Life-history traits and habitat availability shape genomic diversity in birds: implications for conservation.

More than 25% of species assessed by the International Union for Conservation of Nature (IUCN) are threatened with extinction. Understanding how environmental and biological processes have shaped genomic diversity may inform management practices. Using 68 extant avian species, we parsed the effects of habitat availability and life-history traits on genomic diversity over time to provide a baseline for conservation efforts. We used published whole-genome sequence data to estimate overall genomic diversity as indicated by historical long-term effective population sizes (Ne) and current genomic variability (H), then used environmental niche modelling to estimate Pleistocene habitat dynamics for each species. We found that Ne and H were positively correlated with habitat availability and related to key life-history traits (body mass and diet), suggesting the latter contribute to the overall genomic variation. We found that H decreased with increasing species extinction risk, suggesting that H may serve as a leading indicator of demographic trends related to formal IUCN conservation status in birds. Our analyses illustrate that genome-wide summary statistics estimated from sequence data reflect meaningful ecological attributes relevant to species conservation.

Island area, body size and demographic history shape genomic diversity in Darwin's finches and related tanagers.

Genomic diversity is the evolutionary foundation for adaptation to environmental change and thus is essential to consider in conservation planning. Island species are ideal for investigating the evolutionary drivers of genomic diversity, in part because of the potential for biological replicates. Here, we use genome data from 180 individuals spread among 27 island populations from 17 avian species to study the effects of island area, body size, demographic history and conservation status on contemporary genomic diversity. Our study expands earlier work on a small number of neutral loci to the entire genome and from a few species to many. We find significant positive correlation between island size and genomic diversity, a significant negative correlation between body size and genomic diversity, and that historical population declines significantly reduced contemporary genomic diversity. Our study shows that island size is the key factor in determining genomic diversity, indicating that habitat conservation is key to maintaining adaptive potential in the face of global environmental change. We found that threatened species generally had a significantly smaller values of Watterson's theta (θW  = 4Ne µ) compared to nonthreatened species, suggesting that θW may be useful as a conservation indicator for at-risk species. Overall, these findings (a) provide biological insights into how genomic diversity scales with ecological, morphological and demographic factors; and (b) illustrate how population genomic data can be leveraged to better inform conservation efforts.

Prescribed fire and partial overstory removal alter an acorn-rodent conditional mutualism.

In eastern North America, oak (Quercus) regeneration failure has spurred management using silvicultural approaches better aligned with the autecology of oaks. In particular, shelterwood harvests can create favorable intermediate light conditions for oak establishment and prescribed fire is predicted (by the oak-fire hypothesis) to favor oak regeneration. These approaches substantially modify forest structure and may affect crucial trophic interactions including the conditional mutualism between oaks and granivorous rodents that scatterhoard acorns, which shifts along a continuum from antagonistic to mutualistic depending on external factors. We investigated how overwinter survival and dispersal of northern red oak (Quercus rubra) acorns were influenced by location within or outside of group shelterwood harvests (small canopy gaps created throughout an intact forest stand) with and without prescribed fire. We conducted two concurrent experiments to test (1) dispersal and survival of acorns presented on the forest floor and (2) acorn pilferage rates from caches that mimic squirrel scatterhoards in shelterwood gap/group interiors, edges, and the uncut forest matrix in burned and unburned forest stands. In both experiments, acorn survival was generally higher in burned than unburned stands. Acorn survival from forest floor presentations was higher in the unharvested forest matrix than harvest gap interiors; however, there was no effect of proximity to harvest gaps on survival of cached acorns. Survival of cached acorns was associated with understory vegetative cover (-), coarse woody debris cover (-), and distance to nearest tree (+), but uncorrelated with canopy cover above the cache. Our results suggest that reduced understory cover following prescribed fire may increase perceived habitat riskiness for granivores resulting in higher acorn survival up to 2 yr post-fire. These findings unify the oak-fire and oak-granivore conditional mutualism hypotheses, and suggest that the environmental conditions following prescribed fire and group shelterwood harvests may shift the oak-granivore conditional mutualism in a direction beneficial for oak regeneration.

