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# Population viability of recolonizing cougars in midwestern North America

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... In addition, RAMAS/GIS software is well-established and has been used to quantify population trajectories and extinction risk of species affected by factors such as fires, land use change, and climate change (e.g., Akçakaya et al., 2005: Lawson et al., 2010; and to inform conservation planning (Early and Thomas, 2007;Hinrichsen, 2009). For large carnivore species, it has been used to model population dynamics and viability of cougars (Puma concolor; LaRue and Nielsen, 2015), gray wolves (Canis lupus; Maletzke et al., 2015) and Amur tigers (Panthera tigris; Tian et al., 2011). Within this framework, we required three types of information: (1) a model of bear habitat suitability which could be delineated into core habitat patches, (2) demographic parameters including estimates of patch-level carrying capacity, dispersal, and age-based reproduction and survival, and (3) parameters necessary to simulate environmental stochasticity. ...
... We modeled 5000 replications over 10 years with an annual time step, a short time period to reduce error propagation (Akçakaya et al., 2005;LaRue and Nielsen, 2015). We included environmental stochasticity as random variation in fecundities, survivals, and dispersal rates within 10% (i.e. ...
... In addition, our model contained several assumptions, was intentionally conservative, and relatively simple. We chose a short time frame to 1) be able to assume density-independence, and 2) limit error propagation (Haines et al., 2006;LaRue and Nielsen 2015), especially since black bears have life spans over 10 years (Clark et al., 2021). We were conservative in selecting initially occupied patches using bear capture locations at the start of the modeling period, yet other patches could have been occupied. ...
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Despite extensive range contractions, several large carnivore species have recently recolonized portions of their former range. Since the 1990s, American black bear reports have increased in Missouri, USA, corresponding with increasing abundance and distribution. As effective management benefits from sound information on wildlife demographics and spatial distribution, we used a spatially-explicit metapopulation model built from local data to inform a stage matrix, which was used in conjunction with a previously created habitat suitability model to quantify and predict the growth and expansion of the black bear population. We collected demographic data during 2011-2021 from 162 female bears with 159 young from 70 litters. Average litter size was 1.95, 42% of cubs were female, and cub survival to yearling was, on average, 95% for females and 70% for males. The habitat suitability model identified 53 core areas most suitable for bears in Missouri. Increasing from previous local estimates of bear population size (~280 bears in 2012), the model estimated the total bear population inhabiting core habitat patches in year 10 as 732 individuals (598-873, 5th-95th percentiles), with an average annual growth rate of 1.11. The estimated carrying capacity of the core habitat patches (excluding cubs-of-the-year) was ~2700 bears, but adding marginal and highly marginal habitat patches increased the carrying capacity to ~5500 bears. Population estimates were most sensitive to variations in stage matrix parameters though overall patch colonization estimates were relatively stable. Our work is an important step toward understanding the recolo-nization potential of black bears in Missouri and can aid future studies projecting large carnivore density in other areas as well as values and attitudes toward these wide-ranging species. Developing holistic conservation frameworks will be critical if human acceptance toward large carnivores continues increasing and carnivore recolonization of historical range continues.
... Cougars (Puma concolor) have lost over 35% of their historical range throughout North America (Laliberte and Ripple, 2004), yet have been able to persist and recolonize parts of their former range despite habitat fragmentation and human-induced mortality (LaRue et al., 2012;LaRue and Nielsen, 2016). The only currently documented breeding population of cougars in the eastern USA is in Florida (Vickers et al., 2015). ...
... Previous efforts to map potential distribution for cougars in the Midwest were developed mostly using expert opinion (e.g. O'Neil et al., 2014;LaRue and Nielsen, 2016;Smith et al., 2016). In contrast, we use a decade of verified cougar observation reports to model and predict potential distribution, quantify land cover selection ratios, map landscape connectivity, and evaluate the land ownership (i.e., protection status) of predicted core patches and linkages. ...
... Coarse scale = land cover proportions of buffers vs. the study area (Michigan, Minnesota, and Wisconsin however, differences in the distribution of highly suitable patches identified among these studies. For example, northeastern Minnesota has been assessed by experts as highly suitable LaRue and Nielsen, 2016) whereas we suggest it is of low to moderate suitability. It is possible that this area has had inflated suitability in previous studies due to being remote (low human density and road density) rather than representing high-quality cougar habitat. ...
Article
Cougars (Puma concolor) have lost substantial portions of their historical range yet increased sightings suggest potential for re-establishment in some regions; greater understanding of potential distribution and connectivity is necessary to make sound management and policy decisions. Specifically, the Great Lakes region of the USA will likely be an important area for cougar range expansion into the Midwest and Eastern USA. We used verified cougar observations to model and predict potential distribution and connectivity in the Great Lakes region. We compiled all confirmed observations of cougars from Michigan, Minnesota, and Wisconsin (2010–2020); which resulted in 180 reports (154 images/videos, 20 signs, 6 mortalities). We developed an ensemble distribution model (1 km res) based on three machine learning methods. We used weighted cost-distances to identify linkages between core areas and circuit theory to model overall connectivity potential. We calculated selection ratios for land covers (30 m res) at fine and coarse scales. The ensemble distribution model had good performance (ROC of 0.94). Suitability was positively associated with increasing vegetation structure, lower distance to natural cover, and mid-high terrain ruggedness. Forest covers were always selected for regardless of scale, and human development was avoided only at the coarser scale. We identified 191 core patches and 362 linkages connecting them; only 50.1% of them were located within legally protected areas. We identified high regional connectivity in a generally west-east direction. Successful conservation of large carnivores like cougars will depend on conserving not only habitat patches and linkages but also efforts to facilitate long-term coexistence.
... Some large carnivores are currently recovering in human-dominated landscapes (Chapron et al., 2014;Gantchoff and Belant, 2017;LaRue and Nielsen, 2016); thus there is a growing need to predict their establishment and to facilitate management actions that help ensure their long-term viability and mitigate potential conflicts (Redpath et al., 2013). Large carnivore movements are increasingly constrained by human land use, forcing them to occupy or cross human-modified landscapes in search of suitable habitat (Fahrig, 2001;Fahrig, 2007;Tucker et al., 2018). ...
... Suitable habitat patches are often identified from resource selection functions (RSFs) Manly et al., 2007), and the potential for connectivity is evaluated by assessing the cost of passing through different habitat types based on RSF results (Abrahms et al., 2017;Beier et al., 2008;Chetkiewicz and Boyce, 2009;Sawyer et al., 2011). Dispersal is the main driver of both population connectivity and recolonization (Jackson et al., 2016;LaRue and Nielsen, 2016;Morrison et al., 2015). As dispersal events often involve long distance movements (Fahrig, 2007;Tucker et al., 2018), and resource requirements may differ between established and dispersing individuals (Abrahms et al., 2017;Chetkiewicz et al., 2006), connectivity should ideally be assessed using actual dispersal events (Beyer et al., 2010). ...
... The ability of large carnivores to recolonize their former ranges is a central conservation issue in many parts of the world (Chapron et al., 2014;Gantchoff and Belant, 2017;LaRue and Nielsen, 2016;Mattson and Clark, 2010). Therefore, in this study we predicted areas of lynx recolonization based on resource selection functions (RSFs) from the source area, and successfully validated these predictions with data from individuals in the settlement area and with independent monitoring data (Fig. 2). ...
Article
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The conservation of wide-ranging species presents challenges in a world of intensified human land use, forcing animals to occupy and recolonize human-modified landscapes. Although identifying suitable habitat and ensuring connectivity are important in supporting natural recolonization, these actions are rarely validated due to difficulties in monitoring such events. In Sweden, the Eurasian lynx (Lynx lynx) is now recolonizing its former range, after centuries of persecution. We in- vestigated resource selection based on telemetry data from 108 lynx monitored over 20 years. We assessed the differences between the established population in central Sweden and the recolonizing population in southern Sweden, and between established and dispersing individuals. We found that models based on central Sweden successfully identified core habitat patches for establishment in southern Sweden, validated after recolonization. We also found that lynx selected for higher habitat suitability during the recolonization phase, and that dispersing individuals were less selective than established lynx. Using cost-distance analysis, we assessed connectivity between central and southern Sweden, and found that landscape permeability was higher when based on dispersing lynx compared to established lynx. Altogether, our findings suggest that when landscapes are sufficiently similar between source and recolonization areas, resource selection information from an established population can be useful for managers seeking to facilitate recolonization of wide-ranging species. We recommend more frequent use of validation during and after recolonization events, to improve our common understanding of habitat suitability and connectivity modeling, and therefore to enable more active management of recolonization events.
... Much of Midwestern North America has witnessed a rise in the mountain lion sightings as they re-colonize parts of their former range (Cougar Network, 2007;Rosatte, 2011;LaRue et al. 2012), which also increases humancougar interactions (Torres et al. 1996;Sweanor and Logan 2010;LaRue et al. 2012). The cougar population has been extirpated for >100 years, and only about 170 confirmed cougar sightings were reported during 1990-2008 across the entire Midwestern U.S. Recolonization warrants attention because mountain lions can help change ecosystem functioning upon their return (LaRue et al. 2012;LaRue & Nielsen, 2016). The recolonization of wolves, such as in Yellowstone (Ripple & Beschta, 2004;Fortin et al. 2005;LaRue & Nielsen, 2016), has greatly impacted competing carnivore populations through competitive exclusion (LaRue & Nielsen, 2016). ...
