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Wolves (Canis lupus) have been captured with foothold traps for several decades to equip them with radiocollars for population monitoring. However, trapping in most areas is limited to spring, summer, and autumn as cold winter temperatures can lead to frozen appendages in trapped animals. In addition, conflicts arise when domestic dogs encounter these traps in nonwinter seasons. An alternative capture method is the use of cable restraint devices (modified neck snares) in the winter. We evaluated injury scores, movement patterns, and space use of wolves captured in cable restraint devices and foothold traps in north‐central Minnesota, USA, during 2012–2016. Injury scores did not differ between capture techniques; however, movement patterns and space use were different. We found that the movement away from the capture site appeared to plateau by approximately 8–10 days for wolves captured by either foothold traps or cable restraints, but wolves captured in traps travelled farther away. Daily movement rates reached an asymptote approximately 14 days earlier for wolves captured with cable restraints as compared with wolves caught with foothold traps. We found the space use among wolves caught with cable restraint devices plateaued in a shorter time frame than wolves caught with foothold traps whether using days since capture (38 days earlier) or number of locations (149 locations earlier). When we controlled for seasonal effects and the presence of a capture using locational data collected 6 months later, there was no difference in space use. We concluded that wolves captured in cable restraints recovered more quickly from the capture and resumed space use and activity patterns more rapidly than wolves captured with foothold traps. Published 2019. This article is a U.S. Government work and is in the public domain in the USA.

Cross-fostering offspring with nonbiological parents could prove useful to augment populations of endangered carnivores. We used cross-fostering to augment captive-born and wild-born litters for the endangered red wolf (Canis rufus). Between 1987 and 2016, 23 cross-fostering events occurred involving captive-born pups fostered into captive litters (n = 8 events) and captive-born pups fostered into wild recipient litters (n = 15 events). Percentage of pups surviving 3 and 12 months was 91.7% for captive-born pups fostered into captive recipient litters. For pups fostered into wild litters, percentage of pups surviving 5 months was > 94% among fostered pups (pups fostered into a wild red wolf litter or replaced a hybrid litter), pups in recipient litters (wild-born litters receiving fostered pups), and pups in control litters (wild-born litters not in a fostering event) when using pups with known fates. Including pups with unknown fates as deaths, percentage of pups surviving 5 months was > 54% among fostered pups, pups in recipient litters, and pups in control litters. Among wild litters, percentage of pups surviving 12 months was > 82% among fostered pups, pups in recipient litters, and pups in control litters when using pups with known fates. Including pups with unknown fates as deaths, percentage of pups surviving 12 months was > 48% among fostered pups, pups in recipient litters, and pups in control litters. Although survival to 12 months was similar among the groups, average life span was different with pups in control litters living 3.3 years, pups in recipient litters living 4.6 years, and fostered pups living 5.6 years. Of fostered pups surviving > 12 months in the wild, 9 animals whelped or sired 26 litters. Cross-fostering was successful at augmenting litter size for red wolves without any deleterious effects on recipient litters, illustrating fostering as a tool for increasing populations of endangered carnivores.

Polar bears (Ursus maritimus) are experiencing rapid and substantial changes to their environment due to global climate change. Polar bears of the southern Beaufort Sea (SB) have historically spent most of the year on the sea ice. However, recent reports from Alaska indicate that the proportion of the SB subpopulation observed on‐shore during late summer and early fall has increased. Our objective was to investigate whether this on‐shore behavior has developed through genetic inheritance, asocial learning, or through social learning. From 2010 to 2013, genetic data were collected from SB polar bears in the fall via hair snags and remote biopsy darting on‐shore and in the spring from captures and remote biopsy darting on the sea ice. Bears were categorized as either on‐shore or off‐shore individuals based on their presence on‐shore during the fall. Levels of genetic relatedness, first‐order relatives, mother–offspring pairs, and father–offspring pairs were determined and compared within and between the two categories: on‐shore versus off‐shore. Results suggested transmission of on‐shore behavior through either genetic inheritance or social learning as there was a higher than expected number of first‐order relatives exhibiting on‐shore behavior. Genetic relatedness and parentage data analyses were in concurrence with this finding, but further revealed mother–offspring social learning as the primary mechanism responsible for the development of on‐shore behavior. Recognizing that on‐shore behavior among polar bears was predominantly transmitted via social learning from mothers to their offspring has implications for future management and conservation as sea ice continues to decline.

Molecular forensics is an important component of wildlife research and management. Using DNA from noninvasive samples collected at predation sites, we can identify predator species and obtain individual genotypes, improving our understanding of predator–prey dynamics and impacts of predators on livestock and endangered species. To improve sample collection strategies, we tested two sample collection methods and estimated degradation rates of predator DNA on the carcasses of multiple prey species. We fed carcasses of calves (Bos taurus) and lambs (Ovis aires) to three captive predator species: wolves (Canis lupus), coyotes (C. latrans), and mountain lions (Puma concolor). We swabbed the carcass in the field, as well as removed a piece of hide from the carcasses and then swabbed it in the laboratory. We swabbed all tissue samples through time and attempted to identify the predator involved in the depredation using salivary DNA. We found the most successful approach for yielding viable salivary DNA was removing hide from the prey and swabbing it in the laboratory. As expected, genotyping error increased through time and our ability to obtain complete genotypes decreased over time, the latter falling below 50% after 24 h. We provide guidelines for sampling salivary DNA from tissues of depredated carcasses for maximum probability of detection.

