Sub-zero temperatures and large-scale weather patterns induce tooth damage in Icelandic arctic foxes.

Tooth damage in carnivores can reflect shifts in both diet and feeding habits, and in large carnivores, it is associated with increased bone consumption. Variation in tooth condition in Icelandic arctic foxes, a mesocarnivore, was recorded from 854 individual foxes spanning 29 years. We hypothesized that annual climatic variations, which can influence food abundance and accessibility, will influence tooth condition by causing dietary shifts toward less edible prey. We examined tooth condition in relation to four climatic predictors: mean annual winter temperature, indices of both the El Niño anomaly and North Atlantic subpolar gyre (SPG), and the number of rain-on-snow days (ROS). We found unequivocal evidence for a strong effect of annual climate on tooth condition. Teeth of Icelandic foxes were in better condition when winter temperatures were higher, when the SPG was more positive, and when the number of ROS was low. We also found a substantial subregional effect with foxes from northeastern Iceland having lower tooth damage than those from two western sites. Contradicting our original hypothesis that foxes from northeastern Iceland, where foxes are known to scavenge on large mammal remains (e.g., sheep and horses), would show the highest tooth damage, we suggest that western coastal sites exhibited greater tooth damage because cold winter temperatures lowered the availability of seabirds, causing a shift in diet toward abrasive marine subsidies (e.g., bivalves) and frozen beach wrack. Our study shows that monitoring tooth breakage and wear can be a useful tool for evaluating the impact of climate on carnivore populations and that climate change may influence the condition and fitness of carnivores in complex and potentially conflicting ways.

Anthropogenic subsidies influence resource use during a mange epizootic in a desert coyote population.

Colonization of urban areas by synanthropic wildlife introduces novel and complex alterations to established ecological processes, including the emergence and spread of infectious diseases. Aggregation at urban resources can increase disease transfer, with wide-ranging species potentially infecting outlying populations. The garrison at the National Training Center, Fort Irwin, California, USA, was recently colonized by mange-infected coyotes (Canis latrans) that also use the surrounding Mojave Desert. This situation provided an ideal opportunity to examine the effects of urban resources on disease dynamics. We evaluated seasonal space use and determined the influence of anthropogenic subsidies, water sources, and prey density on urban resource selection. We found no difference in home range size between healthy and infected individuals, but infected residents had considerably more spatial overlap with one another than healthy residents. All coyotes selected for anthropogenic subsidies during all seasons, while infected coyotes seasonally selected for urban water sources, and healthy coyotes seasonally selected for urban areas with greater densities of natural prey. These results suggest that while all coyotes were selecting for anthropogenic subsidies, infected resident coyotes demonstrated a greater tolerance for other conspecifics, which could be facilitating the horizontal transfer of sarcoptic mange to non-resident coyotes. Conversely, healthy coyotes also selected for natural prey and healthy residents exhibited a lack of spatial overlap with other coyotes suggesting they were not reliant on anthropogenic subsidies and were maintaining territories. Understanding the association between urban wildlife, zoonotic diseases, and urban resources can be critical in determining effective responses for mitigating future epizootics.

Urbanization's influence on the distribution of mange in a carnivore revealed with multistate occupancy models.

Increasing urbanization and use of urban areas by synanthropic wildlife has increased human and domestic animal exposure to zoonotic diseases and exacerbated epizootics within wildlife populations. Consequently, there is a need to improve wildlife disease surveillance programs to rapidly detect outbreaks and refine inferences regarding spatiotemporal disease dynamics. Multistate occupancy models can address potential shortcomings in surveillance programs by accounting for imperfect detection and the misclassification of disease states. We used these models to explore the relationship between urbanization, slope, and the spatial distribution of sarcoptic mange in coyotes (Canis latrans) inhabiting Fort Irwin, California, USA. We deployed remote cameras across 180 sites within the desert surrounding the populated garrison and classified sites by mange presence or absence depending on whether a symptomatic or asymptomatic coyote was photographed. Coyotes selected flatter sites closer to the urban area with a high probability of use (0.845, 95% credible interval (CRI): 0.728, 0.944); site use decreased as the distance to urban areas increased (standardized [Formula: see text] = - 1.354, 95% CRI - 2.423, - 0.619). The probability of correctly classifying mange presence at a site also decreased further from the urban area and was probably related to the severity of mange infection. Severely infected coyotes, which were more readily identified as symptomatic, resided closer to the urban area and were most likely dependent on urban resources for survival; urban resources probably contributed to sustaining the disease. Multistate occupancy models represent a flexible framework for estimating the occurrence and spatial extent of observable infectious diseases, which can improve wildlife disease surveillance programs.

