Summer predation rates on ungulate prey by a large keystone predator: how many ungulates does a large predator kill?

Estimates of predation rates by large predators can provide valuable information on their potential impact on their ungulate prey populations. This is especially the case for pumas Puma concolor and its main prey, mule deer Odocoileus hemionus . However, only limited information on predation rates of pumas exist where mule deer are the only ungulate prey available. I used VHF telemetry data collected over 24-h monitoring sessions and once daily over consecutive days to derive two independent estimates of puma predation rates on mule deer where they were the only large prey available. For the 24-h data, I had 48 time blocks on female pumas with kittens, 43 blocks on females without kittens and 30 blocks on males. For the daily consecutive data, the average number of consecutive days followed was 51.5±4.2 days. There were data on five female pumas with kittens, five pregnant females and nine females without kittens. Predation rates over an average month of 30 days from the 24-h monitoring sessions were 2.0 mule deer per puma month for males (15.1 days per kill), 2.1 mule deer per puma month (14.3 days per kill) for females without kittens and 2.5 mule deer per puma month (12.0 days per kill) for pregnant females and females with kittens. For the consecutive daily data, females without kittens had an estimated predation rate of 2.1±0.14 mule deer per puma month (14.9±0.90 days per kill). Pregnant and females with kittens had predation rates of 2.7±0.18 and 2.6±0.21 mule deer per puma month, respectively (11.4±0.72 and 12.0±1.1 days per kill, respectively). Predation rates estimated in this study compared with those estimated by energetic demand for pumas in the study area but were lower than other field derived estimates. These data help increase our understanding of predation impacts of large predators on their prey.     

Surplus killing by pumas Puma concolor: rumours and facts

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Reintroduced wolves and hunting limit the abundance of a subordinate apex predator in a multi-use landscape

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.     

Numerical and Demographic Responses of Pumas to Changes in Prey Abundance: Testing Current Predictions

ABSTRACT Information on factors affecting population size of pumas (Puma concolor) can be important because their principal prey over most of the western United States are valued big game species (e.g., mule deer [Odocoileus hemionus], elk [Cervus elaphus], and bighorn sheep [Ovis canadensis]). Based on the hypothesis that puma numbers are limited by their food supply, puma populations should track changes in prey abundance by growing exponentially with increases in prey and by declining with a lag response when prey decreases. Additional predictions proposed by researchers are that body mass of pumas, female productivity, kitten survival, and adult survival should decrease after a prey decline. We used a 15-year database from a hunted population of pumas in southern Idaho and northwestern Utah to test these predictions. During the 15-year time span of the database, a major decline in mule deer abundance occurred. Estimates of puma numbers and demographic characteristics came from intensive capture and radiocollaring efforts. We calculated kitten and adult survival with MICROMORT software. We found that adult puma numbers increased exponentially at r = 0.07 during a period of increasing mule deer numbers. Four years after the mule deer abundance declined, puma numbers decreased at a rate of r = −0.06. Body mass of female pumas was lower after the decline in puma numbers (42.6 ± SE = 1.2 kg, n = 40 vs. 40.1 ± 0.64 kg, n = 34, t = 5.06, P = 0.045). Kitten survival was less after the decline in deer abundance (0.573 ± 0.016, n = 30 vs. 0.856 ± 0.015, n = 25, Z = 2.40, P < 0.01). Survival of resident females was significantly less after the decline in puma numbers (0.783 ± 0.03 vs. 0.929 ± 0.019, U = 55.0, P = 0.009). Female productivity did not differ before or after the decline in deer abundance. Our results supported the majority of the predictions concerning the impact of changing deer abundance, which supported the hypothesis that the abundance of mule deer limited our population of pumas.     

Can't bear the competition: Energetic losses from kleptoparasitism by a dominant scavenger may alter foraging behaviors of an apex predator

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.     

Nuisance Ecology: Do Scavenging Condors Exact Foraging Costs on Pumas in Patagonia?

