David C. Stoner

Cougar Exploitation Levels in Utah: Implications for Demographic Structure, Population Recovery, and Metapopulation Dynamics

Abstract Currently, 11 western states and 2 Canadian provinces use sport hunting as the primary mechanism for managing cougar (Puma concolor) populations. Yet the impacts of sustained harvest on cougar population dynamics and demographic structure are not well understood. We evaluated the effects of hunting on cougar populations by comparing the dynamics and demographic composition of 2 populations exposed to different levels of harvest. We monitored the cougar populations on Monroe Mountain in south-central Utah, USA, and in the Oquirrh Mountains of north-central Utah from 1996 to 2004. Over this interval the Monroe population was subjected to annual removals ranging from 17.6–51.5% (mean ± SE = 35.4 ± 4.3%) of the population, resulting in a >60% decline in cougar population density. Concurrently, the Oquirrh study area was closed to hunting and the population remained stationary. Mean age in the hunted population was lower than in the protected population (F = 9.0; df = 1, 60.3; P = 0.004), and in a pooled sample of all study animals, females were older than males (F = 13.8; df = 1, 60.3; P < 0.001). Females from the hunted population were significantly younger than those from the protected population (3.7 vs. 5.9 yr), whereas male ages did not differ between sites (3.1 vs. 3.4 yr), suggesting that male spatial requirements may put a lower limit on the area necessary to protect a subpopulation. Survival tracked trends in density on both sites. Levels of human-caused mortality were significantly different between sites (χ2 = 7.5; P = 0.006). Fecundity rates were highly variable in the protected population but appeared to track density trends with a 1-year lag on the hunted site. Results indicate that harvest exceeding 40% of the population, sustained for ≥4 years, can have significant impacts on cougar population dynamics and demographic composition. Patterns of recruitment resembled a source–sink population structure due in part to spatially variable management strategies. Based on these observations, the temporal scale of population recovery will most likely be a function of local harvest levels, the productivity of potential source populations, and the degree of landscape connectivity among demes. Under these conditions the metapopulation perspective holds promise for broad-scale management of this species.

Evaluation of Cougar Population Estimators in Utah

Abstract Numerous techniques have been proposed to estimate or index cougar (Puma concolor) populations, but few have been applied simultaneously to populations with reliable estimates of population size. Between 1996 and 2003, we evaluated the relative efficacy and accuracy of multiple estimation and index techniques for populations at 2 locations in Utah, USA: Monroe Mountain and the Oquirrh Mountains. We used radiotagging followed by intensive monitoring and repeated capture efforts to approach a complete enumeration of the populations. We used these benchmarks to evaluate other population estimates (Lincoln–Petersen mark–recapture, helicopter-survey probability sampling, catch-per-unit-effort) and indices (scent-station visits, track counts, hunter harvest). Monitoring over 600 scent-station-nights using different attractants June–September in 1996 and 1997 yielded a single cougar visit. Summer track-based indices reflected a 54–69% reduction in population size on the Monroe site and a numerically stable population on the Oquirrhs, but relationships between indices and the benchmark population estimates varied among techniques. Aerial track surveys required sufficient fresh snowfall accumulations for adequate tracking coverage of a given unit, conditions that were met only once on one study site in each of 3 years. Population estimates derived from helicopter-survey probability sampling exceeded reference population estimates by 120–284%, and bootstrapped estimates of standard error encompassed 25–55% of the population estimates (e.g., 5.6 ± 1.4 cougars/100 km2). Despite poor performance in predicting cougar population sizes, track-based estimates may provide better indices for monitoring large changes in population trends (i.e., with low precision). However, we recommend using multiple indices after determination of a more rigorous initial population estimate for managing populations of conservation concern and when considering connectivity to determine potential refuge sites for regional management (e.g., management by zones).

Long-Distance Dispersal of a Female Cougar in a Basin and Range Landscape

Abstract We used Global Positioning System technology to document distance, movement path, vegetation, and elevations used by a dispersing subadult female cougar (Puma concolor) through the fragmented habitat of the Intermountain West, USA. Over the course of 1 year, female number 31 moved 357 linear km, but an actual distance of 1,341 km from the Oquirrh Mountains, Utah to the White River Plateau, Colorado, constituting the farthest dispersal yet documented for a female cougar. This cougar successfully negotiated 4 major rivers and one interstate highway while traversing portions of 3 states. Our data suggest that transient survival, and therefore total distance moved, may be enhanced when dispersal occurs during the snow-free season due to low hunting pressure and greater access to high elevation habitats. Long-distance movements by both sexes will be required for the recolonization of vacant habitats, and thus inter-state management may be warranted where state boundaries do not coincide with effective dispersal barriers.

Ungulate Reproductive Parameters Track Satellite Observations of Plant Phenology across Latitude and Climatological Regimes.