Elevated surface temperature depresses survival of banner-tailed kangaroo rats: will climate change cook a desert icon?

Modest increases in global temperature have been implicated in causing population extirpations and range shifts in taxa inhabiting colder environs and in ectotherms whose thermoregulation is more closely tied to environmental conditions. Many arid-adapted endotherms already experience conditions at their physiological limits, so it is conceivable that they could be similarly affected by warming temperatures. We explored how climatic variables might influence the apparent survival of the banner-tailed kangaroo rat (Dipodomys spectabilis), a rodent endemic to the Chihuahuan Desert of North America and renowned for its behavioral and physiological adaptations to arid environments. Relative variable weight, strength of variable relationships, and other criteria indicated that summer, diurnal land surface temperature (SD_LST) was the primary environmental driver of apparent survival in these arid-adapted rodents. Higher temperatures had a negative effect on apparent survival, which ranged from 0.15 (SE = 0.04) for subadults to 0.50 (SE = 0.07) for adults. Elevated SD_LST may negatively influence survival through multiple pathways, including increased water loss and energy expenditure that could lead to chronic stress and/or hyperthermia that could cause direct mortality. Land surface temperatures are predicted to increase by as much 6.5°C by 2099, reducing apparent survival of adults to ~0.15 in some regions of the species' range, possibly causing a shift in their distribution. The relationship between SD_LST and survival suggests a mechanism whereby physiological tolerances are exceeded resulting in a reduction to individual fitness that may ultimately cause a shift in the species' range over time.

Pleistocene-Holocene vicariance, not Anthropocene landscape change, explains the genetic structure of American black bear (Ursus americanus) populations in the American Southwest and northern Mexico.

The phylogeography of the American black bear (Ursus americanus) is characterized by isolation into glacial refugia, followed by population expansion and genetic admixture. Anthropogenic activities, including overharvest, habitat loss, and transportation infrastructure, have also influenced their landscape genetic structure. We describe the genetic structure of the American black bear in the American Southwest and northern Mexico and investigate how prehistoric and contemporary forces shaped genetic structure and influenced gene flow. Using a suite of microsatellites and a sample of 550 bears, we identified 14 subpopulations organized hierarchically following the distribution of ecoregions and mountain ranges containing black bear habitat. The pattern of subdivision we observed is more likely a product of postglacial habitat fragmentation during the Pleistocene and Holocene, rather than a consequence of contemporary anthropogenic barriers to movement during the Anthropocene. We used linear mixed-effects models to quantify the relationship between landscape resistance and genetic distance among individuals, which indicated that both isolation by resistance and geographic distance govern gene flow. Gene flow was highest among subpopulations occupying large tracts of contiguous habitat, was reduced among subpopulations in the Madrean Sky Island Archipelago, where montane habitat exists within a lowland matrix of arid lands, and was essentially nonexistent between two isolated subpopulations. We found significant asymmetric gene flow supporting the hypothesis that bears expanded northward from a Pleistocene refugium located in the American Southwest and northern Mexico and that major highways were not yet affecting gene flow. The potential vulnerability of the species to climate change, transportation infrastructure, and the US-Mexico border wall highlights conservation challenges and opportunities for binational collaboration.

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.

Double Allee effects and extinction in the island fox.

An Allee effect (AE) occurs in populations when individuals suffer a decrease in fitness at low densities. If a fitness component is reduced (component AE), per capita population growth rates may decline as a consequence (demographic AE) and extinction risk is increased. The island fox (Urocyon littoralis) is endemic to six of the eight California Channel Islands. Population crashes have coincided with an increase in predation by Golden Eagles (Aquila chrysaetos). We propose that AEs could render fox populations more sensitive and may be a likely explanation for their sharp decline. We analyzed demographic data collected between 1988 and 2000 to test whether fox density (1) influences survival and reproductive rates; (2) interacts with eagle presence and affects fox fitness parameters; and (3) influences per capita fox population trends. A double component AE simultaneously influenced survival (of adults and pups) and proportion of breeding adult females. The adult survival AE was driven by predation by eagles. These component AEs led to a demographic AE. Multiple-component AEs, a predation-driven AE, and the simultaneous occurrence of both component and demographic AEs in a mammal are all previously unreported processes. Populations below 7 foxes/km(2) could have suboptimal population growth rates due to the demographic AE, and AEs may have contributed to the dramatic declines in three fox populations. Because fox densities in critically endangered populations are well below this level, removing Golden Eagles appears necessary to prevent a predation-driven AE. Conservationists should also be aware of AEs when planning the release of captive foxes. More generally, our findings highlight the danger of overlooking AEs in the conservation of populations of rare or threatened species.

