Distribution of Competition Potential Between Native Ungulates and Free-Roaming Equids on Western Rangelands

Free-roaming equids (i.e., feral horses [Equus caballus] and burros [Equus asinus]) are widely distributed and locally abundant across the rangelands of the western United States. The 1971 Wild Free Roaming Horse and Burro Act (WFRHBA) gave the Bureau of Land Management (BLM) and United States Forest Service (USFS) the legal authority to manage these animals on designated public lands. To fulfill this responsibility, federal agencies established an Appropriate Management Level (AML), defined as the number of horses or burros that can be sustained on a given management unit under prevailing environmental conditions and land uses. Although the WFRHBA specifies that feral equids must be managed in ecological balance with other land uses, including conservation of native wildlife, population control measures such as gathers, contraception, and adoptions have failed to keep pace with intrinsic growth rates. Over 80% of federally managed herds currently exceed prescribed population levels, making the potential for competition between native ungulates and feral equids a growing concern among state wildlife agencies. Mule deer (Odocoileus hemionus), pronghorn (Antilocapra americana), elk (Cervus canadensis), and bighorn sheep (Ovis canadensis) are of ecological and economic value to the states where they occur, and all exhibit some degree of distributional, habitat, or dietary overlap with horses or burros. Notwithstanding the scale of the problem, to date there have been no range-wide assessments of competition potential among native and feral ungulates for space, forage, or water. To address this need, we compiled demographic, jurisdictional, and species occurrence data collected from 2010–2019 by federal and state agencies. We used these data to map the distributions of 4 native ungulate species across federal equid management units (FEMUS) in 10 western states (n = 174). We then made within-state rankings of the 50 units that were ≥2 times over AML and encompassed ≥3 native ungulates. Collectively, FEMUs covered approximately 225,000 km², representing 18% of all BLM and USFS lands in affected states. Each FEMU supported ≥1 native ungulate and 14% contained all 4. The degree of overlap between native and feral species varied by state, ranging from <1% for mule deer in Montana, to 40% for bighorn sheep in Nevada. Oregon had the largest proportion of units that supported all 4 native ungulates (58%), whereas Montana and New Mexico had the fewest equids, but all populations were over target densities. Despite the perception that the problem of equid abundance is limited to the Great Basin states, high intrinsic growth rates and social constraints on management practices suggest all affected states should monitor range conditions and native ungulate demography in areas where forage and water resources are limited and expanding equid populations are a concern. © 2021 The Wildlife Society.     

Prey Specialization by Cougars on Feral Horses in a Desert Environment

Natural 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.     

Feral horse impacts on threatened plants and animals in sub‐alpine and montane environments in Victoria,Australia

Feral herbivores are a major driver of biodiversity loss globally and can alter the structure, composition and functioning of ecosystems. The direct impacts of feral herbivores on plant communities are well studied, but the direct and indirect effect they have on wildlife is not well understood. In Victoria (south‐eastern Australia), a large feral Horse (Equus caballus) population coincides with highly sensitive and nationally endangered Alpine Sphagnum Bogs and Associated Fens communities, and several threatened animal species. We assessed the impact of feral horses on this ecological community and the Alpine Water Skink (Eulamprus kosciuskoi) and the Broad‐toothed Rat (Mastacomys fuscus) at 20 sites with varying levels of horse disturbance. We used scat counts to determine an index of feral horse abundance and quantified impacts associated with their presence in the landscape. Active searches were used for Alpine Water Skink and scat and runway surveys for Broad‐toothed Rat. We also measured the vegetation structure and the abundance of different vegetation types (life forms). Our results suggest that feral horses are associated with vegetation types and characteristics that negatively influence the presence or abundance of Alpine Water Skink and Broad‐toothed Rat. Sites with high horse activity had more low‐growing forbs, and the abundance of Alpine Water Skink was negatively related to this vegetation type. Grasses, sedges, rushes and shrubs were also less dense and lower in height in high horse activity sites, and Broad‐toothed Rat was less likely to be present in areas with these habitat attributes. We recommend that feral horses are controlled to protect these threatened vertebrate species and their Sphagnum bog habitat.     

