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Pumas (Puma concolor) were eliminated from most of the eastern USA a century ago. In the past couple of decades, their recovery in the West has increased puma dispersal into the Midwest, with some individuals even traveling to the East Coast. We combined published expert opinion data and a habitat suitability index in an analysis that identified 17 areas in the Upper Midwest, Ozarks, Appalachia, and New England that could potentially host puma populations in the future. Thirteen of these were larger than 10,000 km2 and so likely to ensure a puma population’s long-term genetic health. Further, we quantified patch size, human density, livestock density, percent public land, and a sociocultural index reflecting wildlife values for comparing patches, as well as present a summary of current legislation relevant to puma management in the East. Our work may be useful in identifying suitable areas to restore pumas based not only on the quality of their biophysical habitat, but also on social values conducive to puma-human coexistence.

Kill rates and functional responses are fundamental to the study of predator ecology and the understanding of predatory-prey dynamics. As the most widely distributed apex predator in the western hemisphere, pumas (Puma concolor) have been well studied, yet a synthesis of their kill rates is currently lacking. We reviewed the literature and compiled data on sex- and age-specific kill rate estimates of pumas on ungulates, and conducted analyses aimed at understanding ecological factors explaining the observed spatial variation. Kill rate studies on pumas, while numerous, were primarily conducted in Temperate Conifer Forests (

Pumas (Puma concolor) are solitary large carnivores that exhibit high energetic investments while hunting prey that often take multiple days to consume. Therefore, pumas should behave in a way to maximize their energetic gains, including using caching, which is a behavior used by many mammal species to preserve and store food or to conceal it from conspecifics and scavengers to limit their losses. Yet pumas do not always cache their kills. In order to understand caching behavior, we used variables associated with the kills such as prey mass, search time, climate, and habitat to test 20 ecological models (representing four a priori hypotheses: food perishability, resource pulse, consumption time, and kleptoparasitism deterrence) in an information-theoretic approach of model selection to explore factors related to the caching behavior. Models were run with information from tracked radio-collared pumas in California over a 2.5-year period and identified a total of 352 kills. Overall, we documented pumas caching 61.5% of their kills, including 71.6% of Black-tailed Deer (Odocoileus hemionus columbianus), their primary prey in the study area. The model with a quadratic effect of adjusted mass of prey on puma caching probability had all of the empirical support (w = 1.00). Specifically, pumas were most likely to cache intermediate-sized prey, such as yearling and adult female deer, and also fed from cached kills for longer periods of time. Larger prey may be too large to easily cache, making it less energetically efficient—while small prey can often be consumed quickly enough to not require caching. This suggests that intermediate-sized prey may be the optimal size for caching, allowing a puma to feed for multiple days while not greatly increasing energetic output. The hypotheses we tested were not mutually exclusive and pumas caching their prey may occur for several reasons; nevertheless, our study demonstrated that pumas use caching to extend their foraging time and maximize energetic gains when preying on intermediate-sized prey.

The interspecific interactions of apex predators are integral to the function of ecological communities, but most studies have focused on understanding their top down effects. Kleptoparasitism (the stealing of procured food) by dominant scavengers can have negative effects on populations and behaviors of apex predators. We captured 7 pumas (Puma concolor) and fitted them with GPS collars to investigate potential kill sites (n = 352), some of which we monitored with camera traps (n = 58). We analyzed whether observed kleptoparasitism by American black bears (Ursus americanus) affected puma energetics and foraging behavior. We found that black bears were the most frequent scavenger of puma kills (72.4%), and we documented bears scavenging puma kills during every month. The top model for bear detection of puma kills included prey size, temperature, and canopy cover, with bears more likely to scavenge from adult black-tailed deer (Odocoileus hemionus columbianus) carcasses in warmer temperatures and under dense canopy cover. When black bear scavenging occurred, pumas spent 22% less time at their kill and incurred energetic losses. In response, pumas shortened their inter-kill intervals by 1.3 days thus increasing their kill rates. Our results demonstrate how a dominant scavenger directly mediates the foraging behavior of an apex predator. These results suggest that community interactions do not necessarily start at the top in top-down systems, and the effects of predators on prey populations can only be understood within their respective ecological communities.

Female–female sexual behaviors have been recorded in many species across several taxa, but their infrequency except in a few species has resulted in continued speculation about their function and potential evolutionary consequences. Here, we report two observations of female–female mounting in wild puma populations representing two sub-species from opposite ends of puma range. We believe our observations provided support for the social glue hypothesis and dominance in same-sex hierarchies, but not for surrogacy in the absence of a male or as a means to stimulate and encourage male partners to copulate.

SummaryWe tested for plague (Yersinia pestis) in a puma population in the Greater Yellowstone Ecosystem (GYE) over 9 years, overlapping a case when a boy in the area became infected with plague. Antibodies to Y. pestis were detected in 8 of 17 (47%) pumas tested by complement-enzyme-linked immunosorbent assay, and the organism itself was detected in 4 of 11 (36%) pumas tested after necropsy. Neither puma sex nor age was significantly associated with Y. pestis exposure or mortality, although our sample size was small. The overall prevalence of exposure we recorded was similar to that found along the western slope of Colorado, which is adjacent to the Four Corners region, a known plague hotspot in the USA. This suggests that: (1) Y. pestis may be present at higher levels in the GYE than previously assumed; (2) plague is a significant source of mortality for local pumas (6.6% of sub-adult and adult mortality); and (3) pumas may be a useful sentinel for potential risk of plague exposure to humans throughout the West. We would also emphasize that hunters and others handling pumas in this region should be made aware of the possibility of exposure.

