Support for the size-mediated sensitivity hypothesis within a diverse carnivore community.

Carnivore community dynamics are governed by a complex set of often interacting biotic, abiotic and anthropogenic factors that are increasingly volatile as a result of global change. Understanding how these changing conditions influence carnivore communities is urgent because of the important role carnivores play within ecosystems at multiple trophic levels, and the conservation threats that many carnivores face globally. While a great deal of research attention has historically been focused on large carnivores within ecosystems, the size-mediated sensitivity hypothesis has recently been proposed where the smallest carnivore in a system is likely to be the most responsive to the diverse suite of ongoing environmental and anthropogenic changes within ecological communities. We deployed camera traps at 197 sites over 4 years to monitor a diverse suite of mammalian carnivores within the Blue Ridge Mountains of western North Carolina and then used a two-step occupancy modelling-structural equation modelling framework to investigate the relative support for four primary hypothesized drivers (interspecific competition/predation, habitat complexity, food availability and anthropogenic disturbance) on carnivore occurrence. We found that each of the 10 carnivores in our system responded differently to conditions associated with each of these four hypothesized drivers, but that small and medium-sized carnivores had a greater number of significant (p < 0.05) pathways by which these conditions were influencing occupancy relative to large carnivores. In particular, the smallest carnivore observed in our study was the only species for which we found support for each of the four hypothesized drivers influencing occupancy. Collectively, our study supports the size-mediated sensitivity hypothesis and suggests that small carnivores are ideal sentinel species for global change. We echo recent calls for adopting a middle-out approach to investigations into carnivore community dynamics by refocusing sustained monitoring and research efforts on smaller carnivores within systems.

SNAPSHOT USA 2020: A second coordinated national camera trap survey of the United States during the COVID-19 pandemic.

Managing wildlife populations in the face of global change requires regular data on the abundance and distribution of wild animals, but acquiring these over appropriate spatial scales in a sustainable way has proven challenging. Here we present the data from Snapshot USA 2020, a second annual national mammal survey of the USA. This project involved 152 scientists setting camera traps in a standardized protocol at 1485 locations across 103 arrays in 43 states for a total of 52,710 trap-nights of survey effort. Most (58) of these arrays were also sampled during the same months (September and October) in 2019, providing a direct comparison of animal populations in 2 years that includes data from both during and before the COVID-19 pandemic. All data were managed by the eMammal system, with all species identifications checked by at least two reviewers. In total, we recorded 117,415 detections of 78 species of wild mammals, 9236 detections of at least 43 species of birds, 15,851 detections of six domestic animals and 23,825 detections of humans or their vehicles. Spatial differences across arrays explained more variation in the relative abundance than temporal variation across years for all 38 species modeled, although there are examples of significant site-level differences among years for many species. Temporal results show how species allocate their time and can be used to study species interactions, including between humans and wildlife. These data provide a snapshot of the mammal community of the USA for 2020 and will be useful for exploring the drivers of spatial and temporal changes in relative abundance and distribution, and the impacts of species interactions on daily activity patterns. There are no copyright restrictions, and please cite this paper when using these data, or a subset of these data, for publication.

Empirical evaluation of the spatial scale and detection process of camera trap surveys.

BACKGROUND: Camera traps present a valuable tool for monitoring animals but detect species imperfectly. Occupancy models are frequently used to address this, but it is unclear what spatial scale the data represent. Although individual cameras monitor animal activity within a small target window in front of the device, many practitioners use these data to infer animal presence over larger, vaguely-defined areas. Animal movement is generally presumed to link these scales, but fine-scale heterogeneity in animal space use could disrupt this relationship. METHODS: We deployed cameras at 10 m intervals across a 0.6 ha forest plot to create an unprecedentedly dense sensor array that allows us to compare animal detections at these two scales. Using time-stamped camera detections we reconstructed fine-scale movement paths of four mammal species and characterized (a) how well animal use of a single camera represented use of the surrounding plot, (b) how well cameras detected animals, and (c) how these processes affected overall detection probability, p. We used these observations to parameterize simulations that test the performance of occupancy models in realistic scenarios. RESULTS: We document two important aspects of animal movement and how it affects sampling with passive detectors. First, animal space use is heterogeneous at the camera-trap scale, and data from a single camera may poorly represent activity in its surroundings. Second, cameras frequently (14-71%) fail to record passing animals. Our simulations show how this heterogeneity can introduce unmodeled variation into detection probability, biasing occupancy estimates for species with low p. CONCLUSIONS: Occupancy or population estimates with camera traps could be improved by increasing camera reliability to reduce missed detections, adding covariates to model heterogeneity in p, or increasing the area sampled by each camera through different sampling designs or technologies.

