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 km2 (SD) for Conger harems and 117.47 ± 32.75 km2 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.
Feral horses selected for areas of high herbaceous vegetation and avoided forest but moved to higher elevations in winter. Surface water influenced movement patterns for harem groups, whereas bachelor males moved faster, farther, and were less constrained by water availability than harems. Because of the importance of surface water, strategic placement of water sources where possible may help mitigate horse movement outside Herd Management Area boundaries.
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.
It is challenging to obtain precise and unbiased estimates of feral burro population sizes in the western United States. We compared abundance estimates from helicopter‐based double‐observer sightability surveys and infrared distance sampling surveys to a known minimum population size of burros in central Utah. Both survey types resulted in similar, negatively biased abundance estimates; however, we were able to remove this bias in the double‐observer surveys by including detections of radio‐collared individuals and by analyzing the survey data in a mark‐recapture distance sampling framework.
There are two species of free-roaming feral equids in North America: horses (Equus caballus) and donkeys or "burros" (E. asinus). Both species were introduced as domestic animals to North America in ...the early 1500s and currently inhabit rangelands across the western United States, Canada, and all continents except Antarctica. Despite their global distribution, little is known about their fine scale spatial ecology. Contemporary research tools to assess space use include global positioning system (GPS) tracking collars, but older models were problematic due to stiff collar belting causing poor fit. We tested modern designs of GPS collars on n = 105 horses and n = 60 burros for 4 years in five populations (3 horse, 2 burro) across the western United States, to assess whether collars posed welfare risks to horses or burros. We found no difference in survival of collared versus uncollared mares and jennies, and no difference in survival of their foals. In 4036 of 4307 observations for horses (93.7%) and 2115 of 2258 observations for burros (93.6%), collars were observed symmetrical, maintaining proper fit on the neck. Fur effects from collars (sweaty neck, indented fur, broken fur) were seen in 3% of horse observations and 25% of burro observations. Superficial effects (chafes and marks on skin surface) were seen in 2% of horse observations and 11% of burro observations; no severe effects from collars were seen. Body condition was not affected by collars; mean body condition of collared horses was 4.70 ± 0.54 (mean ± s.d) and 4.71 ± 0.65 for collared burros. Behavior results indicated minimal effects; collared horses stood slightly more than uncollared, and collared burros stood and foraged more in one population, but not in the other. For 6.3% of observations of horses and 6.4% of observations of burros, we found an effect of time wearing a collar on the cumulative sum of fur effects which increased over time (burros: rs = 0.87, P = <0.0001; horses: rs = 0.31, P = 0.002). Burros also showed an increase over time in the number of superficial effects, but horses did not. Collars occasionally moved into the wrong position, shifting forward over the ears; we observed this on 19 horses and 1 burro. Of those, most collars went over the ears in summer (n = 12). All collars were equipped with a remote release mechanism as well as a timed-release mechanism for redundancy, thus removed when observed in wrong position to avoid rubbing or discomfort. Our finding of no consequential physical effects in 98% of horse observations, and 89% of burro observations suggests the consequences of collars on free-roaming equid welfare and survival is biologically insignificant, although collars should be monitored regularly and continue to be equipped with a remote release mechanism to remove a collar if needed. With frequent welfare-driven, visual monitoring, collaring of free-roaming equids can be a safe and useful tool to increase our understanding of their spatial ecology, demography, habitat use, behavior, and interactions with other wildlife.
Plant communities in rangeland ecosystems vary widely in the degree to which they can compensate for losses to herbivores. Ecosystem‐level factors have been proposed to affect this compensatory ...capacity, including timing and intensity of grazing, and availability of soil moisture and nutrients. Arid ecosystems are particularly challenging to predict because of their high degree of temporal variability in moisture inputs. We used a replicated herbivore exclusion experiment to evaluate herbaceous plant responses to grazing by large ungulates to test current theory and identify constraints on plant compensation in a dryland ecosystem. We measured nitrogen (N) yield and herbaceous production in three plant communities: meadows, willow‐associated herbaceous communities, and riparian communities. We implemented grazing exclusion treatments from 2005 to 2008 in areas with elk and bison and areas with only elk. Grazing by large ungulates increased herbaceous production and N yield in herbaceous riparian communities. In willow communities, herbaceous plants displayed equal compensation in response to grazing in total aboveground production and N yield. Our results support the idea that plant compensation in this semiarid system is contingent on soil moisture availability, wherein the most productive sites (that received substantial moisture inputs from subsurface flow) exhibited overcompensation. Although the herbaceous riparian communities we studied are isolated patches of productive grassland in an otherwise shrub‐dominated and minimally productive semiarid landscape, grazing by a combination of bison and elk removed only 44%–53% of aboveground net primary productivity (ANPP) during the growing season, and 25%–38% of production over winter. Consumption by ungulates was a positive linear function of herbaceous production, similar to reported patterns from other temperate and tropical grazing ecosystems. The slope of this relationship was affected by the analytical method used to calculate ANPP and consumption rates, but, regardless of the method, was lower or similar to reported slopes for other intensively grazed systems (Yellowstone, Serengeti, Laikipia) that have sustained high ungulate densities for decades to centuries. Given that the vegetation communities exhibited equal or overcompensation in terms of total herbaceous ANPP in both years, elk and bison population levels during our study period did not appear to occur at densities leading to degradation of herbaceous communities.