Mortality site investigations of telemetered wildlife are important for cause-specific survival analyses and understanding underlying causes of observed population dynamics. Yet, eroding ecoliteracy and a lack of quality control in data collection can lead researchers to make incorrect conclusions, which may negatively impact management decisions for wildlife populations. We reviewed a random sample of 50 peer-reviewed studies published between 2000 and 2019 on survival and cause-specific mortality of ungulates monitored with telemetry devices. This concise review revealed extensive variation in reporting of field procedures, with many studies omitting critical information for the cause of mortality inference. Field protocols used to investigate mortality sites and ascertain the cause of mortality are often minimally described and frequently fail to address how investigators dealt with uncertainty. We outline a step-by-step procedure for mortality site investigations of telemetered ungulates, including evidence that should be documented in the field. Specifically, we highlight data that can be useful to differentiate predation from scavenging and more conclusively identify the predator species that killed the ungulate. We also outline how uncertainty in identifying the cause of mortality could be acknowledged and reported. We demonstrate the importance of rigorous protocols and prompt site investigations using data from our 5-year study on survival and cause-specific mortality of telemetered mule deer (

Noise is the non-target search results that people encounter when searching for a particular topic of interest; it is also the cloud of distracting data that can obscure or deflect conservation communication. Online noise associated with large carnivores is particularly dense because their defining characteristics make them salient. Mountain lions (Puma concolor) exemplify noise associated with multiple vernaculars for a species in the crosshairs of conservation conundrums. We compared internet search results, Google Trends reflecting topic interest, use in science publications and sentiment in print and online media for P. concolor's most frequent vernacular names, “mountain lion,” “cougar,” “puma” and “Florida panther.” Puma and panther exhibited greater noise and salience than cougar or mountain lion, but, results for mountain lion, followed by cougar, yielded the highest biological relevance. Online sentiment negatively correlated with biological relevance, with positive sentiment highest for the noisiest vernaculars, puma and panther. As conservation practitioners, we must recognize that public outreach is part of our scientific agenda and be conscious of crafting communication that reaches and resonates with our intended audiences.

In response to recent discussion about terminology, we propose “tracking science” as a term that is more inclusive than citizen science. Our suggestion is set against a post-colonial political background and large-scale migrations, in which “citizen” is becoming an increasingly contentious term. As a diverse group of authors from several continents, our priority is to deliberate a term that is all-inclusive, so that it could be adopted by everyone who participates in science or contributes to scientific knowledge, regardless of socio-cultural background. For example, current citizen science terms used for Indigenous knowledge imply that such practitioners belong to a sub-group that is other, and therefore marginalized. Our definition for “tracking science” does not exclude Indigenous peoples and their knowledge contributions and may provide a space for those who currently participate in citizen science, but want to contribute, explore, and/or operate beyond its confinements. Our suggestion is not that of an immediate or complete replacement of terminology, but that the notion of tracking science can be used to complement the practice and discussion of citizen science where it is contextually appropriate or needed. This may provide a breathing space, not only to explore alternative terms, but also to engage in robust, inclusive discussion on what it means to do science or create scientific knowledge. In our view, tracking science serves as a metaphor that applies broadly to the scientific community—from modern theoretical physics to ancient Indigenous knowledge.

Human activity and land use change impact every landscape on Earth, driving declines in many animal species while benefiting others. Species ecological and life history traits may predict success in human-dominated landscapes such that only species with "winning" combinations of traits will persist in disturbed environments. However, this link between species traits and successful coexistence with humans remains obscured by the complexity of anthropogenic disturbances and variability among study systems. We compiled detection data for 24 mammal species from 61 populations across North America to quantify the effects of (1) the direct presence of people and (2) the human footprint (landscape modification) on mammal occurrence and activity levels. Thirty-three percent of mammal species exhibited a net negative response (i.e., reduced occurrence or activity) to increasing human presence and/or footprint across populations, whereas 58% of species were positively associated with increasing disturbance. However, apparent benefits of human presence and footprint tended to decrease or disappear at higher disturbance levels, indicative of thresholds in mammal species' capacity to tolerate disturbance or exploit human-dominated landscapes. Species ecological and life history traits were strong predictors of their responses to human footprint, with increasing footprint favoring smaller, less carnivorous, faster-reproducing species. The positive and negative effects of human presence were distributed more randomly with respect to species trait values, with apparent winners and losers across a range of body sizes and dietary guilds. Differential responses by some species to human presence and human footprint highlight the importance of considering these two forms of human disturbance separately when estimating anthropogenic impacts on wildlife. Our approach provides insights into the complex mechanisms through which human activities shape mammal communities globally, revealing the drivers of the loss of larger predators in human-modified landscapes.

