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Habitat selection by a focal predator (Canis lupus) in a multiprey ecosystem of the northern Rockies

Large predators respond to land cover and physiography that maximize the likelihood of encountering prey. Using locations from global positioning system-collared wolves (Canis lupus), we examined whether land cover, vegetation productivity or change, or habitat-selection value for ungulate prey species themselves most influenced patterns of selection by wolves in a large, intact multiprey system of northern British Columbia. Selection models based on land cover, in combination with topographical features, consistently outperformed models based on indexes of vegetation quantity and quality (using normalized difference vegetation index) or on selection value to prey species (moose [Alces americanus], elk [Cervus elaphus], woodland caribou [Rangifer tarandus], and Stone's sheep [Ovis dalli stonei]). Wolves generally selected for shrub communities and high diversity of cover across seasons and avoided conifer stands and non-vegetated areas and west aspects year-round. Seasonal selection strategies were not always reflected in use patterns, which showed highest frequency of use in riparian, shrub, and conifer classes. Patterns of use and selection for individual wolf packs did not always conform to global models, and appeared related to the distribution of land cover and terrain within respective home ranges. Our findings corroborate the biological linkages between wolves and their habitat related to ease of movement and potential prey associations. ?? American 2011 Society of Mammalogists.

Journal of Mammalogy

Estimating detection and density of the Andean cat in the high Andes

The Andean cat ( Leopardus jacobita ) is one of the most endangered, yet least known, felids. Although the Andean cat is considered at risk of extinction, rigorous quantitative population studies are lacking. Because physical observations of the Andean cat are difficult to make in the wild, we used a camera-trapping array to photo-capture individuals. The survey was conducted in northwestern Argentina at an elevation of approximately 4,200 m during October-December 2006 and April-June 2007. In each year we deployed 22 pairs of camera traps, which were strategically placed. To estimate detection probability and density we applied models for spatial capture-recapture using a Bayesian framework. Estimated densities were 0.07 and 0.12 individual/km 2 for 2006 and 2007, respectively. Mean baseline detection probability was estimated at 0.07. By comparison, densities of the Pampas cat ( Leopardus colocolo ), another poorly known felid that shares its habitat with the Andean cat, were estimated at 0.74–0.79 individual/km 2 in the same study area for 2006 and 2007, and its detection probability was estimated at 0.02. Despite having greater detectability, the Andean cat is rarer in the study region than the Pampas cat. Properly accounting for the detection probability is important in making reliable estimates of density, a key parameter in conservation and management decisions for any species.

High Andes

Mortality of Siberian polecats and black-footed ferrets released onto prairie dog colonies

Black-footed ferrets ( Mustela nigripes ) likely were extirpated from the wild in 1985–1986, and their repatriation depends on captive breeding and reintroduction. Postrelease survival of animals can be affected by behavioral changes induced by captivity. We released neutered Siberian polecats ( M. eversmanii ), close relatives of ferrets, in 1989–1990 on black-tailed prairie dog ( Cynomys ludovicianus ) colonies in Colorado and Wyoming initially to test rearing and reintroduction techniques. Captive-born polecats were reared in cages or cages plus outdoor pens, released from elevated cages or into burrows, and supplementally fed or not fed. We also translocated wild-born polecats from China in 1990 and released captive-born, cage-reared black-footed ferrets in 1991, the 1st such reintroduction of black-footed ferrets. We documented mortality for 55 of 92 radiotagged animals in these studies, mostly due to predation (46 cases). Coyotes ( Canis latrans ) killed 31 ferrets and polecats. Supplementally fed polecats survived longer than nonprovisioned polecats. With a model based on deaths per distance moved, survival was highest for wild-born polecats, followed by pen-experienced, then cage-reared groups. Indexes of abundance (from spotlight surveys) for several predators were correlated with mortality rates of polecats and ferrets due to those predators. Released black-footed ferrets had lower survival rates than their ancestral population in Wyoming, and lower survival than wild-born and translocated polecats, emphasizing the influence of captivity. Captive-born polecats lost body mass more rapidly postrelease than did captive-born ferrets. Differences in hunting efficiency and prey selection provide further evidence that these polecats and ferrets are not ecological equivalents in the strict sense.

