Ungulate Use Research Articles

A 2000-year record of fecal biomarkers reveals past herbivore presence and impacts in a catchment in northern Yellowstone National Park, USA.

Molecular biomarkers preserved in lake sediments are increasingly used to develop records of past organism occurrence. When linked with traditional paleoecological methods, analysis of molecular biomarkers can yield new insights into the roles of herbivores and other animals in long-term ecosystem dynamics. We sought to determine whether fecal steroids in lake sediments could be used to reconstruct past ungulate use and dominant taxa in a small catchment in northern Yellowstone National Park. To do so, we characterized the fecal steroid profiles of a selection of North American ungulates historically present in the Yellowstone region (bison, elk, moose, mule deer, and pronghorn) and compared them with those of sediments from a small lake in the Yellowstone Northern Range. Analysis of a set of fecal steroids from herbivore dung (Δ5-sterols, 5α-stanols, 5β-stanols, epi5β-stanols, stanones, and bile acids) differentiated moose, pronghorn, and mule deer, whereas bison and elk were partially differentiated. Our results show that bison and/or elk were the primary ungulates in the watershed over the past c. 2300 years. Fecal steroid influxes reached historically unprecedented levels during the early and middle 20th century, possibly indicating high local use by ungulates. Comparison of fecal steroid influxes with pollen and diatom data suggests that elevated ungulate presence may have contributed to decreased forage taxa (Poaceae, Artemisia, and Salix), relative to long-term averages, and possibly increased lake production. Our results reflect past change within a single watershed, and extending this approach to a network of sites could provide much-needed information on past herbivore communities, use, and environmental influences in Yellowstone National Park and elsewhere.

Ecological dynamics and coexistence patterns of wild and domestic mammals in an abandoned landscape

The issue of agricultural land abandonment in southern Europe has raised concerns about its impact on biodiversity. While abandoned areas can lead to positive developments like creating new habitats and restoring native vegetation, they can also result in human–wildlife conflicts, particularly in areas with extensive farming and free‐ranging livestock. To understand habitat selection and use of livestock and wild ungulates, it is essential to study their spatial and temporal distribution patterns. In this context, we conducted a long‐term large mammal monitoring project using camera traps in the Peneda‐Gerês National Park in northern Portugal. Our primary focus was on exploring habitat preferences, occupancy dynamics, and potential spatial use correlations between domestic and wild species, utilizing dynamic occupancy models. Most wild species exhibited stable area use patterns, while domestic species experienced marginal declines, and the Iberian ibex displayed signs of repopulation. We observed distinct effects of habitat variables on occupancy, colonization, and extinction, revealing species‐specific patterns of habitat utilization. Human disturbance had a notable impact on domestic species but did not affect wild ones. Camera sensitivity emerged as a critical factor, enhancing detection probability for all species. Additionally, habitat and weather variables exerted varying effects on detection probabilities, underscoring the necessity of accounting for these factors in modeling the detection process. We found shared habitat preferences between cattle and horses, both positively correlated with wolves, suggesting potential human–wildlife conflicts. Despite extensive spatial overlap, domestic and wild species seem to exhibit ecological independence possibly due to distinct strategies and low predation pressure. Overall, the study emphasizes the multifaceted factors influencing habitat use. The observed species associations contribute to understanding ecological relationships and potential resource competition, emphasizing the importance of considering environmental variables for effective wildlife conservation and management.

Systematic reduction of natural enemies and competition across variable precipitation approximates buffelgrass invasiveness (Cenchrus ciliaris) in its native range.