Effects of forest management on vertebrates: synthesizing two decades of data from hardwood forests in Missouri, USA.

Modern forest management seeks to balance multiple social, economic, and ecological goals. Different management approaches create different types of disturbances in a forest ecosystem and thus also differ in their impacts on plants, animals, and insects. Understanding these impacts is important for conservation of forest ecosystem function, but challenging due to the large spatial and temporal scale over which management occurs. Most past research has focused on relatively small areas, short time scales, and/or a small number of species. To address this, we examined the effects of two common silvicultural systems (even and uneven aged) on abundance and richness of three vertebrate taxa (birds, small mammals, and herpetofauna) over a two-decade period in a temperate hardwood forest in Missouri, USA. The two systems removed a similar amount of biomass overall, but differed in the intensity, number, and configuration of harvests applied. We found that vertebrate population responses varied by taxa, occurred at multiple spatial scales, and were concentrated in the period following the first harvest entry. Birds generally had the largest changes in relative abundance, both positive and negative, following management. Small mammals and reptiles had smaller, but generally positive, responses; amphibians were mixed. Bird species tended to respond in the same way to both silvicultural systems, while small mammals and herpetofauna did not respond consistently. Thus, for birds, the total amount of harvest disturbance across the landscape drives population responses, while for others the size and configuration of individual harvests is likely more important. Synthesizing results across the vertebrate community at large spatial and temporal scales allows managers to better understand trade-offs when making decisions that will affect wildlife in contrasting ways.

Integrated distance sampling models for simple point counts.

Point counts (PCs) are widely used in biodiversity surveys but, despite numerous advantages, simple PCs suffer from several problems: detectability, and therefore abundance, is unknown; systematic spatiotemporal variation in detectability yields biased inferences, and unknown survey area prevents formal density estimation and scaling-up to the landscape level. We introduce integrated distance sampling (IDS) models that combine distance sampling (DS) with simple PC or detection/nondetection (DND) data to capitalize on the strengths and mitigate the weaknesses of each data type. Key to IDS models is the view of simple PC and DND data as aggregations of latent DS surveys that observe the same underlying density process. This enables the estimation of separate detection functions, along with distinct covariate effects, for all data types. Additional information from repeat or time-removal surveys, or variable survey duration, enables the separate estimation of the availability and perceptibility components of detectability with DS and PC data. IDS models reconcile spatial and temporal mismatches among data sets and solve the above-mentioned problems of simple PC and DND data. To fit IDS models, we provide JAGS code and the new "IDS()" function in the R package unmarked. Extant citizen-science data generally lack the information necessary to adjust for detection biases, but IDS models address this shortcoming, thus greatly extending the utility and reach of these data. In addition, they enable formal density estimation in hybrid designs, which efficiently combine DS with distance-free, point-based PC or DND surveys. We believe that IDS models have considerable scope in ecology, management, and monitoring.

Spatial overlap of gray wolves and ungulate prey changes seasonally corresponding to prey migration.