Pleistocene rewilding: an optimistic agenda for twenty-first century conservation.

Large vertebrates are strong interactors in food webs, yet they were lost from most ecosystems after the dispersal of modern humans from Africa and Eurasia. We call for restoration of missing ecological functions and evolutionary potential of lost North American megafauna using extant conspecifics and related taxa. We refer to this restoration as Pleistocene rewilding; it is conceived as carefully managed ecosystem manipulations whereby costs and benefits are objectively addressed on a case-by-case and locality-by-locality basis. Pleistocene rewilding would deliberately promote large, long-lived species over pest and weed assemblages, facilitate the persistence and ecological effectiveness of megafauna on a global scale, and broaden the underlying premise of conservation from managing extinction to encompass restoring ecological and evolutionary processes. Pleistocene rewilding can begin immediately with species such as Bolson tortoises and feral horses and continue through the coming decades with elephants and Holarctic lions. Our exemplar taxa would contribute biological, economic, and cultural benefits to North America. Owners of large tracts of private land in the central and western United States could be the first to implement this restoration. Risks of Pleistocene rewilding include the possibility of altered disease ecology and associated human health implications, as well as unexpected ecological and sociopolitical consequences of reintroductions. Establishment of programs to monitor suites of species interactions and their consequences for biodiversity and ecosystem health will be a significant challenge. Secure fencing would be a major economic cost, and social challenges will include acceptance of predation as an overriding natural process and the incorporation of pre-Columbian ecological frameworks into conservation strategies.

Coupling stable isotopes with bioenergetics to estimate interspecific interactions.

Interspecific interactions are often difficult to elucidate, particularly with large vertebrates at large spatial scales. Here, we describe a methodology for estimating interspecific interactions by combining stable isotopes with bioenergetics. We illustrate this approach by modeling the population dynamics and species interactions of a suite of vertebrates on Santa Cruz Island, California, USA: two endemic carnivores (the island fox and island spotted skunk), an exotic herbivore (the feral pig), and their shared predator, the Golden Eagle. Sensitivity analyses suggest that our parameter estimates are robust, and natural history observations suggest that our overall approach captures the species interactions in this vertebrate community. Nonetheless, several factors provide challenges to using isotopes to infer species interactions. Knowledge regarding species-specific isotopic fractionation and diet breadth is often lacking, necessitating detailed laboratory studies and natural history information. However, when coupled with other approaches, including bioenergetics, mechanistic models, and natural history, stable isotopes can be powerful tools in illuminating interspecific interactions and community dynamics.

Use of matrix population models to estimate the efficacy of euthanasia versus trap-neuter-return for management of free-roaming cats.

OBJECTIVE: To evaluate the efficacy of trap-neuter-return and trap-euthanatize management strategies for controlling urban free-roaming cat populations by use of matrix population models. DESIGN: Prospective study. SAMPLE POPULATION: Estimates of free-roaming cat populations in urban environments. PROCEDURE: Data from the literature describing the biology of free-roaming cat populations in urban environments were gathered. A matrix population model was developed with a range of high and low survival and fecundity values and all combinations of those values. The response of population growth rate to a range of management actions was assessed with an elasticity analysis. RESULTS: All possible combinations of survival and fecundity values of free-roaming cats led to predictions of rapid, exponential population growth. The model predicted effective cat population control by use of annual euthanasia of > or = 50% of the population or by annual neutering of > 75% of the fertile population. Elasticity analyses revealed that the modeled population was most susceptible to control through euthanasia. CONCLUSIONS AND CLINICAL RELEVANCE: Free-roaming cat populations have a high intrinsic growth rate, and euthanasia is estimated to be more effective at reducing cat populations than trap-neuter-return programs.