Invasive grasses cause devastating losses to biodiversity and ecosystem function directly and indirectly by altering ecosystem processes. Escape from natural enemies, plant-plant competition, and variable resource availability provide frameworks for understanding invasion. However, we lack a clear understanding of how natural stressors interact in their native range to regulate invasiveness. In this study, we reduced diverse guilds of natural enemies and plant competitors of the highly invasive buffelgrass across a precipitation gradient throughout major climatic shifts in Laikipia, Kenya. To do this, we used a long-term ungulate exclosure experiment design across a precipitation gradient with nested treatments that (1) reduced plant competition through clipping, (2) reduced insects through systemic insecticide, and (3) reduced fungal associates through fungicide application. Additionally, we measured the interaction of ungulates on two stem-boring insect species feeding on buffelgrass. Finally, we measured a multiyear smut fungus outbreak. Our findings suggest that buffelgrass exhibits invasive qualities when released from a diverse group of natural stressors in its native range. We show natural enemies interact with precipitation to alter buffelgrass productivity patterns. In addition, interspecific plant competition decreased the basal area of buffelgrass, suggesting that biotic resistance mediates buffelgrass dominance in the home range. Surprisingly, systemic insecticides and fungicides did not impact buffelgrass production or reproduction, perhaps because other guilds filled the niche space in these highly diverse systems. For example, in the absence of ungulates, we showed an increase in host-specific stem-galling insects, where these insects compensated for reduced ungulate use. Finally, we documented a smut outbreak in 2020 and 2021, corresponding to highly variable precipitation patterns caused by a shifting Indian Ocean Dipole. In conclusion, we observed how reducing natural enemies and competitors and certain interactions increased properties related to buffelgrass invasiveness.

Ungulate occurrence in forest harvest blocks is influenced by forage availability, surrounding habitat and silviculture practices

Abstract Forest harvesting causes habitat loss and alteration and can change predator–prey dynamics. In Canada, forest harvesting has shifted the distribution and abundance of ungulates (deer, elk and moose) that prefer early seral forest, resulting in unsustainable caribou predation by shared predators (bears, cougars and wolves). Long‐term solutions for caribou recovery require management to reduce ungulate prey species within caribou ranges. Silviculture practices applied after forest harvesting directly affect the amount of forage available in harvested areas, and therefore influence ungulate distribution, but few studies have completed detailed assessments on how specific treatments of site preparation, planting and stand tending influence ungulate use of harvest blocks. We used camera traps, silviculture data, GIS‐derived habitat and disturbance data, and detailed vegetation data collected at field sites to investigate ungulate occurrence in harvest blocks in west‐central Alberta, Canada. We compared seasonal ungulate occurrence and investigated how site‐specific characteristics, the surrounding habitat and disturbance density, and fine‐scale silviculture treatments influenced ungulate occurrence in blocks. Deer, elk and moose occurrence was higher in summer compared to winter. Elk, moose and white‐tailed deer occurrence was higher in blocks with greater availability of specific forage species. Moose occurrence was higher in blocks with a lower road density in the surrounding area, and white‐tailed deer occurrence was higher in blocks further from seismic lines and with a lower proportion of harvest blocks in the surrounding area. Deer, elk and moose occurrence was higher in younger harvest blocks. Mule deer and white‐tailed deer occurrence was lower in blocks with higher planting densities of lodgepole pine, and mule deer occurrence was also lower in blocks that had been stand tended. Our study provides detailed information on ungulate response to fine‐scale silviculture methods used in Alberta, directly linking wildlife occurrence to forestry practices, and providing practical scientific information to inform sustainable forestry. Translating this research into practical landscape management decisions could benefit boreal biodiversity, including threatened species like caribou, and culturally and economically important species like deer, elk and moose.

Patterns and predictors of ungulate space use across an isolated Miombo woodland reserve

AbstractUnderstanding ecological‐ and management‐related predictors of mammal space use within protected areas is critical for management planning. This is particularly true in small, fenced and isolated reserves where we hypothesised that management‐related activities will influence large herbivore space use more than ecological characteristics. We used camera trap data to assess the space use patterns for 18 ungulate species in the small and isolated Majete Wildlife Reserve (Majete) in the understudied Miombo woodland ecoregion of southern Malawi. In the 2018 dry season, the 691 km2 reserve was systematically surveyed for ungulate presence at 140 camera trap locations. Over a period of 5456 camera days, the survey yielded 11 078 independent detections of 18 ungulate species and three predators. Using a single‐species occupancy modelling framework, the probability of space use of ungulates was assessed in relation to five management (fire exposure, fire frequency, water availability, distance from fence and road) and five ecological (visibility, grass biomass, vegetation type, terrain ruggedness and predator abundance) space use covariates, while accounting for imperfect detection. Top‐ranked models contained multiple covariates for 15 of the 16 species modelled, with only nyala's (Tragelaphus angasii) space use best predicted by vegetation type only. Distance to water, vegetation type, visibility and fire frequency were predictors having strong influences on six or more species each. More management‐related covariates reflected in top models, but ecological covariates had more meaningful effect sizes making us reject the hypothesis. Importantly though, distance from the roads and fence were also identified as prominent predictors. Notably, black rhinoceros' (Diceros bicornis) probability of space use increased with distance from the reserve boundary. Beyond informing habitat management for ungulates in Majete, these results can form the basis for understanding species‐specific space use patterns in other small reserves with similar characteristics and threats.

