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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. [ABSTRACT FROM AUTHOR]

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

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

In total, 177 of 245 terrestrial carnivores are described as solitary, and much of carnivore ecology is built on the assumptions that interactions between adult solitary carnivores are rare. We employed Global Positioning System (GPS) technology and motion-triggered cameras to test predictions of land-tenure territoriality and the resource dispersion hypothesis in a territorial carnivore, the puma Puma concolor. We documented 89 independent GPS interactions, 60% ofwhich occurred at puma kills (n = 53), 59 camera interactions, 11 (17%) ofwhich captured courtship behaviors, and 5 other interactions (1 F-F, 3 M-F, and 1 M-M). Mean minimum weekly contact rates were 5.5 times higher in winter, the season when elk Cervus elaphus were aggregated at lower elevations and during which puma courtship primarily occurred. In winter, contacts rates were 0.6 ± 0.3 (standard deviation (SD)) interactions/week vs. 0.1 ± 0.1 (SD) interactions/week during summer. The preponderance of interactions at food sources supported the resource dispersion hypothesis, which predicts that resource fluxes can explain temporary social behaviors that do not result in any apparent benefits for the individuals involved. Conspecific tolerance is logical when a prey is so large that the predator that killed it cannot consume it entirely, and thus, the costs of tolerating a conspecific sharing the kill are less than the potential costs associated with defending it and being injured. Puma aggregations at kills numbered as high as 9, emphasizing the need for future research on what explains tolerance among solitary carnivores. [ABSTRACT FROM AUTHOR]

The article discusses a study which revealed that the competition between recolonizing wolves and resident cougars in the Southern Yellowstone Ecosystem (SYE) influenced the cougars' prey selection and realized niches. Topics covered include increase in wolves population between 2002 and 2012, the change in cougar prey selection from elk to deer in summer, and the decline in elk and deer population. It also notes the death of several cougars due to starvation between 2005 and 2012.

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We studied survival and causes of mortality of radiocollared cougars (Puma concolor) on the Greater Yellowstone Northern Range (GYNR) prior to (1987-1994) and after wolf (Canis lupus) reintroduction (1998-2005) and evaluated temporal, spatial, and environmental factors that explain variation in adult, subadult, and kitten survival. Using Program MARK and multimodel inference, we modeled cougar survival based on demographic status, season, and landscape attributes. Our best models for adult and independent subadults indicated that females survived better than males and survival increased with age until cougars reached older ages. Lower elevations and increasing density of roads, particularly in areas open to cougar hunting north of Yellowstone National Park (YNP), increased mortality risks for cougars on the GYNR. Indices of ungulate biomass, cougar and wolf population size, winter severity, rainfall, and individual characteristics such as the presence of dependent young, age class, and use of Park or Wilderness were not important predictors of survival. Kitten survival increased with age, was lower during winter, increased with increasing minimum estimates of elk calf biomass, and increased with increasing density of adult male cougars. Using our best model, we mapped adult cougar survival on the GYNR landscape. Results of receiver operating characteristic (ROC) analysis indicated a good model fit for both female (area under the curve [AUC] = 0.81, 95%CI = 0.70-0.92, n = 35 locations) and male cougars (AUC = 0.84, 95%CI = 0.74- 0.94, n = 49 locations) relative to hunter harvest locations in our study area. Using minimum estimates of survival necessary to sustain the study population, we developed a source-sink surface and we identify several measures that resource management agencies can take to enhance cougar population management based on a source-sink strategy. [ABSTRACT FROM AUTHOR]