BACKGROUND: Prey are more vulnerable during migration due to decreased familiarity with their surroundings and spatially concentrated movements. Predators may respond to increased prey vulnerability by shifting their ranges to match prey. Moose (Alces alces) and white-tailed deer (Odocoileus virginianus) are primary gray wolf (Canis lupus) prey and important subsistence species for Indigenous communities. We hypothesized wolves would increase use of ungulate migration corridors during migrations and predicted wolf distributions would overlap primary available prey. METHODS: We examined seasonal gray wolf, moose, and white-tailed deer movements on and near the Grand Portage Indian Reservation, Minnesota, USA. We analyzed GPS collar data during 2012-2021 using Brownian bridge movement models (BBMM) in Migration Mapper and mechanistic range shift analysis (MRSA) to estimate individual- and population-level occurrence distributions and determine the status and timing of range shifts. We estimated proportional overlap of wolf distributions with moose and deer distributions and tested for differences among seasons, prey populations, and wolf sex and pack affiliations. RESULTS: We identified a single migration corridor through which white-tailed deer synchronously departed in April and returned in October-November. Gray wolf distributions overlapped the deer migration corridor similarly year-round, but wolves altered within-range distributions seasonally corresponding to prey distributions. Seasonal wolf distributions had the greatest overlap with deer during fall migration (10 October-28 November) and greatest overlap with moose during summer (3 May-9 October). CONCLUSIONS: Gray wolves did not increase their use of the white-tailed deer migration corridor but altered distributions within their territories in response to seasonal prey distributions. Greater overlap of wolves and white-tailed deer in fall may be due to greater predation success facilitated by asynchronous deer migration movements. Greater summer overlap between wolves and moose may be linked to moose calf vulnerability, American beaver (Castor canadensis) co-occurrence, and reduced deer abundance associated with migration. Our results suggest increases in predation pressure on deer in fall and moose in summer, which can inform Indigenous conservation efforts. We observed seasonal plasticity of wolf distributions suggestive of prey switching; that wolves did not exhibit migratory coupling was likely due to spatial constraints resulting from territoriality.

Effects of lethal management on gray wolf pack persistence and reproduction in Wisconsin, USA.

Direct human-caused mortality accounts for about half of all large mammal mortality in North America. For social species like gray wolves (Canis lupus), the death of pack members can disrupt pack structure and cause pack dissolution, and mortality of breeding adults or wolves during reproduction and pup-rearing can decrease pup recruitment. We estimated minimum and maximum probability of wolf pack persistence in Wisconsin, USA, during biological years (15 April-14 April) 2011-2019 and evaluated the influence of pack size and legal harvest mortality on pack persistence during 2012-2014. Harvests comprised 75-161 mortalities within 194 monitored packs during 2012-2014, with 56-74% of packs having no wolves harvested each year. As an index of reproduction during 2013-2019, we also estimated the proportion of packs where pups responded to howl surveys. We evaluated the influence of pack size, legal harvest, and agency removal on reproduction during 2013-2015. Annual maximum pack persistence probability was uniformly high (0.95-1.00), and annual minimum pack persistence probability ranged from 0.86-0.98 with a possible decline during years of harvest. Reproduction was similar in years following harvest and agency removal (2013-2015, pup response = 0.27-0.40), and years without harvest or agency removal the year prior (2016-2019, pup response = 0.28-0.66). Pack size had a positive effect on pack persistence and reproduction. Total number of wolf mortalities and number of adult male and females removed did not influence pack persistence or reproduction. We suggest that low per-pack mortality, timing of harvest and agency removal, and harvest characteristics during 2012-2014 supported stable pack persistence and reproduction.

Factors influencing frequency and severity of human-American black bear conflicts in New York, USA.

Free-ranging large carnivores are involved in human-wildlife conflicts which can result in economic costs. Understanding factors that lead to human-wildlife conflicts is important to mitigate these negative effects and facilitate human-carnivore coexistence. We used a human-American black bear (Ursus americanus) conflict database maintained by the New York State Department of Environmental Conservation to determine whether drought, conflicts within the Adirondack and Catskill Parks as compared to outside of these parks, mild severity (Class 3) conflicts early in the year (April-June), and bear harvest in the previous year (as an index of bear abundance), were associated with greater frequency of high or moderate severity (Class 1-2) conflicts later in the year (July-September) across New York, USA. During 2006-2019, we obtained 3,782 mild severity conflict records early in the year, and 1,042 high or moderate severity records later in the year. We found that a one standard deviation increase in the cumulative precipitation difference from mean early in the year (about 7.59 cm) coincided with a 20% decrease in conflicts, and that Wildlife Management Units (WMUs) within the parks were predicted to have 5.61 times as many high or moderate severity conflicts as WMUs outside the parks. We also found that a one standard deviation increase in the frequency of mild severity conflicts (equivalent to 5.68 conflicts) early in the year coincided with an increase in the frequency of high or moderate severity conflicts in a WMU later in the year by 49%, while a one standard deviation increase in the bear abundance index in the previous year (0.14 bears/10 km2) coincided with a 23% increase in high or moderate severity conflicts. To reduce the frequency and severity of conflicts to facilitate human-black bear coexistence, we recommend the following measures to be taken in place consistently and build over time in local communities: (i) further reducing black bear access to anthropogenic foods and other attractants, (ii) non-lethal measures including bear-resistant waste management, (iii) electric fencing, and (iv) modifying placement or configuration of field crops.