Attracting Diverse Students to Field Experiences Requires Adequate Pay, Flexibility, and Inclusion.

Access to field experiences can increase participation of diverse groups in the environmental and natural resources (ENR) workforce. Despite a growing interest among the ENR community to attract and retain diverse students, minimal data exist on what factors undergraduate students prioritize when applying for field experiences. Using a nationwide survey of US undergraduate ENR students, we show that attracting most students to field experiences-especially racial or ethnic minority students-will require pay above minimum wage. However, the concurrent landscape of pay in ENR fell short of meeting many students' pay needs. Aside from pay, ENR students valued training in technical field skills and analytical or research skills, working with their desired study species or taxa, and working near school or family. Additional barriers beyond limited pay included incompatible schedules and noninclusive work environments. Our findings provide important insights for attracting a diverse workforce to this critical stage in career advancement for students in ENR.

Predicted climate-induced reductions in scavenging in eastern North America.

Scavenging is an important function within ecosystems where scavengers remove organic matter, reduce disease, stabilize food webs, and generally make ecosystems more resilient to environmental changes. Global change (i.e., changing climate and increasing human impact) is currently influencing scavenger communities. Thus, understanding what promotes species richness in scavenger communities can help prioritize management actions. Using a long-term dataset from camera traps deployed with animal carcasses as bait along a 1881 km latitudinal gradient in the Appalachian Mountains of eastern USA, we investigated the relative impact of climate and humans on the species richness and diversity of vertebrate scavengers. Our most supported models for both mammalian and avian scavengers included climatic, but not human, variables. The richness of mammalian and avian scavengers detected was highest during relatively warm (5-10°C) and dry (100-150 mm precipitation) winters, when food was likely limited and both reliance on and detection of carrion was high. The diversity of mammalian and avian scavengers detected was highest under drier conditions. We then used these results to project the future species richness of scavengers that would be detected within our sampling area and under the climate scenario of 2070 (emissions level RCP8.5). Our predictions suggest up to 80% and 67% reductions, respectively, in the richness of avian and mammalian scavengers that would be detected at baited sites. Climate-induced shifts in behavior (i.e., reduction in scavenging, even if present) at this scale could have cascading implications for ecosystem function, resilience, and human health. Further, our study highlights the importance of conducting studies of scavenger community dynamics within ecosystems across wide spatial gradients within temperate environments. More broadly, these findings build upon our understanding of the impacts of climate-induced adjustments in behavior that can likely have negative impacts on systems at a large scale.

SNAPSHOT USA 2019: a coordinated national camera trap survey of the United States.

With the accelerating pace of global change, it is imperative that we obtain rapid inventories of the status and distribution of wildlife for ecological inferences and conservation planning. To address this challenge, we launched the SNAPSHOT USA project, a collaborative survey of terrestrial wildlife populations using camera traps across the United States. For our first annual survey, we compiled data across all 50 states during a 14-week period (17 August-24 November of 2019). We sampled wildlife at 1,509 camera trap sites from 110 camera trap arrays covering 12 different ecoregions across four development zones. This effort resulted in 166,036 unique detections of 83 species of mammals and 17 species of birds. All images were processed through the Smithsonian's eMammal camera trap data repository and included an expert review phase to ensure taxonomic accuracy of data, resulting in each picture being reviewed at least twice. The results represent a timely and standardized camera trap survey of the United States. All of the 2019 survey data are made available herein. We are currently repeating surveys in fall 2020, opening up the opportunity to other institutions and cooperators to expand coverage of all the urban-wild gradients and ecophysiographic regions of the country. Future data will be available as the database is updated at eMammal.si.edu/snapshot-usa, as will future data paper submissions. These data will be useful for local and macroecological research including the examination of community assembly, effects of environmental and anthropogenic landscape variables, effects of fragmentation and extinction debt dynamics, as well as species-specific population dynamics and conservation action plans. There are no copyright restrictions; please cite this paper when using the data for publication.