Conventional methods for monitoring cougar, Puma concolor, populations involve capture, tagging, and radio-collaring, but these methods are time-consuming, expensive, and logistically challenging. For difficult-to-study species such as cougars, noninvasive genetic sampling (NGS) may be a useful alternative. The ability to identify individuals from samples collected through NGS methods provides many opportunities for developing population-monitoring tools, but the utility of these survey methods is dependent upon collection of samples and accurate genotyping of those samples. In January 2003, we initiated a 3-yr evaluation of NGS methods for cougars using a radio-collared population in Yellowstone National Park (YNP), USA. Our goals were to: 1) determine which DNA collection method, hair snares or snow tracking, provided a better method for obtaining samples for genetic analysis, 2) evaluate reliability of the genetic data derived from hair samples collected in the field, and 3) evaluate the potential of NGS for demographic monitoring of cougar populations. Snow tracking yielded more hair samples and was more cost effective than snagging hair with rub pads. Samples collected from bed sites and natural hair snags (e.g., branch tips, thorn bushes) while snow tracking accurately identified and sexed 22 individuals (9 F, 13 M). The ratio of the count from snow tracking to the count from radio-telemetry was 15:24 in winter 2004, 13:12 in 2005, and 22:29 for both years combined. Annual capture probabilities for obtaining DNA from snow tracking varied considerably between years for females (0.42 in 2004 and 0.88 in 2005) but were more consistent for males (0.77 in 2004 and 0.88 in 2005). Our results indicate that snow tracking can be an efficient, reliable NGS method for cougars in YNP and has potential for estimating demographic and genetic parameters of other carnivore populations in similar climates. [ABSTRACT FROM AUTHOR]

We assessed whether use of 2 methods, intensive very high frequency (VHF) radiotelemetry and Global Positioning System (GPS) cluster sampling, yielded similar estimates of cougar (Puma concolor) kill rates in Yellowstone National Park, 1998-2005. We additionally determined biases (underestimation or overestimation of rates) resulting from each method. We used modeling to evaluate what characteristics of clusters best predicted a kill versus no kill and further evaluated which predictor(s) minimized effort and the number of missed kills. We conducted 16 VHF ground predation sequences resulting in 37 kill intervals (KIs) and 21 GPS sequences resulting in 84 KIs on 6 solitary adult females, 4 maternal females, and 5 adult males. Kill rates (days/kill and biomass [kg] killed/day) did not differ between VHF and GPS predation sampling methods for maternal females, solitary adult females, and adult males. Sixteen of 142 (11.3%) kills detected via GPS clusters were missed through VHF ground-based sampling, and the kill rate was underestimated by an average of 5.2 (95% CI = 3.8-6.6) days/kill over all cougar social classes. Five of 142 (3.5%) kills identified by GPS cluster sampling were incorrectly identified as the focal individual's kill from scavenging, and the kill rate was overestimated within the adult male social class by an average of 5.8 (95% CI = 3.0-8.5) days/ungulate kill. The number of nights (locations between 2000 hours and 0500 hours) a cougar spent at a cluster was the most efficient variable at predicting predation, minimizing the missed kills, and minimizing number of extra clusters that needed to be searched. In Yellowstone National Park, where competing carnivores displaced cougars from their kills, it was necessary to search extra sites where a kill may not have been present to ensure we did not miss small, ungulate prey kills or kills with displacement. Using predictions from models to assign unvisited clusters as no kill, small prey kill, or large prey kill can bias downward the number of kills a cougar made and bias upward kills made by competitors that displace cougars or scavenge cougar kills. Our findings emphasize that field visitation is crucial in determining displacement and scavenging events that can result in biases when using GPS cluster methods in multicarnivore systems. [ABSTRACT FROM AUTHOR]

In a companion article (Beier et al. 2006), we identified 2 sets of unreliable inferences that may compromise efforts to conserve the Florida panther (Puma concolor coryi). In spite of serious flaws in methodology and interpretation, these unreliable conclusions have appeared in prominent, peer-refereed scientific journals and have been repeatedly cited and miscited in support of panther conservation. Future editors and referees may reduce these errors by insisting on adherence to an Introduction, Methods, Results, and Discussion (IMRAD) format; checking improbable assertions attributed to earlier papers; and refusing to allow scientific inference in publication formats not subject to scientific peer review (e.g., editorials). We urge conservation biologists to view science as an adaptive process and to use the method of multiple working hypotheses (Chamberlin 1890) that are now a central feature of adaptive resource management (Walters 1986, Williams et al. 2002). We advocate a workshop approach, similar to that used for analysis of data for the northern spotted owl (Strix occidentalis; Anderson et al. 1999), to deal with scientific disagreement where, as in the case with panthers, stakeholders have entrenched points of view. Finally, we recommend the creation of an independent Scientific Steering Committee to address long-term issues of future research and monitoring of Florida panthers. [ABSTRACT FROM AUTHOR]

