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Mark Hebblewhite

Publications and source records attributed to Mark Hebblewhite.

At least 19 recordsLinked to original sources

Local and landscape-level environmental conditions drive habitat selection across terrestrial mammal species

Aim Animal movements are a fundamental process affecting communities and ecosystems. Quantifying habitat selection across species and habitats is key for understanding how animals respond to environmental change. Currently, we lack comparative studies that examine how habitat selection varies across species traits and landscapes. We aim to quantify global patterns of habitat selection to help understand the fundamental drivers of movement behaviour. Location Global. Time Period Contemporary. Major Taxa Studied Terrestrial mammals. Methods We estimated selection coefficients for terrain ruggedness, vegetation productivity, human population density and distance to roads of individual terrestrial mammals through step-selection analysis of 1344 GPS tracks across 48 species. We quantified intra- and interspecific variation and tested whether selection coefficients were associated with species traits and habitat availability. Results We observe an overall avoidance of roads and areas of high human population density as well as rugged terrain, with a large proportion of individuals selecting for areas of intermediate NDVI. However, we also found large intraspecific variation in habitat selection and show that this variation is predicted by local and landscape-level environmental conditions rather than species traits. Individuals in more remote areas exhibited weaker functional responses to human presence than those in more disturbed areas. Avoidance of rugged terrain is also context-dependent, with stronger avoidance when local ruggedness is high. The only exception to the observed intraspecific variability is consistent species-level responses to road proximity. Main Conclusions Our findings contribute to the understanding of habitat selection by terrestrial mammals, showing that selection is largely shaped by environmental conditions and that animals exhibit high plasticity in their responses. Our results also provide further evidence for the significant impact of roads on animal movement. These insights can help us to understand the potential effects of environmental change on the behaviour of mammal species around the world.

Global Ecology and Biogeography↗

Where will the cat cross the road? Comparing camera and GPS-based models for identifying wildlife corridors

Designing effective wildlife corridors is a critical conservation challenge in fragmented landscapes. GPS-based step selection functions strongly predict dispersal corridors and connectivity, but GPS collaring can be expensive and invasive. Camera-based occupancy models are widely used for connectivity analyses but may involve trade-offs in data resolution. Despite widespread use of both approaches, few studies have directly compared them using concurrent datasets. We developed a stacked single-species, single-season occupancy model and a Circuitscape connectivity surface for mountain lions (Puma concolor) on Washington’s Olympic Peninsula, USA, and compared them with a connectivity surface from an existing integrated step selection function. Both models predicted mountain lion GPS locations well, with binned Spearman rank correlations of 1 for Circuitscape and 0.96 for the step selection function, though step selection better identified habitat use by dispersers. Connectivity predictions were moderately correlated across the landscape ( r = 0.26), but agreement was strongest in human-dominated areas most critical for corridor planning. We conclude that GPS-based approaches are advantageous when data collection is feasible and the focus is on dispersal or fine-scale movement. However, camera-based approaches may be preferable for multi-species monitoring, large spatial and temporal scales, noninvasive sampling, when resources are limited, or when fine-scale or dispersal-specific inference is not required.

Washington↗

Intrinsic and environmental drivers of pairwise cohesion in wild Canis social groups

Animals within social groups respond to costs and benefits of sociality by adjusting the proportion of time they spend in close proximity to other individuals in the group (cohesion). Variation in cohesion between individuals, in turn, shapes important group-level processes such as subgroup formation and fission–fusion dynamics. Although critical to animal sociality, a comprehensive understanding of the factors influencing cohesion remains a gap in our knowledge of cooperative behavior in animals. We tracked 574 individuals from six species within the genus Canis in 15 countries on four continents with GPS telemetry to estimate the time that pairs of individuals within social groups spent in close proximity and test hypotheses regarding drivers of cohesion. Pairs of social canids ( Canis spp.) varied widely in the proportion of time they spent together (5%–100%) during seasonal monitoring periods relative to both intrinsic characteristics and environmental conditions. The majority of our data came from three species of wolves (gray wolves, eastern wolves, and red wolves) and coyotes. For these species, cohesion within social groups was greatest between breeding pairs and varied seasonally as the nature of cooperative activities changed relative to annual life history patterns. Across species, wolves were more cohesive than coyotes. For wolves, pairs were less cohesive in larger groups, and when suitable, small prey was present reflecting the constraints of food resources and intragroup competition on social associations. Pair cohesion in wolves declined with increased anthropogenic modification of the landscape and greater climatic variability, underscoring challenges for conserving social top predators in a changing world. We show that pairwise cohesion in social groups varies strongly both within and across Canis species, as individuals respond to changing ecological context defined by resources, competition, and anthropogenic disturbance. Our work highlights that cohesion is a highly plastic component of animal sociality that holds significant promise for elucidating ecological and evolutionary mechanisms underlying cooperative behavior.