Species assemblages often have a non-random nested organization, which in vertebrate scavenger (carrion-consuming) assemblages is thought to be driven by facilitation in competitive environments. However, not all scavenger species play the same role in maintaining assemblage structure, as some species are obligate scavengers (i.e., vultures) and others are facultative, scavenging opportunistically. We used a database with 177 vertebrate scavenger species from 53 assemblages in 22 countries across five continents to identify which functional traits of scavenger species are key to maintaining the scavenging network structure. We used network analyses to relate ten traits hypothesized to affect assemblage structure with the “role” of each species in the scavenging assemblage in which it appeared. We characterized the role of a species in terms of both the proportion of monitored carcasses on which that species scavenged, or scavenging breadth (i.e., the species “normalized degree”), and the role of that species in the nested structure of the assemblage (i.e., the species “paired nested degree”), therefore identifying possible facilitative interactions among species. We found that species with high olfactory acuity, social foragers, and obligate scavengers had the widest scavenging breadth. We also found that social foragers had a large paired nested degree in scavenger assemblages, probably because their presence is easier to detect by other species to signal carcass occurrence. Our study highlights differences in the functional roles of scavenger species and can be used to identify key species for targeted conservation to maintain the ecological function of scavenger assemblages., ESG, ZMR, JMB and LNA were supported by Generalitat Valenciana (SEJI/2018/024, APOSTD/2019/016, CIDEGENT/2020/030 and ACIF/2019/056, respectively), JMB, JMPG and CGC by Juan de la Cierva contracts (MEC; IJCI-2017-32149, FJCI-2015-25632 and IJC2018-036642-I, respectively), ACA by the Govern de les Illes Balears (PD/039/2017) and ESG and MM by Ramón y Cajal contracts (MEC; RYC-2019-027216-I, RYC-2015-19231). EA was supported by La Caixa-Severo Ochoa International PhD Program 2015. NS was partly supported by the National Science Centre in Poland (2013/08/M/NZ9/00469 and 2016/22/Z/NZ8/00). MK and KJ were supported by the Slovenian Research Agency (P4-0059). Contributions of HG, KLT, EFA, OER, TLD, and JCB were partially supported through funding from U.S. Department of Agriculture and the U.S. Department of Energy under (DE-EM0004391) to the University of Georgia Research Foundation. HG was also supported by the Institute of Environmental Radioactivity at Fukushima University. ALB and JDA were partially supported by Queens College and the Graduate Center at the City University of New York. JDA is currently supported by a Ramón y Cajal contract (RYC-2017-22783). ERB and EF were supported by the USA National Science Foundation Graduate Research Fellowship (1256065). CK completed study with support from Hawk Mountain Sanctuary, The Peregrine Fund, and via Pompeo M. Maresi Memorial Fund via Princeton University. JAS and CCW were supported by the USA National Science Foundation #1255913, the American Association for University Women, and the Gordon and Betty Moore Foundation. HUW acknowledges funding from the California Department of Fish & Wildlife (P0880013). PLP was supported by the Rufford Foundation and University of Wisconsin-Madison. JB and JdT thank the Percy Sladen Memorial Fund and Mr Rodney Fuhr. Several authors were funded by funds from the MEC (CGL2012-40013-C02-01/02, CGL2015-66966-C2-1-R, CGL2015-66966-C2-1-R2, CGL2017-89905-R, RTI2018-099609-B-C21 and RTI2018-099609-B-C22) and from the Junta de Andalucía (RNM-1925). POV was supported by a research contract by the University of Lleida. EES and TMN were funded and supported by Australian Geographic, Bush Heritage Australia, Australian Academy of Sciences, Ecological Society of Australia, NSW Office of Environment and Heritage, NESP Threatened Species Recovery Hub, Emirates Wolgan Valley One and Only Resort. EP, YA, MS and UY completed study under research permit by The Republic of Turkey Ministry of Agriculture and Forestry. SAL thanks PICT (BID) 0725/2014, and IAATE. ALB and JDA would like to thank the Edmund Niles Huyck Preserve, Lyme Adirondack Forest Company, and LandVest Timberland.

© 2017 American Society of Mammalogists. Global positioning system (GPS) technology has revolutionized the study of carnivores. Researchers commonly estimate kill rates with GPS data using the following steps. Firstly, researchers mark individual animals and fit them with GPS collars. Next, they visit a subset of "GPS clusters" (aggregated location data) during field surveys and assign spatiotemporal covariates associated with predation and non-predation events. Lastly, they develop predictive models with data collected in the field to estimate the probability that each cluster they did not visit in the field was a predation event. Such predation models help reduce field efforts and save money; however, these models are prone to error when carnivores eat prey of different sizes or exhibit shorter-than-expected handling times. We simulated reduced field efforts to investigate the reliability of predictive modeling in determining diet composition and detecting predation events for 3 puma (Puma concolor) populations with different prey assemblages and potential effects on handling time of carcasses. We visited a total 1,896 clusters in Chilean Patagonia, Colorado, and California, of which 1,752 clusters (∼92%) were included to build and test predation models. Across all study areas, the total time a puma spent at a cluster was the only reliable predictor of a cluster being a predation event. When we reduced field efforts by selectively removing GPS clusters < 12 and < 24 h in length, model performance improved but produced inaccurate results. Predation models underestimated the number of predation events in California and Colorado and significantly over-or underestimated the number of predation events in Patagonia. Selectively reducing field efforts also reduced the diversity and evenness of prey we recorded in puma diets. Randomly reducing field efforts, in contrast, reduced the precision of model estimates. Our results highlight the importance of conducting intensive fieldwork over predation modeling to measure prey selection and kill rates of carnivores.

© 2018 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd. Humans are primary drivers of declining abundances and extirpation of large carnivores worldwide. Management interventions to restore biodiversity patterns, however, include carnivore reintroductions, despite the many unresolved ecological consequences associated with such efforts. Using multistate capture–mark–recapture models, we explored age-specific survival and cause-specific mortality rates for 134 pumas (Puma concolor) monitored in the Greater Yellowstone Ecosystem during gray wolf (Canis lupus) recovery. We identified two top models explaining differences in puma survivorship, and our results suggested three management interventions (unsustainable puma hunting, reduction in a primary prey, and reintroduction of a dominant competitor) have unintentionally impacted puma survival. Specifically, puma survival across age classes was lower in the 6-month hunting season than the 6-month nonhunting season; human-caused mortality rates for juveniles and adults, and predation rates on puma kittens, were higher in the hunting season. Predation on puma kittens, and starvation rates for all pumas, also increased as managers reduced elk (Cervus elaphus) abundance in the system, highlighting direct and indirect effects of competition between recovering wolves and pumas over prey. Our results emphasize the importance of understanding the synergistic effects of existing management strategies and the recovery of large, dominant carnivores to effectively conserve subordinate, hunted carnivores in human-dominated landscapes.