Abundance estimators are often evaluated with simulations, or by comparing estimates to populations of known size. Advances in noninvasive genetic sampling have fueled an increase in the use of noninvasive genetic sampling-based capture–recapture. However, when working with free-ranging populations of unknown size, managers often lack data necessary to select the appropriate estimator. This leads to uncertainty regarding how choice of estimator or sampling design influence estimates, and managers may select estimators based on funding or logistical constraints. Alternatively, comparing estimates from multiple estimators can provide managers with greater confidence in estimates, or highlight potential differences. We used noninvasive genetic sampling to estimate the abundance of free-ranging kit foxes Vulpes macrotis and coyotes Canis latrans. We generated estimates of abundance with two non-spatial likelihood-based estimators: 1) robust design Huggins capture–recapture models and 2) single-occasion capture with replacement (CAPWIRE) models. We compared these with recently published estimates derived from spatially explicit capture–recapture (SECR) models. For both species, estimates from Huggins models were generally lower than those from SECR. Abundance estimates from CAPWIRE, which was developed specifically for noninvasive genetic sampling and generates estimates from a single sampling occasion, tended to be biased low with high precision. Our results suggest that choice of estimator and sampling design can significantly influence estimates, and that the relationship between estimators varied between species. Our results further suggest that single-occasion sampling often employed with CAPWIRE abundance estimation may produce biased results and be inappropriate for species requiring dispersed sampling strategies.

The increase of global urbanization can have effects on wildlife species, including carnivores such as coyotes ( Canis latrans ). As coyotes continue to settle in more urban areas, reports of human-coyote conflicts, such as attacks on humans or pets, may also increase. Understanding environmental variables that might influence whether or not coyotes and human-coyote conflicts will occur in certain urban areas may assist wildlife officials in creating management plans for urban wildlife. We conducted a survey of 105 urban areas in the United States requesting information on the occurrence of coyotes and human-coyote conflicts. We analyzed the responses with data on human population size, geographic region, land cover, housing density, and precipitation. Larger urban areas were more likely to contain both coyotes and human-coyote conflicts, and were also more likely to have greater numbers of conflicts. Urban areas in the western regions with larger amounts of high-intensity development and less forested and agricultural areas were more likely to have conflicts. Most urban areas considered the management of conflicts to be of low priority and emphasized education of citizens rather than removal of individual coyotes. Our results may assist urban wildlife managers in understanding the geographic and demographic factors correlated with the occurrence of coyotes and human-coyote conflicts. Practices such as education campaigns and landscape design incorporating wildlife habitat modifications (e.g., reducing dense cover) may reduce human-carnivore conflicts in urban ecosystems.

An understanding of how a predator uses the landscape can assist in developing management plans. We modeled resource selection by cougars (Puma concolor) during 2 behavioral states (moving and killing) and 2 seasons (summer and winter) with respect to landscape characteristics using locations from global positioning system (GPS)-collared cougars in the Pryor Mountains, Montana and Wyoming, USA. Furthermore, we examined predation-specific resource selection at 2 scales (fine and coarse). When possible, we backtracked from cache sites to kill sites and used a fine-scale analysis to examine landscape characteristics of confirmed kills. At this fine scale, kill sites had less horizontal visibility, were more likely to be in juniper (Juniperus spp.)-mountain mahogany (Cercocarpus ledifolius), and were less likely to be in grassland vegetation. For the coarse-scale analysis of predation risk, we used the entire dataset of kills by buffering each cache site by 94.9 m, which was the 95% upper cut-off point of the known distances dragged from kill sites to cache sites, thereby creating buffered cache sites that had a high probability of containing the kill site. We modeled seasonal cougar predation site selection by constructing resource selection functions for these buffered cache sites. The top model for summer predation risk consisted of vegetation class, distance to water, and slope. The top model for winter predation risk included vegetation class and elevation. These predation risk models were similar to but simpler than the resource selection models developed from the moving locations. Essentially, the behavioral state (i.e., killing vs. moving) of the cougar had little influence on resource selection, indicating that cougars are generally in hunting mode while moving through their landscape. To potentially reduce cougar predation on mule deer (Odocoileus hemionus) and bighorn sheep (Ovis canadensis) in our study area, managers can intersect the cougar predation-risk resource selection functions with deer and sheep habitat to guide habitat modification efforts aimed at increasing horizontal visibility in risky vegetation classes. © 2016 The Wildlife Society.

Urbanization alters landscapes and ecosystem processes that result in negative impacts for many species. However, urbanization also creates novel environments that certain species, including carnivores, are able to exploit. Coyotes ( Canis latrans ) are 1 example of a species capable of exploiting urban environments throughout North America and, in some cases, becoming involved in human–coyote conflict. As part of a comprehensive study of human–coyote coexistence in the Denver metropolitan area of Colorado, we investigated the spatial ecology of coyotes to determine movement and activity patterns relative to the urban matrix. We examined home-range size, habitat use, and resource selection for 22 coyotes monitored with GPS collars during 2012–2014. Mean (± SD ) home-range size of resident coyotes (11.6±11.0 km2) was smaller than ranges of transient coyotes (200.7±232.4 km2). Home-range size did not vary by season or sex, but resident coyotes during the day (7.2±10.5 km2) had smaller home ranges than during the night (11.3±10.8 km2). Coyotes had high percentages of developed lands (44.5±18.9%) within their home ranges, contrary to previous studies of urban coyotes. However, the percentage of coyote locations in natural lands (48.9±22.4%) was higher than in developed lands (20.6±11.7%). Home-range size of residents was not related to either the percentage of developed lands or altered lands within home ranges. Coyotes selected natural lands over developed lands, and they increased activity at night. Although coyotes were able to thrive in home ranges containing large amounts of development, they continued to avoid areas with high human activity by primarily residing in areas with natural land cover. Similar to urban areas throughout the Northern Hemisphere, coyotes in the Denver metropolitan area have become efficiently adapted to a highly developed landscape, reflecting the flexible nature of this opportunistic carnivore.