Ecology↗

Neonatal antipredator tactics shape female movement patterns in large herbivores

Caring for newborn offspring hampers resource acquisition of mammalian females, curbing their ability to meet the high energy expenditure of early lactation. Newborns are particularly vulnerable, and large herbivores have evolved a continuum of neonatal anti-predator tactics, ranging from immobile hider to highly mobile follower offspring. How these tactics constrain female movements around parturition is unknown, particularly within the current context of increasing habitat fragmentation and earlier plant phenology caused by global warming. Using a analysis across 54 populations of 23 species of large herbivores, we show that mothers adjust their movements to variation in resource productivity and heterogeneity according to their offspring’s neonatal tactic. Mothers with hider offspring are unable to exploit environments where the variability of resources occurs at a broad scale, which might alter resource allocation compared to mothers with follower offspring. Our findings reveal that the overlooked neonatal tactic plays a key role for predicting how species are coping with environmental variation.

Nature Ecology & Evolution↗

Behavioral responses of terrestrial mammals to COVID-19 lockdowns

COVID-19 lockdowns reduced human mobility, providing an opportunity to disentangle its effects on animals from those of landscape modifications. Using GPS data, we compared movements and road avoidance of 2300 terrestrial mammals (43 species) during the 2020 lockdowns to the same period in 2019. Individual responses were variable, with no change in average movements or road avoidance behavior, likely due to variable lockdown conditions. However, under strict lockdowns,10-day, 95th percentile displacements increased by 73%, suggesting increased landscape permeability. In areas of high human footprint, animals' 1-hour, 95th percentile displacements declined by 12%, and animals were 36% closer to roads, indicating reduced avoidance during lockdowns. Overall, lockdowns rapidly altered some spatial behaviors, highlighting the variable but substantial impact of human mobility on wildlife worldwide.

Science↗

Patterns and processes of pathogen exposure in gray wolves across North America

The presence of many pathogens varies in a predictable manner with latitude, with infections decreasing from the equator towards the poles. We investigated the geographic trends of pathogens infecting a widely distributed carnivore: the gray wolf ( Canis lupus ). Specifically, we investigated which variables best explain and predict geographic trends in seroprevalence across North American wolf populations and the implications of the underlying mechanisms. We compiled a large serological dataset of nearly 2000 wolves from 17 study areas, spanning 80° longitude and 50° latitude. Generalized linear mixed models were constructed to predict the probability of seropositivity of four important pathogens: canine adenovirus, herpesvirus, parvovirus, and distemper virus—and two parasites: Neospora caninum and Toxoplasma gondii . Canine adenovirus and herpesvirus were the most widely distributed pathogens, whereas N. caninum was relatively uncommon. Canine parvovirus and distemper had high annual variation, with western populations experiencing more frequent outbreaks than eastern populations. Seroprevalence of all infections increased as wolves aged, and denser wolf populations had a greater risk of exposure. Probability of exposure was positively correlated with human density, suggesting that dogs and synanthropic animals may be important pathogen reservoirs. Pathogen exposure did not appear to follow a latitudinal gradient, with the exception of N. caninum . Instead, clustered study areas were more similar: wolves from the Great Lakes region had lower odds of exposure to the viruses, but higher odds of exposure to N. caninum and T. gondii ; the opposite was true for wolves from the central Rocky Mountains. Overall, mechanistic predictors were more informative of seroprevalence trends than latitude and longitude. Individual host characteristics as well as inherent features of ecosystems determined pathogen exposure risk on a large scale. This work emphasizes the importance of biogeographic wildlife surveillance, and we expound upon avenues of future research of cross-species transmission, spillover, and spatial variation in pathogen infection.