Predation risk describes the energetic cost an animal suffers when making a trade off between maximizing energy intake and minimizing threats to its survival. We tested whether Andean condors (Vultur gryphus) influenced the foraging behaviors of a top predator in Patagonia, the puma (Puma concolor), in ways comparable to direct risks of predation for prey to address three questions: 1) Do condors exact a foraging cost on pumas?; 2) If so, do pumas exhibit behaviors indicative of these risks?; and 3) Do pumas display predictable behaviors associated with prey species foraging in risky environments? Using GPS location data, we located 433 kill sites of 9 pumas and quantified their kill rates. Based upon time pumas spent at a carcass, we quantified handling time. Pumas abandoned >10% of edible meat at 133 of 266 large carcasses after a single night, and did so most often in open grasslands where their carcasses were easily detected by condors. Our data suggested that condors exacted foraging costs on pumas by significantly decreasing puma handling times at carcasses, and that pumas increased their kill rates by 50% relative to those reported for North America to compensate for these losses. Finally, we determined that the relative risks of detection and associated harassment by condors, rather than prey densities, explained puma “giving up times” (GUTs) across structurally variable risk classes in the study area, and that, like many prey species, pumas disproportionately hunted in high-risk, high-resource reward areas.     

Puma population limitation and regulation: What matters in puma management?

Wildlife managers require reliable information on factors that influence animal populations to develop successful management programs, including the puma (Puma concolor), in western North America. As puma populations have recovered in recent decades because of restrictions on human-caused mortality, managers need a clear understanding of the factors that limit or regulate puma populations and how those factors might be manipulated to achieve management objectives, including sustaining puma and other wildlife populations, providing hunting opportunity, and reducing puma interactions with people. I synthesized technical literature on puma populations, behavior, and relationships with prey that have contributed to hypotheses on puma population limitation and regulation. Current hypotheses on puma population limitation include the social limitation hypothesis and the food limitation hypothesis. Associated with each of those are 2 hypotheses on puma population regulation: the social regulation hypothesis and the competition regulation hypothesis. I organize the biological and ecological attributes of pumas reported in the literature under these hypotheses. I discuss the validity of these hypotheses based on the limits of the research associated with the hypotheses and the evolutionary processes theoretically underlying them. I review the management predictions as framed by these hypotheses as they pertain to puma hunting, puma-prey relationships, and human-puma interactions. The food limitation and competition regulation hypotheses explain more phenomena associated with puma and likely would guide more successful management outcomes. © 2019 The Wildlife Society.     

Dietary separation of sympatric carnivores identified by molecular analysis of scats

We studied the diets of four sympatric carnivores in the flooding savannas of western Venezuela by analysing predator DNA and prey remains in faeces. DNA was isolated and a portion of the cytochrome b gene of the mitochondrial genome amplified and sequenced from 20 of 34 scats. Species were diagnosed by comparing the resulting sequences to reference sequences generated from the blood of puma (Puma concolor), jaguar (Panthera onca), ocelot (Leopardus pardalus) and crab-eating fox (Cerdocyon thous). Scat size has previously been used to identify predators, but DNA data show that puma and jaguar scats overlap in size, as do those of puma, ocelot and fox. Prey-content analysis suggests minimal prey partitioning between pumas and jaguars. In field testing this technique for large carnivores, two potential limitations emerged: locating intact faecal samples and recovering DNA sequences from samples obtained in the wet season. Nonetheless, this study illustrates the tremendous potential of DNA faecal studies. The presence of domestic dog (Canis familiaris) in one puma scat and of wild pig (Sus scrofa), set as bait, in one jaguar sample exemplifies the forensic possibilities of this noninvasive analysis. In addition to defining the dietary habits of similar size sympatric mammals, DNA identifications from faeces allow wildlife managers to detect the presence of endangered taxa and manage prey for their conservation.     

Feline immunodeficiency virus in puma: Estimation of force of infection reveals insights into transmission

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Linking anti‐predator behaviour to prey demography reveals limited risk effects of an actively hunting large carnivore

Ecological theory predicts that the diffuse risk cues generated by wide‐ranging, active predators should induce prey behavioural responses but not major, population‐ or community‐level consequences. We evaluated the non‐consumptive effects (NCEs) of an active predator, the grey wolf (Canis lupus), by simultaneously tracking wolves and the behaviour, body fat, and pregnancy of elk (Cervus elaphus), their primary prey in the Greater Yellowstone Ecosystem. When wolves approached within 1 km, elk increased their rates of movement, displacement and vigilance. Even in high‐risk areas, however, these encounters occurred only once every 9 days. Ultimately, despite 20‐fold variation in the frequency of encounters between wolves and individual elk, the risk of predation was not associated with elk body fat or pregnancy. Our findings suggest that the ecological consequences of actively hunting large carnivores, such as the wolf, are more likely transmitted by consumptive effects on prey survival than NCEs on prey behaviour.     