Wildfire and prescribed burning impact moose forage availability and browsing levels in the northern boreal forest

ABSTRACT Fire regimes and herbivore densities have in parts of the boreal been heavily impacted by human activities with consequences for vegetation, forage supply and ungulate use. In this study, we evaluate the effects of natural wildfires and prescribed burning on moose forage availability, use, and browsing in northern Sweden. We studied three wildfire locations, adjacent controls, and five prescribed burns 12 years after fire. To compare forage availability and browsing levels we combined related variables with Principal Component Analyses and used the first axis in a regression analysis. The wildfire locations had a different composition of forage compared with the controls with less forage in the field layer and more woody browse and deciduous saplings. Forage availability was best explained by the occurrence of fire and differed between wildfire and the unburned control areas. Browsing levels were higher in wildfire locations compared with unburned controls. Prescribed burns varied substantially and could resemble both unburnt control and wildfire in terms of forage availability and browsing. We conclude that prescribed burning as a restoration action potentially can improve forage availability, particularly of deciduous species, if they mimic the impact of wildfires in terms of larger area burned and high fire severity.

Water scarcity exacerbates feral ungulate use of ephemeral savanna waterholes in northern Australia

Context Australian savannas evolved in the absence of hooved mammals and are therefore prone to disturbance from exotic ungulates. Several ungulate species have established feral populations in Australia’s northern savannas. Because most ungulate species have high water requirements, seasonal and interannual changes in water availability are likely to affect their behaviour and the extent of their impacts on native ecosystems. However, few studies have investigated how water scarcity affects feral ungulate use of waterpoints in Australia. Aims The aim of this study was to determine whether seasonal and interannual water scarcity affected the visitation behaviours of feral pigs, cattle and water buffalo at ephemeral savanna waterholes within Limmen National Park, Northern Territory, Australia. Methods We used motion-triggered wildlife cameras to study feral ungulate visitation and behaviour at 20 waterholes. Generalised linear mixed-effects models were used to investigate whether the number of visits, duration of visits and number of individuals visiting waterholes varied with year and dry season progression. We also investigated whether these factors affected the amount of time ungulates spent foraging, drinking and wallowing at waterholes. Key results All three species visited waterholes more often, for longer periods and in larger numbers during a drought year compared with an average rainfall year. Cattle and buffalo spent more time drinking from waterholes during the drought, and pigs and cattle spent longer periods foraging. Buffalo also wallowed more during the drought. Responses to dry season progression varied among species. Cattle visited waterholes more frequently, for longer durations and in larger herd sizes as the dry season progressed, whereas buffalo use did not change. Pigs only increased their visitation to waterholes when water scarcity was most extreme, at the end of the dry season during the drought. Conclusions Our results demonstrate that water scarcity exacerbates feral ungulate use of savanna waterholes. Management and control programs for feral ungulates in Australian savannas may benefit from targeting different species under specific water scarcity conditions. Implications Because climate change is predicted to reduce surface water availability in northern Australia, feral ungulate use of savanna waterholes may intensify, risking further biodiversity losses and irreversible ecosystem damage.