Ecosystem engineers create physical changes in abiotic and biotic material, and through this process control the availability of resources for other species. Predators that abandon large portions of their prey may be ecosystem engineers that create habitat for carrion-dependent invertebrates that utilize carcasses during critical life-history periods. Between 04-May-2016 and 04-Oct-2016, we sampled beetle assemblages at 18 carcasses of prey killed by pumas and matching control sites in the southern Greater Yellowstone Ecosystem, USA, to measure the extent to which beetle families utilized these carcass "habitats". We used generalized linear-mixed models and linear-mixed effect models to examine changes in beetle abundance, species richness, and Simpson's Index of Diversity. We estimated kill rates and carrion production rates for individual pumas to better assess the impact of pumas on invertebrate communities. We collected 24,209 beetles representing 215 species. We identified eight beetle families that had significantly higher abundance at carcasses than control sites. Carcasses had a statistically large to very large effect (determined using Cohen's d) on beetle abundance, richness, and diversity for the initial 8 weeks of sampling. Our research revealed strong effects of an ecosystem engineer on beetle assemblages while highlighting the potential role of apex predators in creating and modifying physical habitats for carrion-dependent species. This suggests that there may be consequences for invertebrate communities where apex predators exist at reduced numbers or have been eradicated. The ecological role of invertebrates is often overlooked, yet they are essential taxa that provide critical ecological services upon which we depend.

Carrion promotes biodiversity and ecosystem stability, and large carnivores provide this resource throughout the year. In particular, apex felids subordinate to other carnivores contribute more carrion to ecological communities than other predators. We measured vertebrate scavenger diversity at puma (Puma concolor) kills in the Greater Yellowstone Ecosystem, and utilized a model-comparison approach to determine what variables influenced scavenger diversity (Shannon's H) at carcasses. We documented the highest vertebrate scavenger diversity of any study to date (39 birds and mammals). Scavengers represented 10.9% of local birds and 28.3% of local mammals, emphasizing the diversity of food-web vectors supported by pumas, and the positive contributions of pumas and potentially other subordinate, apex felids to ecological stability. Scavenger diversity at carcasses was most influenced by the length of time the carcass was sampled, and the biological variables, temperature and prey weight. Nevertheless, diversity was relatively consistent across carcasses. We also identified six additional stalk-and-ambush carnivores weighing > 20 kg, that feed on prey larger than themselves, and are subordinate to other predators. Together with pumas, these seven felids may provide distinctive ecological functions through their disproportionate production of carrion and subsequent contributions to biodiversity. We urge conservation managers to increase support for these species, as a means of prioritizing resources to best ensure the persistence of carrion in natural systems.

Ecotourism enhances conservation management, promotes non-consumptive use of wildlife, and increases local community resources over that of select individuals when compared with consumptive uses such as hunting or trapping. The bobcat is a cryptic mesocarnivore widely exploited for pelts across North America, and a species increasingly contributing to ecotourism. Here, we report a conservative, non-consumptive economic value of US$308,105 for a single bobcat in Yellowstone National Park in northwest Wyoming for the 2015–2016 winter season, a figure nearly 1000 times greater than exploitive values of US$315.17 per bobcat trapped or hunted in Wyoming in the same season (US$130.53 per bobcat harvested in revenue earned by the state of Wyoming in trapping license sales + US$184.64 per pelt sold by successful trappers and hunters). In 2016, tourism was the second largest industry in Wyoming and generated US$3.2 billion. Our case study emphasizes that current bobcat regulatory policies across North America do not reflect current cultural values, inclusive of both consumptive and non-consumptive use of wildlife. Therefore, we recommend range-wide regulatory changes to ensure bobcat management is not just sustainable in terms of harvest, but that all people have access to shared resources held in trust.

A Brownian motion whose infinitesimal variance alternates according to a telegraph process is considered. This stochastic process can be employed to model a variety of real-word situations, such as animal movement in ecology and stochastic volatility in mathematical finance. The main goal is to develop an estimation procedure for the underlying model parameters when the process is observed at discrete, possibly irregularly spaced time points. The sequence of observations is not Markov, but the sequence of the state of the telegraph process, if observed, is Markov. The observed sequence is therefore from a hidden Markov model. Likelihood inference is developed via dynamic programming, and is demonstrated to have much higher efficiency than the composite likelihood approach that was applied in an earlier work. The model is applied to model the movement of a mountain lion.

Predator tourism is one strategy to improve tolerance for predators, and support biodiversity and ecosystem health. Torres del Paine National Park (TdP) – a UNESCO Biosphere Reserve in southern Chile - supports productive livestock industries and nascent puma tourism. We compared interviews conducted in the region prior to puma tourism, with results from interviews collected across 45 ranches post-puma tourism. We assessed rancher attitudes regarding pumas, puma-livestock conflict, puma tourism, and linked them with socio-ecological factors. Respondents who viewed pumas as a threat experienced higher livestock losses. Respondents who reported higher sheep losses were inclined to support the lethal removal of livestock-killing pumas, and to initiate a puma hunt, whereas respondents who supported puma tourism disagreed with hunting pumas. Using the Potential for Conflict Index, we found that participants exhibited the highest consensus on the benefit of puma tourism and the lowest consensus over lethal removal of pumas. Our results suggest predator tourism has increased tolerance for pumas but is creating new potential for conflict. Previous to puma tourism, ranchers were almost entirely negative about pumas and unanimously supported illegal puma hunting. Now, most believe that pumas are part of Patagonia's heritage. This divide was best explained by distance to TdP: ranches closer to TdP experienced greater losses to pumas but had neighbours that benefitted most from puma tourism. Therefore, we suggest that tourism revenues supplement community compensation insurance programs that reimburse rancher losses to pumas to mitigate the growing divide between those benefiting from pumas and those experiencing economic hardship.

