Using global remote camera data of a solitary species complex to evaluate the drivers of group formation.

The social system of animals involves a complex interplay between physiology, natural history, and the environment. Long relied upon discrete categorizations of "social" and "solitary" inhibit our capacity to understand species and their interactions with the world around them. Here, we use a globally distributed camera trapping dataset to test the drivers of aggregating into groups in a species complex (martens and relatives, family Mustelidae, Order Carnivora) assumed to be obligately solitary. We use a simple quantification, the probability of being detected in a group, that was applied across our globally derived camera trap dataset. Using a series of binomial generalized mixed-effects models applied to a dataset of 16,483 independent detections across 17 countries on four continents we test explicit hypotheses about potential drivers of group formation. We observe a wide range of probabilities of being detected in groups within the solitary model system, with the probability of aggregating in groups varying by more than an order of magnitude. We demonstrate that a species' context-dependent proclivity toward aggregating in groups is underpinned by a range of resource-related factors, primarily the distribution of resources, with increasing patchiness of resources facilitating group formation, as well as interactions between environmental conditions (resource constancy/winter severity) and physiology (energy storage capabilities). The wide variation in propensities to aggregate with conspecifics observed here highlights how continued failure to recognize complexities in the social behaviors of apparently solitary species limits our understanding not only of the individual species but also the causes and consequences of group formation.

Counting the Capital's cats: Estimating drivers of abundance of free-roaming cats with a novel hierarchical model.

Free-roaming cats are a conservation concern in many areas but identifying their impacts and developing mitigation strategies requires a robust understanding of their distribution and density patterns. Urban and residential areas may be especially relevant in this process because free-roaming cats are abundant in these anthropogenic landscapes. Here, we estimate the occupancy and density of free-roaming cats in Washington D.C. and relate these metrics to known landscape and social factors. We conducted an extended camera trap survey of public and private spaces across D.C. and analyzed data collected from 1483 camera deployments from 2018 to 2020. We estimated citywide cat distribution by fitting hierarchical occupancy models and further estimated cat abundance using a novel random thinning spatial capture-recapture model that allows for the use of photos that can and cannot be identified to individual. Within this model, we utilized individual covariates that provided identity exclusions between photos of unidentifiable cats with inconsistent coat patterns, thus increasing the precision of abundance estimates. This combined model also allowed for unbiased estimation of density when animals cannot be identified to individual at the same rate as for free-roaming cats whose identifiability depended on their coat characteristics. Cat occupancy and abundance declined with increasing distance from residential areas, an effect that was more pronounced in wealthier neighborhoods. There was noteworthy absence of cats detected in larger public spaces and forests. Realized densities ranged from 0.02 to 1.75 cats/ha in sampled areas, resulting in a district-wide estimate of ~7296 free-roaming cats. Ninety percent of cat detections lacked collars and nearly 35% of known individuals were ear-tipped, indicative of district Trap-Neuter-Return (TNR) programs. These results suggest that we mainly sampled and estimated the unowned cat subpopulation, such that indoor/outdoor housecats were not well represented. The precise estimation of cat population densities is difficult due to the varied behavior of subpopulations within free-roaming cat populations (housecats, stray and feral cats), but our methods provide a first step in establishing citywide baselines to inform data-driven management plans for free-roaming cats in urban environments.

High Prevalence of Sarcocystis in a Collapsed Black Rat (Rattus rattus) Population from the Florida Keys.

We collected and screened black rats (Rattus rattus) in Key Largo, Florida, USA, to determine the potential role of disease or parasites in the collapse of the local population. Rats appeared healthy, but 94% (n=15/16) tested positive for Sarcocystis sp. The partial 18S rRNA gene sequence was 98.7-99.7% similar to a strain of Sarcocystis zuoi that is now considered a strain of the newly described Sarcocystis kani within the larger S. zuoi species complex that contains numerous new species. These Sarcocystis spp. use Asian snakes as definitive hosts and rodents, shrews, or tree shrews as intermediate hosts. Pythons are the definitive host for several Sarcocystis spp. in Asia, including a related parasite (Sarcocystis singaporensis) that has been used as a biologic control agent for Rattus spp. in southeast Asia. It is probable that increasing numbers of invasive Burmese pythons (Python bivittatus) in the Florida Keys are contributing to the spread of this parasite in addition to imposing predation pressure on both rodents and native snakes. As such, further surveillance and molecular and morphologic characterization of parasites from rodents and snakes in south Florida should be prioritized.