Ungulate use of an at-grade fauna passage and roadside animal detection system: A pilot study from Southern Sweden

Wildlife-vehicle collisions (WVC) cost millions of euros each year in loss of life and damages. Wildlife crossing structures and fencing are commonly implemented mitigation strategies to reduce WVC and increase landscape connectivity for wildlife. Typically, crossing structures are over- or under-pass structures that allow animals to safely cross the road, while separating them from the road and traffic. An alternative strategy could be an at-grade fauna passage coupled with a Roadside Animal Detection System (RADS). At-grade fauna passages are designated locations where a gap in fencing allows animals to cross over the road, while RADS alert drivers of animals at the upcoming passage, so that they can adjust their driving behaviour accordingly and avoid collisions. In this pilot study, we investigated the use of one at-grade fauna passage by roe deer (Capreolus capreolus), red deer (Cervus elaphus) and wild boar (Sus scrofa) in southern Sweden, and compared changes in the number of WVCs before, during and after the construction of the passage. We collected a total of 326 wildlife crossings from 722 individuals over 1 year of monitoring (24 January 2020–24 January 2021). We found that crossing events tended to be dominated by the time animals spent in the roadside verge from which they approached the at-grade fauna passage, particularly for roe deer that spent a lot of time browsing in the roadside verge during dusk. We also found that animals spent longer in the passage if vehicles were present. In our 1 year of surveys, we only recorded three accidents, and when comparing the annual collision statistics before, during, and after construction of the at-grade fauna passage, we demonstrated an overall reduction in collisions by 66%. While our pilot only evaluates a single site, it does provide promising preliminary results that suggest that at-grade fauna passages can help in efforts to reduce collisions, while maintaining connectivity over medium-sized roads for large ungulates.

Use of Wild Ungulates as Sentinels of TBEV Circulation in a Naïve Area of the Northwestern Alps, Italy.

Wild and domestic animals can be usefully employed as sentinels for the surveillance of diseases with an impact on public health. In the case of tick-borne encephalitis virus (TBEV), the detection of antibodies in animals can be more effective than screening ticks for detecting TBEV foci, due to the patchy distribution of the virus. In the Piedmont region, northwestern Italy, TBEV is considered absent, but an increase in tick densities, of Ixodes ricinus in particular, has been observed, and TBEV is spreading in bordering countries, e.g., Switzerland. Therefore, we collected sera from wild ungulates during the hunting season (October-December) from 2017 to 2019 in the Susa Valley, Italian western Alps, and screened them for TBEV antibodies by a commercial competitive ELISA test. We collected 267 serum samples by endocranial venous sinuses puncture from red deer, roe deer and northern chamois carcasses. The animals were hunted in 13 different municipalities, at altitudes ranging between 750 and 2800 m a.s.l. The serological survey for TBEV yielded negative results. Borderline results for five serum samples were further confirmed as negative for TBEV by a plaque reduction neutralisation test. To date, our results indicate that TBEV is not circulating in western Piedmont. However, monitoring of TBEV should continue since TBEV and its vector are spreading in Europe. The wide-range distribution of wild ungulates and their role as feeding hosts, make them useful indicators of the health threats posed by Ixodid ticks.

Features of the use of ungulates by oxpeckers (Buphagus) in central Kenya

A detailed multi-year observational study of Red-Billed (RB) oxpecker (Buphagus erythrorhynchus) and yellow-billed (YB) oxpecker (B. Africanus) occurrence on a range of ungulate species was performed in two Kenyan national parks. Surprisingly observation of over 2100 individual Kongoni, in group sizes of 1 to > 50, revealed a total absence of oxpeckers, of either species, on Kongoni (Alcephalus buselaphus: subfamily Alcephalinae) in Nairobi National Park (NNP), although it was one of the most common bovid species present. There is no population of A. buselaphus at the second study site in central Kenya, Lake Nakuru National Park (LNNP). By contrast, the Impala (Aepyceros melampus), a smaller species, whose taxonomic status is phylogenetically ambiguous, but is widely considered to have evolved relatively recently in Africa, was a frequent oxpecker host in both NNP (29% of animals observed) and LNNP (12.9% of individual animals observed. Two species of gazelle Thompson’s gazelle (Gazella rufifrons)) and Grant’s gazelle (G. granti), subfamily Gazellinae, which were present in both parks but in relatively low abundance, were also not observed to host oxpeckers. The reason for these bovid subfamilies, apparently not being utilized as oxpecker hosts in central Kenya, is unclear and requires further research.