Spatial and temporal niche overlap of aardwolves and aardvarks in Serengeti National Park, Tanzania.

Species interactions can influence species distributions, but mechanisms mitigating competition or facilitating positive interactions between ecologically similar species are often poorly understood. Aardwolves (Proteles cristata) and aardvarks (Orycteropus afer) are nocturnal, insectivorous mammals that co-occur in eastern and southern Africa, and knowledge of these species is largely limited to their nutritional biology. We used aardwolf and aardvark detections from 105 remote cameras during 2016-2018 to assess their spatial and temporal niche overlap in the grasslands of Serengeti National Park, Tanzania. Using a multispecies occupancy model, we identified a positive interaction between occupancy probabilities for aardwolves and aardvarks. Slope, proportion of grassland and termite mound density did not affect the occupancy probabilities of either species. The probability of aardwolf, but not aardvark, occupancy increased with distance to permanent water sources, which may relate to predation risk avoidance. Diel activity overlap between aardwolves and aardvarks was high during wet and dry seasons, with both species being largely nocturnal. Aardwolves and aardvarks have an important ecological role as termite consumers, and aardvarks are suggested to be ecosystem engineers. Our results contribute to a better understanding of the spatial and temporal niche of insectivores like aardwolves and aardvarks, suggesting high spatial and temporal niche overlap in which commensalism occurs, whereby aardwolves benefit from aardvark presence through increased food accessibility.

Development and validation of an eDNA protocol for monitoring endemic Asian spiny frogs in the Himalayan region of Pakistan.

Wildlife monitoring programs are instrumental for the assessment of species, habitat status, and for the management of factors affecting them. This is particularly important for species found in freshwater ecosystems, such as amphibians, as they have higher estimated extinction rates than terrestrial species. We developed and validated two species-specific environmental DNA (eDNA) protocols and applied them in the field to detect the Hazara Torrent Frog (Allopaa hazarensis) and Murree Hills Frog (Nanorana vicina). Additionally, we compared eDNA surveys with visual encounter surveys and estimated site occupancy. eDNA surveys resulted in higher occurrence probabilities for both A. hazarensis and N. vicina than for visual encounter surveys. Detection probability using eDNA was greater for both species, particularly for A. hazarensis. The top-ranked detection model for visual encounter surveys included effects of both year and temperature on both species, and the top-ranked occupancy model included effects of elevation and year. The top-ranked detection model for eDNA data was the null model, and the top-ranked occupancy model included effects of elevation, year, and wetland type. To our knowledge, this is the first time an eDNA survey has been used to monitor amphibian species in the Himalayan region.

Maternal carryover, winter severity, and brown bear abundance relate to elk demographics.

Ungulates are key components of ecosystems due to their effects on lower trophic levels, role as prey, and value for recreational and subsistence harvests. Understanding factors that drive ungulate population dynamics can inform protection of important habitat and successful management of populations. To ascertain correlates of ungulate population dynamics, we evaluated the effects of five non-exclusive hypotheses on ungulate abundance and recruitment: winter severity, spring nutritional limitation (spring bottleneck), summer-autumn maternal condition carryover, predation, and timber harvest. We used weather, reconstructed brown bear (Ursus arctos) abundance, and timber harvest data to estimate support for these hypotheses on early calf recruitment (calves per 100 adult females in July-August) and population counts of Roosevelt elk (Cervus canadensis roosevelti) on Afognak and Raspberry islands, Alaska, USA, 1958-2020. Increasing winter temperatures positively affected elk abundance, supporting the winter severity hypothesis, while a later first fall freeze had a positive effect on elk recruitment, supporting the maternal carry-over hypothesis. Increased brown bear abundance was negatively associated with elk recruitment, supporting the predation hypothesis. Recruitment was unaffected by spring climate conditions or timber harvest. Severe winter weather likely increased elk energy deficits, reducing elk survival and subsequent abundance in the following year. Colder and shorter falls likely reduced late-season forage, resulting in poor maternal condition which limited elk recruitment more than winter severity or late-winter nutritional bottlenecks. Our results additionally demonstrated potential negative effects of brown bears on elk recruitment. The apparent long-term decline in elk recruitment did not result in a decline of abundance, which suggests that less severe winters may increase elk survival and counteract the potential effects of predation on elk abundance.

