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Spatial patterns of and competition for resources by territorial carnivores are typically explained by two hypotheses: 1) the territorial defence hypothesis and 2) the searching efficiency hypothesis. According to the territorial defence hypothesis, when food resources are abundant, carnivore densities will be high and home ranges small. In addition, carnivores can maximise their necessary energy intake with minimal territorial defence. At medium resource levels, larger ranges will be needed, and it will become more economically beneficial to defend resources against a lower density of competitors. At low resource levels, carnivore densities will be low and home ranges large, but resources will be too scarce to make it beneficial to defend such large territories. Thus, home range overlap will be minimal at intermediate carnivore densities. According to the searching efficiency hypothesis, there is a cost to knowing a home range. Larger areas are harder to learn and easier to forget, so carnivores constantly need to keep their cognitive map updated by regularly revisiting parts of their home ranges. Consequently, when resources are scarce, carnivores require larger home ranges to acquire sufficient food. These larger home ranges lead to more overlap among individuals' ranges, so that overlap in home ranges is largest when food availability is the lowest. Since conspecific density is low when food availability is low, this hypothesis predicts that overlap is largest when densities are the lowest. We measured home range overlap and used a novel method to compare intraspecific home range overlaps for lionsPanthera leo(n=149) and leopardsPanthera pardus(n=111) in Africa. We estimated home range sizes from telemetry location data and gathered carnivore density data from the literature. Our results did not support the territorial defence hypothesis for either species. Lion prides increased their home range overlap at conspecific lower densities whereas leopards did not. Lion pride changes in overlap were primarily due to increases in group size at lower densities. By contrast, the unique dispersal strategies of leopards led to reduced overlap at lower densities. However, when human-caused mortality was higher, leopards increased their home range overlap. Although lions and leopards are territorial, their territorial behaviour was less important than the acquisition of food in determining their space use. Such information is crucial for the future conservation of these two iconic African carnivores.

Aim: Macroecological studies that require habitat suitability data for many species often derive this information from expert opinion. However, expert- based informa-tion is inherently subjective and thus prone to errors. The increasing availability of GPS tracking data offers opportunities to evaluate and supplement expert- based in-formation with detailed empirical evidence. Here, we compared expert- based habitat suitability information from the International Union for Conservation of Nature (IUCN) with habitat suitability information derived from GPS-tracking data of 1,498 individu-als from 49 mammal species.Location: Worldwide.Time period: 1998–2021.Major taxa studied: Forty- nine terrestrial mammal species.Methods: Using GPS data, we estimated two measures of habitat suitability for each individual animal: proportional habitat use (proportion of GPS locations within a habi-tat type), and selection ratio (habitat use relative to its availability). For each individual we then evaluated whether the GPS- based habitat suitability measures were in agree-ment with the IUCN data. To that end, we calculated the probability that the ranking of empirical habitat suitability measures was in agreement with IUCN's classification into suitable, marginal and unsuitable habitat types.Results: IUCN habitat suitability data were in accordance with the GPS data (> 95% probability of agreement) for 33 out of 49 species based on proportional habitat use estimates and for 25 out of 49 species based on selection ratios. In addition, 37 and 34 species had a > 50% probability of agreement based on proportional habitat use and selection ratios, respectively.Main conclusions: We show how GPS-tracking data can be used to evaluate IUCN habitat suitability data. Our findings indicate that for the majority of species in-cluded in this study, it is appropriate to use IUCN habitat suitability data in macro-ecological studies. Furthermore, we show that GPS-tracking data can be used to identify and prioritize species and habitat types for re-evaluation of IUCN habitat suitability data. expert opinion, GPS, habitat suitability, habitat type, habitat use, IUCN, mammals, movement, selection ratio, telemetry

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 (

Anthropogenic impacts on wildlife behavior arise both from the immediate presence of people, which induces fear responses in many species, and the human footprint (i.e., landscape modification such as residential development), which affects animal movement and habitat use. Where both disturbance types co-occur, disentangling their impacts remains a challenge. Disturbance effects may interact such that species respond to increased human footprint by either reducing (habituation) or increasing (sensitization) avoidance of human presence. We experimentally manipulated perceived human presence, using playbacks of people talking, across a gradient of human footprint in California’s Santa Cruz Mountains and used camera traps to quantify the behavioral responses of bobcats (Lynx rufus), striped skunks (Mephitis mephitis), and Virginia opossums (Didelphis virginiana), mesopredators known to exhibit strong fear response to human presence but which vary in their use of developed areas. Bobcats and skunks reduced activity in response to human playbacks but showed no change in responsiveness to playbacks across the gradient of human footprint, suggesting that these species are similarly fearful of humans at all development levels tested. Opossums exhibited a significant interaction between human footprint and playback treatment such that reductions in activity level in response to human playbacks were strongest at higher levels of building density. Our results indicate that, rather than habituating to human presence, some mesopredators retain a strong fear of humans or become more fearful when inhabiting more developed areas. We suggest that consistently high responsiveness to immediate human presence may benefit mesopredators living in human-dominated landscapes by mitigating the risk of anthropogenic mortality.