Habitat complexity and lifetime predation risk influence mesopredator survival in a multi-predator system.

Variability in habitat selection can lead to differences in fitness; however limited research exists on how habitat selection of mid-ranking predators can influence population-level processes in multi-predator systems. For mid-ranking, or mesopredators, differences in habitat use might have strong demographic effects because mesopredators need to simultaneously avoid apex predators and acquire prey. We studied spatially-explicit survival of cheetahs (Acinonyx jubatus) in the Mun-Ya-Wana Conservancy, South Africa, to test hypotheses related to spatial influences of predation risk, prey availability, and vegetation complexity, on mesopredator survival. For each monitored cheetah, we estimated lion encounter risk, prey density, and vegetation complexity within their home range, on short-term (seasonal) and long-term (lifetime) scales and estimated survival based on these covariates. Survival was lowest for adult cheetahs and cubs in areas with high vegetation complexity on both seasonal and lifetime scales. Additionally, cub survival was negatively related to the long-term risk of encountering a lion. We suggest that complex habitats are only beneficial to mesopredators when they are able to effectively find and hunt prey, and show that spatial drivers of survival for mesopredators can vary temporally. Collectively, our research illustrates that individual variation in mesopredator habitat use can scale-up and have population-level effects.

Home range size and resource use by swift foxes in northeastern Montana.

Swift foxes (Vulpes velox) are endemic to the Great Plains of North America, but were extirpated from the northern portion of their range by the mid-1900s. Despite several reintroductions to the Northern Great Plains, there remains a ~350 km range gap between the swift fox population along the Montana and Canada border and that in northeastern Wyoming and northwestern South Dakota. A better understanding of what resources swift foxes use along the Montana and Canada border region will assist managers to facilitate connectivity among populations. From 2016 to 2018, we estimated the home range size and evaluated resource use within the home ranges of 22 swift foxes equipped with Global Positioning System tracking collars in northeastern Montana. Swift fox home ranges in our study were some of the largest ever recorded, averaging (± SE) 42.0 km2 ± 4.7. Our results indicate that both environmental and anthropogenic factors influenced resource use. At the population level, resource use increased by 3.3% for every 5.0% increase in percent grasslands. Relative probability of use decreased by 7.9% and 7.4% for every kilometer away from unpaved roads and gas well sites, respectively, and decreased by 3.0% and 11.3% for every one-unit increase in topographic roughness and every 0.05 increase in normalized difference vegetation index (NDVI), respectively. Our study suggests that, to reestablish connectivity among swift fox populations in Montana, managers should aim to maintain large corridors of contiguous grasslands at a landscape scale, a process that likely will require having to work with multiple property owners.

Using multi-scale behavioral investigations to inform wild pig (Sus scrofa) population management.

Assessing invasive species ecology at multiple scales is needed to understand how to focus ecological monitoring and population control. As a widespread invasive species, wild pigs (Sus scrofa) frequently disrupt land management programs. We provide a detailed, multi-scaled view of the behavior of wild pigs at Fort Hood, Texas, USA by assessing seasonal and daily movement patterns, and diet. First, we quantified movement behavior through assessment of both seasonal home range size and first passage time movement behavior from 16 GPS-collared wild pigs. Home ranges were relatively large (mean: 10.472 km2, SD: 0.472 km2), and Cox proportional hazard models predicted that pigs moved slowest at temperature extremes (15< °C <30), in the spring, in rougher terrain, and in grassland communities. Secondly, we analyzed wild pig stomach contents to determine diet throughout the year. Diet was primarily plant-based, and showed seasonal variation in such items as hard and soft mast, and the foliage of forbs and woody plants. Integration of both movement and diet analyses indicate that wild pigs are more likely to be lured into baited traps in the winter when movement rates are highest and plant-based food resources are likely less abundant. Wild pigs are likely to have the greatest impact on vegetative communities in grassland habitats during the spring season when movement is restricted. Collectively, this multi-scaled approach provided detailed information on wild pig behavioral ecology in this area that would not have been apparent by looking at any single measure individually or only at a large spatial scale (i.e., home range), and could be a useful approach in other invasive species management programs.