The effect of climatically-driven plant phenology on mammalian reproduction is one key to predicting species-specific demographic responses to climate change. Large ungulates face their greatest energetic demands from the later stages of pregnancy through weaning, and so in seasonal environments parturition dates should match periods of high primary productivity. Interannual variation in weather influences the quality and timing of forage availability, which can influence neonatal survival. Here, we evaluated macro-scale patterns in reproductive performance of a widely distributed ungulate (mule deer, Odocoileus hemionus) across contrasting climatological regimes using satellite-derived indices of primary productivity and plant phenology over eight degrees of latitude (890 km) in the American Southwest. The dataset comprised > 180,000 animal observations taken from 54 populations over eight years (2004–2011). Regionally, both the start and peak of growing season (“Start” and “Peak”, respectively) are negatively and significantly correlated with latitude, an unusual pattern stemming from a change in the dominance of spring snowmelt in the north to the influence of the North American Monsoon in the south. Corresponding to the timing and variation in both the Start and Peak, mule deer reproduction was latest, lowest, and most variable at lower latitudes where plant phenology is timed to the onset of monsoonal moisture. Parturition dates closely tracked the growing season across space, lagging behind the Start and preceding the Peak by 27 and 23 days, respectively. Mean juvenile production increased, and variation decreased, with increasing latitude. Temporally, juvenile production was best predicted by primary productivity during summer, which encompassed late pregnancy, parturition, and early lactation. Our findings offer a parsimonious explanation of two key reproductive parameters in ungulate demography, timing of parturition and mean annual production, across latitude and changing climatological regimes. Practically, this demonstrates the potential for broad-scale modeling of couplings between climate, plant phenology, and animal populations using space-borne observations.

Dispersal behaviour of a polygynous carnivore: do cougars Puma concolor follow source-sink predictions?

The source-sink model of population dynamics predicts that density drives emigration of subordinate animals to habitats offering lower competition for resources. Several authors have suggested use of this model as a potential framework for conservation of exploited carnivores when precise enumeration is unfeasible. Dispersal is a critical behavioural mechanism for management based on this model, yet there is a lack of knowledge on the habitat and social conditions that motivate carnivore emigration and settlement. The cougar Puma concolor is a widely distributed and heavily exploited carnivore, indigenous to the western hemisphere. We evaluated patterns in cougar dispersal behaviour from two sites in Utah, differing in terms of management and the level of natural and anthropogenic habitat fragmentation. We used our results to evaluate three predictions with respect to cougar dispersal behaviour: 1) natal population density and maternal reproductive status prompt emigration, 2) movement of dispersing cougars is shaped by habitat configuration and permeability, and 3) dispersers preferentially settle in areas of high habitat quality and low conspecific density. We documented the emigration of 62 individuals and measured movement variables, including sex and site-specific frequency, distance, seasonality, direction and the habitat quality and harvest rates characterizing areas where immigrants settled. Although males and females exhibited pronounced differences in dispersal frequency, we found few differences in distance traveled, season of departure and direction moved. Dispersal occurred most frequently during spring, coinciding with the estrus pulse. Natural and anthropogenic obstacles modified landscape permeability, and therefore dispersal distances were shorter in fragmented habitats than in contiguous ones. Relative to males, females dispersed into habitats of lower productivity with higher mean annual harvest rates. Patterns in male settlement suggested habitat selection based on mating opportunities, whereas female settlement was predicated on avoiding conspecifics. Cougars in this Great Basin ecosystem largely conformed to source-sink predictions. Results can be used to parameterize source-sink models based on animal behaviour and landscape permeability to conserve exploited carnivores, under conditions of population expansion or recolonization of habitats where Allee effects are a limiting factor.

Spatial processes decouple management from objectives in a heterogeneous landscape: predator control as a case study

Predator control is often implemented with the intent of disrupting top-down regulation in sensitive prey populations. However, ambiguity surrounding the efficacy of predator management, as well as the strength of top-down effects of predators in general, is often exacerbated by the spatially implicit analytical approaches used in assessing data with explicit spatial structure. Here, we highlight the importance of considering spatial context in the case of a predator control study in south-central Utah. We assessed the spatial match between aerial removal risk in coyotes (Canis latrans) and mule deer (Odocoileus hemionus) resource selection during parturition using a spatially explicit, multi-level Bayesian model. With our model, we were able to evaluate spatial congruence between management action (i.e., coyote removal) and objective (i.e., parturient deer site selection) at two distinct scales: the level of the management unit and the individual coyote removal. In the case of the former, our results indicated substantial spatial heterogeneity in expected congruence between removal risk and parturient deer site selection across large areas, and is a reflection of logistical constraints acting on the management strategy and differences in space use between the two species. At the level of the individual removal, we demonstrated that the potential management benefits of a removed coyote were highly variable across all individuals removed and in many cases, spatially distinct from parturient deer resource selection. Our methods and results provide a means of evaluating where we might anticipate an impact of predator control, while emphasizing the need to weight individual removals based on spatial proximity to management objectives in any assessment of large-scale predator control. Although we highlight the importance of spatial context in assessments of predator control strategy, we believe our methods are readily generalizable in any management or large-scale experimental framework where spatial context is likely an important driver of outcomes.