Seed dispersal by ungulates in the point calimere wildlife sanctuary: A scientific and perspective analysis

Exotic woody weed plants are a very serious threat to seed dispersed by ungulate in the tropical forest of Asia. The ungulates in Point Calimere Wildlife Sanctuary (PCWS) are a significant role in native indigenous seed dispersal. The exotic woody weed tree Prosopis juliflora prevalence distributed in the PCWS and they might potentially alter the native medicinal plant species. In the present investigation, we have assessed the seed dispersal by ungulates in PCWS from January to March 2017. Four different ungulate species were selected to understand their seed dispersal rate of different plant species in selected sanctuary. This investigation was planned to confirm the seed dispersal by ungulates of blackbuck, spotted deer, wild boar and feral horse. Among the four different ungulates tested, the maximum numbers of pellets collected from blackbuck and no seed found in their pellets. The low quantities of pellets were collected from wild boar and this study has recorded medium-sized ungulates which dispersed variety of plant. However, the dispersal of the seed of medicinal plants were not considerably high and relatively moderate percentage of seeds dispersal occurred in medium-sized ungulates like wild boar and spotted deer. P. juliflora had 100% seed germination rate were observed from the faecal samples of wild boar and feral horse. The control seed achieved maximum seedling rate than the ungulates seeds.     

An emerging infectious disease triggering large-scale hyperpredation

Hyperpredation refers to an enhanced predation pressure on a secondary prey due to either an increase in the abundance of a predator population or a sudden drop in the abundance of the main prey. This scarcely documented mechanism has been previously studied in scenarios in which the introduction of a feral prey caused overexploitation of native prey. Here we provide evidence of a previously unreported link between Emergent Infectious Diseases (EIDs) and hyperpredation on a predator-prey community. We show how a viral outbreak caused the population collapse of a host prey at a large spatial scale, which subsequently promoted higher-than-normal predation intensity on a second prey from shared predators. Thus, the disease left a population dynamic fingerprint both in the primary host prey, through direct mortality from the disease, and indirectly in the secondary prey, through hyperpredation. This resulted in synchronized prey population dynamics at a large spatio-temporal scale. We therefore provide evidence for a novel mechanism by which EIDs can disrupt a predator-prey interaction from the individual behavior to the population dynamics. This mechanism can pose a further threat to biodiversity through the human-aided disruption of ecological interactions at large spatial and temporal scales.     

Phantoms of the forest: legacy risk effects of a regionally extinct large carnivore

The increased abundance of large carnivores in Europe is a conservation success, but the impact on the behavior and population dynamics of prey species is generally unknown. In Europe, the recolonization of large carnivores often occurs in areas where humans have greatly modified the landscape through forestry or agriculture. Currently, we poorly understand the effects of recolonizing large carnivores on extant prey species in anthropogenic landscapes. Here, we investigated if ungulate prey species showed innate responses to the scent of a regionally exterminated but native large carnivore, and whether the responses were affected by human‐induced habitat openness. We experimentally introduced brown bear Ursus arctos scent to artificial feeding sites and used camera traps to document the responses of three sympatric ungulate species. In addition to controls without scent, reindeer scent Rangifer tarandus was used as a noncarnivore, novel control scent. Fallow deer Dama dama strongly avoided areas with bear scent. In the presence of bear scent, all ungulate species generally used open sites more than closed sites, whereas the opposite was observed at sites with reindeer scent or without scent. The opening of forest habitat by human practices, such as forestry and agriculture, creates a larger gradient in habitat openness than available in relatively unaffected closed forest systems, which may create opportunities for prey to alter their habitat selection and reduce predation risk in human‐modified systems that do not exist in more natural forest systems. Increased knowledge about antipredator responses in areas subjected to anthropogenic change is important because these responses may affect prey population dynamics, lower trophic levels, and attitudes toward large carnivores. These aspects may be of particular relevance in the light of the increasing wildlife populations across much of Europe.     