... The cougar population has been extirpated for >100 years, and only about 170 confirmed cougar sightings were reported during 1990-2008 across the entire Midwestern U.S. Recolonization warrants attention because mountain lions can help change ecosystem functioning upon their return (LaRue et al. 2012;LaRue & Nielsen, 2016). The recolonization of wolves, such as in Yellowstone (Ripple & Beschta, 2004;Fortin et al. 2005;LaRue & Nielsen, 2016), has greatly impacted competing carnivore populations through competitive exclusion (LaRue & Nielsen, 2016). ...
... The cougar population has been extirpated for >100 years, and only about 170 confirmed cougar sightings were reported during 1990-2008 across the entire Midwestern U.S. Recolonization warrants attention because mountain lions can help change ecosystem functioning upon their return (LaRue et al. 2012;LaRue & Nielsen, 2016). The recolonization of wolves, such as in Yellowstone (Ripple & Beschta, 2004;Fortin et al. 2005;LaRue & Nielsen, 2016), has greatly impacted competing carnivore populations through competitive exclusion (LaRue & Nielsen, 2016). ...
Article
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The mountain lion range once extended throughout the state of Minnesota. The breeding population has been greatly reduced with time, new roads, and timber harvesting, which have broken large tracts of contiguous forest into isolated patches that are too small and no longer suitable for the breeding mountain lion population. The objective of this study is to use suitability analysis to determine the most suitable habitat to conserve mountain lion populations threatened by habitat fragmentation. To attain our objective, we created three sub models that contribute to the overarching goal of the suitability model. A habitat sub model was developed for finding the best habitat, a food sub model for access to the maximum amount of food needed, and a security sub model focusing on the distance from houses, roads, and urban development. Using the Weighted Sum tool, the three sub models were combined to produce a suitability surface based on the trade-off of the preferences of the goals represented by each sub model. Our suitability model shows large areas of high-quality mountain lion habitat in the northern and north-eastern sections of the state. These areas contain favourable locations for mountain lion habitat, such as forested land cover, low-density populations, steep slopes, short distances to streams, and area unimpeded by major roads. The southern and western parts of the state are characterized by lower slopes, more agricultural land, grassland, developed land, and higher population density, which results in lower quality habitat. The twin cities have the worst mountain lion habitat.
... The United States has more developed lands and higher human densities than Canada, which may reduce the overall availability of potential cougar habitat relative to Canada. Despite this, recolonization events have Continued recolonization is possible from western cougar populations into potential habitats such as we have modeled (LaRue et al. 2016(LaRue et al. , 2019. Although smaller patches of potential habitats were considered too small to maintain a viable population (Beier 1993;Thatcher et al. 2006) existed throughout the study area, these small areas may still be used by dispersing cougars (Beier 1995;Kautz et al. 2006;LaRue and Nielsen 2016). ...
... Despite this, recolonization events have Continued recolonization is possible from western cougar populations into potential habitats such as we have modeled (LaRue et al. 2016(LaRue et al. , 2019. Although smaller patches of potential habitats were considered too small to maintain a viable population (Beier 1993;Thatcher et al. 2006) existed throughout the study area, these small areas may still be used by dispersing cougars (Beier 1995;Kautz et al. 2006;LaRue and Nielsen 2016). ...
Article
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Context Cougars (Puma concolor) have been recolonizing Midwestern North America during the past 3 decades with > 950 cougar confirmations east of established populations. Due to an increase in confirmations east of current breeding populations, evaluation of cougar habitat suitability and connectivity is needed. However, few studies have assessed the habitat potential for cougar recolonization in the eastern portion of their former range. Objectives We used various habitat quality thresholds to model potential cougar habitats and dispersal corridors throughout eastern North America. Methods Based on expert opinion, we used landcover, slope, human density, distance to roads, and distance to water as model variables. Least-cost path methods were used to model dispersal corridors from western populations to potential eastern habitat patches. Results Patches of suitable habitat ranged in size from 3868 km² (Ozark Mountains) to > 2,490,850 km² (central and eastern Canada). Potential habitats were predominantly forest and shrubland, contained little anthropogenic development, and had high stream densities. Dispersal corridors were present throughout the study area. Corridors largely consisted of forested and cultivated landscapes and had higher road densities than habitat patches. Conclusions Our research provides conservationists with insights into areas suitable for cougar recolonization so they may proactively plan for potential cougar populations east of their current range. This work also provides a framework for evaluating multiple levels of landscape suitability for recolonizing species.
... Because of improved wildlife management practices, implementation of game laws and conservation policies, and habitat recovery and restoration during the latter half of the 20 th century, some large carnivores successfully recolonized portions of North America and Europe. Gray wolves (Canis lupus) and brown bears (Ursus arctos), for instance, both naturally recolonized parts of Sweden, Italy, and Greece (Pyare et al. 2004, Fabbri et al. 2014, Votsi et al. 2016, and cougars and American black bears (Ursus americanus) are projected to recolonize localized areas of the United States during coming decades (Smith et al. 2015, LaRue andNielsen 2016). However, the ecological importance and the imperiled status of many large carnivores may necessitate more timely restoration than the rate at which natural recolonization typically occurs for these wide-ranging carnivores. ...
... (Pyare et al. 2004, Fabbri et al. 2014, Votsi et al. 2016. Similar recolonizations will be difficult where human population density is high and habitat is severely degraded, but current predictions indicate regional large carnivore recolonizations may increase in frequency and geographical distribution in portions of the United States and Europe during coming decades (Chapron et al. 2014, Smith et al. 2015, LaRue and Nielsen 2016. ...
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Reintroduced populations are vulnerable to demographic and environmental stochasticity, deleterious genetic effects, and reduced population fitness, all of which can increase extinction probability. Population viability is principle to determining the status of reintroduced populations and for guiding management decisions. To attempt to reestablish black bear (Ursus americanus) populations in the central Appalachians, two reintroductions using small founder groups occurred during the 1990s in the Big South Fork area along the Kentucky-Tennessee border (BSF) and in the Jefferson National Forest along the Kentucky-Virginia border (KVP). My objectives were to estimate demographic and genetic parameters, and to evaluate long-term viability and reintroduction success for the KVP and BSF black bear populations. The KVP grew rapidly to 317–751 bears with a significantly female-biased sex ratio by 2013. Spatially explicit capture-recapture models suggested KVP recolonization may continue to the southwest and northeast along linear mountain ridges. Based on radio-monitoring during 2010–2014, high adult female survival and moderate mean litter sizes were estimated in both populations. All mortality was anthropogenic and males were 4.13 times more likely to die than females. Two-cub litters were most probable in the BSF, whereas the KVP had similar probabilities of two- and three-cub litters. The average annual mortality that occurred during the study period was sustainable and allowed for moderate growth (λKVP = 1.10; λBSF = 1.13). Continued mortality at the higher 2015 rate, however, resulted in probabilities of ≥25% population decline over 10 years of 0.52–0.53 and 0.97–0.98 in the KVP and BSF, respectively. Rapid population growth during the 13–17 years post-reintroduction and the overlapping generations inherent to bears retained genetic diversity. Cumulative findings indicated both reintroductions were successful at establishing viable, self-sustaining populations over the long-term. The anthropogenic mortality rate during 2015, if sustained, could cause precipitous declines in these populations. Reimplementation of annual vital rate monitoring and conservative harvests should be considered. Connectivity may be established between these two reintroduced black bear populations if growth and recolonization continue.
... Based on previous bobcat movement and habitat use studies (Johnson, Walker & Hudson, 2010;Reding et al., 2013;Hughes et al., 2019), we expect natural areas along river corridors or other natural habitats to act as conduits for dispersal in heavily altered (agricultural) areas. Overall, this suite of resource selection and connectivity analyses can help elucidate aspects of bobcat spatial ecology and potential limiting factors to population expansion, as well as provide important information for developing and implementing population models to inform the impacts of potential harvest on population persistence (LaRue & Nielsen, 2016). ...
Article
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Terrestrial carnivores are among the most imperiled species worldwide, yet some species are resilient and are recovering in human-dominated landscapes after decades or centuries of absence. Bobcat ( Lynx rufus ) populations were extirpated from much of Midwestern US in the mid-1800’s, and are currently expanding and recolonizing their former range. In this study, we investigated multi-scale habitat selection for Ohio’s expanding bobcat population, and examined habitat connectivity in order to evaluate the conduits for dispersal statewide. We used citizen observations collected between 1978 and 2019 and logistic regression to evaluate population-level habitat selection, and GPS telemetry data for 20 individuals collected between 2012 and 2014 and a distribution-weighted exponential Resource Selection Function to evaluate individual-level habitat selection within home ranges. At the population level, bobcats selected for higher amounts of forest and pasture (at a 50 km ² scale) and herbaceous vegetation (at 15–50 50 km ² scales), thus overall heterogeneous forested habitat. At individual (home range) level, bobcats selected for forested habitats with low road density and farther away from high traffic roads; they also showed weak selection for open habitat at the home range level. Male home ranges were significantly greater than female home ranges. Lastly, we used the population-level spatial outputs ( i.e . habitat suitability map) to parameterize habitat connectivity models using circuit theory in the program Circuitscape. We tested three relationships between habitat suitability and resistance to movement and used a subset of data on potential dispersing individuals to evaluate which relationship performed best. All three relationships performed almost equally well, and we calculated a weighted averaged connectivity map as our final map. Habitat was highly permeable to movements between core areas of two genetically distinct subpopulations located in southeastern Ohio. We also identified potential dispersal corridors from the core areas to other regions of Ohio dominated by agriculture and suburban development via forested riparian corridors. Overall, our analysis offers new information on habitat selection and connectivity in a rebounding felid population and offers important ecological information for wildlife management strategies. We recommend that the suitability and connectivity models should be periodically updated until the population reaches an equilibrium, and be integrated with data from neighboring states for a comprehensive assessment of a conservation success story.
