The ecology of human-caused mortality for a protected large carnivore.

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.

Mesocarnivores vary in their spatiotemporal avoidance strategies at communications hubs of an apex carnivore.

Mesocarnivores face interspecific competition and risk intraguild predation when sharing resources with apex carnivores. Within a landscape, carnivores across trophic levels may use the same communication hubs, which provide a mix of risks (injury/death) and rewards (gaining information) for subordinate species. We predicted that mesocarnivores would employ different strategies to avoid apex carnivores at shared communication hubs, depending on their trophic position. To test our prediction, we examined how different subordinate carnivore species in the Santa Cruz Mountains of California, USA, manage spatial overlap with pumas (Puma concolor), both at communication hubs and across a landscape-level camera trap array. We estimated species-specific occurrence, visitation rates, temporal overlap, and Avoidance-Attraction Ratios from camera traps and tested for differences between the two types of sites. We found that mesocarnivores generally avoided pumas at communication hubs, and this became more pronounced when pumas scent-marked during their most recent visit. Coyotes (Canis latrans), the pumas' closest subordinate competitor in our system, exhibited the strongest avoidance at communication hubs. Gray foxes (Urocyon cinereoargenteus) avoided pumas the least, which may suggest possible benefits from pumas suppressing coyotes. Overall, mesocarnivores exhibited various spatiotemporal avoidance strategies at communication hubs rather than outright avoidance, likely because they benefit from information gained while 'eavesdropping' on puma activity. Variability in avoidance strategies may be due to differential predation risks, as apex carnivores often interact more aggressively with their closest competitors. Combined, our results show how apex carnivores trigger complex species interactions across the entire carnivore guild and how trophic position determines behavioral responses and subsequent space use of subordinate mesocarnivores across the landscape.

Energetics and fear of humans constrain the spatial ecology of pumas.

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.

Habitat fragmentation reduces survival and drives source-sink dynamics for a large carnivore.

Rigorous understanding of how environmental conditions impact population dynamics is essential for species conservation, especially in mixed-use landscapes where source-sink dynamics may be at play. Conservation of large carnivore populations in fragmented, human-dominated landscapes is critical for their long-term persistence. However, living in human-dominated landscapes comes with myriad costs, including direct anthropogenic mortality and sublethal energetic costs. How these costs impact individual fitness and population dynamics are not fully understood, partly due to the difficulty in collecting long-term demographic data for these species. Here, we analyzed an 11-year dataset on puma (Puma concolor) space use, mortality, and reproduction in the Santa Cruz Mountains, California, USA, to quantify how living in a fragmented landscape impacts individual survival and population dynamics. Long-term exposure to housing density drove mortality risk for female pumas, resulting in an 18-percentage-point reduction in annual survival for females in exurban versus remote areas. While the overall population growth rate appeared stable, reduced female survival in more developed areas resulted in source-sink dynamics across the study area, with 42.1% of the Santa Cruz Mountains exhibiting estimated population growth rates <1. Since habitat selection is often used as a proxy for habitat quality, we also assessed whether puma habitat selection predicted source and sink areas. Patterns of daytime puma habitat selection predicted source areas, while time-of-day-independent habitat selection performed less well as a proxy. These results illuminate the individual- and population-level consequences of habitat fragmentation for large carnivores, illustrating that habitat fragmentation can produce source-sink dynamics that may not be apparent from other metrics of habitat quality. Locally, conserving high-quality source habitat within the Santa Cruz Mountains is necessary to support long-term puma population persistence. More broadly, source-sink dynamics may at play for other carnivore populations in similar fragmented systems, and linking landscape conditions to population dynamics is essential for effective conservation. Caution should be used in inferring habitat quality from habitat selection alone, but these results shed light on metrics of selection that may be better or worse proxies to identify source areas for large carnivores.

Temporal scale of habitat selection for large carnivores: Balancing energetics, risk and finding prey.

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.

Disturbance type and species life history predict mammal responses to humans.

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.

Mixed-species herding levels the landscape of fear.