The ability to non-invasively monitor stress hormone levels in free-ranging animals could significantly aid in conservation and management efforts. Our objective in this study was to demonstrate the effectiveness of assay of fecal corticoid metabolites in detecting a stress response in cougars (Puma concolor). Fecal samples were collected from 9 captive cougars before and after an artificial stressor. Steroid hormones were extracted from the samples. Adrenal corticoid metabolite concentrations of the resulting extracts were quantified using cortisol and corticosterone assays. Results indicated that fecal glucocorticoid metabolite levels increased significantly 24 hours after the stressor in 6 of the 9 individuals. Behavior correlated with the hormonal response; all cougars that displayed a flight response to the stressor also had elevated stress hormone measures. Elevated levels of stress hormones were not observed in individuals that did not attempt to flee. We have demonstrated with this study that measurement of fecal hormone metabolites is sufficiently sensitive to detect an adrenal response to stress in cougars and could be applied in the field to monitor stress levels in free-ranging populations. [ABSTRACT FROM AUTHOR]

Several conceptual models describing patterns of prey selection by predators have been proposed, but such models rarely have been tested empirically, particularly with terrestrial carnivores. We examined patterns of prey selection by sympatric wolves (Canis lupus ) and cougars (Puma concolor ) to determine i) if both predators selected disadvantaged prey disproportionately from the prey population, and ii) if the specific nature and intensity of prey selection differed according to disparity in hunting behavior between predator species. We documented prey characteristics and kill site attributes of predator kills during winters 1999–2001 in Idaho, and located 120 wolf-killed and 98 cougar-killed ungulates on our study site. Elk (Cervus elephus ) were the primary prey for both predators, followed by mule deer (Odocoileus hemionus ). Both predators preyed disproportionately on elk calves and old individuals; among mule deer, wolves appeared to select for fawns, whereas cougars killed primarily adults. Nutritional status of prey, as determined by percent femur marrow fat, was consistently poorer in wolf-killed prey. We found that wolf kills occurred in habitat that was more reflective of the entire study area than cougar kills, suggesting that the coursing hunting behavior of wolves likely operated on a larger spatial scale than did the ambush hunting strategy of cougars. We concluded that the disparity in prey selection and hunting habitat between predators probably was a function of predator-specific hunting behavior and capture success, where the longer prey chases and lower capture success of wolf packs mandated a stronger selection for disadvantaged prey. For cougars, prey selection seemed to be limited primarily by prey size, which could be a function of the solitary hunting behavior of this species and the risks associated with capturing prime-aged prey. [ABSTRACT FROM AUTHOR]

We tested for seasonal differences in cougar (Puma concolor) foraging behaviors in the Southern Yellowstone Ecosystem, a multi-prey system in which ungulate prey migrate, and cougars do not. We recorded 411 winter prey and 239 summer prey killed by 28 female and 10 male cougars, and an additional 37 prey items by unmarked cougars. Deer composed 42.4% of summer cougar diets but only 7.2% of winter diets. Males and females, however, selected different proportions of different prey; male cougars selected more elk (Cervus elaphus) and moose (Alces alces) than females, while females killed greater proportions of bighorn sheep (Ovis canadensis), pronghorn (Antilocapra americana), mule deer (Odocoileus hemionus) and small prey than males. Kill rates did not vary by season or between males and females. In winter, cougars were more likely to kill prey on the landscape as: 1) elevation decreased, 2) distance to edge habitat decreased, 3) distance to large bodies of water decreased, and 4) steepness increased, whereas in summer, cougars were more likely to kill in areas as: 1) elevation decreased, 2) distance to edge habitat decreased, and 3) distance from large bodies of water increased. Our work highlighted that seasonal prey selection exhibited by stationary carnivores in systems with migratory prey is not only driven by changing prey vulnerability, but also by changing prey abundances. Elk and deer migrations may also be sustaining stationary cougar populations and creating apparent competition scenarios that result in higher predation rates on migratory bighorn sheep in winter and pronghorn in summer. Nevertheless, cougar predation on rare ungulates also appeared to be influenced by individual prey selection. [ABSTRACT FROM AUTHOR]