Many species have been increasingly recognized as an aggregate of individual dietary specialists. Previous work has emphasized the importance of individual dietary specialization in driving many ecological processes, but less is known about the relationship between individual diet and habitat use. We used data from 7 cougars and 331 predation events collected in western Colorado from 2010 to 2013 to test whether intraspecific variation in cougar (Puma concolor) diet resulted from chance encounters or from targeting particular prey species. One cougar within our study population (P06) selected for American beaver (Castor canadensis) more than expected based on availability, spent a disproportionate amount of time within beaver habitat, and exhibited reduced travel speeds when near waterways. Our results present evidence that specialist diets in predators are likely reflective of additional differences in foraging behaviors rather than random encounters with prey species (e.g., prey availability).

Prey selection is exhibited by predator populations that kill a prey species disproportionate to its availability, or alternatively, individual predators that select prey disproportionate to the mean selection exhibited by their populations. Prey selection is a simple calculation when one can determine prey availability; however, measuring prey availability is challenging. We compared population- and individual-level prey selection as determined with two measures of prey availability for five ungulate species killed by pumas (Puma concolor (L., 1771)) in the Southern Yellowstone Ecosystem, USA: (1) annual prey counts and (2) total prey killed by marked pumas. We also tested whether individual pumas in the population exhibited a narrower dietary niche breadth compared with their population as a whole. The two methods yielded different estimates of prey availability and highlighted the need to consciously match prey availability estimates with appropriate ecological questions. Prey counts may have overestimated elk (Cervus canadensis (Erxleben, 1777)) abundance and underestimated deer abundance, whereas predation data may have better captured the influence of prey size on puma-specific prey vulnerability and availability. Prey counts were the more appropriate metric for analyzing population-level prey selection or differences in interspecific foraging, whereas total prey killed was the more appropriate metric for intraspecific comparisons.

Top-down effects of apex predators are modulated by human impacts on community composition and species abundances. Consequently, research supporting top-down effects of apex predators occurs almost entirely within protected areas rather than the multi-use landscapes dominating modern ecosystems. Here, we developed an integrated population model to disentangle the concurrent contributions of a reintroduced apex predator, the grey wolf, human hunting and prey abundances on vital rates and abundance of a subordinate apex predator, the puma. Increasing wolf numbers had strong negative effects on puma fecundity, and subadult and adult survival. Puma survival was also influenced by density dependence. Overall, puma dynamics in our multi-use landscape were more strongly influenced by top-down forces exhibited by a reintroduced apex predator, than by human hunting or bottom-up forces (prey abundance) subsidized by humans. Quantitatively, the average annual impact of human hunting on equilibrium puma abundance was equivalent to the effects of 20 wolves. Historically, wolves may have limited pumas across North America and dictated puma scarcity in systems lacking sufficient refugia to mitigate the effects of competition.

Carrion is a rich, ephemeral resource vital to biodiversity and ecosystem health. In temperate ecosystems in which cold temperatures and snowfall influence the accessibility and availability of small prey and seasonal mast crops, carrion may also be a limiting resource for mesocarnivores like red foxes (Vulpes vulpes), which are too small to predate ungulates. Using motion-triggered video cameras and generalized linear mixed models, we studied the spatial and temporal patterns of red fox scavenging at 232 mountain lion kills in the southern Greater Yellowstone Ecosystem (GYE) from 2012–2015. We found that red foxes scavenged mountain lion kills across all habitats throughout the year, however, red fox behaviors varied with season. In winter, we documented red foxes at a greater proportion of mountain lion kills (70.3% in winter vs. 48.9% in summer), and in greater numbers (1.83 foxes per kill in winter vs. 1.16 in summer). In winter, red foxes fed longer (= 102.7 ± 138.3 minutes feeding in winter vs. = 39.7 ± 74.0 in summer), and they more often scavenged while the mountain lion was nearby. We speculated that red foxes may have increased risk taking in winter due to hunger driven by resource scarcity. Our research highlighted an important ecological relationship between red foxes and mountain lions in the GYE. Mountain lions tolerate high levels of scavenging, so the frequency and intensity of red fox scavenging at their kills may not impact mountain lions, but instead facilitate the dispersion and benefits of resources created by this apex predator. Large carnivores, and mid-trophic felids like mountain lions in particular, are essential producers of carrion vital to biodiversity and ecosystem health. In turn, scavengers play critical roles in distributing these resources and increasing the heterogeneity of resources that support biodiversity and ecosystem structure, as well as ecological resilience.