... PVAs have been central to population conservation for decades (Soule 1987;Gerber and González-Suárez 2010;Chirakkal and Gerber 2010;Saunders et al., 2018). PVAs can be conducted aspatially based on demographic vital rates with the use of a standard Leslie matrix life-table analysis and simulation projection models (Kajin et al., 2012), using programs such as Vortex and RAMAS (Lindenmayer et al., 2000;LaRue and Nielsen 2016). PVAs also can be spatialised with individual-based simulation models (Watkins and Rose 2017) such as HexSim (Schumaker and Brookes 2018) or through spatially and temporally explicit population simulations (Visitin et al., 2020). ...
Article
Traditional population viability analysis (PVA) does not address the degree of measurement error or spatial and temporal variability of vital rate parameters, potentially leading to inappropriate conservation decision-making. We provide a methodology of applying Bayesian network (BN) modeling to PVA addressing these considerations, particularly for species with complex stage-class structures. We provide examples of three species from eastern Australia - hip pocket frog (Assa darilingtoni), squirrel glider (Petaurus norfolcensis) and giant burrowing frog (Heleioporus australiacus), comparing traditional matrix-based PVA with BN model analyses of mean stage abundance, quasi-extinction probability, and interval threshold extinction risk. Both approaches project similar population sizes, but BN PVA gave more clearly identifiable thresholds of population changes and extinction levels. The PVA BN uniquely represents complex stage-class structures and in a single network, including variation and uncertainty propagation of vital rates, to better inform conservation management decisions.
... The most common cause of mortality of Texas mountain lions is from lethal trapping for predator control, even for those individuals living on public lands, because they may leave park boundaries (Young et al., 2010). As such, the reported annual survival rates for mountain lions in West Texas are among the lowest reported across the species' range (Harveson et al., 2012;LaRue and Nielsen, 2016;Pittman et al., 1999;van de Kerk et al., 2019;Vickers et al., 2015;Wolfe et al., 2015;Young et al., 2010). Due to controversy regarding the species, collaring mountain lions in the region has been limited and past studies regarding their space use relied on VHF data (Harveson et al., 1999;Pittman et al., 1999;Young et al., 2010). ...
Article
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Local knowledge regarding the movements and space-use of large carnivores can inform species’ management, conservation planning, and strategies for reducing human-wildlife conflict. However, because issues regarding large carnivores can be controversial, obtaining such information may be difficult and therefore lacking in many places. Such is the case for mountain lions (Puma concolor) in West Texas, where most of the land is privately owned. Therefore, we sought to better understand movements and home ranges of mountain lions in the Trans-Pecos region of Texas. We GPS collared 24 mountain lions (15 females, 9 males) between 2011 and 2017. We inspected variograms to distinguish dispersers from residents then we estimated movement metrics and home ranges using continuous-time movement models and autocorrelated kernel density estimation. Using generalized linear mixed models, we tested for seasonal differences in all movement and home range metrics. Of three subadults that dispersed, one had data appropriate for speed estimation; she exhibited faster speeds while dispersing than while exhibiting philopatric movements. Compared to adult resident males, adult resident females moved with slower average speeds (females: 11.82 ± 0.43 km/day; males: 20.36 ± 2.86 km/day), exhibited similar directional persistence in movement (females: 0.95 ± 0.14 hours; males: 1.12 ± 0.21 hours), crossed their home ranges in similar times (females: 5.76 ± 0.98 days; males: 7.07 ± 1.34 days), and used smaller home ranges than males (females: 283.83 ± 37.50 km²: males: 1077.95 ± 219.30 km²). Adults moved with slowest speeds in the cold-dry season; there were no seasonal effects on other estimates. Each individual’s successive seasonal home ranges were highly overlapped (0.95 ± 0.01). The mountain lions did not expand, contract, or shift their home ranges seasonally, which may be due to ample prey-availability and limited seasonal-shifts in prey distribution. Mountain lions used areas larger than most local ranches and tended to move across the landscape quickly. Together, our results indicate that mountain lions do not typically stay on one property for long which may have implications for conservation planning and species management.
... The region also supports suitable habitat for recolonizing cougars (O'Neil et al. 2014, Smith et al. 2016. LaRue and Nielsen (2016) predicted that cougars would likely recolonize the western Great Lakes region by 2040. However, recent (2015-2018 observations of female cougars in Iowa, Missouri, Tennessee, and eastern North Dakota (LaRue et al. 2019) may indicate that cougars will recolonize portions of the western Great Lakes region sooner than predicted. ...
Article
Cougars (Puma concolor) have been recolonizing portions of their historic range over the last few decades and are currently a rare, transient species in the western Great Lakes Region. Anticipatory attitude surveys can identify potential management challenges, stakeholder groups, and conservation and educational priorities for recolonizing cougars. We implemented an anticipatory attitude survey to better understand attitudes towards transient cougars in northern Wisconsin, USA. We distributed the survey in fall and early winter of 2014–15 (n = 423) and 2015–16 (n = 594). We compared attitudes towards transient cougars to attitudes towards other established large carnivores and white‐tailed deer (Odocoileus virginianus). We also explored factors that may affect human attitudes towards cougars, such as hunter identity, livestock ownership, risk perception, and the loss of a domestic animal to wildlife. Despite the rare, transient status of cougars in Wisconsin, many respondents (mean of 29%) reported to have observed a cougar or sign of one in the wild in Wisconsin. Cougars had a mean favorability score similar to coyotes (Canis latrans) and wolves (Canis lupus) and less than bobcats (Lynx rufus), bear (Ursus americanus), and white‐tailed deer. Comparatively, respondents’ risk perception for all categories of risk (i.e., fear for personal safety, the safety of children, and the safety of domestic animals) was higher for cougars than for wolves. Hunter identity and loss of livestock to wildlife were determined by multinomial logistic regression as the most important parameters affecting respondents’ favorability towards cougars (ωi = 0.71). Using the 2015–2016 survey data, we found that as cougar risk perception increased, favorability towards cougars decreased similarly for both hunters and non‐hunters. Hunters were less likely to be favorable towards cougars, more likely to be neutral, and equally likely to be unfavorable towards cougars compared to nonhunters. We identify potential management challenges, stakeholder groups (e.g., hunters and those who have experienced loss of livestock to wildlife), and conservation and educational priorities for recolonizing cougars. © 2021 The Wildlife Society. As cougars recolonize portions of their former range, understanding what influences attitudes towards this large carnivore is a critical step in guiding their conservation. Using anticipatory attitude surveys, this paper provides insights into the factors that shape attitudes towards transient cougars prior to recolonization.
... Thus, large carnivore conservation needs to be well-targeted, while typically being based on scarce data. Spatially explicit metapopulation models provide promising opportunities and can be used to both assess the impact of past conservation interventions or to predict the effect of potential future interventions on large carnivore metapopulation viability (Robinson et al. 2015, LaRue and Nielsen 2016, Ovenden et al. 2019. Moreover, such models are highly informative in predicting which alternative conservation strategies, such as reducing persecution or restoring prey, have a larger impact on metapopulation viability (Chapron et al. 2008). ...
Article
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Large carnivores are currently disappearing from many world regions due to habitat loss, prey depletion, and persecution. Ensuring large carnivore persistence requires safeguarding and sometimes facilitating the expansion of their populations. Understanding which conservation strategies, such as reducing persecution or restoring prey, are most effective to help carnivores to reclaim their former ranges is therefore important. Here, we systematically explored such alternative strategies for the endangered Persian leopard (Panthera pardus saxicolor) in the Caucasus. We combined a rule‐based habitat suitability map and a spatially explicit leopard population model to identify potential leopard sub‐populations (i.e., breeding patches), and to test the effect of different levels of persecution reduction and prey restoration on leopard population viability across the entire Caucasus ecoregion and northern Iran (about 737,000 km2). We identified substantial areas of potentially suitable leopard habitat (~120,000 km2), most of which is currently unoccupied. Our model revealed that leopards could potentially recolonize these patches and increase to a population of > 1,000 individuals in 100 years, but only in scenarios of medium to high persecution reduction and prey restoration. Overall, reducing persecution had a more pronounced effect on leopard metapopulation viability than prey restoration: without conservation strategies to reduce persecution, leopards went extinct from the Caucasus in all scenarios tested. Our study highlights the importance of persecution reduction in small populations, which should hence be prioritized when resources for conservation are limited. We show how individual‐based, spatially explicit metapopulation models can help in quantifying the recolonization potential of large carnivores in unoccupied habitat, designing adequate conservation strategies to foster such recolonizations, and anticipating the long‐term prospects of carnivore populations under alternative scenarios. Our study also outlines how data scarcity, which is typical for threatened range‐expanding species, can be overcome with a rule‐based habitat map. For Persian leopards, our projections clearly suggest that there is a large potential for a viable metapopulation in the Caucasus, but only if major conservation actions are taken towards reducing persecution and restoring prey.
... Although mountain lion hunting occurs in much of western North America (Peebles et al. 2013), laws prohibit hunting in areas with diminished populations (Cardoza and Langlois 2002). Moreover, key prey species have recovered throughout much of the continent (Nowell and Jackson 1996) and suitable dispersal corridors Nielsen 2008, 2011) enable reestablishment of mountain lions in portions of midwestern North America (LaRue et al. 2012(LaRue et al. , 2019LaRue and Nielsen 2016), such as in the Dakotas and Nebraska (Fecske 2003, Wilson et al. 2010. Consequently, range expansion could lead to novel human-mountain lion interactions in areas where people and mountain lions have not lived sympatrically for over a century (Smith et al. 2016). ...