Mammal responses to global changes in human activity vary by trophic group and landscape.

Wildlife must adapt to human presence to survive in the Anthropocene, so it is critical to understand species responses to humans in different contexts. We used camera trapping as a lens to view mammal responses to changes in human activity during the COVID-19 pandemic. Across 163 species sampled in 102 projects around the world, changes in the amount and timing of animal activity varied widely. Under higher human activity, mammals were less active in undeveloped areas but unexpectedly more active in developed areas while exhibiting greater nocturnality. Carnivores were most sensitive, showing the strongest decreases in activity and greatest increases in nocturnality. Wildlife managers must consider how habituation and uneven sensitivity across species may cause fundamental differences in human-wildlife interactions along gradients of human influence.

SNAPSHOT USA 2021: A third coordinated national camera trap survey of the United States.

SNAPSHOT USA is a multicontributor, long-term camera trap survey designed to survey mammals across the United States. Participants are recruited through community networks and directly through a website application (https://www.snapshot-usa.org/). The growing Snapshot dataset is useful, for example, for tracking wildlife population responses to land use, land cover, and climate changes across spatial and temporal scales. Here we present the SNAPSHOT USA 2021 dataset, the third national camera trap survey across the US. Data were collected across 109 camera trap arrays and included 1711 camera sites. The total effort equaled 71,519 camera trap nights and resulted in 172,507 sequences of animal observations. Sampling effort varied among camera trap arrays, with a minimum of 126 camera trap nights, a maximum of 3355 nights, a median 546 nights, and a mean 656 ± 431 nights. This third dataset comprises 51 camera trap arrays that were surveyed during 2019, 2020, and 2021, along with 71 camera trap arrays that were surveyed in 2020 and 2021. All raw data and accompanying metadata are stored on Wildlife Insights (https://www.wildlifeinsights.org/), and are publicly available upon acceptance of the data papers. SNAPSHOT USA aims to sample multiple ecoregions in the United States with adequate representation of each ecoregion according to its relative size. Currently, the relative density of camera trap arrays varies by an order of magnitude for the various ecoregions (0.22-5.9 arrays per 100,000 km2), emphasizing the need to increase sampling effort by further recruiting and retaining contributors. There are no copyright restrictions on these data. We request that authors cite this paper when using these data, or a subset of these data, for publication. Any use of trade, firm, or product names is for descriptive purposes only and does not imply endorsement by the US Government.

Raccoon Roundworm (Baylisascaris procyonis) Undetected in Endangered Key Largo Woodrat (Neotoma floridana smalli) Endemic Range.

Raccoon roundworm (Baylisascaris procyonis) negatively affects woodrat (Neotoma spp.) populations but is not known to occur in the endemic range of endangered Key Largo woodrats (Neotoma floridana smalli). Rectal swabs from 23 raccoons (Procyon lotor) in Key Largo were screened for raccoon roundworm by PCR. All tests were negative, suggesting continued absence.

Human recreation impacts seasonal activity and occupancy of American black bears (Ursus americanus) across the anthropogenic-wildland interface.

Protected areas serve an important role in wildlife conservation, yet most wildlife occur outside these areas, subject to varying degrees of human disturbance. In the Upper Peninsula of Michigan, American black bears (Ursus americanus), a highly mobile, opportunistic species, are common despite an extensive outdoor recreation industry with the potential to affect black bear spatial and temporal activity. We investigated how environmental and anthropogenic factors influence black bear occupancy, detection, and diel activity patterns across the anthropogenic-wildland interface before and after hibernation. Using 30 camera traps deployed across a rural-wildland interface, we captured black bears at 23 camera sites (~ 77%), which exhibited co-occurrence with humans at 10 sites (~ 33%), revealing that human presence and human population density exert negative effects on black bear seasonal occupancy. Bears were more nocturnal during the hunting season, before hibernation. Human recreational activity increased ~ 38% after hibernation, but bear diurnal activity also increased ~ 36%, except when cubs were present. Our results suggest bears prioritize avoiding humans spatially, rather than temporally, except during the hunting season and when cubs are present. Understanding black bear responses to human recreation patterns and environmental variation is essential for minimizing human-mediated disturbance, and fueling conservation efforts of large, charismatic carnivores.