Journal of Mammalogy

Postbreeding resource selection by adult black-footed ferrets in the Conata Basin, South Dakota

We investigated postbreeding resource selection by adult black-footed ferrets ( Mustela nigripes ) on a 452-ha black-tailed prairie dog ( Cynomys ludovicianus ) colony in the Conata Basin of South Dakota during 2007–2008. We used resource selection functions (RSFs) to evaluate relationships between numbers of ferret locations and numbers of prairie dog burrow openings (total or active), distances to colony edges, and connectivity of patches of burrow openings. In both years ferrets selected areas near edges of the prairie dog colony where active burrow openings were abundant. In the interior of the colony ferrets selected areas with low abundance of active burrow openings. At times, prairie dog productivity (i.e., pup abundance) might be greatest at colony edges often characterized by grasses; ferrets are likely to select areas where refuge and vulnerable prey are abundant. Ferrets could have used interior areas with few active burrow openings as corridors between edge areas with many active burrow openings. Also, in areas with few active burrow openings ferrets spend more time aboveground during movements and, thus, are likely to be more easily detected. These results complement previous studies demonstrating importance of refuge and prey in fine-scale resource selection by ferrets and provide insight into factors that might influence edge effects on ferret space use. Conservation and restoration of colonies with areas with high densities of burrow openings and prairie dogs, and corridors between such areas, are needed for continued recovery of the black-footed ferret. RSFs could complement coarse-scale habitat evaluations by providing finer-scale assessments of habitat for the black-footed ferret.

Journal of Mammalogy

Black-footed ferret areas of activity during late summer and fall at Meeteetse, Wyoming

Radiotelemetry was used during 1983 and 1984 to collect information on short-term areas of activity for black-footed ferrets ( Mustela nigripes ) near Meeteetse, Wyoming. This population ultimately provided ferrets for the captive-breeding program that bred and released offspring into the wild since 1991. We fitted 5 adult ferrets and 13 juveniles with radiotransmitters and followed their movements during late summer and fall. Adult males had 7-day areas of activity that were >6 times as large as those of adult females. Activity areas of adult males varied little in coverage or location on a weekly basis, but females sequentially shifted their areas. Unlike juvenile females, juvenile males tended to leave their natal colonies.

Wyoming

Movements and survival of black-footed ferrets associated with an experimental translocation in South Dakota

Black-footed ferrets ( Mustela nigripes ) apparently were extirpated from all native habitats by 1987, and their repatriation requires a combination of captive breeding, reintroductions, and translocations among sites. Improvements in survival rates of released ferrets have resulted from experience in quasi-natural environments during their rearing. Reestablishment of a self-sustaining wild population by 1999 provided the 1st opportunity to initiate new populations by translocating wild-born individuals. Using radiotelemetry, we compared behaviors and survival of 18 translocated wild-born ferrets and 18 pen-experienced captive-born ferrets after their release into a prairie dog colony not occupied previously by ferrets. Translocated wild-born ferrets moved significantly less and had significantly higher short-term survival rates than their captive-born counterparts. Using mark–recapture methods, we also assessed potential impacts to the established donor population of removing 37% of its estimated annual production of kits. Annual survival rates for 30 ferret kits remaining at the donor subcomplex were higher than rates for 54 ferret kits at the control subcomplex (unmanipulated) for males (+82%) and females (+32%). Minimum survival of translocated kits did not differ significantly from survival of those at the control subcomplex. Direct translocation of young, wild-born ferrets from site to site appears to be an efficient method to establish new populations.