Rodent communities have multiple functions including comprising a majority of the mammalian diversity within an ecosystem, providing a significant portion of the available biomass consumed by predators, and contributing to ecosystem services. Despite the importance of rodent communities, few investigations have explored the effects of increasing anthropogenic modifications to the landscape on rodents. Throughout the western United States, the construction of artificial water developments to benefit game species is commonplace. While benefits for certain species have been documented, several researchers recently hypothesized that these developments may cause unintentional negative effects to desert-adapted species and communities. To test this idea, we sampled rodents near to and distant from wildlife water developments over 4 consecutive summers. We employed an asymmetrical before-after-control-impact (BACI) design with sampling over 4 summers to determine if water developments influenced total rodent abundance. We performed an additional exploratory analysis to determine if factors other than free water influenced rodent abundance. We found no evidence that water developments impacted rodent abundance. Rodent abundance was primarily driven by vegetation type and year of sampling. Our findings suggested that water developments on our study area do not represent a significant disturbance to rodent abundance and that rodent abundance was influenced by the vegetative community and temporal factors linked to precipitation and primary plant production. Our findings represent one of the 1st efforts to determine the effects of an anthropogenic activity on the rodent community utilizing a manipulation design.

Scat surveys are commonly used to monitor wildlife populations. For carnivores, surveys are typically conducted along roads and trails. Scats available for detection may not reflect scats deposited and variation in disappearance may bias results. Previous research has investigated natural decay and deterioration, but scats deposited along roads or trails are likely influenced to a greater degree by anthropogenic disturbance in some systems. We used experimental plots to evaluate variation in scat removal for two model carnivores, coyote Canis latrans and kit fox Vulpes macrotis, along roads in the Great Basin Desert, USA. Using parametric survival regression, we predicted scat survival and developed persistence-rate correction factors, which were applied to results from relative abundance scat surveys conducted along 15 transects. Kit fox scats disappeared more rapidly than coyote scats, with 3.3% and 10.6%, respectively, persisting through 42 days. At 14 days, 90.8-41.7% of scats had been removed across ...

article i nfo One of the most endangered species is the red wolf, Canis rufus. Reintroduction of the red wolf began in 1987, but in 1993 hybridization between coyotes (Canis latrans) and wolves was documented. To reduce genetic introgres- sion, coyotes and coyote-wolfhybrids were captured, sterilized, and releasedas "placeholders". Placeholders held territories until either displaced or killed by a wolf, or management personnel removed them before releasing a wolf. We evaluated the placeholder concept by examining the number of animals sterilized and released, likeli- hood of displacement by a wolf, factors influencing displacements, territory fidelity of placeholders, and survival rates and causes of mortality of placeholders and wolves. Of the 182 placeholders, 125 were coyotes and 57 were hybrids. From 1999 to 2013, 51 placeholders were displaced or killed by wolves, and 16 were removed by management personnel. Thus, 37% of the placeholders were displaced leading to occupancy by a wolf. Most dis- placements occurred in winter (43%) and were always by the same sex. Males were more likely to be displaced than females. Home range characteristics influencing the probability of displacement included home-range size (i.e., more placeholders displaced from larger home ranges) and road density (i.e., more placeholders displaced from home ranges with lower road density). Annual survival of placeholders was higher than wolves in 12 of 14 years, with cause-specific mortality similar among wolves and placeholders. Placeholders provided territories for wolves to colonize, yet reduced the production of hybrid litters, thereby limiting genetic introgression to b4% coyote ancestry in the wolf population. Published by Elsevier Ltd.

Harvest indices are used by state wildlife management agencies to monitor population trends and set harvest quotas for furbearer species. Although harvest indices may be readily collected from hunters, the reliability of harvest indices for monitoring demography and abundance of the harvested species is rarely examined, particularly amongst large carnivores. The overall objective of this study was to assess whether cougar (Puma concolor) harvest statistics collected bywildlife managers were correlated with changes in cougar demography, mainly survival rates and abundance. We estimated key demographic parameters for 2 cougar populationsinUtah over17years during whichwemonitored235 radio-collared cougars. Wethencompared these demographic parameters to harvest statistics provided by the Utah Division of Wildlife Resources over the same time period for the Oquirrh-Stansbury (lightly harvested population) and Monroe (heavily harvested population) harvest management units. In the Oquirrh-Stansbury unit, the percent of harvested cougars >6 years old was positively correlated with annual survival, indicative of a population experiencing several years of high survival resulting in an older age structure. Percent of permits filled and cougar abundance were also significantly correlated, suggesting higher hunting success with increased density. In the Monroe management unit, the annual percent of permits filled was correlated with changes in overall annual survival and male and female annual survival. Of utmost importance, pursuit success (cougars treed/day) increased with the number of cougars on the unit suggesting that pursuit indices may be an informative metric for wildlife managers to determine cougar population trends. Because both management units were subjected to contrasting mortality regimes, results provided by this assessment could potentially be applied to additional management areas sharing similar ecological characteristics and harvest metrics. Published 2015. This article is a U.S. Government work and is in the public domain in the USA.

Understanding the interplay between exploitation and natural mortality is essential to guiding sustainable conservation of wildlife. Exploitation of carnivores by humans has long been thought to result in compensatory reductions of natural mortality among survivors. If rates of human exploitation exceed natural mortality, however, such actions will ‘add’ to overall mortality and could imperil the sustainability of such actions. We applied competing risk analyses to P16 years of data for heavily harvested and semi-protected cougar populations in Utah to test the additive and compensatory mortality hypotheses, while accounting for parameter uncertainty. We additionally tested for presence of the two primary mechanisms by which compensatory mortality can arise: density dependence and individual heterogeneity in mortality risks. Despite an opportunity for compensation in the heavily harvested population, we could not reject the additive mortality hypothesis when uncertainty in parameter estimates was accounted for. In the semi-protected population, however, we detected evidence for partial compensation of increased anthropogenic exploitation via reductions in natural mortality. As may be common in carnivore studies, we found that ignoring uncertainty in estimates of cause-specific mortality systematically led to biased conclusions regarding additive and compensatory mortality hypotheses. Efforts should be made to address and minimize this uncertainty in demographic studies of carnivores in order to avoid flawed management recommendations. To attain the necessary sample sizes for making sound inference, this may require that the spatial extent of management units be extended for territorial species with large home-range requirements.