Mitigating human-caused mortality for large carnivores is a pressing global challenge for wildlife conservation. However, mortality is almost exclusively studied at local (within-population) scales creating a mismatch between our understanding of risk and the spatial extent most relevant to conservation and management of wide-ranging species. Here, we quantified mortality for 590 radio-collared mountain lions statewide across their distribution in California to identify drivers of human-caused mortality and investigate whether human-caused mortality is additive or compensatory. Human-caused mortality, primarily from conflict management and vehicles, exceeded natural mortality despite mountain lions being protected from hunting. Our data indicate that human-caused mortality is additive to natural mortality as population-level survival decreased as a function of increasing human-caused mortality and natural mortality did not decrease with increased human-caused mortality. Mortality risk increased for mountain lions closer to rural development and decreased in areas with higher proportions of citizens voting to support environmental initiatives. Thus, the presence of human infrastructure and variation in the mindset of humans sharing landscapes with mountain lions appear to be primary drivers of risk. We show that human-caused mortality can reduce population-level survival of large carnivores across large spatial scales, even when they are protected from hunting.

Puma (Puma concolor) communication with conspecifics is via indirect scent marking behaviours that are important for individuals to advertise their territory and reproductive status, but little is known about how the behaviours develop with age. To examine the development of scent marking behaviours, we monitored the behaviours of adult pumas and dependent kittens. Based on video recordings, we found that the frequency of puma communication behaviours significantly changed over time. Kittens exhibited olfactory investigation more frequently as they aged, but kittens generally did not exhibit scent marking behaviours. Kittens travel with their mothers until they disperse, so there is no need to establish territories or advertise availability to mate, but kittens are at risk of injury or mortality from other pumas. It is possible that there is no functional need for dependent kittens to scent mark until they mature, but there is a need for frequent use of investigative behaviours.

Anellovirus infections are highly prevalent in mammals, however, prior to this study only a handful of anellovirus genomes had been identified in members of the Felidae family. Here we characterise anelloviruses in pumas (Puma concolor), bobcats (Lynx rufus), Canada lynx (Lynx canadensis), caracals (Caracal caracal) and domestic cats (Felis catus). The complete anellovirus genomes (n = 220) recovered from 149 individuals were diverse. ORF1 protein sequence similarity network analysis coupled with phylogenetic analysis, revealed two distinct clusters that are populated by felid-derived anellovirus sequences, a pattern mirroring that observed for the porcine anelloviruses. Of the two-felid dominant anellovirus groups, one includes sequences from bobcats, pumas, domestic cats and an ocelot, and the other includes sequences from caracals, Canada lynx, domestic cats and pumas. Coinfections of diverse anelloviruses appear to be common among the felids. Evidence of recombination, both within and between felid-specific anellovirus groups, supports a long coevolution history between host and virus. ispartof: VIROLOGY vol:562 pages:176-189 ispartof: location:United States status: published

Communication behaviors play a critical role in both an individual’s fitness as well as the viability of populations. Solitary animals use chemical communication (i.e., scent marking) to locate mates and defend their territory to increase their own fitness. Previous research has suggested that striped skunks (Mephitis mephitis) do not perform scent-marking behaviors, despite being best known for using odor as chemical defense. We used video camera traps to document behaviors exhibited by striped skunks at a remote site in coastal California between January 2012 and April 2015. Our camera traps captured a total of 71 visits by striped skunks, the majority of which (73%) included a striped skunk exhibiting scent-marking behaviors. Overall, we documented 8 different scent-marking behaviors. The most frequent behaviors we documented were cheek rubbing (45.1%), investigating (40.8%), and claw marking (35.2%). The behaviors exhibited for the longest durations on average were grooming ( $$ \overline{x} $$ = 34. 4 s) and investigating ( $$ \overline{x} $$ = 21.2 s). Although previous research suggested that striped skunks do not scent mark, we documented that at least some populations do and our findings suggest that certain sites are used for communication via scent marking. Our study further highlights how camera traps allow researchers to discover previously undocumented animal behaviors.