BackgroundInterspecific competition affects species fitness, community assemblages and structure, and the geographic distributions of species. Established dominance hierarchies among species mitigate the need for fighting and contribute to the realized niche for subordinate species. This is especially important for apex predators, many of which simultaneous contend with the costs of competition with more dominant species and the costs associated with human hunting and lethal management.MethodsPumas are a widespread solitary felid heavily regulated through hunting to reduce conflicts with livestock and people. Across their range, pumas overlap with six apex predators (gray wolf, grizzly bear, American black bear, jaguar, coyote, maned wolf), two of which (gray wolf, grizzly bear) are currently expanding in North America following recovery efforts. We conducted a literature search to assess whether pumas were subordinate or dominant with sympatric apex predators, as well as with three felid mesocarnivores with similar ecology (ocelot, bobcat, Canada lynx). We also conducted an analysis of the spatial distributions of pumas and their dominant sympatric competitors to estimate in what part of their range, pumas are dominant versus subordinate.ResultsWe used 64 sources to assess dominance among pumas and other apex predators, and 13 sources to assess their relationships with felid mesocarnivores. Evidence suggested that wolves, grizzly bears, black bears, and jaguars are dominant over pumas, but that pumas are dominant over coyotes and maned wolves. Evidence suggested that pumas are also dominant over all three felid mesocarnivores with which they share range. More broadly, pumas are subordinate to at least one other apex carnivore in 10,799,252 (47.5%) of their 22,735,268 km2range across North and South America.DiscussionSubordinate pumas change their habitat use, suffer displacement at food sources, likely experience increased energetic demands from harassment, exhibit increased starvation, and are sometimes directly killed in competitive interactions with dominant competitors. Nevertheless, we lack research clearly linking the costs of competition to puma fitness. Further, we lack research that assesses the influence of human effects simultaneous with the negative effects of competition with other sympatric carnivores. Until the time that we understand whether competitive effects are additive with human management, or even potentially synergistic, we encourage caution among managers responsible for determining harvest limits for pumas and other subordinate, apex carnivores in areas where they are sympatric with dominant species. This may be especially important information for managers working in regions where wolves and brown bears are recolonizing and recovering, and historic competition scenarios among multiple apex predators are being realized.

Pumas (Puma concolor) and black bears (Ursus americanus) are large carnivores that may influence scavenger population dynamics. We used motion-triggered video cameras deployed at deer carcasses to determine how pumas and black bears affected three aspects of carrion acquisition by scavengers: presence, total feeding time, and mean feeding-bout duration. We found that pumas were unable to limit acquisition of carrion by large carnivores but did limit aspects of carrion acquisition by both birds and mesocarnivores. Through their suppression of mesocarnivores and birds, pumas apparently initiated a cascading pattern and increased carrion acquisition by small carnivores. In contrast, black bears monopolized carrion resources and generally had larger limiting effects on carrion acquisition by all scavengers. Black bears also limited puma feeding behaviors at puma kills, which may require pumas to compensate for energetic losses through increasing their kill rates of ungulates. Our results suggest that pumas provide carrion and selectively influence species acquiring carrion, while black bears limit carrion availability to all other scavengers. These results suggest that the effects of large carnivores on scavengers depend on attributes of both carnivores and scavengers (including size) and that competition for carcasses may result in intraguild predation as well as mesocarnivore release.

One hypothesis for how carnivores with overlapping ecology coexist in natural systems is through heterogeneous competition landscapes, in which subordinates utilize “competition refuges” to mitigate risks associated with dominant competitors. We tested for the effects of American black bear (Ursus americanus) kleptoparasitism on puma (Puma concolor) foraging in 2 systems in North America. We also tested whether partial prey consumption exhibited by pumas in the presence of bears was better explained by rules of optimal foraging or by kleptoparasitism by black bears, and whether pumas utilized spatial competition refuges to mitigate competition with bears over carcass remains. Puma kill rates in ungulates/wk were equivalent across study systems, but 48% greater in the bear season than the no-bear season. Our analyses of handling time did not support the notion that partial prey consumption exhibited by pumas followed patterns of optimal foraging. Rather, puma handling time and prey consumption were better explained by the presence of bears. Surprisingly, pumas did not utilize spatial competition refuges to mitigate competition with black bears, and instead our results suggested they increase their kill rates to compensate for losses. Our results linking high seasonal kill rates of a top predator with kleptoparasitism by a dominant competitor provide strong evidence that the effects of predation can only be understood within a community framework. In particular, we propose that future predation studies should differentiate between relative contributions of predators and competitors on prey dynamics. Further, our results suggest kleptoparasites may indirectly impact prey populations through their effects on top predators.

As technology has improved, our ability to study cryptic animal behavior has increased. Bed site selection is one such example. Among prey species, bed site selection provides thermoregulatory benefits and mitigates predation risk, and may directly influence survival. We conducted research to test whether a subordinate carnivore also selected beds with similar characteristics in an ecosystem supporting a multi-species guild of competing predators. We employed a model comparison approach in which we tested whether cougar (Puma concolor) bed site attributes supported the thermoregulatory versus the predator avoidance hypotheses, or exhibited characteristics supporting both hypotheses. Between 2012–2016, we investigated 599 cougar bed sites in the Greater Yellowstone Ecosystem and examined attributes at two scales: the landscape (second-order,n = 599) and the microsite (fourth order,n = 140). At the landscape scale, cougars selected bed sites in winter that supported both the thermoregulatory and predator avoidance hypotheses: bed sites were on steeper slopes but at lower elevations, closer to the forest edge, away from sagebrush and meadow habitat types, and on southern, eastern, and western-facing slopes. In the summer, bed attributes supported the predator avoidance hypothesis over the thermoregulation hypothesis: beds were closer to forest edges, away from sagebrush and meadow habitat classes, and on steeper slopes. At the microsite scale, cougar bed attributes in both the winter and summer supported both the predator avoidance and thermoregulatory hypotheses: they selected bed sites with high canopy cover, high vegetative concealment, and in a rugged habitat class characterized by cliff bands and talus fields. We found that just like prey species, a subordinate predator selected bed sites that facilitated both thermoregulatory and anti-predator functions. In conclusion, we believe that measuring bed site attributes may provide a novel means of measuring the use of refugia by subordinate predators, and ultimately provide new insights into the habitat requirements and energetics of subordinate carnivores.