Article
Social media is often used to facilitate the exchange of knowledge and engage the public, which can change the public's attitudes and behaviors and may be used to benefit conservation. Moreover, widespread social media use provides an alternative data sourcing platform to inexpensively access countless potential respondents. However, social media data have rarely been used in conservation regardless of the potential benefit to conservation science and practice. We administered a questionnaire via the #CougarOrNot Twitter game, a mountain lion-focused social media campaign, during 26 December 2016 through 16 February 2018, to access a large respondent sample (n = 1,481) and assess a subset of social media users' attitudes toward mountain lions in North America. We then used cumulative link models in an information theoretic approach to gauge the association between respondent level of engagement with #CougarOrNot and other sociodemographic predictors with user attitudes toward mountain lions. Respondent attitudes toward mountain lions were largely positive (83%), with frequent participants in #CougarOrNot, females, pet owners, nonconsumptive recreators, and households with fewer children exhibiting positive attitudes. #CougarOrNot participation had a stronger correlation with respondent attitudes toward mountain lions than commonly used predictors employed in prior studies (i.e. age, education level, livestock ownership). #CougarOrNot may promote positive attitudes toward mountain lions, though additional research is needed to determine the direct effects of playing the game. Stakeholders interested in mountain lion conservation or other conservation topics could be identified via social media networks attached to specific outreach campaigns and possibly mobilized to support conservation actions that aid mountain lions and conservation in general.
... first documented modern appearance of the mountain lion in Nebraska was in 1991 when one was shot near Fort Robinson State Park in Sioux County (Genoways and Freeman 1996). Subsequent sightings have been made statewide (LaRue et al. 2012, LaRue and Nielsen 2016, Wilson 2016 as the excellent website maintained by the Nebraska Game and Parks Commission documents (<https://maps.outdoornebraska.gov/ puma/>). ...
Article
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Information on the distribution and current status of 25 species or subspecies of mammals occurring in Nebraska are presented. The species covered include one shrew, an armadillo, eight bats, 10 rodents (including two subspecies of one species), three carnivores, and one artiodactyl. Distributional information reported includes the first state record for one species (Sorex nanus) and new county records for 18 species. In Nebraska, we know that mammals are shifting their geographic ranges with some extending populations into the state, whereas others are expanding their geographic ranges within the state. The current status of six additional mammalian taxa in Nebraska is documented, providing knowledge important to the conservation of these species during this time of shifting environmental conditions. The concern is that some mammals have suffered population declines since the 1940s and may have a contracting geographic range as well in the state. These species would be endangered and could be lost to the mammalian fauna of the state. Extensive data on reproduction in these mammals are presented and the taxonomy of species is updated from the 1964 Mammals of Nebraska by Jones.
... The loss of this apex predator from the forests of the Northeastern U.S.A. likely had downstream consequences, including deer overpopulation 8,47,48 . Mountain lions may eventually make a return to the Northeast via the slow expansion of Western and Midwestern populations 13,49 . Our work will ultimately characterize the genetic diversity that was permanently lost (if any) from this region with the original extinction, and we hope that our associated outreach activities in the Pennsylvania region will help to raise awareness about the importance of broader ecosystem diversity in advance of the likely return of mountain lions to the Northeast. ...
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Mountain lions ( Puma concolor ) were once endemic across the United States. The Northeastern population of mountain lions has been largely nonexistent since the early 1800s and was officially declared extinct in 2011. This regionally extinct mountain lion is Pennsylvania State University’s official mascot, where it is referred to as the ‘Nittany Lion’. Our goal in this study was to use recent methodological advances in ancient DNA and massively parallel sequencing to reconstruct complete mitochondrial DNA (mtDNA) genomes of multiple Nittany Lions by sampling from preserved skins. This effort is part of a broader Nittany Lion Genome project intended to involve undergraduates in ancient DNA and bioinformatics research and to engage the broader Penn State community in discussions about conservation biology and extinction. Complete mtDNA genome sequences were obtained from five individuals. When compared to previously published sequences, Nittany Lions are not more similar to each other than to individuals from the Western U.S. and Florida. Supporting previous findings, North American mountain lions overall were more closely related to each other than to those from South America and had lower genetic diversity. This result emphasizes the importance of continued conservation in the Western U.S. and Florida to prevent further regional extinctions.
... Female cougars tend to exhibit higher natal philopatry and lower dispersal rates than their male counterparts (Pierce and Bleich 2003), though a >1,000-km dispersal was documented in a global positioning systemtracked female in Utah, USA (Stoner et al. 2008). We would anticipate cougar range expansion to be dependent almost exclusively on female movement dynamics (LaRue et al. 2016) in that closer and more suitable areas are more likely to be occupied than smaller or more distant areas (MacAthur and Wilson 1967). Indeed, the North Dakota Badlands and Nebraska populations were likely recolonized by animals from Wyoming, Montana, and the Black Hills Much of what is known about cougar life history comes from populations in western North America (Hornocker 1970, Hemker et al. 1984, Lindzey et al. 1994, Lambert et al. 2006, Ruth et al. 2011). ...
... However, to our knowledge, few large-scale mid-sized predator distribution models have included urban landscapes within their modeling area (Lewis et al. 2015). As urbanization increases, species distribution models can aid managers in predicting areas at high risk for nuisance wildlife complaints and can serve as the basis for landscape connectivity or population viability analyses (Honda 2009;Thatcher et al. 2009;Rabinowitz and Zeller 2010;LaRue and Nielsen 2016). Additionally, identification of important urban habitat features influential to species-specific occupancy and distribution will aid managers in implementing management actions for medium-sized predator species in urban areas. ...
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Large-scale distribution models are effective predictors of habitat suitability and connectivity across broad landscapes and are useful management tools, though few large-scale species distribution models exist for medium-sized predators in urban landscapes. We modeled the potential distribution of 4 medium-sized predators in a 17,361-km2 portion of the Chicago Metropolitan Area. We applied a maximum entropy algorithm model (MaxEnt) using presence-only data collected via remote cameras from 54 Lake County, Illinois, forest preserves during August–October 2008–2012. Environmental data layers used to model distributions were distances to forest, grassland, barren land, crops, wetlands, developed open space, developed low intensity, developed high intensity, water, primary roads, secondary roads, and tertiary roads. Coyotes (Canis latrans) had the greatest area of potential distribution followed by opossums (Didelphis virginiana), striped skunks (Mephitis mephitis), and raccoons (Procyon lotor). Models for all species had high AUC values (0.90–0.94) indicating strong predictive performance. More than 50% of the study area was predicted to be within the distributional limit for each focal species. Distance to forest was the most important contributory predictor for all species modeled (82% - 96%) and higher probability of presence for all 4 species was indicated closer to forest and further from tertiary roads. However, coyotes and raccoons were predicted to prefer habitat closer to highly-developed areas. Our research indicates medium-sized predators are highly synanthropic and able to persist within the Chicago Metropolitan Area given adequate availability of non-urban land cover, particularly forest, and ample green space linking forest patches within highly-developed areas.
... Population viability analysis (PVA) can be used to estimate the viability, the MVP, and extinction risk of populations (Boyce, 1992). PVAs are becoming increasingly sophisticated, and different softwares can encompass many types of information alongside life history data, including genetic data (VORTEX), spatial information (RAMAS), species interactions, and disease spread (Akcakaya, 2000;Andersen, Sunde, Loeschcke, & Pertoldi, 2015;Bradshaw et al., 2012;Larue & Nielsen, 2016;Olsen et al., 2014;Prowse et al., 2013). ...
Article
The agricultural scene has changed over the past decades, resulting in a declining population trend in many species. It is therefore important to determine the factors that the individual species depend on in order to understand their decline. The landscape changes have also resulted in habitat fragmentation, turning once continuous populations into metapopulations. It is thus increasingly important to estimate both the number of individuals it takes to create a genetically viable population and the population trend. Here, population viability analysis and habitat suitability modeling were used to estimate population viability and future prospects across Europe of the Little Owl Athene noctua, a widespread species associated with agricultural landscapes. The results show a high risk of population declines over the coming 100 years, especially toward the north of Europe, whereas populations toward the southeastern part of Europe have a greater probability of persistence. In order to be considered genetically viable, individual populations must count 1,000-30,000 individuals. As Little Owl populations of several countries count <30,000, and many isolated populations in northern Europe count <1,000 individuals, management actions resulting in exchange of individuals between populations or even countries are probably necessary to prevent losing <1% genetic diversity over a 100-year period. At a continental scale, a habitat suitability analysis suggested Little Owl to be affected positively by increasing temperatures and urban areas, whereas an increased tree cover, an increasing annual rainfall , grassland, and sparsely vegetated areas affect the presence of the owl negatively. However, the low predictive power of the habitat suitability model suggests that habitat suitability might be better explained at a smaller scale. K E Y W O R D S conservation, habitat suitability, management, minimum viable population size, population viability analysis, RAMAS/GIS, VORTEX
... Population viability analysis (PVA) can be used to estimate the viability, the MVP, and extinction risk of populations (Boyce, 1992). PVAs are becoming increasingly sophisticated, and different softwares can encompass many types of information alongside life history data, including genetic data (VORTEX), spatial information (RAMAS), species interactions, and disease spread (Akcakaya, 2000;Andersen, Sunde, Loeschcke, & Pertoldi, 2015;Bradshaw et al., 2012;Larue & Nielsen, 2016;Olsen et al., 2014;Prowse et al., 2013). ...