Alaska, Alberta, Arizona, British Columbia, Michig↗

Mapping out a future for ungulate migrations

Migration of ungulates (hooved mammals) is a fundamental ecological process that promotes abundant herds, whose effects cascade up and down terrestrial food webs. Migratory ungulates provide the prey base that maintains large carnivore and scavenger populations and underpins terrestrial biodiversity (fig. S1). When ungulates move in large aggregations, their hooves, feces, and urine create conditions that facilitate distinct biotic communities. The migrations of ungulates have sustained humans for thousands of years, forming tight cultural links among Indigenous people and local communities. Yet ungulate migrations are disappearing at an alarming rate ( 1 ). Efforts by wildlife managers and conservationists are thwarted by a singular challenge: Most ungulate migrations have never been mapped in sufficient detail to guide effective conservation. Without a strategic and collaborative effort, many of the world's great migrations will continue to be truncated, severed, or lost in the coming decades. Fortunately, a combination of animal tracking datasets, historical records, and local and Indigenous knowledge can form the basis for a global atlas of migrations, designed to support conservation action and policy at local, national, and international levels.

Science↗

Wave-like patterns of plant phenology determine ungulate movement tactics

Animals exhibit a diversity of movement tactics [1]. Tracking resources that change across space and time is predicted to be a fundamental driver of animal movement [2]. For example, some migratory ungulates (i.e., hooved mammals) closely track the progression of highly nutritious plant green-up, a phenomenon called “green-wave surfing” [3-5]. Yet, general principles describing how the dynamic nature of resources determine movement tactics are lacking [6]. We tested emerging theory that predicts surfing and the existence of migratory behavior will be favored in environments where green-up is fleeting and moves sequentially across large landscapes (i.e., wave-like green-up) [7]. Landscapes exhibiting wave-like patterns of green-up facilitated surfing and explained the existence of migratory behavior across 61 populations of four ungulate species on two continents (n=1,696 individuals). At the species level, foraging benefits were equivalent between tactics, suggesting that each movement tactic is fine tuned to local patterns of plant phenology. For decades, ecologists have sought to understand how animals move to select habitat, commonly defining habitat as a set of static patches [8, 9]. Our findings indicate that animal movement tactics emerge as a function of the flux of resources across space and time, underscoring the need to redefine habitat to include its dynamic attributes. As global habitats continue to be modified by anthropogenic disturbance and climate change [10], our synthesis provides a generalizable framework to understand how animal movement will be influenced by altered patterns of resource phenology.

Alberta, British Columbia, Idaho, Wyoming↗

Migrating bison engineer the green wave

Newly emerging plants provide the best forage for herbivores. To exploit this fleeting resource, migrating herbivores align their movements to surf the wave of spring green-up. With new technology to track migrating animals, the Green Wave Hypothesis has steadily gained empirical support across a diversity of migratory taxa. This hypothesis assumes the green wave is controlled by variation in climate, weather, and topography, and its progression dictates the timing, pace, and extent of migrations. However, aggregate grazers that are also capable of engineering grassland ecosystems make some of the world’s most impressive migrations, and it is unclear how the green wave determines their movements. Here we show that Yellowstone’s bison ( Bison bison ) do not choreograph their migratory movements to the wave of spring green-up. Instead, bison modify the green wave as they migrate and graze. While most bison surfed during early spring, they eventually slowed and let the green wave pass them by. However, small-scale experiments indicated that feedback from grazing sustained forage quality. Most importantly, a 6-fold decadal shift in bison density revealed that intense grazing caused grasslands to green up faster, more intensely, and for a longer duration. Our finding broadens our understanding of the ways in which animal movements underpin the foraging benefit of migration. The widely accepted Green Wave Hypothesis needs to be revised to include large aggregate grazers that not only move to find forage, but also engineer plant phenology through grazing, thereby shaping their own migratory movements.

