarid infections than live horses, likely due to malnutrition and corresponding immunodeficiency. Fecal prevalence and intensity of D. arnfieldi and Parascaris spp. were significantly higher in young horses, and in spring versus summer. A higher proportion of fecal samples were positive for strongyle and ascarid eggs using a centrifugal flotation technique on previously frozen feces, as compared to a passive flotation method on fresh feces. Eggs of the tapeworm Paranoplocephala mamillana were present in fecal samples from 28% of live, and 42% of dead, horses in spring. This research represents several new geographic records (S. edentatus, D. arnfieldi, and Eimeria leuckarti), provides insight into unusual patterns of parasite epidemiology in a nutrition-limited environment, and has conservation and biosecurity implications for this unique equine population, as well as for parasite management in domestic horses.

Mortality and Operational Attributes Relative to Feral Horse and Burro Capture Techniques Based on Publicly Available Data From 2010-2019

Management of excessive feral horse (Equus ferus caballus) and burro (Equus asinus) populations in the United States and globally has been a controversial subject for decades. I reviewed all available US federal feral horse and burro daily gather reports from 2010 to 2019 to extract equine species, technique (bait trapping or helicopter gathering), reason (emergency or other), number gathered, number of mortalities, and mortality attributes (acute or chronic/pre-existing condition, specific cause). I found 70 reports (bait trapping burros n = 10, bait trapping horses n = 24, helicopter gathering horses n = 21) from 9 states (AZ, CA, CO, ID, MT, NV, OR, UT, WY) representing 28,821 horses and 2,005 burros. For bait trapping, 100 animals died (4 burros, 96 horses) with 16 acute causes (1 burro, 15 horses) and 84 chronic/pre-existing causes (3 burros, 81 horses). For helicopter gathering, 268 horses died with 62 acute causes and 206 chronic/pre-existing causes. Mortality ratios did not differ by capture technique (P > .05) for broken necks, emaciation, acute causes, or chronic/pre-existing causes. The most common mortality-causing problems were structural deformations, club foot, blindness, and emaciation. The more horses gathered per day resulted in a greater proportion of chronic/pre-existing mortalities for both trapping techniques, but only an increase of acute mortalities for helicopter gathering. The slope suggests 1 acute mortality for every 300 horses gathered. The capture mortality rate across all gathers [1.1% (368 mortalities out of 30,826 horses and burros captured)] is below a general threshold of 2% suggested for wildlife studies.

Evolutionary quantitative genetics of juvenile body size in a population of feral horses reveals sexually antagonistic selection

Inter-individual variation in juvenile body size can have important consequences for individual fitness, population dynamics, and adaptive evolution. In wild vertebrate populations, larger juvenile size is usually expected to be selected for. However, understanding how such selection may translate into adaptive evolution requires an understanding of the genetic underpinnings of early development and the factors modulating selection. In this study, we characterised the genetic basis of and selection pressures acting upon juvenile body size in a large insular population of feral horses on Sable Island, Canada, to gain insights into the evolution of juvenile body size in wild vertebrate populations. We used pedigree-based quantitative genetic ‘animal models’ to quantify the sources of phenotypic variation in withers-knee length, and assessed the influence of maternal age, sex, and temporal (birth year) and spatial environmental heterogeneity in modulating overwinter survival selection. We found that withers-knee length is moderately heritable and that there was a significant positive genetic correlation between males and females. There was no indication of directional selection in a pooled-sex analysis, but we did find evidence for significant sexually antagonistic selection, with a tendency for smaller body size to be favoured in males and larger body size to be favoured in females. These results suggest that juvenile body size has the potential to evolve in this population, and that selection on juvenile size may play an important role in modulating sex-specific contributions to population dynamics. However, our results also suggest that there is unlikely to be evolutionary change in the mean body size of Sable Island foals.

Environmental and demographic drivers of male mating success vary across sequential reproductive episodes in a polygynous breeder.

Ecological and social factors underpinning the inequality of male mating success in animal societies can be related to sex ratio, sexual conflict between breeders, effects of nonbreeders, resource dispersion, climatic conditions, and the various sequential stages of mating competition that constitute the sexual selection process. Here, we conducted an individual‐based study to investigate how local resource availability and demography interact with annual climate conditions to determine the degree of male mating inequality, and thus opportunity for sexual selection across two sequential reproductive episodes (harem and subsequent mate acquisition) in a naturally regulated (feral) horse population in Sable Island National Park Preserve, Canada. Using a 5‐year, spatially explicit, mark‐resight dataset and hierarchical mixed‐effects linear modeling, we evaluated the influence of adult sex ratio (ASR) on mating success and then tested for effects of freshwater availability, density, unpaired male abundance, and precipitation during each breeding season. Unpaired male abundance, freshwater availability, and ASR differed in their effects on male mating success according to year and selection episode. Opportunity for sexual selection in males associated with harem acquisition increased with ASR, and unpaired male abundance further explained weather‐related interannual variation after accounting for ASR. In contrast, once a harem was secured, ASR had little effect on male mating inequality in regard to acquiring additional females, while interannual variation in mating inequality increased with decreasing freshwater availability. Our findings show that local demography, resource availability, and weather effect opportunity for sexual selection in males differently depending on selection episode, and can attenuate or accentuate effects of ASR.

Decreased female fidelity alters male behavior in a feral horse population managed with immunocontraception

In social species like the feral horse (Equus caballus), changes in individual behavior are likely to affect associated animals. On Shackleford Banks, North Carolina, USA, mares treated with the contraceptive agent porcine zona pellucida (PZP) demonstrate decreased fidelity to their band stallions. Here, we assess the effects of such decreased mare fidelity on male behavior and address potential interactions with habitat visibility, a component of the environment shown to significantly affect feral horse behavior. We compared the frequency and escalation of male-male contests, rates of aggressive and reproductive behaviors directed toward females, and the percentage of time spent vigilant among males experiencing varying levels of mare group changing behavior. We found that regardless of habitat visibility, males experiencing more female group changes engaged in contests at a higher rate (P = 0.003) and escalation (P = 0.029) and spent more time vigilant (P = 0.014) than males experiencing fewer group changes. However, while visibility had a positive effect on aggression directed by stallions toward mares (P = 0.013), female group changing behavior did not influence male-female aggressive or reproductive behaviors (P > 0.1), showing that decreases in mare fidelity altered male-male but not male-female interactions. These results have important implications for feral horse management; PZP-contracepted mares demonstrating prolonged decreases in stallion fidelity may have a disproportionate effect on male behavior. Moreover, our results shed light on the relative influences of female behavior and environmental factors like habitat visibility on male behavior. Such findings can ultimately improve our understanding of how the social and physical environments interact to shape male-male and male-female interactions.