Variable effects of wolves on niche breadth and density of intraguild competitors.

The parallel niche release hypothesis (PNR) indicates that reduced competition with dominant competitors results in greater density and niche breadth of subordinate competitors and which may support an adaptive advantage.We assessed support for the PNR by evaluating relationships between variation in niche breadth and intra- and interspecific density (an index of competition) of wolves (Canis lupus) coyotes (C. latrans), and bobcats (Lynx rufus).We estimated population density (wolf track surveys, coyote howl surveys, and bobcat hair snare surveys) and variability in space use (50% core autocorrelated kernel density home range estimators), temporal activity (hourly and overnight speed), and dietary (isotopic δ13C and δ15N) niche breadth of each species across three areas of varying wolf density in the Upper Peninsula of Michigan, USA, 2010-2019.Densities of wolves and coyotes were inversely related, and increased variability in space use, temporal activity, and dietary niche breadth of coyotes was associated with increased coyote density and decreased wolf density supporting the PNR. Variability in space use and temporal activity of wolves and dietary niche breadth of bobcats also increased with increased intraspecific density supporting the PNR.Through demonstrating decreased competition between wolves and coyotes and increased coyote niche breadth and density, our study provides multidimensional support for the PNR. Knowledge of the relationship between niche breadth and population density can inform our understanding of the role of competition in shaping the realized niche of species.

Quantifying anthropogenic wolf mortality in relation to hunting regulations and landscape attributes across North America.

Understanding the types and magnitude of human-caused mortality is essential for maintaining viable large carnivore populations. We used a database of cause-specific mortality to examine how hunting regulations and landscape configurations influenced human-caused mortality of North American gray wolves (Canis lupus). Our dataset included 21 studies that monitored the fates of 3564 wolves and reported 1442 mortalities. Human-caused mortality accounted for 61% of mortality overall, with 23% due to illegal harvest, 16% due to legal harvest, and 12% the result of management removal. The overall proportion of anthropogenic wolf mortality was lowest in areas with an open hunting season compared to areas with a closed hunting season or mixed hunting regulations, suggesting that harvest mortality was neither fully additive nor compensatory. Proportion of mortality from management removal was reduced in areas with an open hunting season, suggesting that legal harvest may reduce human-wolf conflicts or alternatively that areas with legal harvest have less potential for management removals (e.g., less livestock depredation). Proportion of natural habitat was negatively correlated with the proportion of anthropogenic and illegal harvest mortality. Additionally, the proportion of mortality due to illegal harvest increased with greater natural habitat fragmentation. The observed association between large patches of natural habitat and reductions in several sources of anthropogenic wolf mortality reiterate the importance of habitat preservation to maintain wolf populations. Furthermore, effective management of wolf populations via implementation of harvest may reduce conflict with humans. Effective wolf conservation will depend on holistic strategies that integrate ecological and socioeconomic factors to facilitate their long-term coexistence with humans.

Factors influencing lion movements and habitat use in the western Serengeti ecosystem, Tanzania.