Developed landscapes are increasingly important movement habitat for many large carnivore populations, despite fragmentation and heightened anthropogenic risks. The availability of vegetation cover is a key factor mediating carnivore use of human-dominated landscapes. Restoring or modifying networks of vegetation patches may therefore provide an important tool for enhancing the connectivity value of developed areas, but requires understanding how vegetation patch networks are functionally linked by carnivore movement decisions, which occur at scales considerably finer than those typically addressed by connectivity analyses. We investigated the factors driving fine-scale movement decisions by pumas (Puma concolor) in fragmented habitats and applied our results to enhancing puma connectivity through human-dominated landscapes. We used high-resolution data on vegetation cover and puma locations from central California to model puma habitat selection at the scale of individual movements between vegetation patches. These results informed network-based connectivity models comparing the benefits of specific wildlife corridor restoration actions (e.g., revegetation). Puma movements between vegetation patches were driven by patch size, vegetation type, and spatial arrangement relative to sources of anthropogenic risk (buildings). Pumas avoided buildings but accepted higher building densities as patch area increased or inter-patch travel distances decreased. Connectivity modeling revealed that the strategic placement of vegetation patches can substantially reduce resistance to puma movement across an otherwise high resistance developed landscape by diversifying movement options. Our results reveal the factors mediating large carnivore use of human-dominated landscapes and provide a generalizable tool for increasing movement potential via the manipulation of vegetation cover.

A growing body of evidence has documented how wildlife alter their behavior in response to human encroachment. For carnivores, behaviors related to reproduction and communication are particularly sensitive to human disturbance and can provide an early warning indicator of development’s negative impacts. Despite the important role carnivores play in an ecosystem, few tools have been developed to anticipate how future human development impacts these behaviors. We developed a set of models to understand spatial relationships between anthropogenic development and puma (Puma concolor) habitat selection for two critical reproductive behaviors: nursery habitat for raising young, and sites for communication with mates. Using geospatial location data from the Santa Cruz Mountains in California, USA, we found that female pumas use small nursery home ranges (9 km2 ± 1.72 SE) of predominantly natural habitat, potentially with low levels of human development (

While the functional response of predators is commonly measured, recent work has revealed that the age and sex composition of prey killed is often a better predictor of prey population dynamics because the reproductive value of adult females is usually higher than that of males or juveniles.Climate is often an important mediating factor in determining the composition of predator kills, but we currently lack a mechanistic understanding of how the multiple facets of climate interact with prey abundance and demography to influence the composition of predator kills.Over 20 winters, we monitored 17 wolf packs in Yellowstone National Park and recorded the sex, age and nutritional condition of kills of their dominant prey—elk—in both early and late winter periods when elk are in relatively good and relatively poor condition, respectively.Nutritional condition (as indicated by per cent marrow fat) of wolf‐killed elk varied markedly with summer plant productivity, snow water equivalent (SWE) and winter period. Moreover, marrow was poorer for wolf‐killed bulls and especially for calves than it was for cows.Wolf prey composition was influenced by a complex set of climatic and endogenous variables. In early winter, poor plant growth in either year t or t − 1, or relatively low elk abundance, increased the odds of wolves killing bulls relative to cows. Calves were most likely to get killed when elk abundance was high and when the forage productivity they experienced in utero was poor. In late winter, low SWE and a relatively large elk population increased the odds of wolves killing calves relative to cows, whereas low SWE and poor vegetation productivity 1 year prior together increased the likelihood of wolves killing a bull instead of a cow.Since climate has a strong influence on whether wolves prey on cows (who, depending on their age, are the key reproductive components of the population) or lower reproductive value of calves and bulls, our results suggest that climate can drive wolf predation to be more or less additive from year to year., The impacts of large carnivores on their prey are importantly impacted by the composition of the prey killed. Here the authors show that climate and elk population size interact in important ways to determine when wolves kill cows, bulls or calves.