Competition and coexistence in sympatric bobcats and pumas

Space use and diets of sympatric bobcats Lynx rufus and pumas Puma concolor were compared using sign surveys and scat analysis during 1997–2002 in south-eastern Arizona, USA. Bobcats appeared to use grassland, scrub, riparian and woodland habitats equally, but pumas had higher activity in riparian and woodland habitats. There was little evidence that bobcats avoided pumas in space use. Bobcats ate primarily rodents (33% of items in scats), lagomorphs (32%) and ungulates (16%), whereas pumas ate primarily ungulates (69%) and carnivores (21%). Pumas had a narrower dietary niche breadth than bobcats, and puma diet overlapped bobcat diet by 56%, suggesting that pumas may be more vulnerable to changes in prey density than bobcats. Pumas also killed and consumed bobcats, indicating that interference competition may be manifesting through intraguild predation.     

Accounting for tagging‐to‐harvest mortality in a Brownie tag‐recovery model by incorporating radio‐telemetry data

The Brownie tag‐recovery model is useful for estimating harvest rates but assumes all tagged individuals survive to the first hunting season; otherwise, mortality between time of tagging and the hunting season will cause the Brownie estimator to be negatively biased. Alternatively, fitting animals with radio transmitters can be used to accurately estimate harvest rate but may be more costly. We developed a joint model to estimate harvest and annual survival rates that combines known‐fate data from animals fitted with transmitters to estimate the probability of surviving the period from capture to the first hunting season, and data from reward‐tagged animals in a Brownie tag‐recovery model. We evaluated bias and precision of the joint estimator, and how to optimally allocate effort between animals fitted with radio transmitters and inexpensive ear tags or leg bands. Tagging‐to‐harvest survival rates from >20 individuals with radio transmitters combined with 50–100 reward tags resulted in an unbiased and precise estimator of harvest rates. In addition, the joint model can test whether transmitters affect an individual's probability of being harvested. We illustrate application of the model using data from wild turkey, Meleagris gallapavo, to estimate harvest rates, and data from white‐tailed deer, Odocoileus virginianus, to evaluate whether the presence of a visible radio transmitter is related to the probability of a deer being harvested. The joint known‐fate tag‐recovery model eliminates the requirement to capture and mark animals immediately prior to the hunting season to obtain accurate and precise estimates of harvest rate. In addition, the joint model can assess whether marking animals with radio transmitters affects the individual's probability of being harvested, caused by hunter selectivity or changes in a marked animal's behavior.     

Drivers of jaguar (Panthera onca) and puma (Puma concolor) predation on endangered primates within a transformed landscape in southern Mexico

Human pressures have increasingly placed keystone species, such as large cats, under threat. Together with forest loss, prey depletion is one of the main threats to the survival of jaguars (Panthera onca) and pumas (Puma concolor) throughout the Neotropics. Generally, primates are not considered main prey for jaguar and puma, and their inclusion in the diet could be indicative of ongoing prey species decline. Here, we investigate the effect of habitat type and disturbance on primate predation by large cats. Surveys took place during the dry seasons (March to June) of 2010 and 2011, covering a total of 608.5 km across 24 localities in the Uxpanapa Valley, Mexico. We found 65 felid scat samples with the aid of a wildlife scat detection dog, and then examined them to identify predator species and classify the prey remains they contained. Primates represented the most frequent prey (35%) for both jaguar and puma in our study site and constituted approximately half of the biomass consumed by these felines in the area. Primate remains were more likely to be found in scats surrounded by the lowest percentage of conserved forest or in areas surrounded by more villages, showing the potential effects of human activities on these species' populations. The high proportion of primates found in scats within our study site could be an early indication that populations of ungulates and other “typical” prey are beginning to collapse, and urgent conservation interventions are needed for both large cats and primates before they become locally extinct. Abstract in Spanish is available with online material.     