Five-Year Effects of Introduced Mountain Goats and Recreation on Plant Communities and Species of Conservation Concern in an Alpine Sky Island

Alpine ecosystems in the arid southwestern United States are vulnerable to climate change. Many of these ecosystems are experiencing increased recreational use and introductions of nonindigenous ungulates. In the La Sal Mountains of southeastern Utah, alpine plant communities support an endemic plant species, Erigeron mancus, 11 plant and two animal species of conservation concern, and a United States Department of Agriculture (USDA) Forest Service Research Natural Area. Nonindigenous mountain goats (Oreamnos americanus) were introduced by the Utah Division of Wildlife Resources (UDWR) in 2013 and 2014, and recreational use has increased in the intervening period. To evaluate potential effects of mountain goat and recreational use, vegetation monitoring was initiated by the USDA Forest Service in 2016. We used 5 y monitoring data (2016–2020) and generalized linear mixed models to analyze the separate and interacting effects of year, recreational use, and mountain goat use on: (1) populations of plant species of conservation concern; (2) vascular and nonvascular plant cover; and (3) ground cover. Our analyses revealed decreases in proportional frequency of E. mancus at the end of the monitoring. Proportional cover of dominant plant growth forms, especially forbs, declined, while proportional cover of litter and ineffective ground cover (bare soil and pavement) increased. Recreational or goat use were a factor in several of the observed changes, although weather and climate likely also influenced the results. Management complexity arises because of the different missions of USDA Forest Service and UDWR, but adjustments in mountain goat numbers and recreational use may be needed to maintain ecologically resilient ecosystems. Continued monitoring can provide the basis for adaptive management.

Habitat use and activity patterns of ungulates in a tropical rainforest of southern México

Baird’s tapir (Tapirella bairdii), white-lipped peccary (Tayassu pecari), and collared peccary (Dicotyles tajacu) sympatrically occur in the Lacandon Forest of Chiapas, México. These species contribute to maintain ecosystem dynamics through herbivory, seed dispersal, and seed predation, constituting important prey for large carnivores and hunters. We analyzed activity patterns and habitat use of the three focal species to assess the degree of temporal and spatial habitat segregation among them in Montes Azules Biosphere Reserve (REBIMA) and surrounding communities. Between February and October 2015 we deployed camera-traps during 8,463 camera-trap days to estimate the presence and activity of tapirs and peccaries in two habitat types: “conserved” (REBIMA), and “transformed” (community forests; AFC). Habitat use and activity patterns of tapirs and peccaries were assessed through logistic regression models. We found that Baird’s tapir was almost exclusively nocturnal with a trend towards crepuscular activity, while both peccary species were diurnal, therefore showing a high daily temporal segregation from the tapir. Both peccary species were similarly active in the two study sites, while tapirs were more active in continuous forest within the protected area. The occurrence of tapirs and white-lipped peccaries depended on the presence and proximity of water sources and roads. Our results suggest that spatial segregation of the habitat allows coexistence of both peccary species in the study area. AFC have potential for maintaining populations of tapirs and peccaries in the Lacandon Forest. Conserving these forests by local communities is essential to ensure the persistence of these mammals.

Persistence of a locally endangered mouse-deer amidst the re-emergence of two larger ungulates in small urban rainforest fragments