Article
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The agricultural scene has changed over the past decades, resulting in a declining population trend in many species. It is therefore important to determine the factors that the individual species depend on in order to understand their decline. The landscape changes have also resulted in habitat fragmentation, turning once continuous populations into metapopulations. It is thus increasingly important to estimate both the number of individuals it takes to create a genetically viable population and the population trend. Here, population viability analysis and habitat suitability modeling were used to estimate population viability and future prospects across Europe of the Little Owl Athene noctua, a widespread species associated with agricultural landscapes. The results show a high risk of population declines over the coming 100 years, especially toward the north of Europe, whereas populations toward the southeastern part of Europe have a greater probability of persistence. In order to be considered genetically viable, individual populations must count 1,000–30,000 individuals. As Little Owl populations of several countries count <30,000, and many isolated populations in northern Europe count <1,000 individuals, management actions resulting in exchange of individuals between populations or even countries are probably necessary to prevent losing <1% genetic diversity over a 100-year period. At a continental scale, a habitat suitability analysis suggested Little Owl to be affected positively by increasing temperatures and urban areas, whereas an increased tree cover, an increasing annual rainfall, grassland, and sparsely vegetated areas affect the presence of the owl negatively. However, the low predictive power of the habitat suitability model suggests that habitat suitability might be better explained at a smaller scale.
... These long-range dispersal movements are of particular interest in North Dakota, where mountain lions have naturally recolonized the state, but remain geographically semiisolated from other mountain lion populations by large expanses of agricultural and grassland habitat (NDGFD 2006). Furthermore, because the North Dakota population primary range occurs on the eastern edge of the current mountain lion distribution, the population represents a possible source of dispersing individuals contributing to continued species range expansion eastward , LaRue and Nielsen 2015, Juarez et al. 2016). ...
... Our results indicate that assumption can sometimes fail. With increasingly variable and extreme weather patterns in temperate regions due to climate change (Seneviratne et al. 2012), increasing predator populations (Bluett 2013), and novel predator communities due to recolonization and expansion of predator ranges into novel habitats (Gompper 2002;LaRue and Nielsen 2016;Smith et al. 2016), wildlife managers should be aware that historical patterns (e.g., similar overwinter survival in fawns and adult females) may no longer hold true. These interacting environmental trends indicate that periodic research to update estimates of agespecific survival may be needed to responsively manage deer populations. ...
Article
Mortality rates of white-tailed deer (Odocoileus virginianus (Zimmermann, 1780)) fawns have been quantified throughout North America. Few studies, however, have assessed cause-specific mortality of fawns after the first 3 months of life or during a severe weather event. During 2010–2014, we captured and radio-tracked 93 fawns in southern and central Illinois and recorded 18 mortality events. In order of importance, survival rates were affected by days since capture, year of drought, age at capture, week after capture (1 or 0 indicator), and region. Estimated overwinter (fall through spring) survival rate (±SE) of fawns in both regions during 2010–2014 was 0.83 ± 0.04. However, estimated overwinter survival rates were depressed during 2012–2013, following the severe drought of 2012 (0.63 ± 0.11 or 0.66 ± 0.11 depending on the model). Main causes of mortality were capture-related and predation, though some dead deer also showed signs of hemorrhagic disease. We suspect that the extreme drought of 2012 created favorable conditions for fall–spring mortality of fawns, due to elevated disease transmission and lower forage quality and quantity for deer. In addition, drought may have contributed to predation by reducing abundance of alternative prey. Our results suggest that severe weather conditions during summer can substantially impact overwinter fawn survival.
... survival/mortality, recruitment, longevity, sex ratio) obtained from long-term monitoring of predators are used widely for assessing population viability and predicting population change under alternative scenarios (e.g. habitat fragmentation, [19,20]; disease, [21], climate change, [22]; and poaching/harvesting/ trophy hunting, [23,24,25]. These parameters are relatively well-known for a number of large felids (e.g. ...
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In this study, we estimate life history parameters and abundance for a protected jaguar population using camera-trap data from a 14-year monitoring program (2002–2015) in Belize, Central America. We investigated the dynamics of this jaguar population using 3,075 detection events of 105 individual adult jaguars. Using robust design open population models, we estimated apparent survival and temporary emigration and investigated individual heterogeneity in detection rates across years. Survival probability was high and constant among the years for both sexes (φ = 0.78), and the maximum (conservative) age recorded was 14 years. Temporary emigration rate for the population was random, but constant through time at 0.20 per year. Detection probability varied between sexes, and among years and individuals. Heterogeneity in detection took the form of a dichotomy for males: those with consistently high detection rates, and those with low, sporadic detection rates, suggesting a relatively stable population of ‘residents’ consistently present and a fluctuating layer of ‘transients’. Female detection was always low and sporadic. On average, twice as many males than females were detected per survey, and individual detection rates were significantly higher for males. We attribute sex-based differences in detection to biases resulting from social variation in trail-walking behaviour. The number of individual females detected increased when the survey period was extended from 3 months to a full year. Due to the low detection rates of females and the variable ‘transient’ male subpopulation, annual abundance estimates based on 3-month surveys had low precision. To estimate survival and monitor population changes in elusive, wide-ranging, low-density species, we recommend repeated surveys over multiple years; and suggest that continuous monitoring over multiple years yields even further insight into population dynamics of elusive predator populations.
... Small prey (e.g. non-ungulates) formed a large portion of dispersing puma diets of both sexes (Fig. 1).Knopff et al. (2010)also noted a higher proportion of small prey in young pumas, and this stage-dependent foraging pattern might be important in describing the ecology of the species, as well as in modeling potential dispersal habitat that could see their expansion east in North America (e.g.LaRue & Nielsen, 2016) or across hostile matrices where their primary prey has been reduced through anthropogenic effects (Stoner et al., 2013). ...
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Predators likely assess their risk of injury with regards to hunting different prey types, while deciding whether to initiate an attack or to avoid dangerous prey. Risk-taking is age-, stage- and state-dependent, and foraging theory predicts that juvenile predators, individuals of lower social rank, and less-experienced predators that are hungry are among those most likely to engage dangerous prey. In carnivores that optimally hunt prey larger than themselves, however, juveniles may also select for smaller, suboptimal prey as they refine their hunting skills. Thus, in the case of predators that exhibit age-specific prey selection, there may be multiple factors influencing decision-making about prey selection. We compared puma (Puma concolor) predation on dangerous adult ungulates and safer juvenile ungulates, as well as two similar-sized rodent species: the more dangerous North American porcupine (Erethizon dorsatum) and the more vulnerable American beaver (Castor canadensis). We found that dispersing, subadult pumas with limited hunting experience and without territories avoided dangerous but optimally sized adult ungulates, and instead hunted dangerous, suboptimal porcupines more than resident, territorial adults. In contrast, there was no difference in puma predation on beavers between dispersers and residents. Small prey (e.g. not ungulates) formed a large portion of dispersing puma diets of both sexes, and this stage-dependent foraging pattern might be important in describing the ecology of the species, as well as in modeling potential dispersal habitat that could see puma expansion east in North America across hostile matrices where their primary prey has been reduced through anthropogenic effects.
... Although wildlife management practices, conservation policies, and habitat improvement implemented during the last half of the twentieth century facilitated some large carnivore recolonizations in parts of Europe and North America (e.g., gray wolf [Canis lupus] and brown bear [Ursus arctos]), this process occurs relatively slowly for these animals (Pyare et al. 2004, Fabbri et al. 2014, Votsi et al. 2016. Similar recolonizations will be difficult where human population density is high and habitat is severely degraded, but current predictions indicate regional large carnivore recolonizations may increase in frequency and geographical distribution in portions of the United States and Europe during coming decades (Chapron et al. 2014, LaRue and Nielsen 2016, Smith et al. 2016. ...
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Many large carnivores are recolonizing range as a result of improved management and conservation policy, habitat restoration, and reintroduction programs. American black bears (Ursus americanus) are projected to recolonize portions of the United States, but few studies have characterized or provided practical methods for monitoring this process. We used noninvasive hair sampling at 4 proximal study areas along the Kentucky–Virginia, USA, border during 2012–2013 to estimate demographics and population genetics, and investigate recolonization patterns of an American black bear population that was founded by 55 bears reintroduced to a fragmented mountainous landscape during the 1990s and subjected to harvest 6 years post-reintroduction. Using spatially explicit capture–recapture (SECR) models, we estimated a density of 0.26 bear/km2, or minimum abundance of 482 bears, distributed among 2 primary core areas previously identified by occupancy analysis: a southern and northern core area. The southern core area was established by a founder adult female that exhibited post-release dispersal, but moderate asymmetrical gene flow (Nm = 6 bears) from the northern core area mitigated deleterious genetic consequences typical of such founder events. Effective number of breeders (NB = 62 bears) was similar to the number of founders, suggesting that genetically, the population remains mostly the product of reintroduction. Despite limited connectivity with other populations in the region, genetic diversity (HE = 0.78) was retained because of rapid population growth during the 16 years post-reintroduction (λ = 1.14/year). This bear population exhibited demographic characteristics indicative of continued recolonization, including a significantly female-biased sex ratio (0.53M:1.00F) and female density decreasing with increasing distance from the reintroduction release areas in the northern core. Few bear detections at 2 peripheral study areas and results from SECR model detection function transformation suggested recolonization may continue to the southwest and northeast along prominent linear mountain ridges. Although the population has grown and is genetically stable, because of relatively low population density and recolonization direction, we suggest monitoring demographic vital rates to evaluate harvest sustainability and population viability. Our study demonstrates the utility of noninvasive genetic sampling in conjunction with SECR models to characterize and monitor recolonizing bear populations, which may also be useful for management of expanding populations of other large carnivores.