California↗

Factors influencing elk recruitment across ecotypes in the Western United States

Ungulates are key components in ecosystems and economically important for sport and subsistence harvest. Yet the relative importance of the effects of weather conditions, forage productivity, and carnivores on ungulates are not well understood. We examined changes in elk ( Cervus canadensis ) recruitment (indexed as age ratios) across 7 states and 3 ecotypes in the northwestern United States during 1989–2010, while considering the effects of predator richness, forage productivity, and precipitation. We found a broad‐scale, long‐term decrease in elk recruitment of 0.48 juveniles/100 adult females/year. Weather conditions (indexed as summer and winter precipitation) showed small, but measurable, influences on recruitment. Forage productivity on summer and winter ranges (indexed by normalized difference vegetation index [NDVI] metrics) had the strongest effect on elk recruitment relative to other factors. Relationships between forage productivity and recruitment varied seasonally and regionally. The productivity of winter habitat was more important in southern parts of the study area, whereas annual variation in productivity of summer habitat had more influence on recruitment in northern areas. Elk recruitment varied by up to 15 juveniles/100 adult females across the range of variation in forage productivity. Areas with more species of large carnivores had relatively low elk recruitment, presumably because of increased predation. Wolves ( Canis lupus ) were associated with a decrease of 5 juveniles/100 adult females, whereas grizzly bears ( Ursus arctos ) were associated with an additional decrease of 7 juveniles/100 adult females. Carnivore species can have a critical influence on ungulate recruitment because their influence rivals large ranges of variation in environmental conditions. A more pressing concern, however, stems from persistent broad‐scale decreases in recruitment across the distribution of elk in the northwestern United States, irrespective of carnivore richness. Our results suggest that wildlife managers interested in improving recruitment of elk consider the combined effects of habitat and predators. Efforts to manage summer and winter ranges to increase forage productivity may have a positive effect on recruitment.

Journal of Wildlife Management↗

Moving in the Anthropocene: Global reductions in terrestrial mammalian movements

Animal movement is fundamental for ecosystem functioning and species survival, yet the effects of the anthropogenic footprint on animal movements have not been estimated across species. Using a unique GPS-tracking database of 803 individuals across 57 species, we found that movements of mammals in areas with a comparatively high human footprint were on average one-half to one-third the extent of their movements in areas with a low human footprint. We attribute this reduction to behavioral changes of individual animals and to the exclusion of species with long-range movements from areas with higher human impact. Global loss of vagility alters a key ecological trait of animals that affects not only population persistence but also ecosystem processes such as predator-prey interactions, nutrient cycling, and disease transmission.

Science↗

Scaling-up camera traps: monitoring the planet's biodiversity with networks of remote sensors

Countries committed to implementing the Convention on Biological Diversity's 2011–2020 strategic plan need effective tools to monitor global trends in biodiversity. Remote cameras are a rapidly growing technology that has great potential to transform global monitoring for terrestrial biodiversity and can be an important contributor to the call for measuring Essential Biodiversity Variables. Recent advances in camera technology and methods enable researchers to estimate changes in abundance and distribution for entire communities of animals and to identify global drivers of biodiversity trends. We suggest that interconnected networks of remote cameras will soon monitor biodiversity at a global scale, help answer pressing ecological questions, and guide conservation policy. This global network will require greater collaboration among remote-camera studies and citizen scientists, including standardized metadata, shared protocols, and security measures to protect records about sensitive species. With modest investment in infrastructure, and continued innovation, synthesis, and collaboration, we envision a global network of remote cameras that not only provides real-time biodiversity data but also serves to connect people with nature.

Frontiers in Ecology and the Environment↗

Summer habitat selection by Dall’s sheep in Wrangell-St. Elias National Park and Preserve, Alaska

Sexual segregation occurs frequently in sexually dimorphic species, and it may be influenced by differential habitat requirements between sexes or by social or evolutionary mechanisms that maintain separation of sexes regardless of habitat selection. Understanding the degree of sex-specific habitat specialization is important for management of wildlife populations and the design of monitoring and research programs. Using mid-summer aerial survey data for Dall’s sheep ( Ovis dalli dalli ) in southern Alaska during 1983–2011, we assessed differences in summer habitat selection by sex and reproductive status at the landscape scale in Wrangell-St. Elias National Park and Preserve (WRST). Males and females were highly segregated socially, as were females with and without young. Resource selection function (RSF) models containing rugged terrain, intermediate values of the normalized difference vegetation index (NDVI), and open landcover types best explained resource selection by each sex, female reproductive classes, and all sheep combined. For male and all female models, most coefficients were similar, suggesting little difference in summer habitat selection between sexes at the landscape scale. A combined RSF model therefore may be used to predict the relative probability of resource selection by Dall’s sheep in WRST regardless of sex or reproductive status.