Models of regional habitat quality and connectivity for pumas (Puma concolor) in the southwestern United States.

The impact of landscape changes on the quality and connectivity of habitats for multiple wildlife species is of global conservation concern. In the southwestern United States, pumas (Puma concolor) are a well distributed and wide-ranging large carnivore that are sensitive to loss of habitat and to the disruption of pathways that connect their populations. We used an expert-based approach to define and derive variables hypothesized to influence the quality, location, and permeability of habitat for pumas within an area encompassing the entire states of Arizona and New Mexico. Survey results indicated that the presence of woodland and forest cover types, rugged terrain, and canyon bottom and ridgeline topography were expected to be important predictors of both high quality habitat and heightened permeability. As road density, distance to water, or human population density increased, the quality and permeability of habitats were predicted to decline. Using these results, we identified 67 high quality patches across the study area, and applied concepts from electronic circuit theory to estimate regional patterns of connectivity among these patches. Maps of current flow among individual pairs of patches highlighted possible pinch points along two major interstate highways. Current flow summed across all pairs of patches highlighted areas important for keeping the entire network connected, regardless of patch size. Cumulative current flow was highest in Arizona north of the Colorado River and around Grand Canyon National Park, and in the Sky Islands region owing to the many small habitat patches present. Our outputs present a first approximation of habitat quality and connectivity for dispersing pumas in the southwestern United States. Map results can be used to help target finer-scaled analyses in support of planning efforts concerned with the maintenance of puma metapopulation structure, as well as the protection of landscape features that facilitate the dispersal process.

Characterization of 15 tetranucleotide microsatellite markers in the ringtail (Bassariscus astutus).

We characterized 15 polymorphic tetranucleotide microsatellite markers for the ringtail, Bassariscus astutus. We tested these loci in 21 individuals captured in Arizona and Texas and found six to 19 alleles per locus. Observed and expected heterozygosities ranged from 0.381 to 1.000 and from 0.381 to 0.941, respectively. All loci were in Hardy-Weinberg equilibrium, and none were in linkage disequilibrium. These markers may be used to investigate population genetics and mating patterns in this species.

Does the order of invasive species removal matter? The case of the eagle and the pig.

BACKGROUND: Invasive species are recognized as a primary driver of native species endangerment and their removal is often a key component of a conservation strategy. Removing invasive species is not always a straightforward task, however, especially when they interact with other species in complex ways to negatively influence native species. Because unintended consequences may arise if all invasive species cannot be removed simultaneously, the order of their removal is of paramount importance to ecological restoration. In the mid-1990s, three subspecies of the island fox Urocyon littoralis were driven to near extinction on the northern California Channel Islands owing to heightened predation by golden eagles Aquila chrysaetos. Eagles were lured to the islands by an abundant supply of feral pigs Sus scrofa and through the process of apparent competition pigs indirectly facilitated the decline in foxes. As a consequence, both pigs and eagles had to be removed to recover the critically endangered fox. Complete removal of pigs was problematic: removing pigs first could force eagles to concentrate on the remaining foxes, increasing their probability of extinction. Removing eagles first was difficult: eagles are not easily captured and lethal removal was politically distasteful. METHODOLOGY/PRINCIPAL FINDINGS: Using prey remains collected from eagle nests both before and after the eradication of pigs, we show that one pair of eagles that eluded capture did indeed focus more on foxes. These results support the premise that if the threat of eagle predation had not been mitigated prior to pig removal, fox extinction would have been a more likely outcome. CONCLUSIONS/SIGNIFICANCE: If complete eradication of all interacting invasive species is not possible, the order in which they are removed requires careful consideration. If overlooked, unexpected consequences may result that could impede restoration.

Golden eagles, feral pigs, and insular carnivores: how exotic species turn native predators into prey.

Island ecosystems are particularly vulnerable to exotic species. Here we show how an introduced prey has led to the wholesale restructuring of an island food web, including the near extinction of an endemic carnivore. Introduced pigs, by providing abundant food, enabled golden eagles to colonize the California Channel Islands. Eagles preyed heavily on the island fox, whose resulting decline toward extinction released populations of the competitively inferior island skunk. The presence of exotic pigs led to major ecosystem shifts by indirectly causing predation to replace competition as the dominant force shaping these island communities.