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.

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.

Seasonal flow dynamics exacerbate overlap between artisanal fisheries and imperiled Ganges River dolphins.

Here we quantify the effects of artisanal fisheries on the ecology of a small cetacean, the Ganges River dolphin (Platanista gangetica gangetica, GRD), in a large river system of Nepal. We examine the size-classes of fisheries' catches, behavioural changes in GRD in response to fishing activities, and diel overlap between GRD and fishing activity. We observed high human exploitation rates (> 60% of the total catch per effort) of GRD-preferred prey sizes, indicating risks of high resource competition and dietary overlap, especially during the low water season when resource availability is reduced. Competitive interactions in the feeding niches during the low water season, plus temporal overlap between the peak exploitation and critical life-history events (e.g., reproduction), likely have ecological consequences. Furthermore, we detected 48% (95% CI 43-52%) increase in the chance of behavioural changes among dolphins exposed to anthropopressure (fishing activity), risking social behaviour impairment in exposed dolphins. The higher diel overlap and increased diel coefficient as the surveys progressed towards the monsoon season suggest temporal shifts in GRD socio-behavioural states and seasonal effects on resource partitioning, respectively. This work identifies drivers of small cetaceans-fisheries interactions and their consequences, and can be used to help reduce biologically significant fishing impacts on small cetaceans. Mitigation strategies, together with river sanctuary and distanced-based approaches, should be urgently included in a framework of ecosystem-based management.

Land management strategies can increase oil palm plantation use by some terrestrial mammals in Colombia.

While the conservation role of remaining natural habitats in anthropogenic landscapes is clear, the degree to which agricultural matrices impose limitations to animal use is not well understood, but vital to assess species' resilience to land use change. Using an occupancy framework, we evaluated how oil palm plantations affect the occurrence and habitat use of terrestrial mammals in the Colombian Llanos. Further, we evaluated the effect of undergrowth vegetation and proximity to forest on habitat use within plantations. Most species exhibited restricted distributions across the study area, especially in oil palm plantations. Habitat type strongly influenced habitat use of four of the 12 more widely distributed species with oil palm negatively affecting species such as capybara and naked-tailed armadillo. The remaining species showed no apparent effect of habitat type, but oil palm and forest use probabilities varied among species. Overall, generalist mesocarnivores, white-tailed deer, and giant anteater were more likely to use oil palm while the remaining species, including ocelot and lesser anteater, showed preferences for forest. Distance to nearest forest had mixed effects on species habitat use, while understory vegetation facilitated the presence of species using oil palm. Our findings suggest that allowing undergrowth vegetation inside plantations and maintaining nearby riparian corridors would increase the likelihood of terrestrial mammals' occurrence within oil palm landscapes.

Co-occurrence dynamics of endangered Lower Keys marsh rabbits and free-ranging domestic cats: Prey responses to an exotic predator removal program.

The Lower Keys marsh rabbit (Sylvilagus palustris hefneri) is one of many endangered endemic species of the Florida Keys. The main threats are habitat loss and fragmentation from sea-level rise, development, and habitat succession. Exotic predators such as free-ranging domestic cats (Felis catus) pose an additional threat to these endangered small mammals. Management strategies have focused on habitat restoration and exotic predator control. However, the effectiveness of predator removal and the effects of anthropogenic habitat modifications and restoration have not been evaluated. Between 2013 and 2015, we used camera traps to survey marsh rabbits and free-ranging cats at 84 sites in the National Key Deer Refuge, Big Pine Key, Florida, USA. We used dynamic occupancy models to determine factors associated with marsh rabbit occurrence, colonization, extinction, and the co-occurrence of marsh rabbits and cats during a period of predator removal. Rabbit occurrence was positively related to freshwater habitat and patch size, but was negatively related to the number of individual cats detected at each site. Furthermore, marsh rabbit colonization was negatively associated with relative increases in the number of individual cats at each site between survey years. Cat occurrence was negatively associated with increasing distance from human developments. The probability of cat site extinction was positively related to a 2-year trapping effort, indicating that predator removal reduced the cat population. Dynamic co-occurrence models suggested that cats and marsh rabbits co-occur less frequently than expected under random conditions, whereas co-detections were site and survey-specific. Rabbit site extinction and colonization were not strongly conditional on cat presence, but corresponded with a negative association. Our results suggest that while rabbits can colonize and persist at sites where cats occur, it is the number of individual cats at a site that more strongly influences rabbit occupancy and colonization. These findings indicate that continued predator management would likely benefit endangered small mammals as they recolonize restored habitats.

