Large “hypercarnivorous” felids are recognized for their role as apex predators and hence as key elements in food webs and ecosystem functioning through competition and depredation. Here we show that cougars (Puma concolor), one of the largest and the most widely ranging apex felid predators with a strictly carnivorous diet, could also be effective secondary long distance seed dispersers, potentially establishing direct and non-herbivore mediated interactions with plant species at the bottom of the food web. Cougars accidently ingest and disseminate large amounts of seeds (31,678 seeds in 123 scats) of plant species initially consumed by their main prey, the Eared Dove Zenaida auriculata. The germination potential of seeds for the three plant species most abundantly found in cougar scats (19,570 seeds) was not significantly different from that observed in seeds obtained from dove gizzards, indicating that seed passage through cougar guts did not affect seed germination. Considering the estimated cougar density in our study area, dispersal of seeds by cougars could allow a mean, annual seed spread of ~5,000 seeds per km(2). Our results demonstrate that strictly carnivorous, felid predators could have broad and overlooked ecological functions related to ecosystem structuring and functioning.
Cost-benefit trade-offs for individuals participating in social behaviors are the basis for current theories on the evolution of social behaviors and societies. However, research on social strategies has largely ignored solitary animals, in which we assume that rare interactions are explained by courtship or territoriality or, in special circumstances, resource distributions or kinship. We used directed network analysis of conspecific tolerance at food sources to provide evidence that a solitary carnivore, the puma (Puma concolor), exhibited adaptive social strategies similar to more social animals. Every puma in our analysis participated in the network, which featured densely connected communities delineated by territorial males. Territorial males also structured social interactions among pumas. Contrary to expectations, conspecific tolerance was best characterized by direct reciprocity, establishing a fitness benefit to individuals that participated in social behaviors. However, reciprocity operated on a longer time scale than in gregarious species. Tolerance was also explained by hierarchical reciprocity, which we defined as network triangles in which one puma (generally male) received tolerance from two others (generally females) that also tolerated each other. Hierarchical reciprocity suggested that males might be cheating females; nevertheless, we suspect that males and females used different fitness currencies. For example, females may have benefited from tolerating males through the maintenance of social niches that support breeding opportunities. Our work contributes evidence of adaptive social strategies in a solitary carnivore and support for the applicability of theories of social behavior across taxa, including solitary species in which they are rarely tested.
The importance of examining multiple hierarchical levels when modeling resource use for wildlife has been acknowledged for decades. Multi-level resource selection functions have recently been promoted as a method to synthesize resource use across nested organizational levels into a single predictive surface. Analyzing multiple scales of selection within each hierarchical level further strengthens multi-level resource selection functions. We extend this multi-level, multi-scale framework to modeling resistance for wildlife by combining multi-scale resistance surfaces from two data types, genetic and movement. Resistance estimation has typically been conducted with one of these data types, or compared between the two. However, we contend it is not an either/or issue and that resistance may be better-modeled using a combination of resistance surfaces that represent processes at different hierarchical levels. Resistance surfaces estimated from genetic data characterize temporally broad-scale dispersal and successful breeding over generations, whereas resistance surfaces estimated from movement data represent fine-scale travel and contextualized movement decisions. We used telemetry and genetic data from a long-term study on pumas (Puma concolor) in a highly developed landscape in southern California to develop a multi-level, multi-scale resource selection function and a multi-level, multi-scale resistance surface. We used these multi-level, multi-scale surfaces to identify resource use patches and resistant kernel corridors. Across levels, we found puma avoided urban, agricultural areas, and roads and preferred riparian areas and more rugged terrain. For other landscape features, selection differed among levels, as did the scales of selection for each feature. With these results, we developed a conservation plan for one of the most isolated puma populations in the U.S. Our approach captured a wide spectrum of ecological relationships for a population, resulted in effective conservation planning, and can be readily applied to other wildlife species.
Pumas (Puma concolor) live in diverse, often rugged, complex habitats. The energy they expend for hunting must account for this complexity but is difficult to measure for this and other large, cryptic carnivores. We developed and deployed a physiological SMART (species movement, acceleration, and radio tracking) collar that used accelerometry to continuously monitor energetics, movements, and behavior of free-ranging pumas. This felid species displayed marked individuality in predatory activities, ranging from low-cost sit-and-wait behaviors to constant movements with energetic costs averaging 2.3 times those predicted for running mammals. Pumas reduce these costs by remaining cryptic and precisely matching maximum pouncing force (overall dynamic body acceleration = 5.3 to 16.1g) to prey size. Such instantaneous energetics help to explain why most felids stalk and pounce, and their analysis represents a powerful approach for accurately forecasting resource demands required for survival by large, mobile predators.