Urbanisation has threatened native forests globally with remaining forests often existing as small isolated patches in a harsh landscape matrix. Devoid of large predators, forest patches may continue to support ungulate populations, but their conservation is challenging as unchecked population growth can bring about ecosystem imbalances and threaten other native species. Three ungulate species exist in small rainforest fragments of urbanised Singapore: the locally endangered lesser mouse-deer (Tragulus kanchil), and the re-emergent wild boar (Sus scrofa) and sambar deer (Rusa unicolor). Using an array of camera traps, we estimated the densities of the lesser mouse-deer, wild boar and sambar deer in rainforest patches, and determined the influence of habitat and landscape factors on their occupancies. We also studied the interactions of the wild boar and sambar deer with the lesser mouse-deer spatially and temporally. Estimates of the lesser mouse-deer density showed that the population has recovered since the 1990s. Its density was comparable to the wild boar, but higher than sambar deer. Detections of lesser mouse-deer were significantly higher in primary forests compared to old secondary forests, and increasing distance away from the nearest road or public trail was a strong predictor of its occupancy. There was no evidence that the wild boar and sambar deer competitively excluded the lesser mouse-deer, rather, co-occupancy was observed. We demonstrated that temporal niche partitioning between the lesser mouse-deer and the two other ungulates likely explained their co-existence. Notwithstanding this, there are indications of understorey alteration due to the wild boar occurrences. Thus, continued monitoring of ungulate populations and conservation efforts focused on maintaining the health of the primary and secondary rainforest patches, and reducing disturbance, are critical to sustain future populations of lesser mouse-deer. Our study has shed new light on the habitat use and species interactions of tropical ungulates in urban predator-free ecosystems and highlight the importance in gaining a deeper understanding of their dynamics to more effectively guide conservation and management decisions.

Annual movement strategy predicts within-season space use by moose

The heterogeneity of resource availability shapes animal movements at different spatio-temporal scales. Given that movements at various scales are assumed to be linked, the space use of temperate ungulates within seasonal ranges (winter, summer) should be related to their movement patterns at the annual scale. In this study, we aimed to evaluate the level of stationarity of moose (Alces alces) within their seasonal ranges and to link annual movement patterns to within-season space use. We analysed the ranging behaviour of 32 moose fitted with GPS collars from two study areas in Eastern Poland, where at the annual scale a fraction of individuals migrate between summer and winter ranges (partial migration). Our results revealed that moose stationarity within seasonal home ranges expressed remarkable variation. The probability of moose stationarity within seasonal ranges was significantly higher (by 23%), and the mean home range size tended to be lower (9.7 km2) among individuals that seasonally migrated than among non-migratory moose (14.3 km2). In addition, we found that (i) in summer, moose were significantly more stationary (by 19%) and exhibited a smaller mean home range size than in winter (9.0 and 15.9 km2, respectively) and (ii) the mean seasonal home range size of males (19.6 km2) was remarkably greater than that of females (9.6 km2). Given the significant link between annual and seasonal scales of animal movements, any environmental change (e.g. climate warming) affecting an animal’s annual movement strategy could alter within-season animal space use and presumably individual fitness.Significance statementTo maximize their fitness, animals adjust their movements to deal with variations in resource distribution in the landscape. The scale of spatio-temporal variation causes different types of migratory behaviours, ranging from year-round stationarity to migration, when individuals establish spatially separated seasonal ranges. Studies on ungulates suggest that the stability and the size of seasonal home ranges can be linked to annual movement behaviour. Using the locations of GPS-tracked moose, we demonstrate in this study that migratory individuals were more prone to establishing stable seasonal home ranges (especially in summer) than moose that occupied the same area throughout the year. Moreover, stable seasonal home ranges were remarkably smaller in summer than in winter, which may suggest a season-specific spatial distribution and a renewability of moose forage. Our results show a clear link between different temporal scales of animal movements.