Context dependency of top-down, bottom-up and density-dependent influences on cheetah demography.

Research on drivers of demographic rates has mostly focused on top predators and their prey, and comparatively less research has considered the drivers of mesopredator demography. Of those limited studies, most focused on top-down effects of apex predators on mesopredator population dynamics, whereas studies investigating alternative mechanisms are less common. In this study, we tested hypotheses related to top-down, bottom-up and density-dependent regulation of demographic rates in an imperilled mesopredator, the cheetah (Acinonyx jubatus). We used a 25-year dataset of lion density, cheetah density and prey density from the Mun-Ya-Wana Conservancy in South Africa and assessed the effects of top-down, bottom-up and density-dependent drivers on cheetah survival and reproduction. In contrast to the top-down and bottom-up predictions, both adult and juvenile cheetahs experienced the lowest survival during months with high prey densities and low lion densities. We observed support only for a density-dependent response in juvenile cheetahs, where they had a higher probability of reaching independence during times with low cheetah density and low prey density. We did not identify any strong drivers of litter size. Collectively, our results indicate that high apex predator abundance might not always have negative effects on mesopredator populations, and suggest that context dependency in top-down, bottom-up and density-dependent factors may regulate demographic rates of cheetahs and other mesopredators. Our results highlight the complexities of population-level drivers of cheetah demographic rates and the importance of considering multiple hypotheses of mesopredator population regulation.

Variation in regional and landscape effects on occupancy of temperate bats in the southeastern U.S.

Habitat loss, wind energy development, and the disease white-nose syndrome are major threats contributing to declines in bat populations in North America. In the southeastern US in particular, the recent arrival of white-nose syndrome and changes in landscape composition and configuration have driven shifts in bat species populations and distributions. Effective management strategies which address these large-scale, community-level threats require landscape-scale analyses. Our objective was to model the relationship between ecoregional and landscape factors and occupancy by all bat species in South Carolina, USA, during summer. We conducted acoustic surveys from mid-May through July 2015 and 2016 and evaluated temporally dynamic occupancy models for eight bat species or species groups at the 100 km2 level. We found significant effects of landscape factors such as ecoregion and forest edge density for three species, but habitat condition effects were not statistically significant for five other species. Thus, for some species, site-use analyses may be more appropriate than larger scale occupancy analyses. However, our occupancy predictions generally matched statewide historical distributions for all species, suggesting our approach could be useful for monitoring landscape-level trends in bat species. Thus, while our scale of study was likely too coarse for assessing fine-scale habitat associations for all bat species, our findings can improve future monitoring efforts, inform conservation priorities, and guide subsequent landscape-scale studies for bat species and community-level responses to global change.

Integrating physiological stress into the movement ecology of migratory ungulates: a spatial analysis with mule deer.