Monitoring mammalian carnivores at den sites with human observers to document behavior, fecundity, litter size, and natal survival is commonplace when compared with monitoring den sites with cameras. However, no published studies exist comparing the effectiveness of human observers versus cameras in a quantitative manner. Obtaining complete and reliable counts of adult and pup kit foxes (Vulpes macrotis) can be crucial for estimating population parameters and life-history traits. In the summers of 2010 and 2011, we made weekly counts of adult and pup kit foxes at active den sites using 2 methods: 1) an observer equipped with spotting scope and/or binoculars and 2) a remote infrared motion-sensitive camera. We accumulated 71 and 29 counts at active den sites, respectively. The median number of adults observed at den sites with a camera and observer differed and were 2 adults versus 1 adult, respectively. Similarly, the median number of pups observed at den sites with a remote camera versus an observer differed and was 2 pups versus 1 pup, respectively. We observed these differences despite the similar effort required to manage cameras and conduct observations. We recommend future surveys aimed at ascertaining more reliable counts of kit foxes and similar species at den sites use a survey methodology employing remote cameras or video over a multi-day period. 2013 The Wildlife Society.

Resource selection studies are common in the wildlife ecology literature and typically rely on the comparison of locations used by wildlife and locations assumed to be available for use but where use was not observed. While standard use-availability designs are helpful for establishing general patterns of species occurrence, they are limited in their ability to help researchers understand the underlying behavioural mechanisms that lead to observed space-use patterns. Based on spatially-explicit behavioural observations from coyotes Canis latrans in Yellowstone National Park, Wyoming, we estimated resource selection for specific behaviours (i.e. predatory, laying and travelling) and for all used locations irrespective of behaviour, to test whether resource selection is behaviour-specific and not generalizable across behaviours. Behaviour-specific models differed significantly from the model not partitioned by behaviour. In particular, the predatory model identified selection for mesic-meadows which have pr...

Snowshoe hares (Lepus americanus) are a primary prey species for Canada lynx (Lynx canadensis) in western North America. Lynx management plans require knowledge of potential prey distribution and abundance in the western United States. Whether even-aged regenerating forests or multi-storied forests contain more snowshoe hares is currently unknown. During 2006–2008, we estimated snowshoe hare density in 3 classes of 30–70-year-old lodgepole pine (Pinus contorta) and 4 classes of late seral multi-storied forest with a spruce (Picea engelmannii)-fir (Abies lasiocarpa) component in the Bridger-Teton National Forest, Wyoming. We recorded physiographic variables and forest structure characteristics to understand how these factors influence abundance of snowshoe hares. In many instances, snowshoe hares were more abundant in late seral multi-storied forests than regenerating even-aged forests. Forest attributes predicting hare abundance were often more prevalent in multi-storied forests. Late seral multi-storied forests with a spruce–fir component and dense horizontal cover, as well as 30–70-year-old lodgepole pine with high stem density, were disproportionately influential in explaining snowshoe hare densities in western Wyoming. In order to promote improved habitat conditions for snowshoe hares in this region, management agencies should consider shifting their focus towards maintaining, enhancing, and promoting multi-storied forests with dense horizontal cover, as well as developing 30–70-year-old lodgepole pine stands with high stem density that structurally mimic multi-storied forests. © 2012 The Wildlife Society.

Snowshoe hares (Lepus americanus) are an important prey species for Canada lynx (Lynx canadensis) and are considered critical for lynx population persistence. Determination of snowshoe hare distribution and abundance is needed by land management agencies for lynx conservation. An accepted approach for estimating snowshoe hare abundance is the use of fecal-pellet plot counts. Locally derived regression equations are preferred for accurate calibration of pellet counts to snowshoe hare density due to local differences in pellet deposition and decomposition. We used linear regression to examine correlations between snowshoe hare density, as determined by mark–recapture estimates, and pellet plot counts on both uncleared plots and annually cleared plots on the Bridger-Teton National Forest, western Wyoming, USA. We found significant correlations between snowshoe hare density estimates and fecal pellet counts for both uncleared and annually cleared pellet counts; however, the relationship was stronger ...

Jaguars (Panthera onca) often prey on livestock, resulting in conflicts with humans. To date, kill rates and predation patterns by jaguars have not been well documented. We studied the foraging ecology of jaguars in an area with both livestock and native prey and documented kill rates, characteristics of prey killed, patterns of predation, and the influence of prey size on the duration at kill sites and the time interval between kills. Between October 2001 and April 2004 we monitored 10 jaguars equipped with global positioning system (GPS) collars. We collected 11,787 GPS locations and identified 1,105 clusters of locations as sites of concentrated use (e.g., kill sites, bed sites, and dens). Of these, we found prey remains at 415 kill sites and documented 438 prey items. Kills were composed of 31.7% cattle (9.8% adults and 21.9% calves), 24.4% caiman (Caiman crocodilus yacare), 21.0% peccaries (mostly Tayassu pecari), 4.1% feral hogs (Sus scrofa), 3.9% marsh deer (Blastocerus dichotomus), 3.2% giant anteaters (Myrmecophaga tridactyla), 2.0% capybaras (Hydrochoeris hydrochaeris), 1.6% brocket deer (Mazama americana and M. gouazoubira), and other avian, mammalian, and reptilian species. Individual jaguars differed in the proportion of each species they killed and the proportion of native prey versus cattle. Although all 10 cats killed cattle, 5 killed a high proportion of cattle (.35% of kills), and 3 killed few cattle (,15%). Males (27%) and females (35%) killed cattle in similar proportions. In contrast, male jaguars killed a higher proportion of peccaries than did females, and female jaguars killed more caiman than did males. The mean kill rate for all jaguars was 4.3 days 6 4.4 SD between known consecutive kills. The time interval to the next subsequent kill by jaguars increased with increasing prey size. Jaguars also increased the length of time at a carcass as prey size increased. Jaguar kill rates on peccaries steadily increased over the 4year study. In contrast, kill rates on cattle decreased during the same period. Rainfall, and subsequent water levels on the Pantanal, was the main driver of seasonal kill rates by jaguars on cattle and caiman. As water levels increased, predation on caiman increased as caiman became more distributed throughout the landscape. Conversely, as water levels fell, caiman became less plentiful, and cattle were moved out into pastures thereby increasing their availability to more jaguars. DOI: 10.1644/09-MAMM-A-171.1.