Estimating wildlife activity curves: comparison of methods and sample size.

Camera traps and radiotags commonly are used to estimate animal activity curves. However, little empirical evidence has been provided to validate whether they produce similar results. We compared activity curves from two common camera trapping techniques to those from radiotags with four species that varied substantially in size (~1 kg-~50 kg), diet (herbivore, omnivore, carnivore), and mode of activity (diurnal and crepuscular). Also, we sub-sampled photographs of each species with each camera trapping technique to determine the minimum sample size needed to maintain accuracy and precision of estimates. Camera trapping estimated greater activity during feeding times than radiotags in all but the carnivore, likely reflective of the close proximity of foods readily consumed by all species except the carnivore (i.e., corn bait or acorns). However, additional analyses still indicated both camera trapping methods produced relatively high overlap and correlation to radiotags. Regardless of species or camera trapping method, mean overlap increased and overlap error decreased rapidly as sample sizes increased until an asymptote near 100 detections which we therefore recommend as a minimum sample size. Researchers should acknowledge that camera traps and radiotags may estimate the same mode of activity but differ in their estimation of magnitude in activity peaks.

Do coyotes Canis latrans influence occupancy of prey in suburban forest fragments?

With the extirpation of apex predators from many North American systems, coyotes Canis latrans have become the de facto top predator and are ubiquitous members of most ecosystems. Keystone predators aid in maintaining ecosystem function by regulating the mammal community through direct predation and instilling the landscape of fear, yet the value of coyotes regulating systems to this capacity is understudied and likely variable across environments. Since coyotes are common in the Midwestern United States, we utilized camera traps and occupancy analyses to assess their role in regulating the distribution of mammalian herbivores in a fragmented suburban ecosystem. Forest cover was a strong positive predictor of white-tailed deer Odocoileus virginianus detection, while coyote occurrence had a negative effect. Coyotes exerted a negative effect on squirrel (Sciurus spp.) and eastern cottontail rabbit Sylvilagus floridanus occurrence, while urban cover was a positive predictor for the prey species' occurrence. These results suggest all 3 species behaviorally avoid coyotes whereby deer seek denser forest cover and squirrels and cottontails mitigate risk by increasing use of urban areas. Although previous studies reveal limited influence of coyote on the rest of the carnivore guild in suburban systems, we suggest coyotes play an important role in regulating the herbivorous mammals and hence may provide similar ecological benefits in urban/suburban forest fragments through trophic cascades. Furthermore, since hunting may not be allowed in urban and suburban habitats, coyotes might also serve as the primary regulator of nuisance species occurring at high abundance such as white-tailed deer and squirrels.

The role of fragmentation and landscape changes in the ecological release of common nest predators in the Neotropics.

Loss of large mammalian carnivores may allow smaller mesopredators to become abundant and threaten other community members. There is considerable debate about mesopredator release and the role that other potential factors such as landscape variables and human alterations to land cover lead to increased mesopredator abundance. We used camera traps to detect four mesopredators (tayra, Eira barbara; white-nosed coati, Nasua narica; northern raccoon, Procyon lotor; and common opossum, Didelphis opossum) in a biological corridor in Costa Rica to estimate habitat covariates that influenced the species' detection and occurrence. We selected these mesopredators because as semi-arboreal species they might be common nest predators, posing a serious threat to resident and migratory songbirds. Pineapple production had a pronounced positive effect on the detectability of tayras, while forest cover had a negative effect on the detection of coatis. This suggests that abundance might be elevated due to the availability of agricultural food resources and foraging activities are concentrated in forest fragments and pineapple edge habitats. Raccoon and opossum models exhibited little influence on detection from habitat covariates. Occurrence models did not suggest any significant factors influencing site use by nest predators, revealing that all four species are habitat generalists adapted to co-existing in human altered landscapes. Furthermore, fragmentation and land cover changes may predispose nesting birds, herpetofauna, and small mammals to heightened predation risk by mesopredators in the Neotropics.