Protected areas that restrict human activities can enhance wildlife habitat quality. Efficacy of protected areas can be improved with increased protection from illegal activities and presence of buffer protected areas that surround a core protected area. Habitat value of protected areas also can be affected by seasonal variation in anthropogenic pressures. We examined seasonal space use by African lions (Panthera leo) within a core protected area, Serengeti National Park, Tanzania, and surrounding buffer protected areas with varying protection strengths. We used lion locations in logistic regression models during wet and dry seasons to estimate probability of use in relation to protection strength, distance to protected area edge, human and livestock density, distance to roads and rivers, and land cover. Lions used strongly protected buffer areas over the core protected area and unprotected areas, and moved away from protected area boundaries toward the core protected area when buffer protected areas had less protection. Lions avoided high livestock density in the wet season and high human density in the dry season. Increased strength of protection can decrease edge effects on buffer areas and help maintain habitat quality of core protected areas for lions and other wildlife species.

Reducing Baylisascaris procyonis roundworm larvae in raccoon latrines.

Baylisascaris procyonis roundworms, a parasite of raccoons, can infect humans, sometimes fatally. Parasite eggs can remain viable in raccoon latrines for years. To develop a management technique for parasite eggs, we tested anthelmintic baiting. The prevalence of eggs decreased at latrines, and larval infections decreased among intermediate hosts, indicating that baiting is effective.

A penalized likelihood for multispecies occupancy models improves predictions of species interactions.

Multispecies occupancy models estimate dependence among multiple species of interest from patterns of co-occurrence, but problems associated with separation and boundary estimates can lead to unreasonably large estimates of parameters and associated standard errors when species are rarely observed at the same site or when data are sparse. In this paper, we overcome these issues by implementing a penalized likelihood, which introduces a small bias in parameter estimates in exchange for a potentially large reduction in variance. We compare parameter estimates obtained from both penalized and unpenalized multispecies occupancy models fit to simulated data that exhibit various degrees of separation and to a real-word data set of bird surveys with little apparent overlap between potentially interacting species. Our simulation results demonstrate that penalized multispecies occupancy models did not exhibit boundary estimates and produced lower bias, lower mean squared error, and improved inference relative to unpenalized models. When applied to real-world data, our penalized multispecies occupancy model constrained boundary estimates and allowed for meaningful inference related to the interactions of two species of conservation concern. To facilitate the use of our penalized multispecies occupancy model, the techniques demonstrated in this paper have been integrated into the unmarked package in R programing language.

Landscape transformations produce favorable roosting conditions for turkey vultures and black vultures.

Recent increases in turkey vulture (Cathartes aura) and black vulture (Coragyps atratus) populations in North America have been attributed in part to their success adapting to human-modified landscapes. However, the capacity for such landscapes to generate favorable roosting conditions for these species has not been thoroughly investigated. We assessed the role of anthropogenic and natural landscape elements on roosting habitat selection of 11 black and 7 turkey vultures in coastal South Carolina, USA using a GPS satellite transmitter dataset derived from previous research. Our dataset spanned 2006-2012 and contained data from 7916 nights of roosting. Landscape fragmentation, as measured by land cover richness, influenced roosting probability for both species in all seasons, showing either a positive relationship or peaking at intermediate values. Roosting probability of turkey vultures was maximized at intermediate road densities in three of four seasons, and black vultures showed a positive relationship with roads in fall, but no relationship throughout the rest of the year. Roosting probability of both species declined with increasing high density urban cover throughout most of the year. We suggest that landscape transformations lead to favorable roosting conditions for turkey vultures and black vultures, which has likely contributed to their recent proliferations across much of the Western Hemisphere.

American martens use vigilance and short-term avoidance to navigate a landscape of fear from fishers at artificial scavenging sites.