Rapid climate and human land-use change may limit the ability of long-distance migratory herbivores to optimally track or 'surf' high-quality forage during spring green-up. Understanding how anthropogenic and environmental stressors influence migratory movements is of critical importance because of their potential to cause a mismatch between the timing of animal movements and the emergence of high-quality forage. We measured stress hormones (fecal glucocorticoid metabolites; FGMs) to test hypotheses about the effects of high-quality forage tracking, human land-use and use of stopover sites on the physiological state of individuals along a migratory route. We collected and analysed FGM concentrations from 399 mule deer (Odocoileus hemionus) samples obtained along a 241-km migratory route in western Wyoming, USA, during spring 2015 and 2016. In support of a fitness benefit hypothesis, individuals occupying areas closer to peak forage quality had decreased FGM levels. Specifically, for every 10-day interval closer to peak forage quality, we observed a 7% decrease in FGMs. Additionally, we observed support for both an additive anthropogenic stress hypothesis and a hypothesis that stopovers act as physiological refugia, wherein individuals sampled far from stopover sites exhibited 341% higher FGM levels if in areas of low landscape integrity compared to areas of high landscape integrity. Overall, our findings indicate that the physiological state of mule deer during migration is influenced by both anthropogenic disturbances and their ability to track high-quality forage. The availability of stopovers, however, modulates physiological responses to those stressors. Thus, our results support a recent call for the prioritization of stopover locations and connectivity between those locations in conservation planning for migratory large herbivores.

Correction: Seeing the Forest through the Trees: Considering Roost-Site Selection at Multiple Spatial Scales.

[This corrects the article DOI: 10.1371/journal.pone.0150011.].

Seeing the Forest through the Trees: Considering Roost-Site Selection at Multiple Spatial Scales.

Conservation of bat species is one of the most daunting wildlife conservation challenges in North America, requiring detailed knowledge about their ecology to guide conservation efforts. Outside of the hibernating season, bats in temperate forest environments spend their diurnal time in day-roosts. In addition to simple shelter, summer roost availability is as critical as maternity sites and maintaining social group contact. To date, a major focus of bat conservation has concentrated on conserving individual roost sites, with comparatively less focus on the role that broader habitat conditions contribute towards roost-site selection. We evaluated roost-site selection by a northern population of federally-endangered Indiana bats (Myotis sodalis) at Fort Drum Military Installation in New York, USA at three different spatial scales: landscape, forest stand, and individual tree level. During 2007-2011, we radiotracked 33 Indiana bats (10 males, 23 females) and located 348 roosting events in 116 unique roost trees. At the landscape scale, bat roost-site selection was positively associated with northern mixed forest, increased slope, and greater distance from human development. At the stand scale, we observed subtle differences in roost site selection based on sex and season, but roost selection was generally positively associated with larger stands with a higher basal area, larger tree diameter, and a greater sugar maple (Acer saccharum) component. We observed no distinct trends of roosts being near high-quality foraging areas of water and forest edges. At the tree scale, roosts were typically in American elm (Ulmus americana) or sugar maple of large diameter (>30 cm) of moderate decay with loose bark. Collectively, our results highlight the importance of considering day roost needs simultaneously across multiple spatial scales. Size and decay class of individual roosts are key ecological attributes for the Indiana bat, however, larger-scale stand structural components that are products of past and current land use interacting with environmental aspects such as landform also are important factors influencing roost-tree selection patterns.

Toward a mechanistic understanding of animal migration: incorporating physiological measurements in the study of animal movement.

Movements are a consequence of an individual's motion and navigational capacity, internal state variables and the influence of external environmental conditions. Although substantial advancements have been made in methods of measuring and quantifying variation in motion capacity, navigational capacity and external environmental parameters in recent decades, the role of internal state in animal migration (and in movement in general) is comparatively little studied. Recent studies of animal movement in the wild illustrate how direct physiological measurements can improve our understanding of the mechanisms underlying movement decisions. In this review, we synthesize and provide examples of how recent technical advances in the physiology-related fields of energetics, nutrition, endocrinology, immunology and ecotoxicology provide opportunities for direct measurements of physiological state in the study of animal movement. We then propose a framework for practitioners to enable better integration of studies of physiological state into animal movement ecology by assessing the mechanistic role played by physiology as both a driver and a modulator of movement. Finally, we highlight the current limitations and research priorities for better integration of direct measurements of animal physiological state into the study of animal movement.

Physiological stress and refuge behavior by African elephants.