Feral horses (Equus ferus caballus) have become abundant on public lands in the American West, particularly over the past 10 yr. In areas where they are overabundant, there is risk of habitat ...degradation. Most previous studies on diet and habitat use of feral horses were conducted more than 20 yr ago; rangelands have changed considerably in that time, so it is useful to revisit horse diets. We conducted a study to examine the diet of feral horses using noninvasive methods and subjectively compare diet analysis techniques. We collected feral horse fecal samples from a sagebrush/pinyon-juniper ecosystem in Colorado in May, August, and October 2014. We analyzed 30 fecal samples from each collection session by both microhistology and plant DNA barcoding. Both microhistology and plant DNA barcoding results indicated horse diet consisted primarily of graminoids (78.5% and 68.8%, respectively, both of which are in greater proportion than availability based on ecological site descriptions); however, the two methods differed in species composition of grasses. Similar to other studies, microhistological analyses underestimated the proportion of forbs in the diet compared with plant DNA barcoding analyses, which showed a surprisingly high contribution of forbs to the diet compared with previous studies. Our results suggest plant DNA barcoding analyses have great potential, although both methods have inherent biases.
Aerial thermal infrared (TIR) surveys are an attractive option for estimating abundances of large mammals inhabiting extensive and heterogeneous terrain. Compared to standard helicopter or fixed-wing ...aerial surveys, TIR flights can be conducted at higher altitudes translating into greater spatial coverage and increased observer safety; however, monetary costs are much greater. Further, there is no consensus on whether TIR surveys offer improved detection. Consequently, we performed a study to compare results of a TIR and helicopter survey of bison (Bison bison) on the Powell Plateau in Grand Canyon National Park, USA. We also compared results of both surveys to estimates obtained using a larger dataset of bison helicopter detections along the entire North Rim of the Grand Canyon. Observers in the TIR survey counted fewer individual bison than helicopter observers (101 to 127) and the TIR survey cost was 367% higher. Additionally, the TIR estimate was 18.8% lower than the estimate obtained using a larger dataset, while the comparative helicopter survey was 9.3% lower. Despite our small sample size, we found that helicopter surveys are currently the best method for estimating bison abundances in dense canopy cover sites due to ostensibly more accurate estimates and lower cost compared to TIR surveys. Additional research will be needed to evaluate the efficacy of these methods, as well as very high resolution satellite imagery, for bison populations in more open landscapes.
Abstract
Feral horse (
Equus caballus
) population management is a challenging problem around the world because populations often exhibit density‐independent growth, can exert negative ecological ...effects on ecosystems, and require great cost to be managed. However, strong value‐based connections between people and horses cause contention around management decisions. To help make informed decisions, natural resource managers might benefit from more detailed understanding of how horse management alternatives, including combinations of removals and fertility control methods, could achieve objectives of sustainable, multiple‐use ecosystems while minimizing overall horse handling and fiscal costs. Here, we describe a modeling tool that simulates horse management alternatives and estimates trade‐offs in predicted metrics related to population size, animal handling, and direct costs of management. The model considers six management actions for populations (removals for adoption or long‐term holding; fertility control treatment with three vaccines, intrauterine devices, and mare sterilization), used alone or in combination. We simulated 19 alternative management scenarios at 2‐, 3‐, and 4‐year management return intervals and identified efficiency frontiers among alternatives for trade‐offs between predicted population size and six management metrics. Our analysis identified multiple alternatives that could maintain populations within target population size ranges, but some alternatives (e.g., removal and mare sterilization, removal and GonaCon treatment) performed better at minimizing overall animal handling requirements and management costs. Cost savings increased under alternatives with more effective, longer lasting fertility control techniques over longer management intervals compared with alternatives with less‐effective, shorter lasting fertility control techniques. We built a user‐friendly website application,
PopEquus
, that decision makers and interested individuals can use to simulate management alternatives and evaluate trade‐offs among management and cost metrics. Our results and website application provide quantitative trade‐off tools for horse population management decisions and can help support value‐based management decisions for wild or feral horse populations and ecosystems at local and regional scales around the world.
Non‐native species can affect ecosystems by influencing native predator‐prey dynamics. Therefore, management interventions designed to remove non‐natives may inadvertently lead to increased predation ...on native species. Feral horses are widely distributed throughout the arid parts of western North America. A growing body of research indicates that horses can be an important prey species to mountain lions in ecosystems where they overlap. In December 2020, the Bureau of Land Management removed 455 horses from the Delamar Mountains, Nevada, USA. We leveraged this management intervention to implement a before–after–control–impact study to test hypotheses about predation on horses and native ungulates. We predicted (1) that horses would comprise an important part of the diet in this mixed‐prey community, (2) following removal, the proportion of horses in the diet would decrease and native ungulates would increase, and (3) mountain lion home ranges overlapping the treatment areas would increase in response to decreased prey availability. From 2018 to 2022, we investigated 1360 clusters from 29 GPS‐collared lions and identified 1056 prey items. To model the probability of a predation event (a kill), we fit a mixed‐effects logistic regression model for ungulate prey as a function of lion sex, treatment area (in/out), and treatment period (pre‐/post‐removal). We used a log‐linear regression model to evaluate changes in home range size. The most common prey were mule deer (55%), feral horses (32%), and coyotes (4%). Twenty‐two of 29 lions consumed horses, although the rate of horse consumption was highly variable across individuals. Horses of both sexes and all age classes were predated. In contrast to predictions, our models detected no effect of removals on diet composition (βinteraction = 0.30 ± 1.1), nor did the removal influence home range size (βinteraction = 0.02 ± 0.02). Despite a 46% reduction in horse abundance, we found no evidence for prey‐switching following the horse removal treatment. Removal magnitude, rapid horse immigration, and/or behavioral specialization of individual mountain lions may help explain these results. Our findings have important implications for mountain lion and feral horse management in arid environments characterized by high prey diversity, but low prey abundance.
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