South Dakota

Conflicting research on the demography, ecology, and social behavior of Gunnison's prairie dogs ( Cynomys gunnisoni )

Gunnison's prairie dogs ( Cynomys gunnisoni ) are rare, diurnal, colonial, burrowing, ground-dwelling squirrels. Studies of marked individuals living under natural conditions in the 1970s, 1980s, and 1990s showed that males are heavier than females throughout the year; that adult females living in the same territory are consistently close kin; and that females usually mate with the sexually mature male(s) living in the home territory. Research from 2007 through 2010 challenges all 3 of these findings. Here we discuss how different methods might have led to the discrepancies.

Journal of Mammalogy

Evaluating interactions between river otters and muskrats at bridge crossings in Kentucky

hreatened or endangered. Muskrat populations have been reduced in some streams where North American river otters (Lontra canadensis) were reintroduced, and it has been hypothesized that otter reintroduction could be used as a tool for conservation of mussels. We used occupancy estimation methods to evaluate the ecological relationship between muskrats and otters by collecting presence–absence data based on field sign found at bridge crossings in eastern and central Kentucky. Mean detection probabilities (ps) and occupancy probabilities (ψs) for muskrats were 0.692 (SE = 0.045) and 0.723 (SE = 0.071) and for otters were 0.623 (SE = 0.036) and 0.662 (SE = 0.069), respectively. Otter occupancy was related negatively to distance from release sites, which suggests that the otter population is still expanding its range. A 2-species interaction model indicated that the occupancy by muskrats and river otters was independent, and we conclude that river otter reintroduction would not be an effective strategy for conserving mussels.

Journal of Mammalogy

Roost selection by western long-eared myotis (Myotis evotis) in burned and unburned piñon–juniper woodlands of southwestern Colorado

All 16 species of bats known to occur in western Colorado are found at Mesa Verde National Park (MVNP) in the southwestern United States. Since 1996, wildfires have burned more than 70% of MVNP (> 15,000 ha), potentially altering food and roosting resources for bats. During the summers of 2006–2007, we investigated roost use by reproductive female western long-eared myotis ( Myotis evotis ). We located 33 bat roosts in rock crevices and 1 in a juniper snag. All but 2 of the roosts were in unburned habitat. Bats roosted alone or in small groups (≤3 individuals) and switched roosts frequently (1–7 roosts per bat, median = 1.5 roosts per bat, SE = 0.5 roosts per bat). We compared occupied roosts with randomly selected unoccupied crevices and used an information theoretic approach to determine which variables were most important in determining roost use at microhabitat and landscape scales. At the microhabitat scale, maternity roosts were higher above the ground and deeper than random, unoccupied rock crevices. At the landscape scale, roosts were closer to water and farther from burned habitat than random crevices, providing reproductive female M. evotis with the best opportunities to drink and forage for insects. Tree roosts are apparently not a vital resource for reproductive female M. evotis during the summer months at our study site, presumably because of the extensive availability of rock crevices. Understanding site-specific roosting behavior is important for proper management of bat populations because differences can exist between geographic regions, even among areas with similar plant communities.

Colorado

Diversity of nitrogen isotopes and protein status in caribou: implications for monitoring northern ungulates

Nutritional condition is an important determinant of productivity and survival in caribou (Rangifer tarandus). We used samples of excreta (n = 1,150) to estimate diet composition from microhistology and 2 isotopic proxies of protein status for 2 ecotypes of caribou in 4 herds in late winter (2006–2008). Isotopes of nitrogen (δ 15 N in parts per thousand [‰]) from excreta samples (urea, diet, and body N) were used to estimate indexes of protein status: the proportion of urea N derived from body N (p-UN) and the difference between the δ15N of the body and urinary urea (Δbody-urea). We examined dietary and terrain characteristics, δ 15 N, p-UN, and Δ body-urea by ecotype, herd, year, and foraging site. Multiple regression and an information-theoretic approach were used to evaluate correlates of protein status at each foraging site. The dietary and terrain characteristics of foraging sites did not vary by ecotype or herd (P > 0.108); diets were dominated by lichens (68% ± 14.1% SD). The δ 15 N of urea, diet, body N, p-UN, and Δ body-urea varied among foraging sites within each herd (P < 0.001). Although highly variable, the δ 15 N of urinary urea was typically low (−4.68‰ ± 2.67‰ SD). Dietary N also had low δ 15 N (−4.18‰ ± 0.92‰ SD), whereas body N was generally heavier in 15 N (2.20‰ ± 1.56‰ SD) than urinary urea or the diet. Both measures of protein status were similarly diverse between ecotypes and among herds, which limited their applicability to monitor protein status at the population level. Although we observed limitations to interpreting estimates of p-UN from highly vagile ungulates, the Δbody-urea may prove to be a useful index of protein status at smaller spatial and temporal scales. Indeed, a portion of the observed variance (r 2 = 0.26) in Δbody-urea at each foraging site was explained by the proportion of shrubs in the winter diet. There remains potential in using δ 15 N in excreta as a noninvasive tool for evaluating protein status in northern ungulates; however, considerable analytical and sampling challenges remain for applying these isotopic approaches at large scales.