Impacts of feral horses in the Australian Alps and evidence‐based solutions

New evidence of impacts by feral horses in Australia's alpine parks systems confirms they endanger threatened species and extensively damage critically endangered bog communities that could take millennia to recover. These impacts are not confounded by effects of deer and accumulate over time, even when only a small number of feral horses (~100) are present. With protected areas representing only a small proportion of the area of the Australian states of New South Wales (9.3%) and Victoria (17%), allowing feral horses to degrade reserves is not a reasonable management compromise, is contrary to the purpose of the protected area system and conflicts with international obligations. Modelling and decades of management experience indicate that trapping alone does not control feral horse numbers. Trapping and fertility control can work in small populations, but not when there are several thousand horses in remote areas. Aerial culling is needed to cost‐effectively and humanely control feral horse populations. The relatively small amount of suffering feral horses experience during a cull is outweighed by (i) avoiding suffering and death of horses from starvation and thirst, (ii) avoiding the suffering of native animals displaced by horses and (iii) avoiding the ethical concerns of driving threatened species towards extinction. Objections to aerial culling on welfare and cultural grounds are contradicted by evidence. Improving knowledge in the general community about what is at stake is long overdue because without this knowledge, small groups with vested interests and unfounded claims have been able to dominate debate and dictate management actions. As a result of ineffective management, horse populations are now expanding and causing well‐documented damage to Australia's alpine parks, placing at risk almost $10M spent on restoration after livestock grazing ended. The costs of horse control and restoration escalate the longer large horse populations remain in the alpine parks. It is crucial that feral horse numbers are rapidly reduced to levels where ecosystems begin to recover. Aerial culling is needed as part of the toolbox to achieve that reduction.     

An assessment of feral horse impacts on treeless drainage lines in the Australian Alps

The feral Horse (Equus caballus) is widespread across the Australian Alps. Feral horses degrade alpine and sub‐alpine ecosystems and damage habitat of a range of threatened species. Despite this, there is little published work to document the extent and severity of these impacts. This study investigated impacts of feral horses on treeless drainage lines at 186 sites across the Australian Alps. The study included sites in the Australian Capital Territory, New South Wales and Victoria. We assessed nine variables related to soil and stream stability and vegetation cover, which in turn influence ecosystem function and habitat quality. We found significant differences among horse‐occupied and horse‐free sites for all soil and stream stability variables assessed. For all variables assessed, the average score (and hence, condition) was worse in horse‐occupied areas. The sites in poorest condition were occupied by horses. Impacts from other mammalian herbivores species appeared to be minor. Management intervention is necessary if these impacts of feral horses are to be addressed.     

Causes and consequences of lags in basic and applied research into feral wildlife ecology: the case for feral horses

The biomass of feral wildlife is eclipsing that of native wildlife in many parts of the world. Consequently, feral species are playing an increasingly important role in ecological community dynamics. Artificially selected life-history traits of wild but once domesticated species can elicit population dynamics that differ substantially from that of native species. Yet, we continue to lag in our understanding of the ecology and evolution of feral species with direct consequences to resource management and biodiversity conservation. In part, this is because basic and applied research into the ecology of feral wildlife is fraught with social and political challenges unique to science. Feral populations of companion animals or livestock, especially, can evoke strong emotional reactions among advocacy groups, particularly around issues of animal welfare and management policy. Managers tasked with controlling feral populations are often bound by social license, including legislative restrictions, incomparable to that of other wildlife, and harassment or litigation of researchers and managers is not uncommon. Further, research and management of feral species is often delegated to agricultural instead of wildlife government agencies with clear differences in mandate, staff education, and training. Using examples primarily from feral horses in North America, we show how scientists conducting research independent of the management process can find themselves placed between managers, advocates, and opponents of feral species, implicitly tasked with satisfying multiple and often contradictory interests of stakeholders, sometimes with direct and litigious interference. These barriers are exacerbated by inter-disciplinary tendencies to dismiss the importance of basic and applied ecological research into feral species, despite its relevance to sound decision-making. Feral species therefore possess politically and biologically facilitated asymmetries that favor persistence, growth, and expansion relative to native wildlife, while the timely study of these characteristics in nature continues to suffer from ideological opposition.     