Investigations into mechanisms of resource partitioning are particularly suited to systems where nascent interactive behaviors are observable. Wolf (Canis lupus) recolonization of the Greater Yellowstone Ecosystem provided such a system, and we were able to identify behaviors influencing the partitioning of resources by coyotes (Canis latrans) and wolves. We observed coyote-wolf interactions immediately after wolf recolonization, when reemergent behaviors mediating the outcome of competitive interactions were detectable and mechanisms of spatial avoidance were identifiable. Although coyotes used the same space as wolves, they likely minimized risk of encounter by making adaptive changes in resource selection based on perception of wolf activity and potential scavenging opportunities. When exploiting carrion subsidies (i.e., wolf-killed ungulates), coyotes relied on social behaviors (i.e., numerical advantage in concert with heightened aggression) to mitigate escalating risk from wolves and increase resource-holding potential. By adapting behaviors to fluctuating risk, coyotes might reduce the amplitude of competitive asymmetries. We concluded coyotes do not perceive wolves as a threat requiring generalized spatial avoidance. Rather, the threat of aggressive interactions with wolves is spatially discrete and primarily contained to areas adjacent to carrion resources.

The Pantanal of Brazil is an important area for the conservation of jaguars (Panthera onca). As the size of traditional large ranches in the Pantanal decreases, human access to jaguar habitat increases, resulting in humanaltered landscapes that may influence patterns of resource selection and space use by jaguars. We used global positioning system radiocollars to study jaguars in the southern Pantanal. We radiocollared 10 jaguars (6 males and 4 females), obtained 11,787 locations, and examined their space use, movement rates, and social interactions between October 2001 and April 2004. Estimates of 90% kernel home ranges varied among animals and seasons (range: 34.1–262.9 km 2 ). Core areas (50% isopleth) of both females and males did not differ in size between seasons, but home ranges (90% isopleth) during the dry season were generally larger than during the wet season. The stability of home ranges varied among seasons and individuals. Some females maintained 80% of their home ranges from 1 season to the next, whereas other females used 50% of their home ranges from the previous season. Site fidelity within individual home ranges also varied; 70% of the core areas of some females were located in different sites within their home ranges during different seasons. Locations of females suggested a pattern of spatial avoidance among females during the wet season. Home-range overlap among males was extensive, both in the wet and dry seasons, suggesting that males did not maintain exclusive ranges. Overlap between males and females occurred both in the wet and dry seasons, and movements by females were not restricted within the ranges of individual males. Jaguars were located ,200 m apart more often than expected, suggesting some degree of sociality. The reproductive profiles of females suggested either a low conception rate, a low survival rate of young, or that jaguars may be more social than previously thought. Interactions among males also suggested some degree of sociality.

Minimizing risk of predation from multiple predators can be difficult, particularly when the risk effects of one predator species may influence vulnerability to a second predator species. We decomposed spatial risk of predation in a 2-predator, 2-prey system into relative risk of encounter and, given an encounter, conditional relative risk of being killed. Then, we generated spatially explicit functions of total risk of predation for each prey species (elk [Cervus elaphus] and mule deer [Odocoileus hemionus]) by combining risks of encounter and kill. For both mule deer and elk, topographic and vegetation type effects, along with resource selection by their primary predator (cougars [Puma concolor] and wolves [Canis lupus], respectively), strongly influenced risk of encounter. Following an encounter, topographic and vegetation type effects altered the risk of predation for both ungulates. For mule deer, risk of direct predation was largely a function of cougar resource selection. However, for elk, risk of direct predation was not only a function of wolf occurrence, but also of habitat attributes that increased elk vulnerability to predation following an encounter. Our analysis of stage-based (i.e., encounter and kill) predation indicates that the risk effect of elk shifting to structurally complex habitat may ameliorate risk of direct predation by wolves but exacerbate risk of direct predation by cougars. Information on spatiotemporal patterns of predation will be become increasingly important as state agencies in the western United States face pressure to integrate predator and prey management.

The reproductive biology of wild Canis species is often described as unique among mammals because an unusual combination of behavioral and physiological characteristics including a seasonally monestrous cycle, copulatory lock or tie, obligatory pseudopregnancy, social monogamy, and biparental care of the young. We investigated social behavior, endocrine profiles, and vaginal cytology of female coyotes (Canis latrans) during 4 breeding seasons, 2000–2003. Blood levels of estradiol, progesterone, prolactin, and relaxin were measured, and mating behavior and changes in vaginal epithelium were documented. After aligning the data from each individual to her estimated day of ovulation, we compared pregnant coyotes with nonpregnant females and evaluated temporal relationships among hormone levels, behavior, and vaginal cytology. We found that patterns of proceptive and receptive behaviors correlated with the secretion of steroid hormones, as did vaginal epithelial cytomorphosis. In addition, although progesterone levels of pregnant and pseudopregnant coyotes were indistinguishable, prolactin demonstrated a discernible intergroup difference and relaxin was only detectable in pregnant females. Although this study included characteristics not previously published for this species, it also showed how key aspects of reproduction were correlated temporally, and emphasized the importance of an integrated perspective when addressing the reproductive biology of coyotes, or other wild species of canids.