Prey anti-predator behaviours are influenced by perceived predation risk in a landscape and social information gleaned from herd mates regarding predation risk. It is well documented that high-quality social information about risk can come from heterospecific herd mates. Here, we integrate social information with the landscape of fear to quantify how these landscapes are modified by mixed-species herding. To do this, we investigated zebra vigilance in single- and mixed-species herds across different levels of predation risk (lion versus no lion), and assessed how they manage herd size and the competition-information trade-off associated with grouping behaviour. Overall, zebra performed higher vigilance in high-risk areas. However, mixed-species herding reduced vigilance levels. We estimate that zebra in single-species herds would have to feed for approximately 35 min more per day in low-risk areas and approximately 51 min more in high-risk areas to compensate for the cost of higher vigilance. Furthermore, zebra benefitted from the competition-information trade-off by increasing the number of heterospecifics while keeping the number of zebra in a herd constant. Ultimately, we show that mixed-species herding reduces the effects of predation risk, whereby zebra in mixed-species herds, under high predation risk, perform similar levels of vigilance compared with zebra in low-risk scenarios.

How climate impacts the composition of wolf-killed elk in northern Yellowstone National Park.

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.

Fear of humans as apex predators has landscape-scale impacts from mountain lions to mice.

Apex predators such as large carnivores can have cascading, landscape-scale impacts across wildlife communities, which could result largely from the fear they inspire, although this has yet to be experimentally demonstrated. Humans have supplanted large carnivores as apex predators in many systems, and similarly pervasive impacts may now result from fear of the human 'super predator'. We conducted a landscape-scale playback experiment demonstrating that the sound of humans speaking generates a landscape of fear with pervasive effects across wildlife communities. Large carnivores avoided human voices and moved more cautiously when hearing humans, while medium-sized carnivores became more elusive and reduced foraging. Small mammals evidently benefited, increasing habitat use and foraging. Thus, just the sound of a predator can have landscape-scale effects at multiple trophic levels. Our results indicate that many of the globally observed impacts on wildlife attributed to anthropogenic activity may be explained by fear of humans.

Scavenging in the Anthropocene: Human impact drives vertebrate scavenger species richness at a global scale.

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.

Fear of the human 'super predator' reduces feeding time in large carnivores.

Large carnivores' fear of the human 'super predator' has the potential to alter their feeding behaviour and result in human-induced trophic cascades. However, it has yet to be experimentally tested if large carnivores perceive humans as predators and react strongly enough to have cascading effects on their prey. We conducted a predator playback experiment exposing pumas to predator (human) and non-predator control (frog) sounds at puma feeding sites to measure immediate fear responses to humans and the subsequent impacts on feeding. We found that pumas fled more frequently, took longer to return, and reduced their overall feeding time by more than half in response to hearing the human 'super predator'. Combined with our previous work showing higher kill rates of deer in more urbanized landscapes, this study reveals that fear is the mechanism driving an ecological cascade from humans to increased puma predation on deer. By demonstrating that the fear of humans can cause a strong reduction in feeding by pumas, our results support that non-consumptive forms of human disturbance may alter the ecological role of large carnivores.

The importance of motivation, weapons, and foul odors in driving encounter competition in carnivores.

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.

The comparative effects of large carnivores on the acquisition of carrion by scavengers.

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.

Top carnivores increase their kill rates on prey as a response to human-induced fear.

The fear induced by predators on their prey is well known to cause behavioural adjustments by prey that can ripple through food webs. Little is known, however, about the analogous impacts of humans as perceived top predators on the foraging behaviour of carnivores. Here, we investigate the influence of human-induced fear on puma foraging behaviour using location and prey consumption data from 30 tagged individuals living along a gradient of human development. We observed strong behavioural responses by female pumas to human development, whereby their fidelity to kill sites and overall consumption time of prey declined with increasing housing density by 36 and 42%, respectively. Females responded to this decline in prey consumption time by increasing the number of deer they killed in high housing density areas by 36% over what they killed in areas with little residential development. The loss of food from declines in prey consumption time paired with increases in energetic costs associated with killing more prey may have consequences for puma populations, particularly with regard to reproductive success. In addition, greater carcass availability is likely to alter community dynamics by augmenting food resources for scavengers. In light of the extensive and growing impact of habitat modification, our study emphasizes that knowledge of the indirect effects of human activity on animal behaviour is a necessary component in understanding anthropogenic impacts on community dynamics and food web function.