Alaska

A new species of Lophostoma d'Orbigny, 1836 (Chiroptera: Phyllostomidae) from Panama

We report the discovery of a new species of Lophostoma from Panama, which we name L. kalkoae . This new species resembles L. carrikeri and L. yasuni in possessing a white venter, but is distinguishable from both by external and cranial characteristics. The new species is similar in size to L. carrikeri and L. schulzi . Lophostoma sp. nov. can be most easily recognized by its combination of white venter, postauricular patches connected by a thin line of pale hair to the white fur on the chest, elongated clitoris and swollen labia, less strongly developed lateral projection of mastoid processes, well-marked indentation on the lingual cingulum of the upper canine, well-developed P3, well-developed posterior lingual cusp on the cingulum of P4, and parastyle absent on M1 and M2. We present a dichotomous key for the genus Lophostoma and a map showing all the localities where white-bellied Lophostoma have been recorded.

Journal of Mammalogy

Using hand proportions to test taxonomic boundaries within the Tupaia glis species complex (Scandentia, Tupaiidae)

Treeshrews (order Scandentia) comprise 2 families of squirrel-sized terrestrial, arboreal, and scansorial mammals distributed throughout much of tropical South and Southeast Asia. The last comprehensive taxonomic revision of treeshrews was published in 1913, and a well-supported phylogeny clarifying relationships among all currently recognized extant species within the order has only recently been published. Within the family Tupaiidae, 2 widely distributed species, the northern treeshrew, Tupaia belangeri (Wagner, 1841), and the common treeshrew, T. glis (Diard, 1820), represent a particularly vexing taxonomic complex. These 2 species are currently distinguished primarily based on their respective distributions north and south of the Isthmus of Kra on the Malay Peninsula and on their different mammae counts. This problematic species complex includes 54 published synonyms, many of which represent putative island endemics. The widespread T. glis and T. belangeri collectively comprise a monophyletic assemblage representing the sister lineage to a clade composed of the golden-bellied treeshrew, T. chrysogaster Miller, 1903 (Mentawai Islands), and the long-footed treeshrew, T. longipes (Thomas, 1893) (Borneo). As part of a morphological investigation of the T. glis – T. belangeri complex, we studied the proportions of hand bones, which have previously been shown to be useful in discriminating species of soricids (true shrews). We measured 38 variables from digital X-ray images of 148 museum study skins representing several subspecies of T. glis , T. belangeri , T. chrysogaster , and T. longipes and analyzed these data using principal components and cluster analyses. Manus proportions among these 4 species readily distinguish them, particularly in the cases of T. chrysogaster and T. longipes . We then tested the distinctiveness of several of the populations comprising T. glis and T. longipes . T. longipes longipes and T. l. salatana Lyon, 1913, are distinguishable from each other, and populations of T. "glis" from Bangka Island and Sumatra are distinct from those on the Malay Peninsula, supporting the recognition of T. salatana , T. discolor Lyon, 1906, and T. ferruginea Raffles, 1821 as distinct species in Indonesia. These relatively small, potentially vulnerable treeshrew populations occur in the Sundaland biodiversity hotspot and will require additional study to determine their appropriate conservation status.