Wide-ranging large carnivores pose myriad challenges for conservation, especially in highly fragmented landscapes. Over a 13-year period, we combined monitoring of radio collared pumas (Puma concolor) with complementary multi-generational genetic analyses to inform puma conservation in southern California, USA. Our goals were to generate survivorship estimates, determine causes of mortality, identify barriers to movement, and determine the genetic and demographic challenges to puma persistence among >20,000,000 people and extensive urban, suburban, and exurban development. Despite protection from hunting, annual survival for radio collared pumas was surprisingly low (55.8%), and humans caused the majority of puma deaths. The most common sources of mortality were vehicle collisions (28% of deaths), and mortalities resulting from depredation permits issued after pumas killed domestic animals (17% of deaths). Other human-caused mortalities included illegal shootings, public safety removals, and human-caused wildfire. An interstate highway (I-15) bisecting this study area, and associated development, have created a nearly impermeable barrier to puma movements, resulting in severe genetic restriction and demographic isolation of the small puma population (n ~ 17-27 adults) in the Santa Ana Mountains west of I-15. Highways that bisect habitat or divide remaining “conserved” habitat, and associated ongoing development, threaten to further subdivide this already fragmented puma population and increase threats to survival. This study highlights the importance of combining demographic and genetic analyses, and illustrates that in the absence of effective measures to reduce mortality and enhance safe movement across highways, translocation of pumas, such as was done with the endangered Florida panther (P. c. coryi), may ultimately be necessary to prevent further genetic decline and ensure persistence of the Santa Ana Mountains population.
Pumas (Puma concolor; also known as mountain lions and cougars) in southern California live among a burgeoning human population of roughly 20 million people. Yet little is known of the consequences of attendant habitat loss and fragmentation, and human-caused puma mortality to puma population viability and genetic diversity. We examined genetic status of pumas in coastal mountains within the Peninsular Ranges south of Los Angeles, in San Diego, Riverside, and Orange counties. The Santa Ana Mountains are bounded by urbanization to the west, north, and east, and are separated from the eastern Peninsular Ranges to the southeast by a ten lane interstate highway (I-15). We analyzed DNA samples from 97 pumas sampled between 2001 and 2012. Genotypic data for forty-six microsatellite loci revealed that pumas sampled in the Santa Ana Mountains (n = 42) displayed lower genetic diversity than pumas from nearly every other region in California tested (n = 257), including those living in the Peninsular Ranges immediately to the east across I-15 (n = 55). Santa Ana Mountains pumas had high average pairwise relatedness, high individual internal relatedness, a low estimated effective population size, and strong evidence of a bottleneck and isolation from other populations in California. These and ecological findings provide clear evidence that Santa Ana Mountains pumas have been experiencing genetic impacts related to barriers to gene flow, and are a warning signal to wildlife managers and land use planners that mitigation efforts will be needed to stem further genetic and demographic decay in the Santa Ana Mountains puma population.
Large carnivores can be particularly sensitive to the effects of habitat fragmentation on genetic diversity [1, 2]. The Santa Monica Mountains (SMMs), a large natural area within Greater Los Angeles, is completely isolated by urban development and the 101 freeway to the north. Yet the SMMs support a population of mountain lions (Puma concolor), a very rare example of a large carnivore persisting within the boundaries of a megacity. GPS locations of radio-collared lions indicate that freeways are a near-absolute barrier to movement. We genotyped 42 lions using 54 microsatellite loci and found that genetic diversity in SMM lions, prior to 2009, was lower than that for any population in North America except in southern Florida, where inbreeding depression led to reproductive failure [3-5]. We document multiple instances of father-daughter inbreeding and high levels of intraspecific strife, including the unexpected behavior of a male killing two of his offspring and a mate and his son killing two of his brothers. Overall, no individuals from the SMMs have successfully dispersed. Gene flow is critical for this population, and we show that a single male immigrated in 2009, successfully mated, and substantially enhanced genetic diversity. Our results imply that individual behaviors, most likely caused by limited area and reduced opportunities to disperse, may dominate the fate of small, isolated populations of large carnivores. Consequently, comprehensive behavioral monitoring can suggest novel solutions for the persistence of small populations, such as the transfer of individuals across dispersal barriers.