Evaluation of Potential Factors Predisposing Livestock to Predation by Jaguars

ABSTRACT Depredation of livestock by large carnivores is an important but poorly understood source of human-carnivore conflict. We examined patterns of livestock depredation by jaguars (Panthera onca) and pumas (Puma concolor) on a ranch-wildlife reserve in western Brazil to assess factors contributing to prey mortality. We predicted jaguars would kill a greater proportion of calves than yearling and adult cattle and that proximity to suitable habitat would increase mortality risk. We further speculated that exposure to predation risk would promote livestock grouping and increased movement distance. We recorded 169 cattle mortality incidents during 2003–2004, of which 19% were due to predation by jaguars and pumas. This level of mortality represented 0.2–0.3% of the total livestock holdings on the ranch. Jaguars caused most (69%) cattle predation events, and survival in allotments was lower for calves than for other age classes. Forest proximity was the only variable we found to explain patterns of livestock mortality, with predation risk increasing as distance to forest cover declined. Due to low predation risk, cattle movement patterns and grouping behavior did not vary relative to level of spatial overlap with radiocollared jaguars. The overall effect of predation on cattle was low and livestock likely constituted an alternative prey for large cats in our study area. However, selection of calves over other age cohorts and higher predation risk among cattle in proximity to forest cover is suggestive of selection of substandard individuals. Cattle ranchers in the Pantanal region may reduce cattle mortality rates by concentrating on losses due to nonpredation causes that could be more easily controlled.     

Food habits of three carnivores in a mosaic landscape of São Paulo state,Brazil

São Paulo is the most developed state in Brazil, but despite intense land use changes, some mammal species still inhabit remnant fragments of native vegetation of the state. We used fecal analyses to investigate prey diversity, trophic niche breadth, and trophic niche overlap of pumas, ocelots, and maned wolves in protected and non-protected areas in São Paulo state. During 18 months, we inspected more than 10,000 km of roads and trails and collected 233 fecal samples. Maned wolves showed generalist food habit, ocelots were predominantly carnivores, and pumas were strict carnivores. The European hare, an exotic species which had not yet been reported in the study area, was identified in the feces of a puma. Trophic niche breadth was the lowest for maned wolves (BA?=?0.339) and the highest for pumas (BA?=?0.723). Trophic niche overlap was high between maned wolves and ocelots (O_jk?=?0.765) and between pumas and ocelots (O_jk?=?0.639). Significantly different diets of maned wolves and ocelots were observed between dry and rainy seasons. Our findings indicate that these three species may adapt their diets to the disturbed landscapes. This is the second study characterizing food habits of ocelots in Cerrado areas but the first considering a large number of samples.     

The role of exotic mammals in the diet of native carnivores from South America

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Modeling connectivity to identify current and future anthropogenic barriers to movement of large carnivores: A case study in the American Southwest

This study sought to identify critical areas for puma (Puma concolor) movement across the state of Arizona in the American Southwest and to identify those most likely to be impacted by current and future human land uses, particularly expanding urban development and associated increases in traffic volume. Human populations in this region are expanding rapidly, with the potential for urban centers and busy roads to increasingly act as barriers to demographic and genetic connectivity of large‐bodied, wide‐ranging carnivores such as pumas, whose long‐distance movements are likely to bring them into contact with human land uses and whose low tolerance both for and from humans may put them at risk unless opportunities for safe passage through or around human‐modified landscapes are present. Brownian bridge movement models based on global positioning system collar data collected during bouts of active movement and linear mixed models were used to model habitat quality for puma movement; then, a wall‐to‐wall application of circuit theory models was used to produce a continuous statewide estimate of connectivity for puma movement and to identify pinch points, or bottlenecks, that may be most at risk of impacts from current and future traffic volume and expanding development. Rugged, shrub‐ and scrub‐dominated regions were highlighted as those offering high quality movement habitat for pumas, and pinch points with the greatest potential impacts from expanding development and traffic, although widely distributed, were particularly prominent to the north and east of the city of Phoenix and along interstate highways in the western portion of the state. These pinch points likely constitute important conservation opportunities, where barriers to movement may cause disproportionate loss of connectivity, but also where actions such as placement of wildlife crossing structures or conservation easements could enhance connectivity and prevent detrimental impacts before they occur.     