The distribution and abundance of swift foxes (Vulpes velox) has declined from historic levels. Causes for the decline include habitat loss and fragmentation, incidental poisoning, changing land use practices, trapping, and predation by other carnivores. Coyotes (Canis latrans) overlap the geographical distribution of swift foxes, compete for similar resources, and are a significant source of mortality amongst many swift fox populations. Current swift fox conservation and management plans to bolster declining or recovering fox populations may include coyote population reduction to decrease predation. However, the role of coyote predation in swift fox population dynamics is not well-understood. To better understand the interactions of swift foxes and coyotes, we compared swift fox population demographics (survival rates, dispersal rates, reproduction, density) between areas with and without coyote population reduction. On the Pinon Canyon Maneuver Site, Colorado, USA, we monitored 141 swift foxes for 65,226 radio-days from 15 December 1998 to 14 December 2000 with 18,035 total telemetry locations collected. Juvenile swift fox survival rate was increased and survival was temporarily prolonged in the coyote removal area. Adult fox survival patterns were also altered by coyote removal, but only following late-summer coyote removals and, again, only temporarily. Coyote predation remained the main cause of juvenile and adult fox mortality in both areas. The increase in juvenile fox survival in the coyote removal area resulted in a compensatory increase in the juvenile dispersal rate and an earlier pulse in dispersal movements. Adult fox dispersal rate was more consistent throughout the year in the coyote removal area. Coyote removal did not influence the reproductive parameters of the swift foxes. Even though juvenile survival increased, swift fox density remained similar between the areas due to the compensatory dispersal rate among juvenile foxes. We concluded that the swift fox population in the area was saturated. Although coyote predation appeared additive in the juvenile cohort, it was compensatory with dispersal.

Numerous studies have documented how prey may use antipredator strategies to reduce the risk of predation from a single predator. However, when a recolonizing predator enters an already complex predator–prey system, specific antipredator behaviors may conflict and avoidance of one predator may enhance vulnerability to another. We studied the patterns of prey selection by recolonizing wolves (Canis lupus) and cougars (Puma concolor) in response to prey resource selection in the northern Madison Range, Montana, USA. Elk (Cervus elaphus) were the primary prey for wolves, and mule deer (Odocoileus hemionus) were the primary prey for cougars, but elk made up an increasingly greater proportion of cougar kills annually. Although both predators preyed disproportionately on male elk, wolves were most likely to prey on males in poor physical condition. Although we found that the predators partitioned hunting habitats, structural complexity at wolf kill sites increased over time, whereas complexity of couga...

Wolves (Canis lupus) have expanded their distribution into areas of the midwest United States that have not had wolves for several decades. With recolonization of wolves into agricultural areas, there is increasing concern of wolf–livestock conflicts. To assess the risk wolves may pose to livestock, we initiated a 3-year study investigating the activity patterns, movements, habitat use, visitation to livestock pastures by wolves, and the occurrence of depredation events in an agricultural–wildland matrix in northwestern Minnesota, USA. From June 1997 to November 1999, we captured 23 wolves, including pups, from 3 packs; we radiocollared 16 of these wolves. We tracked radioed wolves intensively on a 24-hour basis during the spring, summer, and autumn of 1998 and 1999. We found wolves passed directly through a pasture containing cattle on 28% of the nights of tracking; 58% and 95% of the wolf locations were ≤1 km and ≤5 km from a pasture, respectively. Space use of wolves showed that while they visited livestock pastures during the 24-hour tracking sessions, they apparently were passing through these pastures with cattle and not preying on livestock. When compared to random simulations of movements, wolves appeared to encounter livestock pastures randomly. Thirty percent of random movements passed directly through a pasture; 65% and 95% of random movements were within ≤1 km and ≤5 km of a pasture, respectively. Wolves were more active at night than during the day. Wolves avoided pastures during the day and visited pastures at night when depredations were most likely (i.e., human presence was low). Visitation of livestock pastures was not related to any discernible characteristics of the pastures (i.e., pasture size, cattle density, distance to human habitation, percent forest cover, index of deer abundance). However, pastures in which livestock were killed by wolves contained more cattle than pastures without depredations, but in 1998 only. While the risk of wolf predation on livestock was potentially high (wolves were within ≤1 km of a pasture on 58% of nights), few livestock were actually killed. During the 3-year study, only 8 animals (all young or vulnerable livestock) were depredated by wolves. Maintaining healthy wild prey populations, removing offending wolves that kill livestock, and encouraging effective and proper husbandry practices (e.g., disposal of carcasses) among livestock producers, should allow for the persistence of wolves in northwestern Minnesota, USA, while minimizing their impact to farmers in this agriculture–wildland matrix.

The United States Fish and Wildlife Service (USFWS) developed guidelines for the composition and role of endangered species recovery implementation teams, but few teams have been established and their success has not been evaluated. Using the recovery program of the red wolf (Canis rufus) as a model, we describe the genesis, function, and success of the Red Wolf Recovery Implementation Team (RWRIT) in helping guide the establishment of a viable red wolf population in eastern North Carolina. In operation since 1999, the RWRIT meets bi-annually to review USFWS progress and provide recommendations aimed at maximizing success of species recovery. The team is comprised of 8 research scientists from disciplines including population genetics, canid ecology, population ecology, veterinary medicine, and captive management. Representation from each of these disciplines is deemed necessary for proper evaluation of recovery progress and assessment of future needs. Meeting attendance by the USFWS field manage...