Acquiring reliable data on large felid populations is crucial for effective conservation and management. However, large felids, typically solitary, elusive and nocturnal, are difficult to survey. Tagging and following individuals with VHF or GPS technology is the standard approach, but costs are high and these methodologies can compromise animal welfare. Such limitations can restrict the use of these techniques at population or landscape levels. In this paper we describe a robust technique to identify and sex individual pumas from footprints. We used a standardized image collection protocol to collect a reference database of 535 footprints from 35 captive pumas over 10 facilities; 19 females (300 footprints) and 16 males (235 footprints), ranging in age from 1-20 yrs. Images were processed in JMP data visualization software, generating one hundred and twenty three measurements from each footprint. Data were analyzed using a customized model based on a pairwise trail comparison using robust cross-validated discriminant analysis with a Ward’s clustering method. Classification accuracy was consistently > 90% for individuals, and for the correct classification of footprints within trails, and > 99% for sex classification. The technique has the potential to greatly augment the methods available for studying puma and other elusive felids, and is amenable to both citizen-science and opportunistic/local community data collection efforts, particularly as the data collection protocol is inexpensive and intuitive.
A mosaic of intact native and human-modified vegetation use can provide important habitat for top predators such as the puma (Puma concolor), avoiding negative effects on other species and ecological processes due to cascade trophic interactions. This study investigates the effects of restoration scenarios on the puma’s habitat suitability in the most developed Brazilian region (São Paulo State). Species Distribution Models incorporating restoration scenarios were developed using the species' occurrence information to (1) map habitat suitability of pumas in São Paulo State, Southeast, Brazil; (2) test the relative contribution of environmental variables ecologically relevant to the species habitat suitability and (3) project the predicted habitat suitability to future native vegetation restoration scenarios. The Maximum Entropy algorithm was used (Test AUC of 0.84 ± 0.0228) based on seven environmental non-correlated variables and non-autocorrelated presence-only records (n = 342). The percentage of native vegetation (positive influence), elevation (positive influence) and density of roads (negative influence) were considered the most important environmental variables to the model. Model projections to restoration scenarios reflected the high positive relationship between pumas and native vegetation. These projections identified new high suitability areas for pumas (probability of presence >0.5) in highly deforested regions. High suitability areas were increased from 5.3% to 8.5% of the total State extension when the landscapes were restored for ≥ the minimum native vegetation cover rule (20%) established by the Brazilian Forest Code in private lands. This study highlights the importance of a landscape planning approach to improve the conservation outlook for pumas and other species, including not only the establishment and management of protected areas, but also the habitat restoration on private lands. Importantly, the results may inform environmental policies and land use planning in São Paulo State, Brazil.
BACKGROUND: The Chilean temperate rainforest has been subjected to dramatic fragmentation for agriculture and forestry exploitation. Carnivore species are particularly affected by fragmentation and the resulting resource use conflicts with humans. This study aimed at understanding values and human-animal relationships with negatively perceived threatened carnivores through the disclosure of local stories and Mapuche traditional folktales. METHODS: Our mixed approach comprised the qualitative analysis of 112 stories on the kodkod cat (Leopardus guigna) and the puma (Puma concolor) collected by students (9-14 years) from 28 schools in the Araucania region within their family contexts, 10 qualitative in-depth interviews with indigenous Mapuche people, and 35 traditional Mapuche legends and the significance of naming found in ethnographic collections. RESULTS: We revealed a quasi-extinction of traditional tales in the current knowledge pool about pumas and kodkods, local anecdotes, however, were present in significant numbers. Values associated to both felids were manifold, ranging from negativistic to positive values. While pumas played an important role in people’s spirituality, negative mythological connotations persisted in kodkod stories. Four prominent relationships were derived: (1) Both felids represent threats to livestock, pumas even to life (2) both felids are symbols for upcoming negative events, (3) pumas are spiritual creatures, and (4) kodkods are threatened by humans. Recommendations are provided for stimulating new ways of perceiving unpopular and threatened carnivores among those who live in vicinity to them.