We employed global positioning system (GPS) locations of 18 marked cougars and genetic material from 68 cougars in the Southern Yellowstone Ecosystem to test our current assumptions about the social organization of this non-cooperative, solitary carnivore. We tested whether variable numbers of spatial associations over 7 years of our project could be explained by increasing numbers of wolves in the system, decreasing numbers of prey, changes in cougar density, the mean genetic relatedness between interacting individuals, or the timing of the breeding season. We documented 92 spatial associations and an additional 190 incidents of spatial overlap. Our models suggested only season influenced the number of associations in any given year, and a greater frequency of associations occurred in the breeding season. Nevertheless, the proportion of associations that were male–female (M–F) remained consistent across breeding and non-breeding seasons, suggesting M–F associations may not entirely be driven by mating opportunities; alternatively, the frequency of associations may have been driven by snow depths and the distributions of cougar prey in winter. The mean r value for female–female spatial associations was 0.087 (defined as “unrelated”) and did not significantly differ from M–F or M–M associations. In conclusion, our genetic research revealed not only matrilineal lines in the Southern Yellowstone Ecosystem but also immigration by new females. Subadult males primarily emigrated out of the system, but one male remained philopatric. Our results highlighted the notion that solitary cougars are associating, and potentially interacting, on the landscape with regularity and predictability. Further, our results raised critical questions: If cougars are associating more frequently than previously believed, why do they do so and are these interactions typically agonistic?

Density estimates are critical to proper population management and conservation, yet difficult to obtain for many wide-ranging or cryptic species. One proven method used to quantify carnivore density, especially species difficult to individually identify from photos taken by camera traps, utilizes overlapping home ranges of individual animals in the study area. This method, however, may be particularly prone to residency and extrapolation biases. Residency bias occurs when the reference area for the density estimate is incorrect, and extrapolation bias occurs when scaling a density to a different spatial extent than that of the study area upon which the estimate was based. We used a simulation approach based on GPS locations to diagnose potential biases in published densities of pumas Puma concolor from Patagonia, where Franklin et al. 1999 (Biol. Conserv. 90: 33–40) reported ‘minimum’ densities of 6 and 30 pumas per 100 km2, and Elbroch and Wittmer 2012a (Mammal. Biol. 77: 377–384) reported densities of ...

The gray fox (Urocyon cinereoargenteus) is an under-studied species in North America. Here we provide data gathered through observations over 17 d at a gray fox den in San Mateo County, California. We recorded prey indices through direct observation, as well as through indirect observation with photos recorded by motion-triggered cameras. The largest prey was a mule deer fawn, which we determined was killed by a gray fox. This finding is the first record of gray fox predation on mule deer. Lagomorphs and rodents formed the majority of prey items. We also recorded behavior that both contradicted and corroborated previous literature. We observed the male bringing food items to the den, a behavior that previous researchers have disagreed about. We also observed allogrooming between the adult pair, as well as one instance among pups where leg-lifting accompanied by presentation of the genitalia was clearly used as an aggressive dominant behavior rather than a submissive behavior, as reported in previ...

We compared injuries resulting from two different capture methods, i.e. trailing hounds and foot snares, for pumas Puma concolor in a mixed landscape with open grasslands and limited ‘retreat’ habitat in southern Chilean Patagonia. Injury scores were not significantly different for the two methods, although the small sample size for captures made with snares likely influenced our findings. Based on a potential range of 0-400, the mean injury score for pumas caught using hounds was 56.3 ± 132.9 (SD). The mean puma injury score for five animals caught in snares was 3.8 ± 1.1. Pumas were injured and/or killed in 86% of captures using trailing hounds. The number of hounds used in a capture attempt did not predict the likelihood of successfully catching a puma (P = 0.35), whereas there was a strong relationship between the number of dogs and the likelihood of a fight with the puma (P < 0.0001). The odds ratio calculation predicted a 14.7% increase in the likelihood of a fight between puma and hounds with the a...

Conservation interventions increasingly involve active management of relative species abundances, especially when taxa of conservation concern are threatened by complex food web interactions. Unfortunately, the complexity of such interspecific interactions means that well-meaning management interventions can have unexpected, sometimes detrimental, effects on the species they are intended to protect. Here we report that the abrupt removal of an abundant non-native prey species (domestic sheep) and the cessation of predator control, actions intended to protect huemul deer Hippocamelus bisulcus in the future Patagonia National Park, appear to have had negative effects on this Endangered ungulate. During and following the changes in predator–prey management, predation of huemul fawns and females older than 1 year by native culpeo foxes Lycalopex culpaeus and pumas Puma concolor increased 3- and 5-fold, respectively. Predictions from demographic models suggest that elevated mortality rates of female huemul older than 1 year will, on average, cause this population of huemul to decline. These results highlight risks of unintended consequences when aggressive management actions are taken to protect taxa embedded in complex food webs. They also suggest that careful consideration of both inter- and intra-trophic level effects among all species in a system is warranted before conservation interventions are undertaken.