Physiological stress responses allow individuals to adapt to changes in their status or surroundings, but chronic exposure to stressors could have detrimental effects. Increased stress hormone secretion leads to short-term escape behavior; however, no studies have assessed the potential of longer-term escape behavior, when individuals are in a chronic physiological state. Such refuge behavior is likely to take two forms, where an individual or population restricts its space use patterns spatially (spatial refuge hypothesis), or alters its use of space temporally (temporal refuge hypothesis). We tested the spatial and temporal refuge hypotheses by comparing space use patterns among three African elephant populations maintaining different fecal glucocorticoid metabolite (FGM) concentrations. In support of the spatial refuge hypothesis, the elephant population that maintained elevated FGM concentrations (iSimangaliso) used 20% less of its reserve than did an elephant population with lower FGM concentrations (Pilanesberg) in a reserve of similar size, and 43% less than elephants in the smaller Phinda reserve. We found mixed support for the temporal refuge hypothesis; home range sizes in the iSimangaliso population did not differ by day compared to nighttime, but elephants used areas within their home ranges differently between day and night. Elephants in all three reserves generally selected forest and woodland habitats over grasslands, but elephants in iSimangaliso selected exotic forest plantations over native habitat types. Our findings suggest that chronic stress is associated with restricted space use and altered habitat preferences that resemble a facultative refuge behavioral response. Elephants can maintain elevated FGM levels for ≥ 6 years following translocation, during which they exhibit refuge behavior that is likely a result of human disturbance and habitat conditions. Wildlife managers planning to translocate animals, or to initiate other management activities that could result in chronic stress responses, should consider the potential for, and consequences of, refuge behavior.

Field evaluation of imidacloprid as a systemic approach to flea control in black-tailed prairie dogs, Cynomys ludovicianus.

Epizootic outbreaks of sylvatic plague have dramatically influenced prairie dog (Cynomys sp.) populations across North America. While a great deal of debate surrounds the cause and persistence of plague, flea control can stop the spread of plague epizootic outbreaks and even increase prairie dog survival under non-epizootic conditions. We investigated a newly-developed imidacloprid-treated grain bait that could potentially reduce flea infestations and mitigate the effects of plague on black-tailed prairie dogs (C. ludovicianus). We used a study design involving randomly assigned experimental and control study plots to assess the effectiveness of the systemic flea control product. We observed a significant difference in flea prevalence and abundance between experimental and control sites on three of the four sites treated with a single application of imidacloprid-treated grain bait for up to 90 days post-treatment. We observed an even greater reduction in flea infestations following the double application of treatment bait on two of three additional experimental sites. While we were unable to reduce flea infestations to the extent reported for more commonly used topical insecticides containing deltamethrin, imidacloprid might still be effective at reducing the risk of plague and halting epizootics. In addition, this systemic product can be more rapidly applied than topical insecticides, providing managers with a tool to quickly reduce flea infestations. Future research is needed to evaluate the effectiveness of different application timing and rates, the utility of the product in limiting plague, and the potential effects on non-target species that might also consume the treated bait.

Seasonal and among-site variation in the occurrence and abundance of fleas on California ground squirrels (Otospermophilus beecheyi).

An improved understanding of the ecology of fleas on California ground squirrels, Otospermophilus beecheyi, is warranted given the role of fleas in the transmission, and perhaps persistence, of the plague-causing bacterium Yersinia pestis. We sampled O. beecheyi on a seasonal basis from three study sites, each representing a different land use type (preserve, pasture, and agriculture) in the San Joaquin Valley, CA. Overall, the abundance of fleas on squirrels was greatest in spring at the preserve site, in summer at the agriculture and pasture sites, and in winter at the pasture site. Hoplopsyllus anomalus, the species most frequently found on squirrels, was most abundant in spring at the preserve site and in summer at the agriculture and pasture sites. Oropsylla montana was most abundant in winter at the pasture site and on adult squirrels. Echidnophaga gallinacea was most abundant in fall on juvenile squirrels at the preserve site. All three flea species we encountered are known to be potential vectors of Y. pestis. Future efforts to predict flea species occurrence and abundance (and plague risk) at sites of concern should consider seasonal microclimatic conditions and the potential influence of human land use practices.