Lentic Meadows and Riparian Functions Impaired After Horse and Cattle Grazing

ABSTRACTPerennially and intermittently available water in lentic, spring‐fed herbaceous meadows supports wetland plants that build and maintain absorbent soil organic matter, capture sediment, prevent erosion, and thereby sustain riparian ecosystems. Enhanced forage and available water attract a diversity of wildlife and grazing animals that can degrade riparian vegetation through prolonged overuse. We tested novel monitoring methods and adapted existing methods to obtain quantified vegetation data that have been consistently linked to riparian functionality and compared these data to ungulate species use. We quantified ungulate use with time‐lapse cameras at 10 Nevada spring‐fed herbaceous meadows where core greater sage‐grouse (Centrocercus urophasianus) habitat, public land grazing allotments, and free‐roaming horse (Equus caballus) herd management areas (Bureau of Land Management) or territories (U.S. Department of Agriculture Forest Service) overlapped. Free‐roaming horses used most study sites over longer periods and grazed more intensely than cattle or domestic sheep. Native ungulate grazing intensity was comparatively negligible. Free‐roaming horses selected spring‐fed meadow sites on average 51 (±33 [SE]) times more than the surrounding rangelands, and cattle selected them 30 ± 23 times more. Within the wettest area of a site, study sites with longer duration of use and higher intensity of use had more soil alteration and bare ground. Riparian ecosystem degradation occurred where meadows had limited rest or recovery time from grazing during the growing season. At all sites we noted loss of riparian function, especially within the hydric areas with the highest potential to grow abundant cover of stabilizing wetland plant species. Transects for monitoring soil alteration and vegetation cover in this hydric location can provide a leading indicator of riparian recovery or degradation and help in adjusting free‐roaming horse population levels or livestock management strategies. © 2021 The Wildlife Society.

Fear of large carnivores is tied to ungulate habitat use: evidence from a bifactorial experiment

The fear large carnivores inspire in large ungulates has been argued to have cascading effects down food webs. However, a direct link between ungulate habitat use and their fear of large carnivores has not been experimentally tested. To fill this critical gap, we conducted a bi-factorial experiment in an African savanna. We removed shrub cover and broadcast large carnivore vocalizations (leopard, hyena, dog) or non-threatening control vocalizations in both experimentally cleared and shrubby control sites. We recorded the proactive (frequency of visitation) and reactive (fleeing or vigilance) responses of multiple prey (impala, warthog, nyala and bushbuck). Critically, we found a significant proactive–reactive interaction. Ungulates were 47% more likely to run after hearing a predator vocalization in shrubby control sites than experimental clearings, demonstrating that ungulates perceived less fear from large carnivores in open habitat (clearings). Consistent with this finding, ungulates visited clearings 2.4 times more often than shrubby control sites and visited shrubby control sites less often at night, when large carnivores are most active. Combined with results from previous experiments demonstrating that the disproportionate use of available habitats by large ungulates can alter ecosystem properties, our experiment provides critical evidence that the fear large carnivores inspire in large ungulates can cause trophic cascades.

Disentangling landscape features and vegetation composition effects on wild ungulate use of hay meadows in high Pyrenees

Wild ungulates seek out food resources at the bottom of mountain valleys in winter and spring, where their high density can negatively impact the production of hay meadows. This study determines the factors that explain the meadow use of four sympatric species: red-, roe-, and fallow-deer, and wild boar. In 2017, we counted the individuals of each species in 94 meadows for eight nights, between March and June, and estimated the frequency of meadow use as the proportion of positive censuses over the total censuses per meadow. We also recorded the presence of domestic ungulates. We inventoried the floristic composition in June. Species tended to use different meadows but only fallow- and roe-deer significantly segregated, indicated by the negative C-score index. As the ungulate frequency was skewed to 0, we used a generalized additive model (GAMLSS) to simultaneously simulate the probability of animals being absent and the frequency of positive censuses. Landscape features contributed to explaining the absence probability, while plant richness and legume cover explained the frequency of meadow use. The presence of domestic ungulates did not affect wild ungulate behavior. All studied species, except roe-deer, avoided exposed spots, preferentially using large meadows close to the forest. At meadow level, each species seemed to have different forage requirements. The frequency of fallow deer increased with the meadow forage quality, indicating a more grasses-based diet than the other two deer, and probably displacing roe-deer to suboptimum meadows. Red-deer used less plant-rich meadows, suggesting flexibility in their dietary choices.