Temporal Overlap Among Feral Horses,Cattle, and Native Ungulates at Water Sources

Feral horse (Equus ferus caballus) populations on public rangelands in the western United States threaten forage production for livestock and wildlife habitat. Interference competition between feral horses and heterospecifics at watering sources can have negative effects on livestock and wildlife. Researchers have documented altered timing and behavior of wild ungulates at water sources when horses were present. The few studies examining these interactions have infrequently occurred within areas specifically managed for feral equids and have not occurred in sites with cattle. We used motion-sensitive cameras at 8 watering sources to document watering activity patterns and construct indices of temporal overlap among feral horses, cattle, elk (Cervus canadensis), mule deer (Odocoileus hemionus), and pronghorn (Antilocapra americana) within the Adobe Town Herd Management Area in southern Wyoming, USA, between June and September 2018 and 2019. Feral horses, cattle, and pronghorn exhibited a high degree of temporal overlap (>79%) in water use, with feral horses and pronghorn exhibiting the highest estimated percent overlap (88.1%, 95% CI = 86.5–89.6%). Mule deer and elk watering activity also overlapped with horses and cattle but to a lesser degree (<55%). Feral horses spent a mean of 16.7 ± 30.5 (SD) minutes during a watering event and were present at a given water source on average 4.5 ± 6.3% and up to 34.9% of the day, which is less than reported in previous studies. Cattle spent on average 23.5 ± 44.9 minutes during a watering event, and were present on average 4.2 ± 7.7% and up to 42.4% of the day at a single water source. Results of generalized linear mixed-effects models indicated that number of conspecifics was the strongest predictor of visit duration for pronghorn and horses; hour of the day and group size of heterospecifics were informative, but less important, variables. There was no difference in peak visitation time for any species between sites of high versus low horse or cattle use. Despite temporal overlap, we did not find evidence of interference competition between feral horses, cattle, and pronghorn. We recommend future examination of interference competition and its biological consequences between introduced and native ungulates at water sources of varying size across sites, equid population levels, and livestock stocking rates. © 2020 The Wildlife Society.     

Relative influence of habitat structure,species interactions and rainfall on the post‐fire population dynamics of ground‐dwelling vertebrates

The long‐term impacts of wildfires on animal populations are largely unknown. We used time‐series data based on a tracking index, from coastal NSW spanning 28 years after a wildfire, to investigate the relative influence of habitat structure, species interactions and climate on post‐fire animal population dynamics. The fire had an immediate impact on habitat structure, reducing and simplifying vegetation cover, which then underwent post‐fire successional change including an increase and plateau in tree canopy cover; an increase, stabilization and then decline in shrub cover; and an increase in ground litter cover. Population changes of different animal species were influenced by different components of successional change, but there was also evidence that species interactions were important. For example, bandicoots (Isoodon obesulus and Perameles nasuta combined) increased concurrent with an increase in shrub cover then declined at a faster rate than a direct association with senescing shrub cover would suggest, while the feral cat (Felis catus) population changed with the bandicoot population, suggesting a link between these species. Potoroos (Potorous tridactylus) increased 10 years after the fire concurrent with the closing tree canopy, but there was also evidence of a negative association with feral foxes (Vulpes vulpes). Variation in rainfall did not have significant effects on the population dynamics of any species. Our results suggest that changes in habitat structure play a key role in the post‐fire dynamics of many ground‐dwelling animals and hence different fire regimes are likely to influence animal dynamics through their effects on habitat structure. However, the role of predator–prey interactions, particularly with feral predators, is less clear and further study will require manipulative experiments of predators in conjunction with fire treatments to determine whether feral predator control should be integrated with fire management to improve outcomes for some native species.     

Do rabbits eat voles? Apparent competition,habitat heterogeneity and large‐scale coexistence under mink predation

Habitat heterogeneity is predicted to profoundly influence the dynamics of indirect interspecific interactions; however, despite potentially significant consequences for multi-species persistence, this remains almost completely unexplored in large-scale natural landscapes. Moreover, how spatial habitat heterogeneity affects the persistence of interacting invasive and native species is also poorly understood. Here we show how the persistence of a native prey (water vole, Arvicola terrestris ) is determined by the spatial distribution of an invasive prey (European rabbit, Oryctolagus cuniculus ) and directly infer how this is defined by the mobility of a shared invasive predator (American mink, Neovison vison ). This study uniquely demonstrates that variation in habitat connectivity in large-scale natural landscapes creates spatial asynchrony, enabling coexistence between apparent competitive native and invasive species. These findings highlight that unexpected interactions may be involved in species declines, and also that in such cases habitat heterogeneity should be considered in wildlife management decisions.     