Journal of Mammalogy

Spatial genetic structure and asymmetrical gene flow within the Pacific walrus

Pacific walruses ( Odobenus rosmarus divergens ) occupying shelf waters of Pacific Arctic seas migrate during spring and summer from 3 breeding areas in the Bering Sea to form sexually segregated nonbreeding aggregations. We assessed genetic relationships among 2 putative breeding populations and 6 nonbreeding aggregations. Analyses of mitochondrial DNA (mtDNA) control region sequence data suggest that males are distinct among breeding populations (&Phi; ST =0.051), and between the eastern Chukchi and other nonbreeding aggregations (&Phi; ST =0.336–0.449). Nonbreeding female aggregations were genetically distinct across marker types (microsatellite F ST =0.019; mtDNA &Phi; ST =0.313), as was eastern Chukchi and all other nonbreeding aggregations (microsatellite F ST =0.019–0.035; mtDNA &Phi; ST =0.386–0.389). Gene flow estimates are asymmetrical from St. Lawrence Island into the southeastern Bering breeding population for both sexes. Partitioning of haplotype frequencies among breeding populations suggests that individuals exhibit some degree of philopatry, although weak. High levels of genetic differentiation among eastern Chukchi and all other nonbreeding aggregations, but considerably lower genetic differentiation between breeding populations, suggest that at least 1 genetically distinct breeding population remained unsampled. Limited genetic structure at microsatellite loci between assayed breeding areas can emerge from several processes, including male-mediated gene flow, or population admixture following a decrease in census size (i.e., due to commercial harvest during 1880–1950s) and subsequent recovery. Nevertheless, high levels of genetic diversity in the Pacific walrus, which withstood prolonged decreases in census numbers with little impact on neutral genetic diversity, may reflect resiliency in the face of past environmental challenges.

Journal of Mammalogy

Taxonomic status and relationships of Sorex obscurus parvidens Jackson, 1921, from California

The San Bernardino shrew, Sorex obscurus parvidens Jackson, 1921, is a population inhabiting the San Bernardino and San Gabriel mountains of southern California. For the past 9 decades, this population has been considered either a subspecies of S. obscurus Merriam, 1895, S. vagrans Baird, 1857, or S. monticola Merriam, 1890; or an undifferentiated population of S. ornatus Merriam, 1895. Aside from the changing taxonomic landscape that contextualizes the genus Sorex, previous study of S. obscurus parvidens has been retarded by the perception of limited available samples (typically, fewer than 8 specimens); misinterpretation of the provenance of specimens identified as S. obscurus parvidens; misunderstanding of the type locality; and inclusion of specimens of this taxon in the type series of another species with which S. obscurus parvidens has been both contrasted and allied at different times. My investigation of S. obscurus parvidens indicates that it is a distinctive population that is morphologically closest to S. ornatus, and it corresponds to the Southern Clade of that species. However, the appropriate names for deep clades within S. ornatus remain uncertain. Until this uncertainty is resolved, S. obscurus parvidens should be considered a distinctive population within S. ornatus; for conservation purposes, it should be recognized as S. ornatus parvidens.

California

Population-level impact of white-nose syndrome on the endangered Indiana bat

Establishing status and trend for an endangered species is critical to recovery, especially when it is faced with a nascent extinction agent. We calculated, with hierarchical log-linear change-point models, hibernaculum-level population trends between 1983 and 2009 for the endangered Indiana bat ( Myotis sodalis ) now subjected to the fast-spreading fungal disease white-nose syndrome. We combined trends from 222 wintering populations before and after onset of the disease to determine trend for clusters of interacting wintering populations, recovery units, and the species. Before onset of the disease, a west-to-east gradient in trends existed, with westernmost populations declining and easternmost populations increasing in abundance. The species as a whole, however, was stationary between 1983 and 2005 (-0.5% mean annual change; 95% confidence interval [ CI ] = -2.8, +1.8%). Estimated mean population size in 2009 was 377,124 bats (195,398-957,348), with large variance apparently caused by white-nose syndrome. With the onset of white-nose syndrome (2006-2009), the species exhibited a 10.3% annual decline (95% CI = -21.1, +2.0%). White-nose syndrome is having an appreciable influence on the status and trends of Indiana bat populations, stalling and in some cases reversing population gains made in recent years.