Comparison of Effects of Shed Antler Hunting and Helicopter Surveys on Ungulate Movements and Space Use

ABSTRACTShed antler hunting (i.e., collecting cast cervid antlers) has increased in popularity during the past decade, but little is known about how this recreational activity affects ungulate movements and space use. We placed geographic positioning system (GPS)‐collars on 133 female and male bighorn sheep (Ovis canadensis), bison (Bison bison), and mule deer (Odocoileus hemionus) to quantify their movements and space use during shed antler hunts compared with those behaviors during helicopter surveys in Utah, USA, from 2012 to 2015. For each species, we calculated means and 95% confidence intervals for distance moved during 90‐minute segments (16 points/day) pre‐event (control, 7 consecutive days prior to event), event (1–2 days), and post‐event (7 consecutive days after event) for shed antler hunts and helicopter surveys. We also compared use of space for each species during these events. Female bighorn sheep did not increase distance moved or substantially change space use during shed antler hunts and helicopter surveys. Male bighorn sheep increased distance moved 41% on average during shed antler hunts and by 2.02 times during helicopter surveys but did not change space use during those events. Female bison increased distance moved 15% on average during shed antler hunts and 30% during helicopter surveys. Mule deer increased distance moved and altered space use the most during shed antler hunts; females increased distance moved 97%, and 54% of females moved a mean distance of 742 ± 642 (SD) m (range = 9–3,778 m) outside of their home ranges during those hunts for a mean of 9.2 ± 9.4 hours (range = 1.5 to 41 hr). Male mule deer increased distance moved by 2.10 times on average during shed antler hunts, and 82% of males moved a mean distance of 1,264 ± 732 m (range = 131–3,637 m) outside of their home ranges during those hunts for a mean of 12.6 ± 7.6 hours (range = 4.5–33 hr). Our results provide timely information about how legal shed antler hunting affects movements and space use of female and male ungulates, especially mule deer, and can guide the conservation of ungulate populations and their habitat. © 2021 The Wildlife Society.

Sex-Specific Elk Resource Selection during the Anthrax Risk Period.

Anthrax, caused by the spore-forming bacterium Bacillus anthracis, is a zoonosis affecting animals and humans globally. In the United States, anthrax outbreaks occur in wildlife and livestock, with frequent outbreaks in native and exotic wildlife species in Texas, livestock outbreaks in the Dakotas, and sporadic mixed outbreaks in Montana. Understanding where pathogen and host habitat selection overlap is essential for anthrax management. Resource selection and habitat use of ungulates may be sex-specific and lead to differential anthrax exposure risks across the landscape for males and females. We evaluated female elk (Cervus canadensis) resource selection in the same study areas as male elk in a previous anthrax risk study to identify risk of anthrax transmission to females and compare transmission risk between females and males. We developed a generalized linear mixed-effect model to estimate resource selection for female elk in southwest Montana during the June to August anthrax transmission risk period. We then predicted habitat selection of female and male elk across the study area and compared selection with the distribution of anthrax risk to identify spatial distributions of potential anthrax exposure for the male and female elk. Female and male elk selected different resources during the anthrax risk period, which resulted in different anthrax exposure areas for females and males. The sex-specific resource selection and habitat use could infer different areas of risk for anthrax transmission, which can improve anthrax and wildlife management and have important public health and economic implications.

Ungulate use of locally infectious zones in a re-emerging anthrax risk area.

Environmentally mediated indirect pathogen transmission is linked to host movement and foraging in areas where pathogens are maintained in the environment. In the case of anthrax, spores of the causative bacterium Bacillus anthracis are released into the environment following host death and create locally infectious zones (LIZs) around carcass sites; by grazing at LIZs, herbivores are potentially exposed to spores. Here, we used camera traps to assess how ungulate species use carcass sites in southwestern Montana and evaluated how these behaviours may promote indirect anthrax transmission, thus providing, to our knowledge, the first detailed documentation and study of the fine-scale mechanisms underlying foraging-based disease transmission in this ecosystem. We found that carcasses at LIZs significantly increased aboveground biomass of vegetation and concentrations of sodium and phosphorus, potentially making these sites more appealing to grazers. Host behavioural responses to LIZs varied depending on species, sex, season and carcass age; but, overall, our results demonstrated that carcasses or carcass sites serve as an attractant to herbivores in this system. Attraction to LIZs probably represents an increased risk of exposure to B. anthracis and, consequently, increased anthrax transmission rates. Accordingly, continued anthrax surveillance and control strategies are critical in this system.