The golden age of bio-logging: how animal-borne sensors are advancing the frontiers of ecology.

Great leaps forward in scientific understanding are often spurred by innovations in technology. The explosion of miniature sensors that are driving the boom in consumer electronics, such as smart phones, gaming platforms, and wearable fitness devices, are now becoming available to ecologists for remotely monitoring the activities of wild animals. While half a century ago researchers were attaching balloons to the backs of seals to measure their movement, today ecologists have access to an arsenal of sensors that can continuously measure most aspects of an animal's state (e.g., location, behavior, caloric expenditure, interactions with other animals) and external environment (e.g., temperature, salinity, depth). This technology is advancing our ability to study animal ecology by allowing researchers to (1) answer questions about the physiology, behavior, and ecology of wild animals in situ that would have previously been limited to tests on model organisms in highly controlled settings, (2) study cryptic or wide-ranging animals that have previously evaded investigation, and (3) develop and test entirely new theories. Here we explore how ecologists are using these tools to answer new questions about the physiological performance, energetics, foraging, migration, habitat selection, and sociality of wild animals, as well as collect data on the environments in which they live.

Using grizzly bears to assess harvest-ecosystem tradeoffs in salmon fisheries.

Implementation of ecosystem-based fisheries management (EBFM) requires a clear conceptual and quantitative framework for assessing how different harvest options can modify benefits to ecosystem and human beneficiaries. We address this social-ecological need for Pacific salmon fisheries, which are economically valuable but intercept much of the annual pulse of nutrient subsidies that salmon provide to terrestrial and aquatic food webs. We used grizzly bears, vectors of salmon nutrients and animals with densities strongly coupled to salmon abundance, as surrogates for "salmon ecosystem" function. Combining salmon biomass and stock-recruitment data with stable isotope analysis, we assess potential tradeoffs between fishery yields and bear population densities for six sockeye salmon stocks in Bristol Bay, Alaska, and British Columbia (BC), Canada. For the coastal stocks, we find that both bear densities and fishery yields would increase substantially if ecosystem allocations of salmon increase from currently applied lower to upper goals and beyond. This aligning of benefits comes at a potential cost, however, with the possibility of forgoing harvests in low productivity years. In contrast, we detect acute tradeoffs between bear densities and fishery yields in interior stocks within the Fraser River, BC, where biomass from other salmon species is low. There, increasing salmon allocations to ecosystems would benefit threatened bear populations at the cost of reduced long-term yields. To resolve this conflict, we propose an EBFM goal that values fisheries and bears (and by extension, the ecosystem) equally. At such targets, ecosystem benefits are unexpectedly large compared with losses in fishery yields. To explore other management options, we generate tradeoff curves that provide stock-specific accounting of the expected loss to fishers and gain to bears as more salmon escape the fishery. Our approach, modified to suit multiple scenarios, provides a generalizable method to resolve conflicts over shared resources in other systems.

Deer, predators, and the emergence of Lyme disease.

Lyme disease is the most prevalent vector-borne disease in North America, and both the annual incidence and geographic range are increasing. The emergence of Lyme disease has been attributed to a century-long recovery of deer, an important reproductive host for adult ticks. However, a growing body of evidence suggests that Lyme disease risk may now be more dynamically linked to fluctuations in the abundance of small-mammal hosts that are thought to infect the majority of ticks. The continuing and rapid increase in Lyme disease over the past two decades, long after the recolonization of deer, suggests that other factors, including changes in the ecology of small-mammal hosts may be responsible for the continuing emergence of Lyme disease. We present a theoretical model that illustrates how reductions in small-mammal predators can sharply increase Lyme disease risk. We then show that increases in Lyme disease in the northeastern and midwestern United States over the past three decades are frequently uncorrelated with deer abundance and instead coincide with a range-wide decline of a key small-mammal predator, the red fox, likely due to expansion of coyote populations. Further, across four states we find poor spatial correlation between deer abundance and Lyme disease incidence, but coyote abundance and fox rarity effectively predict the spatial distribution of Lyme disease in New York. These results suggest that changes in predator communities may have cascading impacts that facilitate the emergence of zoonotic diseases, the vast majority of which rely on hosts that occupy low trophic levels.