Alaska↗

Annual elk calf survival in a multiple carnivore system

The realized effect of multiple carnivores on juvenile ungulate recruitment may depend on the carnivore assemblage as well as compensation from forage and winter weather severity, which may mediate juvenile vulnerability to predation in ungulates. We used a time-to-event approach to test for the effects of risk factors on annual elk ( Cervus canadensis ) calf survival and to estimate cause-specific mortality rates for 2 elk populations in adjacent study areas in the southern Bitterroot Valley, Montana, USA, during 2011–2014. We captured and radio-tagged 286 elk calves: 226 neonates, and 60 6-month-old calves. Summer survival probability was less variable than winter ( P = 0.12) and averaged 0.55 (95% CI = 0.47–0.63), whereas winter survival varied more than summer and significantly across study years ( P = 0.003) and averaged 0.73 (95% CI = 0.64–0.81). During summer, elk calf survival increased with biomass of preferred forage biomass, and was slightly lower following winters with high precipitation; exposure to mountain lion ( Puma concolor ) predation risk was unimportant. In contrast, during winter, we found that exposure to mountain lion predation risk influenced survival, with a weak negative effect of winter precipitation. We found no evidence that forage availability or winter weather severity mediated vulnerability to mountain lion predation risk in summer or winter (e.g., an interaction), indicating that the effect of mountain lion predation was constant regardless of spatial variation in forage or weather. Mountain lions dominated known causes of elk calf mortality in summer and winter, with estimated cause-specific mortality rates of 0.14 (95% CI = 0.09–0.20) and 0.12 (95% CI = 0.07–0.18), respectively. The effect of carnivores on juvenile ungulate recruitment varies across ecological systems depending on relative carnivore densities. Mountain lions may be the most important carnivore for ungulates, especially where grizzly bears ( Ursus arctos ) and wolves ( Canis lupus ) are rare or recovering. Finally, managers may need to reduce adult female harvest of elk as carnivores recolonize to balance carnivore and ungulate management objectives, especially in less productive habitats for elk.

Journal of Wildlife Management↗

Gravel-bed river floodplains are the ecological nexus of glaciated mountain landscapes

Gravel-bed river floodplains in mountain landscapes disproportionately concentrate diverse habitats, nutrient cycling, productivity of biota, and species interactions. Although stream ecologists know that river channel and floodplain habitats used by aquatic organisms are maintained by hydrologic regimes that mobilize gravel-bed sediments, terrestrial ecologists have largely been unaware of the importance of floodplain structures and processes to the life requirements of a wide variety of species. We provide insight into gravel-bed rivers as the ecological nexus of glaciated mountain landscapes. We show why gravel-bed river floodplains are the primary arena where interactions take place among aquatic, avian, and terrestrial species from microbes to grizzly bears and provide essential connectivity as corridors for movement for both aquatic and terrestrial species. Paradoxically, gravel-bed river floodplains are also disproportionately unprotected where human developments are concentrated. Structural modifications to floodplains such as roads, railways, and housing and hydrologicaltering hydroelectric or water storage dams have severe impacts to floodplain habitat diversity and productivity, restrict local and regional connectivity, and reduce the resilience of both aquatic and terrestrial species, including adaptation to climate change. To be effective, conservation efforts in glaciated mountain landscapes intended to benefit the widest variety of organisms need a paradigm shift that has gravel-bed rivers and their floodplains as the central focus and that prioritizes the maintenance or restoration of the intact structure and processes of these critically important systems throughout their length and breadth.

Science Advances↗

Linking resource selection and mortality modeling for population estimation of mountain lions in Montana