Influence of interspecific competition on mule deer birthing and rearing site selection

Ungulates often alter behavior and space use in response to interspecific competition. Despite observable changes in behavior caused by competitive interactions, research describing the effects of competition on survival or growth is lacking. We used spatial modeling to determine if habitat use by female mule deer (Odocoileus hemionus) was affected by other ungulate species prior to, during, and after parturition. We conducted our study in the Book Cliffs region of eastern Utah, USA, during 2019 and 2020. We used resource selection function (RSF) analysis to model space use of 4 ungulate species that potentially competed with mule deer: bison (Bos bison), cattle, elk (Cervus canadensis), and feral horses. We incorporated RSF models for competing species into a random forest analysis to determine if space use by mule deer was influenced by these other ungulate species. We used survival and growth data from neonate mule deer to directly assess potential negative effects of other ungulates. Habitat use by elk was an important variable in predicting use locations of mule deer during birthing and rearing. The relationship was positive, suggesting interference competition was not occurring. Survival of neonate mule deer increased as the probability of use by elk increased (hazard ratio = 0.185 ± 0.497 [SE]). Further, probability of use by elk in rearing habitat had no influence on growth of neonate mule deer from birth to 6 months of age, suggesting that exploitative competition was not occurring.     

Drones are an effective tool to assess the impact of feral horses in an alpine riparian environment

Protected areas can be impacted by the presence and proliferation of feral species. Effective management of feral species requires reliable tools to monitor their population size and ecological impacts. Here, we used drone-based image analysis to assess evidence of feral horses and horse-specific ecological impacts on alpine riparian habitat. Valleys with low (0), medium (1–16) and high (>16) horse abundances were chosen for drone imagery analysis based on independent aerial counts of horses. Data collection trips were carried out pre- and post-2019/2020 wildfires, which unexpectedly burnt valleys with low horse presence. Drone-based RGB orthomosaic imagery was sufficient to identify seven indicators of horse presence and determine the severity of feral horse impacts. Despite the impact of fire, drone-derived classifications were able to accurately detect a gradient of horse impacts, showing a significant difference in indicators from low presence valleys compared with medium and high presence valleys, which did not differ significantly from each other. The significance of differences between valleys reveals that regions routinely inhabited by feral horses will display significant environmental impacts. Our results clearly indicated significant differences between valleys with low horse presence compared with either medium or high horse presence regions (0.01 for differences between both low and medium and low and high horse presence). This was evident both before and after the 2019/2020 fires, suggesting that wildfires did not significantly impact horse populations or distribution in the sampled region. Overall, it was evident that feral horses have a clear and definable impact on alpine riparian vegetation, and drone surveying can be used to routinely monitor potential spread and the outcome of management actions.     

Lack of feral livestock interference with native guanaco during the dry season in a South American desert

Analyzing coexistence of exotic and native ungulates in arid areas is important from both a theoretical and a species conservation perspective. We assess the habitat use patterns and possible interference between guanaco (Lama guanicoe) and feral livestock (donkey and cattle) in arid environments of South America. To determine habitat use and niche overlap between exotic and native ungulate species, ten sites with different habitats and six natural waterholes were selected. Plots (20 at each site, ten around each waterhole) were randomly set up and characterized by environmental variables and relative use by cattle, donkey and guanaco through faecal pellet counts. Aggregation, niche breadth and niche overlap of the three herbivores were analyzed at habitat level (mesoscale). A direct redundancy analysis was used to examine the relationships between abundance of herbivore faeces and environmental variables at microscale. Mesoscale analyses showed (i) an extensive use of the area by all three species, with guanaco having the highest niche breadth followed by donkey and cattle and (ii) a large, broad guanaco–donkey and donkey–cattle habitat overlap. However, results at a finer scale showed high spatial aggregation of feral livestock species and an independent use of territory by guanacos. This study is the first to provide information about habitat partitioning between guanacos and feral livestock in the hyper-arid Monte Desert biome and points to an apparent lack of negative effects on the native ungulate.     