The natural recolonization of gray wolves (Canis lupus) into parts of their former range in the upper Midwest of the United States has led to wolves establishing territories in semi- agricultural areas containing livestock. As part of a study on wolf-livestock relationships in a northwestern Minnesota agricultural area, we surveyed rural landowners within and outside of wolf range to assess perceptions regarding the risks wolves pose to livestock (mainly cattle). The mean response score for rural landowners to the statement "I think wolves should be allowed to exist in northwest Minnesota" was between neutral and dis- agree. There was no difference in mean response scores between rural residents living within wolf range and residents living adjacent to but outside of wolf range. The rural res- idents' mean response score to the statement "Wolves are causing unacceptable levels of damage to northwest Minnesota's livestock industry" was between neutral and agree. Although there was a statistical difference in mean response scores of residents living within wolf range and residents living outside of wolf range, the scores were not sub- stantially different from each other. While landowners felt wolves were a threat to their livelihood, other factors (market fluctuations, laws and government, diseases, extreme weather, flooding) were ranked as greater threats to the agricultural community. Rural residents both within and outside of wolf range harbored similar negative attitudes toward wolves, even though residents outside of wolf range have not had a population of wolves in their area for >100 years, indicating little change in cultural attitudes toward wolves.

We examined social group formation, movements and denning relative to other group members, home-range use, and the response to social group disruption among 188 radiocollared swift foxes (Vulpes velox) in Colorado. We found that during the initial stages of pair formation mated foxes shared dens more frequently than during the remainder of their pair-bond. The average distance between mated pairs was influenced by season and time of day, with the greatest proximity in the breeding season (727.2 m ± 42.3 SE), and during diurnal hours (463.7 ± 34.7 m). Female foxes spent more time in the core area of the group home range (60.71% of locations versus 54.66% for males) and males spent more time on the range boundary (19.34% of locations versus 15.61% for females). Home-range use was influenced by season, with females spending more time in the core area in the breeding and pup-rearing seasons than in the dispersal season. Males also spent the highest proportion of their time in the core area during the breeding season, but used the boundary area more frequently than the core area in the pup-rearing season. A sex difference also was found in the response of a swift fox to the death or disappearance of its mate. All females maintained their territory in the event of mate death or disappearance; however, 50% of males emigrated from their range when the female mate died or disappeared. These differences in space use between social group members provide important insights into the territorial behavior and mating system of the swift fox and indicate that the 2 sexes likely play different roles in care of young and home-range defense.

Predation by coyotes (Canis latrans) on domestic sheep is a problem for many livestock producers throughout the United States Intermountain West. We examined whether surgical sterilization of coyote packs would modify their predatory behavior and reduce predation rates on domestic sheep as compared to coyote packs with pups. From June 1997 to December 1997, we gathered baseline information on coyote pack size and movements. In winter 1998, we surgically sterilized and radiocollared members of 5 coyote packs. We also captured and radiocollared members of 6 packs that remained intact (i.e., reproductive). During summer 1998, only 1 sterile pack killed a lamb, while 3 intact packs killed 11 lambs. When only sheep-killing packs were included, sterile packs killed an average of 0.35 lambs/week, while intact packs killed 1.53 lambs/week in 1998. During winter 1999, we monitored 4 sterile and 8 intact packs. In summer 1999, 3 sterile packs killed 3 lambs, while 4 intact packs killed 22 lambs. Considering only sheep-killing packs, sterile packs killed on average 0.38 lambs/week, while intact packs killed an average of 2.95 lambs/week in 1999. Coyotes were more likely to kill lambs that were on the edges of coyote territories as compared to core areas. Lambs of less than average weight were also more likely to be killed by coyotes. The available rodent biomass in each territory was not an influence on the differential kill rates exhibited between sterile and intact packs, nor did the amount of available alternate prey influence annual coyote predation rates on sheep. We conclude that we could use surgical sterilization to modify the predatory behavior of coyotes associated with pup production and provisioning of pups. Sterilization successfully reduced, but did not eliminate, coyote predation on domestic sheep. The amount of losses averted in the first year exceeded the costs associated with surgically sterilizing a coyote pack, which indicates that surgical sterilization could prove beneficial on small-scale livestock operations.

We tracked 72 radio-collared coyotes (Canis latrans) for 13 consecutive biological seasons spanning 4 years. Coyotes displayed 2 behavior modes based on home-range characteristics: resident and transient. Resident and transient coyotes comprised 78 and 22% of the population, respectively. The mean annual home-range size was 11.3 km2 (range = 2.8-32.0 km2) and 106.5 km2 (range = 60.9-185.3 km2) for resident and transient coyotes, respectively. Yearlings and very old coyotes (>8 yr) accounted for 68% of the transient cohort. Resident annual home ranges in canyon, hill, pinyon (Pinus edulis)-juniper (Juniperus monosperma)prairie, and prairie habitats averaged 5.5, 6.6, 11.1, and 16.5 km2, respectively. Coyotes preferred pinyonjuniper woodlands and shrub-grasslands but used open grassland habitat less than expected. Regression analysis showed that 65% of the variation in resident home ranges could be explained by the amount of available pinyon-juniper cover. Coyotes with access to little or no pinyon-juniper cover used shrub-grasslands. Similarily, 47% of the variation of home ranges for coyotes inhabiting the prairie could be explained by the amount of available shrub-grassland habitat within an animal's home range. J. WILDL. MANAGE. 52(4):640-646 Coyote home ranges have been studied throughout the United States (Gipson and Sealander 1972, Berg and Chesness 1978, Andelt and Gipson 1979, Springer 1982, Woodruff and Keller 1982, Andelt 1985). Most studies pooled home-range sizes from their study area, reported a mean size, and compared their estimate to other studies. To date, no one has attempted to explain the variance in home-range sizes. Laundre, and Keller (1984) concluded that future studies should examine coyote home-range use in relation to physical, faunal, and vegetal characteristics. We describe home-range use and size relative to habitat use by coyotes in southeastern Colorado. We examined the relationship between home-range size and availability of vegetative components. We thank P. A. Terletzky, B. A. Abel, L. A. H ugh, G. B. Perlmutter, F. M. Lentsch, and C. C. Gordon for field assistance; the Western En rgy and Land Use Team (WELUT) and K. M. Firchow for habitat mapping; J. R. Cary for computer assistance; and M. D. Samuel for staistical assistance. J. A. Bissonette, J. W. Laundr6, and an anonymous reviewer provided critical reviews of the manuscript. This study was funded by the Environmental, Energy, and Natural Resources Division, U.S. Army, Fort Carson, Colorado, through the U.S. Fish and Wildlife Service (USFWS), Colorado Fish and Wildlife Assistance Office, Golden, Colorado, and the USFWS Wisconsin Cooperative Wildlife Re' Present address: U.S. Fish and Wildlife Service, Creston Fisheries Center, Kalispell, MT 59901. This content downloaded from 157.55.39.243 on Tue, 04 Oct 2016 06:06:53 UTC All use subject to http://about.jstor.org/terms J. Wildl. Manage. 52(4):1988 COYOTE HOME RANGE AND HABITAT USE * Gese et al. 641 search Unit, the College of Agricultural and Life Sciences, and the Graduate School, University of Wisconsin, Madison.