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Pumas ( Puma concolor ) were eliminated from most of the eastern USA a century ago. In the past couple of decades, their recovery in the West has increased puma dispersal into the Midwest, with some individuals even traveling to the East Coast. We combined published expert opinion data and a habitat suitability index in an analysis that identified 17 areas in the Upper Midwest, Ozarks, Appalachia, and New England that could potentially host puma populations in the future. Thirteen of these were larger than 10,000 km ² and so likely to ensure a puma population’s long-term genetic health. Further, we quantified patch size, human density, livestock density, percent public land, and a sociocultural index reflecting wildlife values for comparing patches, as well as present a summary of current legislation relevant to puma management in the East. Our work may be useful in identifying suitable areas to restore pumas based not only on the quality of their biophysical habitat, but also on social values conducive to puma-human coexistence.
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Se analiza la distribución y en algunos casos abundancia de los grandes depredadores a nivel mundial, así como 3 de estos grandes depredadores en México y algunos temas como etnobiología, y el papel de los zoos en la conservación
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Understanding landscape patterns in mortality risk is crucial for promoting recovery of threatened and endangered species. Humans affect mortality risk in large carnivores such as wolves (Canis lupus), but spatiotemporally varying density dependence can significantly influence the landscape of survival. This potentially occurs when density varies spatially and risk is unevenly distributed. We quantified spatiotemporal sources of variation in survival rates of gray wolves (C. lupus) during a 21-year period of population recovery in the Upper Peninsula of Michigan, USA. We focused on mapping risk across time using Cox Proportional Hazards (CPH) models with time-dependent covariates, thus exploring a shifting mosaic of survival. Extended CPH models and time-dependent covariates revealed influences of seasonality, density dependence and experience, as well as individual-level factors and landscape predictors of risk. We used results to predict the shifting landscape of risk at the beginning, middle, and end of the wolf recovery time series. Survival rates varied spatially and declined over time. Long-term change was density-dependent, with landscape predictors such as agricultural land cover and edge densities contributing negatively to survival. Survival also varied seasonally and depended on individual experience, sex, and resident versus transient status. The shifting landscape of survival suggested that increasing density contributed to greater potential for human conflict and wolf mortality risk. Long-term spatial variation in key population vital rates is largely unquantified in many threatened, endangered, and recovering species. Variation in risk may indicate potential for source-sink population dynamics, especially where individuals preemptively occupy suitable territories, which forces new individuals into riskier habitat types as density increases. We encourage managers to explore relationships between adult survival and localized changes in population density. Density-dependent risk maps can identify increasing conflict areas or potential habitat sinks which may persist due to high recruitment in adjacent habitats.
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The decline of top carnivores has released large herbivore populations around the world, incurring socioeconomic costs such as increased animal-vehicle collisions. Attempts to control overabundant deer in the Eastern U.S. have largely failed, and deer-vehicle collisions (DVCs) continue to rise at alarming rates. We present the first valuation of an ecosystem service provided by large carnivore recolonization, using DVC reduction by cougars as a case study. Our coupled deer population models and socioeconomic valuations revealed that cougars could reduce deer densities and DVCs by 22% in the Eastern US, preventing 21,400 human injuries, 155 fatalities, and $2.13 billion in avoided costs within 30 years of establishment. Recently established cougars in South Dakota prevent$1.1 million in collision costs annually. Large carnivore restoration could provide valuable ecosystem services through such socio-ecological cascades, and these benefits could offset the societal costs of coexistence. This article is protected by copyright. All rights reserved
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We report the long-distance dispersal of a subadult male cougar (Puma concolor) from South Dakota to Milford, Connecticut, where it was struck and killed by a vehicle. Genetic samples suggest this animal originated from the Black Hills of South Dakota while isotope analysis and physical inspection revealed no evidence that the animal had been held in captivity. We detected this dispersing individual at 5 locations along its route (Minnesota, 3 times in Wisconsin and New York) with DNA from fecal or hair samples, and with multiple photographs from citizen-run camera traps (3 in Wisconsin and 1 in Michigan). The > 2,450 km straight-line distance (Black Hills of South Dakota to Connecticut) traveled by the cougar is the longest dispersal documented for the species. We propose a likely route of > 2,700 km over 2 years based on topography and our confirmed records. We suggest that this excessive movement was motivated by the absence of female cougars along the route. The documentation of such a rare biological event not only shows the great dispersal potential for male cougars but also highlights our ability to detect these movements with verifiable voucher DNA and photographic records. Evidence collected for this one animal, and complete absence of verifiable data from most anecdotal reports of cougars in the east, further confirms the lack of a breeding population in the region.
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There is little information on the spatiotemporal pattern of dispersal of juvenile cougars (Felis concolor) and no data on disperser use of habitat corridors. I investigated dispersal of radio-tagged juvenile cougars (8 M, 1 F) in a California landscape containing 3 corridors (1.5, 4.0, and 6.0 km long) and several habitat peninsulas created by urban growth. Dispersal was usually initiated by the mother abandoning the cub near an edge of her home range. The cub stayed within 300 m of that site for 13-19 days and then dispersed in the direction opposite that taken by the mother. Mean age at dispersal was 18 months (range 13-21 months). Each disperser traveled from its natal range to the farthest part of the urban-wildland edge. Dispersing males occupied a series of small (<30% the area used by ad M in the same time span), temporary (10-298 days) home ranges, usually near the urban-wildland interface, and often with its longest border along that edge. Each of the 3 corridors was used by 1-3 dispersers, 5 of the 9 dispersers found and successfully used corridors, and 2 dispersers entered but failed to traverse corridors. Dispersing cougars will use corridors that are located along natural travel routes, have ample woody cover, include an underpass integrated with roadside fencing at high-speed road crossings, lack artificial outdoor lighting, and have <1 dwelling unit/16 ha.
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Carnivore guilds play a vital role in ecological communities by cascading trophic effects, energy and nutrient transfer, and stabilizing or destabilizing food webs. Consequently, the structure of carnivore guilds can be critical to ecosystem patterns. Body size is a crucial influence on intraguild interactions, because it affects access to prey resources, effectiveness in scramble competition, and vulnerability to intraguild predation. Coyotes (Canis latrans), bobcats (Lynx rufus), gray foxes (Urocyon cinereoargenteus), raccoons (Procyon lotor), red foxes (Vulpes vulpes), and striped skunks (Mephitis mephitis) occur sympatrically throughout much of North America and overlap in resource use, indicating potential for interspecific interactions. Although much is known about the autecology of the individual species separately, little is known about factors that facilitate coexistence and how interactions within this guild influence distribution, habitat use, and temporal activity of the smaller carnivores. To assess how habitat autecology and interspecific interactions affect the structure of this widespread carnivore guild, we conducted a large-scale, non-invasive carnivore survey using an occupancy modeling framework. We deployed remote cameras during 3-week surveys to detect carnivores at 1,118 camera locations in 357 2.6-km 2 sections (3–4 cameras/section composing a cluster) in the 16 southernmost counties of Illinois (16,058 km 2
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Understanding the impact of habitat fragmentation, roads, and other anthropogenic influences on cougars (Puma concolor) requires quantitative assessment of habitat selection at multiple scales. We calculated annual and multiyear home ranges using a fixed-kernel (FK) estimator of home range for 13 adult female and 2 adult male radiotagged cougars that were monitored October 1986 through December 1992 in the Santa Ana Mountain Range of southern California, USA. Using compositional analysis, we assessed diurnal use of vegetation types and areas near roads at 2 orders of selection (second- and third-order; Johnson 1980). Mean annual and multiyear 85% FK home ranges for males were larger than those reported by previous studies in California. Mean wet-season 85% FK home ranges were significantly larger than those of the dry season. At both scales of selection and across seasons, cougars preferred riparian habitats and avoided human-dominated habitats. Grasslands were the most avoided natural vegetation type at both scales of selection. Although cougar home ranges tended to be located away from high- and low-speed 2-lane paved roads (second-order avoidance), cougars did not avoid roads within their home range, especially when roads were in preferred riparian areas. Protection of habitat mosaics that include unroaded riparian areas is critical to the conservation of this cougar population.
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Background Recent findings indicate that cougars (Puma concolor) are expanding their range into the midwestern United States. Confirmed reports of cougar in Michigan, Minnesota, and Wisconsin have increased dramatically in frequency during the last five years, leading to speculation that cougars may re-establish in the Upper Great Lakes (UGL) region, USA. Recent work showed favorable cougar habitat in northeastern Minnesota, suggesting that the northern forested regions of Michigan and Wisconsin may have similar potential. Recolonization of cougars in the UGL states would have important ecological, social, and political impacts that will require effective management. Methodology/Principal Findings Using Geographic Information Systems (GIS), we extended a cougar habitat model to Michigan and Wisconsin and incorporated primary prey densities to estimate the capacity of the region to support cougars. Results suggest that approximately 39% (>58,000 km2) of the study area could support cougars, and that there is potential for a population of approximately 500 or more animals. An exploratory validation of this habitat model revealed strong association with 58 verified cougar locations occurring in the study area between 2008 and 2013. Conclusions/Significance Spatially explicit information derived from this study could potentially lead to estimation of a viable population, delineation of possible cougar-human conflict areas, and the targeting of site locations for current monitoring. Understanding predator-prey interactions, interspecific competition, and human-wildlife relationships is becoming increasingly critical as top carnivores continue to recolonize the UGL region.