Population dynamical consequences of reduced predator switching at low total prey densities

Several models of rapid switching by a predator in a two-prey environment are analyzed. The goal is to determine how the dynamics of the system and the potential indirect effects between prey are affected by the dependence of switching on total prey density. In exploring this question, the difference between the population-level consequences of switching in stable and cycling predator-prey systems is also examined. We concentrate on reduced switching at low densities, a feature that is likely because of the difficulty of distinguishing between two very low densities. The main findings are: (1) switching in unstable systems can produce positive indirect effects of one prey species on the other; and (2) reduced switching at low densities can greatly alter the dynamics of the system and the indirect effects between prey. Both of the possibilities are only evident in cycling systems. Reduced switching at low total prey densities leads to heavier predation on the slower-growing prey when both prey species are rare. As a consequence, there is a lag in the recovery of the slower-growing prey species after predator densities fall, and the dynamics of the two prey become desynchronized. The net result is increased indirect interactions between prey, and a greater likelihood of exclusion of the slower growing prey. The analysis of these models suggests a need for more empirical work to determine whether switching is reduced by very low total prey densities, and to study the long-term dynamics that occur in systems with switching predators.     

Native and Non-Native Plants Provide Similar Refuge to Invertebrate Prey,but Less than Artificial Plants

Non-native species introductions are widespread and can affect ecosystem functioning by altering the structure of food webs. Invading plants often modify habitat structure, which may affect the suitability of vegetation as refuge and could thus impact predator-prey dynamics. Yet little is known about how the replacement of native by non-native vegetation affects predator-prey dynamics. We hypothesize that plant refuge provisioning depends on (1) the plant’s native status, (2) plant structural complexity and morphology, (3) predator identity, and (4) prey identity, as well as that (5) structurally similar living and artificial plants provide similar refuge. We used aquatic communities as a model system and compared the refuge provided by plants to macroinvertebrates (Daphnia pulex, Gammarus pulex and damselfly larvae) in three short-term laboratory predation experiments. Plant refuge provisioning differed between plant species, but was generally similar for native (Myriophyllum spicatum, Ceratophyllum demersum, Potamogeton perfoliatus) and non-native plants (Vallisneria spiralis, Myriophyllum heterophyllum, Cabomba caroliniana). However, plant refuge provisioning to macroinvertebrate prey depended primarily on predator (mirror carp: Cyprinus carpio carpio and dragonfly larvae: Anax imperator) and prey identity, while the effects of plant structural complexity were only minor. Contrary to living plants, artificial plant analogues did improve prey survival, particularly with increasing structural complexity and shoot density. As such, plant rigidity, which was high for artificial plants and one of the living plant species evaluated in this study (Ceratophyllum demersum), may interact with structural complexity to play a key role in refuge provisioning to specific prey (Gammarus pulex). Our results demonstrate that replacement of native by structurally similar non-native vegetation is unlikely to greatly affect predator-prey dynamics. We propose that modification of predator-prey interactions through plant invasions only occurs when invading plants radically differ in growth form, density and rigidity compared to native plants.     

The occurrence of the Broad‐toothed Rat Mastacomys fuscus in relation to feral Horse impacts