Seasonal occupancy of sympatric larger carnivores in the southern San Andres Mountains,south-central New Mexico,USA

We investigated factors facilitating coexistence of pumas (Puma concolor), coyotes (Canis latrans), and bobcats (Lynx rufus) in the arid San Andres Mountains of south-central New Mexico, during the season (winter and spring, prior to the annual monsoon) of greatest resource stress. We established a camera-trapping grid in the San Andres, 2007–2011, and modeled occupancy of the three carnivores as a function of habitat, prey, and presence of the other carnivore species. Species interaction factors were >1.3 for each pair of carnivores, and the presence of the other carnivore species never significantly influenced occupancy of any other carnivore. Similarly, occupancy of the San Andres landscape was positively correlated among all carnivores. Occupancy of pumas was most influenced by proximity of water; coyote occupancy was influenced by terrain ruggedness and presence of medium (primarily lagomorph) prey, and bobcat occupancy was influence primarily small prey and proximity to water. The three carnivores also did not show temporal partitioning in use of habitats. Rather than segregation driven by competition, predation, or despotism, our results appeared to reflect preferences for differing habitat characteristics between ambush and cursorial predators and preferred habitats for travel.     

Interspecific interactions between wild felids vary across scales and levels of urbanization

Ongoing global landscape change resulting from urbanization is increasingly linked to changes in species distributions and community interactions. However, relatively little is known about how urbanization influences competitive interactions among mammalian carnivores, particularly related to wild felids. We evaluated interspecific interactions between medium‐ and large‐sized carnivores across a gradient of urbanization and multiple scales. Specifically, we investigated spatial and temporal interactions of bobcats and pumas by evaluating circadian activity patterns, broad‐scale seasonal interactions, and fine‐scale daily interactions in wildland–urban interface (WUI), exurban residential development, and wildland habitats. Across levels of urbanization, interspecific interactions were evaluated using two‐species and single‐species occupancy models with data from motion‐activated cameras. As predicted, urbanization increased the opportunity for interspecific interactions between wild felids. Although pumas did not exclude bobcats from areas at broad spatial or temporal scales, bobcats responded behaviorally to the presence of pumas at finer scales, but patterns varied across levels of urbanization. In wildland habitat, bobcats avoided using areas for short temporal periods after a puma visited an area. In contrast, bobcats did not appear to avoid areas that pumas recently visited in landscapes influenced by urbanization (exurban development and WUI habitat). In addition, overlap in circadian activity patterns between bobcats and pumas increased in exurban development compared to wildland habitat. Across study areas, bobcats used sites less frequently as the number of puma photographs increased at a site. Overall, bobcats appear to shape their behavior at fine spatial and temporal scales to reduce encounters with pumas, but residential development can potentially alter these strategies and increase interaction opportunities. We explore three hypotheses to explain our results of how urbanization affected interspecific interactions that consider activity patterns, landscape configuration, and animal scent marking. Altered competitive interactions between animals in urbanized landscapes could potentially increase aggressive encounters and the frequency of disease transmission.     

Differences in diet and trophic interactions of Patagonian carnivores between areas with mostly native or exotic prey

The prey base for Patagonian carnivores has been altered greatly over the past 150 years due to widespread overgrazing by livestock, invasions by exotic wildlife, and hunting. On ranches in northern Patagonia carnivores consume mostly exotic species, and native herbivores are ecologically extinct in their role as prey. In this study we compare diets of the culpeo (Lycalopex culpaeus), chilla (L. griseus), puma (Puma concolor), Geoffroy's cat (Leopardus geoffroyi), colocolo (L. colocolo), and hog-nosed skunks (Conepatus chinga and C. humboldtii) in a reserve with a mostly native prey base to their diets on the ranches, and evaluate how differences in prey bases affect trophic interactions among carnivores. Carnivores in the reserve consumed mostly native prey. Dietary overlap among carnivores was not significant on the reserve, but was highly significant on the ranches. This homogenization of diets where densities of native species are reduced could lead to stronger negative interactions among carnivores, altering the composition of the carnivore assemblage to the detriment of the more specialized species. Study of carnivore diets may be a relatively quick way to evaluate the conservation status and ecological functionality of prey assemblages in Patagonia and other areas where these have been altered.