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 (

Wildlife populations are increasingly challenged by human activities that disrupt landscape connectivity, animal movement, population dynamics and population persistence. Yet modified habitats may provide resource subsidies for generalist species resulting in increased selection of disturbed areas. Understanding how species adjust their space use and activity in human-modified landscapes is fundamental to conserving wildlife populations globally. To test three competing hypotheses explaining spatiotemporal responses to human activity, we investigated coarse (3 hour)- and fine (20 minute)- scale habitat selection, activity patterns, and measured home ranges in a human-dominated landscape. We GPS-collared 25 adults and subadult caracals (Caracal caracal) from three contiguous subpopulations in urban and wildland-dominated regions. Caracals in the wildland-dominated region avoided proximity to urban areas while caracals in the urban-dominated region selected for close proximity to urban areas. Selection responses for vegetative cover, freshwater, vineyards, the coast, and burned areas was dependent on age class, diel period, and whether individuals were in close proximity to urban areas. Large home ranges demonstrated habitat selection results were not a result of limited landscape use. Caracals did not shift their temporal activity to avoid humans, but selected for closer proximity to urban areas at night, suggesting they seek spatial refugia during the day. Behavioral plasticity whereby individuals seek out spatial refugia promotes the coexistence of this carnivore in a human-dominated landscape. These data are essential to our understanding of what is viable wildlife habitat to a generalist mesocarnivore and provide opportunities to guide land acquisition and conservation by local management agencies.

Home range size is a fundamental measure of animal space use, providing insight into habitat quality, animal density, and social organization. Human impacts increasingly are affecting wildlife, especially among wide-ranging species that encounter anthropogenic disturbance. Leopards (Panthera pardus) provide a useful model for studying this relationship because leopards coexist with people at high and low human densities and are sensitive to human disturbance. To compare leopard home range size across a range of human densities and other environmental conditions, we combined animal tracking data from 74 leopards in multiple studies with new analytical techniques that accommodate different sampling regimes. We predicted that home ranges would be smaller in more productive habitats and areas of higher human population density due to possible linkage with leopard prey subsidies from domestic species. We also predicted that male leopards would have larger home ranges than those of females. Home ranges varied in size from 14.5 km2 in India to 885.6 km2 in Namibia, representing a 60-fold magnitude of variation. Home range stability was evident for 95.2% of nontranslocated individuals and 38.5% of translocated individuals. Leopard home range sizes were negatively correlated with landscape productivity, and males used larger areas than females. Leopards in open habitats had a predicted negative correlation in home range size with human population density, but leopards in closed habitats used larger home ranges in areas with more people.

When navigating heterogeneous landscapes, large carnivores must balance trade-offs between multiple goals, including minimizing energetic expenditure, maintaining access to hunting opportunities and avoiding potential risk from humans. The relative importance of these goals in driving carnivore movement likely changes across temporal scales, but our understanding of these dynamics remains limited. Here we quantified how drivers of movement and habitat selection changed with temporal grain for two large carnivore species living in human-dominated landscapes, providing insights into commonalities in carnivore movement strategies across regions. We used high-resolution GPS collar data and integrated step selection analyses to model movement and habitat selection for African lions Panthera leo in Laikipia, Kenya and pumas Puma concolor in the Santa Cruz Mountains of California across eight temporal grains, ranging from 5 min to 12 hr. Analyses considered landscape covariates that are related to energetics, resource acquisition and anthropogenic risk. For both species, topographic slope, which strongly influences energetic expenditure, drove habitat selection and movement patterns over fine temporal grains but was less important at longer temporal grains. In contrast, avoiding anthropogenic risk during the day, when risk was highest, was consistently important across grains, but the degree to which carnivores relaxed this avoidance at night was strongest for longer term movements. Lions and pumas modified their movement behaviour differently in response to anthropogenic features: lions sped up while near humans at fine temporal grains, while pumas slowed down in more developed areas at coarse temporal grains. Finally, pumas experienced a trade-off between energetically efficient movement and avoiding anthropogenic risk. Temporal grain is an important methodological consideration in habitat selection analyses, as drivers of both movement and habitat selection changed across temporal grain. Additionally, grain-dependent patterns can reflect meaningful behavioural processes, including how fitness-relevant goals influence behaviour over different periods of time. In applying multi-scale analysis to fine-resolution data, we showed that two large carnivore species in very different human-dominated landscapes balanced competing energetic and safety demands in largely similar ways. These commonalities suggest general strategies of landscape use across large carnivore species.

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.