To be most effective, the scale of wildlife management practices should match the range of a particular species’ movements. For this reason, combined with our inability to rigorously or regularly census mountain lion populations, several authors have suggested that mountain lions be managed in a source-sink or metapopulation framework. We used a combination of resource selection functions, mortality estimation, and dispersal modeling to estimate cougar population levels in Montana statewide and potential population level effects of planned harvest levels. Between 1980 and 2012, 236 independent mountain lions were collared and monitored for research in Montana. From these data we used 18,695 GPS locations collected during winter from 85 animals to develop a resource selection function (RSF), and 11,726 VHF and GPS locations from 142 animals along with the locations of 6343 mountain lions harvested from 1988–2011 to validate the RSF model. Our RSF model validated well in all portions of the State, although it appeared to perform better in Montana Fish, Wildlife and Parks (MFWP) Regions 1, 2, 4 and 6, than in Regions 3, 5, and 7. Our mean RSF based population estimate for the total population (kittens, juveniles, and adults) of mountain lions in Montana in 2005 was 3926, with almost 25% of the entire population in MFWP Region 1. Estimates based on a high and low reference population estimates produce a possible range of 2784 to 5156 mountain lions statewide. Based on a range of possible survival rates we estimated the mountain lion population in Montana to be stable to slightly increasing between 2005 and 2010 with lambda ranging from 0.999 (SD = 0.05) to 1.02 (SD = 0.03). We believe these population growth rates to be a conservative estimate of true population growth. Our model suggests that proposed changes to female harvest quotas for 2013–2015 will result in an annual statewide population decline of 3% and shows that, due to reduced dispersal, changes to harvest in one management unit may affect population growth in neighboring units where smaller or even no changes were made. Uncertainty regarding dispersal levels and initial population density may have a significant effect on predictions at a management unit scale (i.e. 2000 km 2 ), while at a regional scale (i.e. 50,000 km 2 ) large differences in initial population density result in relatively small changes in population growth rate, and uncertainty about dispersal may not be as influential. Doubling the presumed initial density from a low estimation of 2.19 total animals per 100 km 2 resulted in a difference in annual population growth rate of only 2.6% statewide when compared to high density of 4.04 total animals per 100 km 2 (low initial population estimate λ = 0.99, while high initial population estimate λ = 1.03). We suggest modeling tools such as this may be useful in harvest planning at a regional and statewide level.

Ecological Modelling↗

Modeling multi-scale resource selection for bear rubs in northwestern Montana

Both black ( Ursus americanus ) and grizzly bears ( U. arctos ) are known to rub on trees and other objects, producing a network of repeatedly used and identifiable rub sites. In 2012, we used a resource selection function to evaluate hypothesized relationships between locations of 887 bear rubs in northwestern Montana, USA, and elevation, slope angle, density of open roads and distance from areas of heightened plant-productivity likely containing forage for bears. Slope and density of open roads were negatively correlated with rub presence. No other covariates were supported as explanatory variables. We also hypothesized that bear rubs would be more strongly associated with closed roads and developed trails than with game trails. The frequencies of bear rubs on 30 paired segments of developed tracks and game trails were not different. Our results suggest bear rubs may be associated with bear travel routes, and support their use as “random” sampling devices for non-invasive spatial capture–recapture population monitoring.

Montana↗

A test of the compensatory mortality hypothesis in mountain lions: a management experiment in West-Central Montana

Mountain lions ( Puma concolor ) are widely hunted for recreation, population control, and to reduce conflict with humans, but much is still unknown regarding the effects of harvest on mountain lion population dynamics. Whether human hunting mortality on mountain lions is additive or compensatory is debated. Our primary objective was to investigate population effects of harvest on mountain lions. We addressed this objective with a management experiment of 3 years of intensive harvest followed by a 6-year recovery period. In December 2000, after 3 years of hunting, approximately 66% of a single game management unit within the Blackfoot River watershed in Montana was closed to lion hunting, effectively creating a refuge representing approximately 12% (915 km 2 ) of the total study area (7,908 km 2 ). Hunting continued in the remainder of the study area, but harvest levels declined from approximately 9/1,000 km 2 in 2001 to 2/1,000 km 2 in 2006 as a result of the protected area and reduced quotas outside. We radiocollared 117 mountain lions from 1998 to 2006. We recorded known fates for 63 animals, and right-censored the remainder. Although hunting directly reduced survival, parameters such as litter size, birth interval, maternity, age at dispersal, and age of first reproduction were not significantly affected. Sensitivity analysis showed that female survival and maternity were most influential on population growth. Life-stage simulation analysis (LSA) demonstrated the effect of hunting on the population dynamics of mountain lions. In our non-hunted population, reproduction (kitten survival and maternity) accounted for approximately 62% of the variation in growth rate, whereas adult female survival accounted for 30%. Hunting reversed this, increasing the reliance of population growth on adult female survival (45% of the variation in population growth), and away from reproduction (12%). Our research showed that harvest at the levels implemented in this study did not affect population productivity (i.e., maternity), but had an additive effect on mountain lion mortality, and therefore population growth. Through harvest, wildlife managers have the ability to control mountain lion populations.

Montana↗