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We monitored size and composition of a southern Utah cougar (Felis concolor) population during 1979-87 to document the dynamics of this unhunted population and to test the hypothesis that cougars would regulate their density at a level below that set by prey abundance alone (Seidensticker et al. 1973). We captured cougars when detected during ongoing searches for sign in the study area. Resident adult cougar density remained relatively constant (0.37/100 km2) for the first 7 years but increased slightly in the last 2 years. Mule deer (Odocoileus hemionus), the cougar's primary prey, increased over the 9 years, but magnitude of this increase was unknown. Results supported the hypothesis that cougar density is set by environmental features other than prey abundance alone. Adult resident females bred as young as 17 months and produced litters that averaged 2.4 kittens at an interval of 24.3 months.
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Twenty-two cougars (Felis concolor) were monitored by radiotelemetry between January 1979 and July 1981 in southern Utah. The population, comprised of resident, transient, and juvenile cougars, remained relatively constant during the study. Densities (0.3-0.5 cougars/100 km2) were considerably lower and home-area size of four resident females (685 km2, SE = 257, range = 396-1,454) and a single resident male (826 km2) were larger than reported for other areas. Home areas of resident females overlapped, but with the exception of family groups, close spatial associations were rare. Dispersal of male cubs appeared independent of resident adult density. Density of resident cougars was apparently regulated by a social pattern based on land tenure, but limited by the abundance of mule deer (Odocoileus hemionus), their principal prey.
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We monitored survival of resident mountain lions (Felis concolor) during a radio-telemetry study between 1980 and 1986 in southern Utah. Yearly survival of resident adults ranged from 52 to 100% (x̄ = 74%). Causes of death included intraspecific killing, injury related to prey capture, trapping, and starvation. Deaths of dispersing offspring were human-related.
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The Amur tiger, a flagship species of the boreal forest ecosystem in Russian Far East and northeastern China, has declined dramatically in population and geographic distribution due to human caused habitat fragmentation and poaching over the past century. The fate of this largest feline species will also be influenced by the worsening impacts of climate change. In this paper we assess the possible effects of climate change (three scenarios from the 2007 IPCC Report) on the Amur tiger by integrating species distribution modeling (SDM) and population viability analysis (PVA). We projected the potential and realized suitable habitat distributions to examine the impacts from anthropogenic factors, and evaluated the changes of suitable habitat and extinction risk for 100 years under climate change. The realized suitable habitat was projected to be more severely fragmented than the potential suitable habitat because of human-related factors. The potential suitable habitat would expand northward under all climate change scenarios considered. However, the tiger population would suffer the largest decline and highest extinction risk in the next 100 years under the worst climate change scenario (A1B) even though the size of potential habitat would be greatest. Under climate change, the tiger population could persist for the next century only if the size and quality of current habitat patches would remain intact. In addition, our study demonstrated that using SDM alone could grossly overestimate the geographic distribution of the Amur tiger, and that coupling SDM and PVA could provide important insights into conservation planning to mitigate the effects of climate change.
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We studied a cougar (Puma concolor) population from May 1985 to November 1987 on a 540-km2 study area in southeastern British Columbia (BC) to collect detailed ecological data to assist in refining regional species management. We used capture-recapture and radiotelemetry techniques, observed 34 different cougars, and estimated winter population densities of 3.5 cougars/100 km2 (1985-86) and 3.7 cougars/100 km2 (1986-87). Sex ratios of 20 kittens and 12 adults did not differ significantly from equality (P > 0.05). Kittens made up 55-58% of the population each winter. Mean litter size was 3.1 ± 1.2 [SD] kittens/litter (n = 7). Birth intervals were 15-23 months. Eight juveniles dispersed 12-163 km from the study area. Mortality was 5.3% of the population the first winter, and 15% the second winter. Four of 7 mortalities were human-related (vehicular) despite the area being closed to hunting. Home range sizes of 4 resident females were 31 ± 10 km2 [SD] in winter, 35 km2 ± 14 in summer, and 55 km2 ± 25 yearly with low or no home range overlap. Two resident males yearly home range averaged 151 km2. No male home range overlap was documented.
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Breeding populations of wolves (Canis lupus) were absent from the western United States for about 50 years following their extirpation by humans in the 1930s. Here we describe the recolonization by wolves of northwestern Montana and southeastern British Columbia, from the initial production of a litter by a pair of wolves in 1982 through the mid-1990s when 3-4 packs produced litters. Sex ratio of captured wolves favored females (38/54 = 70%; χ 2 = 8.96, 1 df, P < 0.005). Litter size in early summer (x̄ = 5.3, SE = 0.4, n = 26) and in December (x̄ = 4.5, SE = 0.5, n = 26) were relatively high compared to similar counts in established populations elsewhere. Pack size in May was unrelated to litter size in June (rs = -0.13, 23 df, P = 0.25) or the following December (rs = -0.12, 23 df, P = 0.28). Annual adult survival rate (0.80) was relatively high in this semi-protected population and was higher among residents (0.84) than among wolves that dispersed (0.66) from the study area (Z = 2.24, P = 0.025). Although dispersal was common among radiocollared wolves (19/43 = 44%), population growth within the study area averaged 20% per year from 1982 to 1995. Low human-caused mortality rates and maintenance of connectivity for wolves between this small population in the United States and larger populations in Canada will enhance the probability of persistence and expansion of this population.
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Population characteristics were estimated from June 1981 to July 1983 for a hunted mountain lion (Felis concolor) population occupying a $741-\text{km}^{2}$ study area in the Big Horn Mountains, Wyoming. Based on the capture-recapture of 46 lions and radio-telemetry, snow-tracking, and harvest data, winter population densities were estimated at 29 km2/lion (1981-82) and 22 km2/lion (1982-83). Sex ratios of 28 kittens and 22 adults did not differ (P > 0.05) from equality. Kittens, born primarily in autumn, comprised about 50% of the population each winter, and 11 postnatal litters averaged 2.7 kittens. Some juveniles dispersed at about 12-15 months of age; 5 were recovered 9-274 km from their natal areas. Two resident females bred at 13- and 19-month intervals. The age structure of both sexes was young, the oldest adult being about 7 years old. Observed mortality the 1st year was 27% of the total population and 0% the 2nd year; immigration apparently compensated for mortalities. Home areas of 4 resident females averaged 67 km2 and overlapped almost completely. Those of the 2 resident males overlapped slightly and averaged 320 km2. Male home areas overlapped several female home areas.
Article
We translocated wild cougars (Puma concolor) with known social and behavioral histories to evaluate the feasibility of translocation as a management tool to reestablish populations, relieve the threat of inbreeding in isolated populations, and manage problem individuals. Eight female and 5 male cougars 16-108 months old were translocated a mean distance of 477 km from the San Andres Mountains (SAM) study area in southcentral New Mexico to 9 release sites in northeastern New Mexico (NENM) from December 1990 through June 1991 and then radiomonitored through January 1993. Another male cougar was translocated 338 km from the SAM to northwestern New Mexico in April 1989 and monitored until May 1990. Initial movement directions away from release sites were uniformly distributed about a 360° circle. Eight (4 F, 4 M) of 14 cougars had movement end points >80 km from their release sites, and end point directions were generally south, suggesting they homed toward the source population; 2 male cougars returned to their original home ranges. Dispersing cougars from a SAM reference population showed no southerly tendencies. Distances moved from release sites to end points ranged from 3 to 494 km; 4 cougars (3 F, 1 M) established home ranges ≤84 km from release sites. Mean pretranslocation home ranges were generally smaller than mean posttranslocation use areas. Nine of 14 translocated cougars died during the study. Annual survival rates of translocated cougars did not differ by sex and were lower in 1992 than survival rates of cougars from the reference area. We concluded that translocation was most successful with cougars 12-27 months old.
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Dispersal plays a vital role in cougar (Puma concolor) population ecology, creating genetic viability and maintaining gene flow between populations. The naturally recolonized cougar population in the Black Hills is at the edge of the species' range in North America and completely surrounded by the grasslands of the Northern Great Plains. Our objective was to document dispersal movements and possible range expansion of subadult cougars captured within the Black Hills ecosystem of southwestern South Dakota and eastern Wyoming. Twenty-four (n=14 males, n=10 females) subadult cougars were captured in the Black Hills. Independence of cougars from females averaged 13.5 months (range = 10-16 months) from parturition; dispersal occurred 1-3 months post independence. Males dispersed (mean=274.7 km SE 88.3) farther than females (mean = 48.0 km SE 10.9), with females exhibiting 40% philopatry. We documented several (n = 6) long-distance dispersal movements (>250 km) of male cougars and hypothesize that males making long-distance movements were in search of available mates. The longdistance cougar dispersal movements documented by our study indicate that range expansion and habitat recolonization are occurring and further suggest proactive efforts to increase public knowledge of cougar ecology in areas where cougars are recolonizing previously occupied range.
Article
Reliable estimates of survival and mortality rates for mountain lions (Puma concolor) have been difficult to obtain because of their low densities and secretive behavior We estimated annual survival and cause-specific mortality rates for a heavily exploited mountain lion population in southeastern Arizona from February 1991 to April 1994. We monitored 24 adult radio-collared mountain lions weekly. We used MICROMORT to determine daily, annual, and overall survival rates. Radio-collared mountain lions had low annual survival (0.62). Depredation control was the leading cause of mortality. Survival rates of female (0.67) and male (0.58) lions did not differ. Because of the high mortality in the area of depredation control, this area may represent a mortality sink.