Background: Wildlife populations are increasingly challenged by human activities that disrupt landscape connectivity and animal movement, and thus population dynamics and persistence. Yet modified habitats may provide resource subsidies for generalist species resulting in increased selection of disturbed areas. Understanding how species adjust their space use and activity in human-modified landscapes is increasingly critical to conserving wildlife populations globally. Methods: To test three competing hypotheses explaining spatiotemporal responses to human activity, we GPS-collared 26 individual adults and subadults from three contiguous subpopulations in wildland and urban-dominated regions. We investigated coarse (3 hour)- and fine (20 minute)- scale movement-explicit habitat selection using step selection functions and tested for increased nocturnality of caracals (Caracal caracal) in a human-dominated landscape in South Africa. Results: Caracals in a wildland-dominated landscape were on average > 8.0 km from the urban-natural boundary and strongly selected for proximity to the coastline. Caracals in the urban-dominated landscape were on average Conclusions: Behavioral plasticity whereby individuals seek out spatial, but not temporal refugia, appears to promote the coexistence of this generalist carnivore in a human-dominated landscape. Both age class and degree of exposure to anthropogenic activities influence habitat selection, while some individuals have learned to successfully exploit the riskiest areas at night. These data are essential to our understanding of what is viable wildlife habitat in this rapidly urbanizing landscape and provide opportunities to guide land acquisition and conservation guidelines by local management agencies.

Energetic demands and fear of predators are considered primary factors shaping animal behavior, and both are likely drivers of movement decisions that ultimately determine the spatial ecology of wildlife. Yet energetic constraints on movement imposed by the physical landscape have only been considered separately from those imposed by risk avoidance, limiting our understanding of how short-term movement decisions scale up to affect long-term space use. Here, we integrate the costs of both physical terrain and predation risk into a common currency, energy, and then quantify their effects on the short-term movement and long-term spatial ecology of a large carnivore living in a human-dominated landscape. Using high-resolution GPS and accelerometer data from collared pumas (Puma concolor), we calculated the short-term (i.e., 5-min) energetic costs of navigating both rugged physical terrain and a landscape of risk from humans (major sources of both mortality and fear for our study population). Both the physical and risk landscapes affected puma short-term movement costs, with risk having a relatively greater impact by inducing high-energy but low-efficiency movement behavior. The cumulative effects of short-term movement costs led to reductions of 29% to 68% in daily travel distances and total home range area. For male pumas, long-term patterns of space use were predominantly driven by the energetic costs of human-induced risk. This work demonstrates that, along with physical terrain, predation risk plays a primary role in shaping an animal's "energy landscape" and suggests that fear of humans may be a major factor affecting wildlife movements worldwide.

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.

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 widely studied yet a biogeographical 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 variation across their range. Kill rate studies on pumas, while numerous, were primarily conducted in Temperate Conifer Forests (< 10% of puma range), revealing a dearth of knowledge across much of their range, especially from tropical and subtropical habitats. Across studies, kill rates in ungulates/week were highest for adult females with kitten(s) (1.24 ± 0.41 ungulates/week) but did not vary significantly between adult males (0.84 ± 0.18) and solitary adult females (0.99 ± 0.26). Kill rates in kg/day did not differ significantly among reproductive classes. Kill rates of adult pumas increased with ungulate density. Ungulate species richness had a weak negative association with adult male kill rates. Neither scavenger richness, the proportion of non-ungulate prey in the diet, nor regional human population density had a significant effect on ungulate kill rates. Our results had a strong temperate-ecosystem bias highlighting the need for further research across the diverse biomes pumas occupy in order to make species level inferences. Data from more populations would also allow for multivariate analyses providing deeper inference into the ecological and behavioural factors driving kill rates and functional responses of pumas, and apex predators in general.

Background Under current scenarios of climate change and habitat loss, many wild animals, especially large predators, are moving into novel energetically challenging environments. Consequently, changes in terrain associated with such moves may heighten energetic costs and effect the decline of populations in new localities. Methods To examine locomotor costs of a large carnivorous mammal moving in mountainous habitats, the oxygen consumption of captive pumas (Puma concolor) was measured during treadmill locomotion on level and incline (6.8°) surfaces. These data were used to predict energetic costs of locomotor behaviours of free-ranging pumas equipped with GPS/accelerometer collars in California’s Santa Cruz Mountains. Results Incline walking resulted in a 42.0% ± 7.2 SEM increase in the costs of transport compared to level performance. Pumas negotiated steep terrain by traversing across hillsides (mean hill incline 17.2° ± 0.3 SEM; mean path incline 7.3° ± 0.1 SEM). Pumas also walked more slowly up steeper paths, thereby minimizing the energetic impact of vertical terrains. Estimated daily energy expenditure (DEE) based on GPS-derived speeds of free-ranging pumas was 18.3 MJ day− 1 ± 0.2 SEM. Calculations show that a 20 degree increase in mean steepness of the terrain would increase puma DEE by less than 1% as they only spend a small proportion (10%) of their day travelling. They also avoided elevated costs by utilizing slower speeds and shallower path angles. Conclusions While many factors influence survival in novel habitats, we illustrate the importance of behaviours which reduce locomotor costs when traversing new, energetically challenging environments, and demonstrate that these behaviours are utilised by pumas in the wild.