Journal of Mammalogy

Interface between black-footed ferret research and operational conservation

Questions and problems that emerged during operational conservation of black-footed ferrets ( Mustela nigripes ) have been addressed by a wide variety of studies. Early results from such studies often were communicated orally during meetings of recovery groups and in written form using memoranda, unpublished reports, and theses. Typically, implementation of results preceded their publication in widely distributed journals. Many of these studies eventually were published in journals, and we briefly summarize the contents of 8 volumes and special features of journals that have been dedicated to the biology of ferrets and issues in ferret recovery. This year marks the 30th anniversary of rediscovery of the black-footed ferret, and the 7 papers of the following Special Feature summarize data collected over nearly that span of time.

Journal of Mammalogy

Evidence of cryptic individual specialization in an opportunistic insectivorous bat

Habitat use and feeding behaviors of cryptic animals are often poorly understood. Analyses of stable isotope ratios in animal body tissues can help reveal an individual's location and resource use during tissue growth. We investigated variation in stable isotope ratios of 4 elements (H, C, N, and S) in the hair of a sedentary species of insectivorous bat (Eptesicus fuscus) inhabiting a chemically complex urban landscape. Our objective was to quantify population-level isotopic variation and test for evidence of resource specialization by individuals. Bats were sampled over 3 annual molt cycles at maternity roosts in buildings and variance components analysis was used to test whether intraindividual isotopic variation among molts differed from interindividual variation, after controlling for year and roost-group effects. Consistent with prior evidence that E. fuscus is opportunistic in its habitat use and foraging at the population level, we observed wide population-level variation for all isotopes. This variation likely reflects the chemical complexity of the urban landscape studied. However, isotopic variation among years within marked individuals was lower than variation among marked individuals within year for all isotopes, and carbon signatures indicated resource specialization by roost groups and individuals. This is the 1st study to examine variation in stable isotope ratios of individual wild bats over multiple years. Although our results suggest this population tends toward opportunistic habitat use or prey selection, or both, during molt periods, results also indicate that individuals and groups of bats composing the population might be habitat or dietary specialists—a novel finding for insectivorous bats.

Journal of Mammalogy

Tools for quantifying isotopic niche space and dietary variation at the individual and population level.

Ecologists are increasingly using stable isotope analysis to inform questions about variation in resource and habitat use from the individual to community level. In this study we investigate data sets from 2 California sea otter ( Enhydra lutris nereis ) populations to illustrate the advantages and potential pitfalls of applying various statistical and quantitative approaches to isotopic data. We have subdivided these tools, or metrics, into 3 categories: IsoSpace metrics, stable isotope mixing models, and DietSpace metrics. IsoSpace metrics are used to quantify the spatial attributes of isotopic data that are typically presented in bivariate (e.g., δ 13 C versus δ 15 N) 2-dimensional space. We review IsoSpace metrics currently in use and present a technique by which uncertainty can be included to calculate the convex hull area of consumers or prey, or both. We then apply a Bayesian-based mixing model to quantify the proportion of potential dietary sources to the diet of each sea otter population and compare this to observational foraging data. Finally, we assess individual dietary specialization by comparing a previously published technique, variance components analysis, to 2 novel DietSpace metrics that are based on mixing model output. As the use of stable isotope analysis in ecology continues to grow, the field will need a set of quantitative tools for assessing isotopic variance at the individual to community level. Along with recent advances in Bayesian-based mixing models, we hope that the IsoSpace and DietSpace metrics described here will provide another set of interpretive tools for ecologists.

Journal of Mammalogy