Where two sympatric species compete for the same resource and one species is dominant, there is potential for the subordinate species to be affected through interference competition or energetic costs of avoiding predation. Fishers (Pekania pennanti) and American martens (Martes americana) often have high niche overlap, but fishers are considered dominant and potentially limiting to martens. We observed presence and vigilance of fishers and martens at winter carcass sites using remote cameras in Michigan, USA, to test the hypothesis that interference competition from fishers creates a landscape of fear for martens. Within winters, fishers co-occupied 78-88% of sites occupied by martens, and martens co-occupied 79-88% of sites occupied by fishers. Fishers displaced martens from carcasses during 21 of 6117 marten visits, while martens displaced fishers during 0 of 1359 fisher visits. Martens did not alter diel activity in response to fisher use of sites. Martens allocated 37% of time to vigilance compared to 23% for fishers, and martens increased vigilance up to 8% at sites previously visited by fishers. Fishers increased vigilance by up to 8% at sites previously visited by martens. Our results indicate that fishers were dominant over martens, and martens had greater baseline perception of risk than fishers. However, fishers appeared to be also affected as the dominant competitor by putting effort into scanning for martens. Both species appeared widespread and common in our study area, but there was no evidence that fishers spatially or temporally excluded martens from scavenging at carcasses other than occasional short-term displacement when a fisher was present. Instead, martens appeared to mitigate risk from fishers by using vigilance and short-term avoidance. Multiple short-term anti-predator behaviors within a landscape of fear may facilitate coexistence among carnivore species.

Marginal support for a trophic cascade among sympatric canids in peripheral wolf range.

Trophic cascades reportedly structure ecological communities through indirect species interactions. Though the predator-herbivore-autotroph relationship has received much attention, mechanistic evidence supporting intraguild trophic cascades is rare. We established 348 remote camera sites (1 August-5 September 2019) across seven study areas of varying wolf (Canis lupus) density including one study area where wolves were absent in northern Michigan, USA. Using multi-species occupancy modeling at species-relevant spatial scales, we evaluated the hypothesis that increased wolf occurrence suppresses coyote (C. latrans) occurrence with corresponding increased red fox (Vulpes vulpes) occurrence mediated by land cover edge density, human presence, and temporal partitioning. Remote cameras recorded >600,000 images and included 6,370, 10,137, and 4,876 detections of wolves, coyotes, and foxes, respectively. Fox occupancy probability was more than three times as high (0.29) at camera sites where wolves were present, relative to sites wolves were absent (0.09). Pairwise species interactions supported expected size-based dominance patterns among canids and insignificant effects were directionally consistent with reported reduced strength of top-down effects in peripheral wolf range. Increased edge density also increased co-occurrence of coyote and wolves, likely a function of increased prey availability and refugia for coyotes. Though foxes occurred in spatial proximity to wolves, competition was limited by greater temporal partitioning than observed between coyotes and foxes that were spatially segregated. Collectively, our results provide marginal support for the reported trophic cascade among wolves, coyotes, and foxes wherein top-down effects may be reduced near the edge of current wolf distributions. As predators continue to recolonize portions of their historic range, knowledge of the effects on intraguild predators has implications for species management and predicting prey population responses.

Responses of sympatric canids to human development revealed through citizen science.

Measuring wildlife responses to anthropogenic activities often requires long-term, large-scale datasets that are difficult to collect. This is particularly true for rare or cryptic species, which includes many mammalian carnivores. Citizen science, in which members of the public participate in scientific work, can facilitate collection of large datasets while increasing public awareness of wildlife research and conservation. Hunters provide unique benefits for citizen science given their knowledge and interest in outdoor activities. We examined how anthropogenic changes to land cover impacted relative abundance of two sympatric canids, coyote (Canis latrans), and red fox (Vulpes vulpes) at a large spatial scale. In order to assess how land cover affected canids at this scale, we used citizen science data from bow hunter sighting logs collected throughout New York State, USA, during 2004-2017. We found that the two species had contrasting responses to development, with red foxes positively correlated and coyotes negatively correlated with the percentage of low-density development. Red foxes also responded positively to agriculture, but less so when agricultural habitat was fragmented. Agriculture provides food and denning resources for red foxes, whereas coyotes may select forested areas for denning. Though coyotes and red foxes compete in areas of sympatry, we did not find a relationship between species abundance, likely a consequence of the coarse spatial resolution used. Red foxes may be able to coexist with coyotes by altering their diets and habitat use, or by maintaining territories in small areas between coyote territories. Our study shows the value of citizen science, and particularly hunters, in collection of long-term data across large areas (i.e., the entire state of New York) that otherwise would unlikely be obtained.