Understanding the distribution of biodiversity across the Earth is one of the most challenging questions in biology. Much research has been directed at explaining the species latitudinal pattern showing that communities are richer in tropical areas; however, despite decades of research, a general consensus has not yet emerged. In addition, global biodiversity patterns are being rapidly altered by human activities. Here, we aim to describe large-scale patterns of species richness and diversity in terrestrial vertebrate scavenger (carrion-consuming) assemblages, which provide key ecosystem functions and services. We used a worldwide dataset comprising 43 sites, where vertebrate scavenger assemblages were identified using 2,485 carcasses monitored between 1991 and 2018. First, we evaluated how scavenger richness (number of species) and diversity (Shannon diversity index) varied among seasons (cold vs. warm, wet vs. dry). Then, we studied the potential effects of human impact and a set of macroecological variables related to climatic conditions on the scavenger assemblages. Vertebrate scavenger richness ranged from species-poor to species rich assemblages (4–30 species). Both scavenger richness and diversity also showed some seasonal variation. However, in general, climatic variables did not drive latitudinal patterns, as scavenger richness and diversity were not affected by temperature or rainfall. Rainfall seasonality slightly increased the number of species in the community, but its effect was weak. Instead, the human impact index included in our study was the main predictor of scavenger richness. Scavenger assemblages in highly human-impacted areas sustained the smallest number of scavenger species, suggesting human activity may be overriding other macroecological processes in shaping scavenger communities. Our results highlight the effect of human impact at a global scale. As species-rich assemblages tend to be more functional, we warn about possible reductions in ecosystem functions and the services provided by scavengers in human-dominated landscapes in the Anthropocene., SG, JMPG, and ACA were supported by Juan de la Cierva contracts (Ministerio de Economía y Competitividad, MEC; IJCI‐2015‐24947, FJCI‐2015‐25632, and IJCI‐2014‐20744, respectively). ESG was also supported by Generalitat Valenciana (SEJI/2018/024), ACA by the Govern de les Illes Balears (PD/039/2017), and MM by a Ramón y Cajal contract (MEC; 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). SAL thanks PICT (BID) 0725/2014. MK and KJ were supported by the Slovenian Research Agency (P4‐0059). Contributions of KT and JCB were partially supported through funding from U.S. Department of Agriculture, the U.S. Department of Energy under (DE‐EM0004391) to the University of Georgia Research Foundation. EB and EF were supported by the USA National Science Foundation. CK completed study under research permit NCST/5/002/R/448 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 and the Gordon and Betty Moore Foundation. Several authors were funded by funds from the MEC (CGL2012‐40013‐C02‐01/02, CGL2015‐66966‐C2‐1‐R and CGL2017‐89905‐R) and from the Junta de Andalucía (RNM‐1925).

Understanding the behavioral responses of large carnivores to human activity in protected areas is important for conserving top predators. Roads and associated vehicle traffic have a range of impacts on wildlife, including mortality from vehicle collisions and behavioral changes from increasing traffic levels. Roads concentrate human activities and may be particularly impactful when located adjacent to high‐quality habitat for wildlife. However, people often overlook road impacts in protected areas because of relatively low road densities. From 1979 to 2017, annual visits to Yellowstone National Park increased from 1.9 to 4.1 million, with many visitors in the last 25 yr focusing on the opportunity to view wild wolves (Canis lupus) in their natural habitat from the roadway. To better understand how human activity interacts with landscape attributes and prey availability to shape wolf habitat use, we developed seasonal and diel‐specific step selection functions (SSF) for wolves. Wolves responded to increased human activity by using areas farther from roadways during the day and during peak visitation in summer. Prey availability, as estimated by an elk SSF, did not significantly alter habitat selection patterns by wolves. The strength of habitat selection in relation to roads varied among wolf packs. The most heavily viewed wolf packs exhibited less road avoidance, suggesting increased tolerance, which could lead to increased vulnerability to human harvest if they leave the park. Federal and state managers have implemented several measures to mitigate disturbance effects to wolves and curtail habituation. These results may inform adaptive management strategies that seek to continue to conserve natural wolf behavior.