How territoriality and sociality influence the habitat selection and movements of a large carnivore.

While territoriality is one of the key mechanisms influencing carnivore space use, most studies quantify resource selection and movement in the absence of conspecific influence or territorial structure. Our analysis incorporated social information in a resource selection framework to investigate mechanisms of territoriality and intra-specific competition on the habitat selection of a large, social carnivore. We fit integrated step selection functions to 3-h GPS data from 12 collared African wild dog packs in the Okavango Delta and estimated selection coefficients using a conditional Poisson likelihood with random effects. Packs selected for their neighbors' 30-day boundary (defined as their 95% kernel density estimate) and for their own 90-day core (defined as their 50% kernel density estimate). Neighbors' 30-day boundary had a greater influence on resource selection than any habitat feature. Habitat selection differed when they were within versus beyond their neighbors' 30-day boundary. Pack size, pack tenure, pup presence, and seasonality all mediated how packs responded to neighbors' space use, and seasonal dynamics altered the strength of residency. While newly-formed packs and packs with pups avoided their neighbors' boundary, older packs and those without pups selected for it. Packs also selected for the boundary of larger neighboring packs more strongly than that of smaller ones. Social structure within packs has implications for how they interact with conspecifics, and therefore how they are distributed across the landscape. Future research should continue to investigate how territorial processes are mediated by social dynamics and, in turn, how territorial structure mediates resource selection and movement. These results could inform the development of a human-wildlife conflict (HWC) mitigation tool by co-opting the mechanisms of conspecific interactions to manage space use of endangered carnivores.

Season, prey availability, sex, and age explain prey size selection in a large solitary carnivore.

Prey selection is a fundamental aspect of ecology that drives evolution and community structure, yet the impact of intraspecific variation on the selection for prey size remains largely unaccounted for in ecological theory. Here, we explored puma (Puma concolor) prey selection across six study sites in North and South America. Our results highlighted the strong influence of season and prey availability on puma prey selection, and the smaller influence of puma age. Pumas in all sites selected smaller prey in warmer seasons following the ungulate birth pulse. Our top models included interaction terms between sex and age, suggesting that males more than females select larger prey as they age, which may reflect experiential learning. When accounting for variable sampling across pumas in our six sites, male and female pumas killed prey of equivalent size, even though males are larger than females, challenging assumptions about this species. Nevertheless, pumas in different study sites selected prey of different sizes, emphasizing that the optimal prey size for pumas is likely context-dependent and affected by prey availability. The mean prey weight across all sites averaged 1.18 times mean puma weight, which was less than predicted as the optimal prey size by energetics and ecological theory (optimal prey = 1.45 puma weight). Our results help refine our understanding of optimal prey for pumas and other solitary carnivores, as well as corroborate recent research emphasizing that carnivore prey selection is impacted not just by energetics but by the effects of diverse ecology.

Anthropogenic activities and age class mediate carnivore habitat selection in a human-dominated landscape.

Human activities increasingly challenge wild animal populations by disrupting ecological connectivity and population persistence. Yet, human-modified habitats can provide resources, resulting in selection of disturbed areas by generalist species. To investigate spatial and temporal responses of a generalist carnivore to human disturbance, we investigated habitat selection and diel activity patterns in caracals (Caracal caracal). We GPS-collared 25 adults and subadults in urban and wildland-dominated subregions in Cape Town, South Africa. Selection responses for landscape variables were dependent on subregion, animal age class, and diel period. Contrary to expectations, caracals did not become more nocturnal in urban areas. Caracals increased their selection for proximity to urban areas as the proportion of urban area increased. Differences in habitat selection between urban and wildland caracals suggest that individuals of this generalist species exhibit high behavioral flexibility in response to anthropogenic disturbances that emerge as a function of habitat context.