Populations of generalist foragers may in fact be composed of individuals that select different prey. We monitored 9 pumas (Puma concolor) in Chilean Patagonia using Argos–global positioning system (Argos-GPS) technology for a mean of 9.33 months 6 5.66 SD. We investigated 694 areas where puma location data were spatially aggregated, called GPS clusters, at which we identified 433 kill sites and 6 acts of scavenging. Pumas as a population specialized upon guanacos (Lama guanicoe), whereas only 7 of 9 individual pumas specialized upon guanacos. One puma specialized upon domestic sheep (Ovis aries) and 1 upon European hares (Lepus europaeus) in terms of numbers of prey killed. Male and female pumas selected different distributions of prey and pumas exhibited prey selection at both the individual and population level. Three of 9 pumas exhibited selection when we compared individual prey use to prey availability within individual pumas’ home ranges. One puma selected endangered huemul (Hippocamelus bisulcus) and 2 selected sheep. When we compared individual prey use to prey use at the population level, 5 of 9 pumas differed from the population norm. Whereas pumas did not select huemul at the population level, 2 individuals did select huemul. Two individuals also selected domestic sheep, and the influence of these 2 pumas was substantial enough to result in a population-level effect. Our research highlights the need to determine whether pumas exhibit individual foraging variation throughout their range, the extrinsic factors associated with (and possibly influencing) such variation, and how pumas that select rare and less abundant species in multiprey systems impact recovering prey populations.

Apparent competition is an indirect interaction between 2 or more prey species through a shared predator, and it is increasingly recognized as a mechanism of the decline and extinction of many species. Through case studies, we evaluated the effectiveness of 4 management strategies for species affected by apparent competition: predator control, reduction in the abundances of alternate prey, simultaneous control of predators and alternate prey, and no active management of predators or alternate prey. Solely reducing predator abundances rapidly increased abundances of alternate and rare prey, but observed increases are likely short-lived due to fast increases in predator abundance following the cessation of control efforts. Substantial reductions of an abundant alternate prey resulted in increased predation on endangered huemul (Hippocamelus bisulcus) deer in Chilean Patagonia, which highlights potential risks associated with solely reducing alternate prey species. Simultaneous removal of predators and alternate prey increased survival of island foxes (Urocyon littoralis) in California (U.S.A.) above a threshold required for population recovery. In the absence of active management, populations of rare woodland caribou (Rangifer tarandus caribou) continued to decline in British Columbia, Canada. On the basis of the cases we examined, we suggest the simultaneous control of predators and alternate prey is the management strategy most likely to increase abundances and probabilities of persistence of rare prey over the long term. Knowing the mechanisms driving changes in species' abundances before implementing any management intervention is critical. We suggest scientists can best contribute to the conservation of species affected by apparent competition by clearly communicating the biological and demographic forces at play to policy makers responsible for the implementation of proposed management actions.

© 2016 by the Ecological Society of America. Encounter competition is interference competition in which animals directly contend for resources. Ecological theory predicts the trait that determines the resource holding potential (RHP), and hence the winner of encounter competition, is most often body size or mass. The difficulties of observing encounter competition in complex organisms in natural environments, however, has limited opportunities to test this theory across diverse species. We studied the outcome of encounter competition contests among mesocarnivores at deer carcasses in California to determine the most important variables for winning these contests. We found some support for current theory in that body mass is important in determining the winner of encounter competition, but we found that other factors including hunger and species-specific traits were also important. In particular, our top models were "strength and hunger" and "size and hunger," with models emphasizing the complexity of variables influencing outcomes of encounter competition. In addition, our wins above predicted (WAP) statistic suggests that an important aspect that determines the winner of encounter competition is species-specific advantages that increase their RHP, as bobcats (Lynx rufus) and spotted skunks (Spilogale gracilis) won more often than predicted based on mass. In complex organisms, such as mesocarnivores, species-specific adaptations, including strategic behaviors, aggressiveness, and weapons, contribute to competitive advantages and may allow certain species to take control or defend resources better than others. Our results help explain how interspecific competition shapes the occurrence patterns of species in ecological communities.

Solitary felids are commonly associated with structurally complex habitats, where their foraging success is attributed to stealth and remaining undetected by competitive scavengers. Research in North America suggests that pumas (Puma concolor), a wide-ranging species found throughout the Americas, conform to the general characteristics of solitary felids and avoid open grasslands with aggregating prey. Researchers hypothesize that pumas are limited to structurally complex habitats in North America because of pressures from other large, terrestrial competitors. We explored the spatial ecology of pumas in open habitat with aggregating prey in Chilean Patagonia, where pumas lack large, terrestrial competitors. We tracked 11 pumas over 30 months (intensive location data for 9 pumas with GPS collars for 9.33 ± 5.66 months each) in an area where mixed steppe grasslands composed 53% of the study area and carried 98% of available prey biomass, to track resource use relative to availability, assess daily movements, quantify home ranges and calculate their density. As determined by location data and kill sites, Patagonia pumas were primarily associated with open habitats with high prey biomass, but at finer scales, preferentially selected for habitat with complex structure. On average, pumas traveled 13.42 ± 2.50 km per day. Estimated 95% fixed kernel home ranges averaged 98 ± 31.8 km2 for females and 211 ± 138.8 km2 for males, with high spatial overlap within and between the sexes. In a multivariate analysis, available prey biomass was the strongest predictor of variation in the size of an individual puma’s home range. Finally, we determined a total puma density of 3.44 pumas/100 km 2 , a significantly smaller estimate than previously reported for

Observations of pumas (Puma concolor) swimming across large bodies of water remain anecdotal in the scientific literature. Here we report long-distance swimming by a foraging male puma in Chilean Patagonia, as revealed by Argos and GPS technology. Our observation raises the question as to what might constitute a barrier to puma movements and gene flow in southern South America. Pumas are a flagship species used in identifying wildlife corridors and in landscape-scale conservation efforts in North America, and we feel a better understanding of puma metapopulation dynamics in South America is essential to both future conservation of the species and landscape-scale conservation efforts in southern South America as well.