Article
Large carnivores are recolonizing portions of midwestern North America given harvest protection and the presence of suitable habitat in the region. Perhaps more so than other species, the successful management and viability of large carnivore populations is as dependent on social acceptance as on biological factors. However, knowledge of human attitudes and perceptions toward large carnivores in much of the Midwest remains unknown. We assessed attitudes and perceptions of Illinois citizens about black bears (Ursus americanus), cougars (Puma concolor), and gray wolves (Canis lupus) via a mail survey to provide wildlife managers with an understanding of residents' views prior to formulating carnivore management plans. The survey instrument consisted of questions about large carnivores regarding knowledge and beliefs, experiences and encounters, attitudes toward carnivores and management, and demographic questions and behavioral characteristics. We surveyed residents statewide; the sample drawn was stratified by geographic region (northern, central, and southern) and urban or rural county designation within regions. Because we observed differences in demographic variables between respondents (by mail) and non-respondents (telephone interviewees), we did not pool responses from the 2 groups for analysis; the final response rate for the survey was 13%. More than 70% of survey respondents (n = 791) were male and their average age was 60; 55% were hunters. Approximately 40% were unsure about the population status of large carnivores in Illinois; of the remaining respondents, most (ranging from 20% for black bears to 41% for cougars) believed the presence of all 3 focal species had increased over the past decade. More residents supported protection (43%) and increasing numbers of large carnivores (39%) than opposed them (26%); however, support for black bears was slightly higher than for cougars and wolves. Rural residents and livestock owners were the most likely to want carnivore numbers to decrease and least likely to support their protection; higher levels of education corresponded to positive attitudes toward large carnivores. Our research provides the foundation for well-informed management plans, policy decisions, and educational initiatives for large carnivores in midwestern states where these species appear to be recolonizing following decades of absence. © 2014 The Wildlife Society.
Article
Cougar (Puma concolor) populations have expanded in many western areas of the United States. The Black Hills cougar population naturally recolonized the area and cohabitates within an ecosystem heavily dissected by roads and human activity. Assessing mortality characteristics and determining survival of cougar populations is critical for managing and conserving this species. Our objectives were to assess cause-specific mortality and estimate survival of the Black Hills cougar population, in addition to assessing annual mortality through ancillary opportunistic methods. We captured and radiocollared cougars during 1999–2005 to assess survival and cause-specific mortality. In addition to cause-specific mortality, we also documented all known cougar mortality opportunistically throughout the study area in conjunction with the South Dakota Department of Game, Fish and Parks. We captured 31 independent-aged cougars (n = 12 M, 19 F) for analyses. We opportunistically documented 85 mortality events of cougars in South Dakota during 1998–2005. Despite protection from hunting during our study, 61.5% of mortality was human-induced, in contrast to other studies of unhunted cougar populations that generally attribute natural mortality as the primary mortality source. Male and female cougars exhibited relatively high survival through the course of the study; however, low sample sizes precluded rigorous comparisons of annual survival rates between sex and age cohorts. Proportionally, the largest contributors to cougar mortality were lethal removal by the state agency (e.g., depredation, human-safety concerns) and vehicular trauma. Continued assessment of cause-specific mortality and survival will be useful for evaluating effects of future manipulations of this population. The potential effects of human-caused cougar mortalities should be considered when evaluating management strategies for cougars in landscapes with high propensity for human–cougar interactions.
Article
Although cougars (Puma concolor) were extirpated from much of midwestern North America around 1900, hard evidence of cougar presence has increased and populations have become established in the upper portions of the Midwest during the past 20 years. Recent occurrences of cougars in the Midwest are likely due to dispersal of subadult cougars into the region from established western populations, and may be indicative of further recolonization and range expansion. We compiled confirmed locations of cougars (i.e., via carcasses, tracks, photos, video, and DNA evidence) collected during 1990–2008 in 14 states and provinces of midwestern North America. We separated our study area into 2 regions (east and west), calculated number and types of confirmations, and assessed trends in confirmations during the study period. We recorded 178 cougar confirmations in the Midwest and the number of confirmations increased during the study period (r = 0.79, P ≤ 0.001). Confirmations by state or province ranged from 1 (Kansas, Michigan, and Ontario) to 67 (Nebraska). Carcasses were the most prevalent confirmation type (n = 56). Seventy-six percent of known-sex carcass confirmations were males, consistent with predominantly male-biased dispersal in cougars. More confirmations (P = 0.05) were recorded in the western region than the eastern region . Seventy-nine percent of cougar confirmations occurred within 50 km of highly suitable habitat (i.e., forest areas with steep terrain and low road and human densities). Given the number of cougar confirmations, the increasing frequency of occurrences, and that long-distance dispersal has been documented via radiocollared individuals, our research suggests that cougars are continuing to recolonize midwestern North America. © 2012 The Wildlife Society.
Article
We examined cougar ( Puma concolor) dispersal, emigration, and immigration in the San Andres Mountains, New Mexico, from 1985 to 1995 to quantify the effects of dispersal on the local population and surrounding subpopulations. We captured, tagged, and radio-collared animals to detect the arrival of new immigrants and dispersal characteristics of progeny. We found that cougars in southern New Mexico exhibited a metapopulation structure in which cougar subpopulations were separated by expanses of noncougar habitat and linked by dispersers. Of 43 progeny (n = 20 males, 23 females) studied after independence, only 13 females exhibited philopatric behavior. Males dispersed significantly farther than females, were more likely to traverse large expanses of noncougar habitat, and were probably most responsible for nuclear gene flow between habitat patches. We estimated that an average of 8.5 progeny (i.e., cougars born in the study area) successfully emigrated from and 4.3 cougars successfully immigrated to the San Andres Mountains each year. Concurrently, an average of 4.1 progeny were recruited into the San Andres cougar population. Protected cougar subpopulations can contribute to metapopulation persistence by supplying immigrants to surrounding subpopulations that are affected by fragmentation or offtake by humans. Cougar population dynamics and dispersal behavior dictate that cougar management and conservation should be considered on a regional scale.
Article
We examined the genetic structure of 5 Wyoming cougar (Puma concolor) populations surrounding the Wyoming Basin, as well as a population from southwestern Colorado. When using 9 microsatellite DNA loci, observed heterozygosity was similar among populations (HO = 0.49–0.59) and intermediate to that of other large carnivores. Estimates of genetic structure (FST = 0.028, RST = 0.029) and number of migrants per generation (Nm) suggested high gene flow. Nm was lowest between distant populations and highest among adjacent populations. Examination of these data, plus Mantel test results of genetic versus geographic distance (P ≤ 0.01), suggested both isolation by distance and an effect of habitat matrix. Bayesian assignment to population based on individual genotypes showed that cougars in this region were best described as a single panmictic population. Total effective population size for cougars in this region ranged from 1,797 to 4,532 depending on mutation model and analytical method used. Based on measures of gene flow, extinction risk in the near future appears low. We found no support for the existence of metapopulation structure among cougars in this region.
Article
Cougars (Puma concolor) are of considerable interest to wildlife biologists and the general public in midwestern North America, yet no researchers have modelled potential habitat in the region. We created a model of potential cougar habitat in 9 midwestern states using geospatial data, expert-opinion surveys, the analytical hierarchy process, and a GIS. About 8% of the study region contained highly favorable habitat (with favorability scores ≥75%) for cougars; the states of Arkansas (19%) and Missouri (16%) contained the highest proportions of potentially favorable habitat. We identified 6 large (≥2500km2 in size), contiguous areas of highly favorable habitat for cougars. Model testing procedures indicated a valid model when compared to an independent set of cougar locations, a null dataset, and similar studies. Our model is useful as a planning tool to proactively address future human–cougar conflicts should cougars re-colonize the Midwest via subadult dispersal.
Article
We present an analysis of the metapopulation dynamics of the federally threatened coastal California Gnatcatcher (Polioptila c. californica) for an approximately 850 km2 region of Orange County, California. We developed and validated a habitat suitability model for this species using data on topography, vegetation, and locations of gnatcatcher pair observation. Using this habitat model, we calculated the spatial structure of the metapopulation, including size and location of habitat patches and the distances among them. We used data based on field studies to estimate parameters such as survival, fecundity, dispersal, and catastrophes, and combined these parameters with the spatial structure to build a stage-structured, stochastic, spatially-explicit metapopulation model. The model predicted a fast decline and high risk of population extinction with most combinations of parameters. Results were most sensitive to density-dependent effects, the probability of weather-related catastrophes, adult survival, and adult fecundity. Based on data used in the model, the greatest difference in results was given when the simulation's time horizon was only a few decades, suggesting that modeling based on longer or shorter time horizons may underestimate the effects of alternative management actions.
Article
Studies of the interaction of wolves and their prey during the past twenty years have demonstrated that ungulates are the primary prey of wolves both in winter and summer and that predation in summer tends to be concentrated on the young of the year while in winter it is concentrated on animals in older-age classes. There appears to be intrinsic control of wolf numbers and there is evidence which suggests that a wolf per 10 square miles may approach the maximum density that is attainable in most ranges. The adaptations between ungulates and their predators may have evolved in relatively stable forest environments which could not support high-density prey populations. This could explain why wolves do not appear to be capable of controlling moose and deer populations in environments that have been drastically altered by man.