We investigated the response of coyotes (Canis latrans) to military activity on the Pifion Canyon Maneuver Site (PCMS), Colorado, during 1984-86. Sixteen coyotes responded to military activity by expanding, contracting, abandoning, or not changing their home range during military maneuvers compared to before and after maneuvers. Three coyotes abandoned their home ranges, with 1 animal returning to its original (i.e., pre-military activity) home range 1 week after maneuvers. Most coyotes that expanded or contracted their home range during military maneuvers resumed their original home range after military maneuvers ceased. Responses appeared to be related to the amount of available cover, topography, and intensity of military activity in a coyote's home range. Coyote activity patterns during the day increased, while activity at sunrise, sunset, and night remained the same during military activity. J. WILDL. MANAGE. 53(2):334-339 The response of wildlife populations to human disturbances is of concern to wildlife professionals and the public. Most research has concentrated on the response of big game species to logging (Peek and Hieb 1976, Van Dyke et al. 1986), mining (Amstrup 1978, Helms 1978, Kuck et al. 1985), recreation (Dorrance et al. 1975, Richens and Lavigne 1978, Ferguson and Keith 1982), and development (Van Dyke et al. 1986, Knight et al. 1988). The response of protected species (e.g., raptors) to recreation and human activity has also been documented (Stalmaster and Newman 1978, Knight and Knight 1984, Andersen et al. 1986). The response of coyotes to increased human activity is unknown. We documented the effect of military activity on coyote home-range size, geometric centers of activity, and activity patterns. We thank P. A. Terletzky, G. B. Perlmutter, B. A. Abel, F. M. Lentsch, and J. L. Williamson for field assistance; the Western Energy and Land Use Team and K. M. Firchow for habitat mapping; J. R. Cary for computer and statistical assistance; and N. S. Smith, D. E. Andersen, T. L. Warren, A. P. Pfister, and S. R. Emmons for review of the manuscript. This study was funded by the Environmental, Energy, and Natural Resources Division, U.S. Army, Fort Carson, Colorado, through the U.S. Fish and Wildlife Service (USFWS), Colorado Fish and Wildlife Assistance Office, Golden, Colorado, and the USFWS Wisconsin Cooperative Wildlife Research Unit, the College of Agricultural and Life Sciences, and the Graduate School, University of Wisconsin, Madison.

We investigated the diet of coyotes (Canis latrans) on the Pifion Canyon Maneuver Site (PCMS) in southeastern Colorado from February 1983 to June 1986. Based on regression analysis, 71% of the variation in the volume of large prey (e.g., adult ungulates) in coyote scats could be explained by coyote group size; 77% of this variation could be explained by coyote group size and snow depth. Snow depth alone explained 28% of the variation in the volume of large prey in the scats. Coyote group size explained 19% of the variation in the volume of medium size prey (e.g., lagomorphs and ungulate fawns) and 38% of the variation in the volume of small prey (e.g., rodents). Coyote groups were largest in winter and smallest in summer. Most juveniles dispersed before the largest group sizes were observed. Group foraging and snow depth may influence prey selection by coyotes in southeastern Colorado. J. WILDL. MANAGE. 52(4):647-653 Diets of coyotes have been examined (Fichter et al. 1955, Gipson 1974, MacCracken 1981) but few studies have examined the relationship of prey selection and coyote social organization. In Jasper National Park, Alberta, during summer, when rodents were the major food, mean coyote group size was 1.4 (Bowen 1978, 1981). In winter, when ungulates were the major food, group size averaged 2.0. Mean group size in Wyoming was 1.3 coyotes in summer, when rodents were the major resource consumed; in winter, when ungulate carrion dominated the diet, coyote group size was 1.8 (Bekoff and Wells 1980). Bekoff and Wells (1980) noted that during 3 successive winters in Rocky Mountain National Park, Colorado, when no ungulate carrion was available, coyotes preyed on small mammals throughout the year. Mean group size for coyotes was 1.1 in summer and winter. Bowen (1978, 1981) and Bekoff and Wells (1980) concluded that when coyotes were feeding on large prey, group sizes were correspondingly larger, which increased foraging efficiency. They further observed that delayed dispersal of pups resulted in the formation of larger packs; however, seasonal variation in the diet still accounted for much of the variation in the number of pack members observed together. Messier and Barrette (1982) found no relationship between prey size and coyote group size in Quebec; they concluded that larger coyote groups could be explained by delayed dispersal of juveniles. The obj ctive of our study was to document the diet of coyotes in southeastern Colorado and examine the relationship between prey size, pack sociality, time of juvenile dispersal, and group