Feral Horse (Equus caballus) impacts in northern Kosciuszko National Park, New South Wales, Australia are directly occurring in habitat of the nationally threatened Broad‐toothed Rat (Mastacomys fuscus). This species is endemic primarily to the mountain regions of south‐eastern mainland Australia and Tasmania, with a disjunct population at Barrington Tops. The Broad‐toothed Rat's preferred habitat is being increasingly impacted by browsing and trampling associated with the expansion of feral horse populations. This study surveyed 180 sites supporting preferred habitat for this species to determine Broad‐toothed Rat presence and relative abundance in relation to the level of feral horse impacts within the reserve. There was a significant negative relationship between feral horse impacts and both Broad‐toothed Rat presence and abundance. No scats were identified at localities where feral horse impacts were severe, and at moderate horse impact sites, there was a proportion (34%) without scats found. Locations with low horse impacts had little impact on Broad‐toothed Rat occurrence. As feral horse populations increase, Broad‐toothed Rat populations may be further impacted. Such impacts will be due to the loss of vegetation cover from feral horse trampling and grazing, making animals more vulnerable to predation by predators or impacting on their ability to disperse to more suitable habitat. Habitat remnants and vegetation corridors along drainage lines require protection from feral horses to prevent localized extinctions of Broad‐toothed Rat.     

Consequences of behavioral dynamics for the population dynamics of predator-prey systems with switching

This article explores how different mechanisms governing the rate of change of the predators preference alter the dynamics of predator-prey systems in which the predator exhibits positive frequency-dependent predation. The models assume that individuals of the predator species adaptively adjust a trait that determines their relative capture rates of each of two prey species. The resulting switching behavior does not instantaneously attain the optimum for current prey densities, but instead lags behind it. Several mechanisms producing such lags are discussed and modeled. In all cases examined, our question is whether a realistic behavioral lag can significantly change the dynamics of the system relative to an analogous case in which the predators switching is effectively instantaneous. We also explore whether increasing the rate parameters of dynamic models of behavior results in convergence to the population dynamics of analogous models with instantaneous switching, and whether different behavioral models produce similar population dynamics. The analysis concentrates on systems that undergo endogenously generated predator-prey cycles in the absence of switching behavior. The average densities and the nature of indirect interactions are often sensitive to the rate of behavioral change, and are often qualitatively different for different classes of behavioral models. Dynamics and average densities can be very sensitive to small changes in parameters of either the prey growth or predator switching functions. These differences suggest that an understanding of switching in natural systems will require research into the behavioral mechanisms that govern lags in the response of predator preference to changes in prey density.     

Habitat selection and space use overlap between feral horses,pronghorn, and greater sage-grouse in cold arid steppe

Populations of feral horses (Equus ferus caballus) in the western United States have increased during the past decade, consequently affecting co-occurring wildlife habitat. Feral horses may influence 2 native wildlife species, greater sage-grouse (Centrocercus urophasianus; sage-grouse) and pronghorn (Antilocapra americana) through mechanisms of habitat alteration and competition. Wyoming, USA, contains the largest populations of pronghorn and sage-grouse of any state and also has the highest degree of range overlap between feral horses and these species. Consequently, the effects that horses may have on pronghorn and sage-grouse populations in Wyoming have implications at local, state, and population-wide levels. Managers need information concerning habitat selection and space use overlap among these species to develop appropriate management strategies; yet this information is absent for most feral horse management areas. To address this knowledge need, we attached global positioning system (GPS) transmitters to horses, pronghorn, and sage-grouse within the greater Bureau of Land Management–Adobe Town Herd Management Area in southern Wyoming and northern Colorado, USA, between 2017 and 2021 to evaluate habitat selection and space use of all species during 3 biologically relevant seasons: spring (Apr–Jun; sage-grouse breeding, nesting, and early-brood rearing; pronghorn late gestation and early parturition), summer (Jul–Oct; sage-grouse summer and late-brood rearing; pronghorn late parturition and breeding), and winter (Nov–Mar; non-breeding season). Feral horses selected flatter slopes and shorter mean shrub height across all seasons and were closer to water in spring and summer. Pronghorn habitat selection was similar to horses, but they also avoided oil and gas well pads year-round. During spring, sage-grouse selected greater herbaceous cover, flatter slopes, and areas farther from well pads. In summer, sage-grouse selected greater mean shrub height, flatter slopes, and were closer to water. In winter, sage-grouse selected flatter slopes and areas with greater vegetation production during the preceding summer. Our results indicate strong year-round overlap in space use between horses and pronghorn, whereas overlap between horses and sage-grouse is greatest during the summer in this region. Consequently, managers should recognize the potential for horses to influence habitat quality of pronghorn and sage-grouse in the region.