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Taxonomic revision of Phascogale tapoatafa (Meyer, 1793) (Dasyuridae; Marsupialia), including descriptions of two new subspecies and confirmation of P. pirata Thomas, 1904 as a ‘Top End’ endemic
The Australian Brush-tailed Phascogale ( Phascogale tapoatafa sensu lato ) has a broad but highly fragmented distribution around the periphery of the Australian continent and all populations are under significant ongoing threat to survival. A new appraisal of morphological and molecular diversity within the group reveals that the population in the ‘Top End’ of the Northern Territory is specifically distinct from all others, including those in the Kimberley region of Western Australia to the west and on Cape York of Queensland to the east. The name P. pirata Thomas, 1904 is available for the ‘Top End’ taxon. Three geographically disjunct populations are distinguished at subspecies level within P. tapoatafa on a suite of external and cranio-dental features; these are found in southeast Australia from South Australia to mid-coastal Queensland (nominotypical tapoatafa ), southwest Western Australia ( wambenger subsp. nov.), and the Kimberley region of Western Australia ( kimberleyensis subsp. nov.). A potential fourth subspecies occurs on Cape York but remains too poorly represented in collections for adequate characterization. Molecular divergence estimates based on partial sequences of the mitochondrial cytochrome b gene indicate that the range disjunction across southern Australia probably dates from the Late Pliocene, with the multiple disjunctions across northern Australia being more recent though almost certainly exceeding 400,000 years. An argument is made for the continued use of the subspecies rank in Australian mammalogy, despite a general lack of consistency in its current application.
Search path of a fossorial herbivore, Geomys bursarius , foraging in structurally complex plant communities
The influence of habitat patchiness and unpalatable plants on the search path of the plains pocket gopher ( Geomys bursarius ) was examined in outdoor enclosures. Separate experiments were used to evaluate how individual animals explored (by tunnel excavation) enclosures free of plants except for one or more dense patches of a palatable plant ( Daucus carota ), a dense patch of an unpalatable species ( Pastinaca sativa ) containing a few palatable plants ( D. carota ), or a relatively sparse mixture of palatable ( D. carota ) and unpalatable ( Raphanus sativus ) species. Only two of eight individuals tested showed the predicted pattern of concentrating search effort in patches of palatable plants. The maintenance of relatively high levels of effort in less profitable sites may reflect the security afforded food resources by the solitary social system and fossorial lifestyle of G. bursarius . Unpalatable plants repelled animals under some conditions, but search paths in the sparsely planted mixed-species treatment suggest animals can use visual or other cues to orient excavations. Evidence supporting area-restricted search was weak. More information about the use of visual cues by G. bursarius and the influence of experience on individual search mode is needed for refining current models of foraging behavior in this species.
Effects of experience and cage enrichment on predatory skills of black-footed ferrets (Mustela nigripes)
We studied ontogeny of predatory skills of growing black-footed ferrets (Mustela nigripes) raised under different captive conditions. To test effects of maturation, experience, and cage enrichment on predatory behavior, we exposed 70 juvenile black-footed ferrets to different numbers of live hamsters (Mesocricetus auratus) throughout development. Both maturation and experience increased the likelihood of a black-footed ferret making a successful kill. Black-footed ferrets exposed to greater environmental complexity (enriched cage, including encouragement of food-searching behaviors) also were more likely to kill than ferrets raised in a deprived environment. Behavioral studies of captive-raised black-footed ferrets can help refine current breeding techniques and ultimately enhance efforts to recover this endangered carnivore.
Traveling to thermal refuges during stressful temperatures leads to foraging constraints in a central-place forager
Central-place foragers can be constrained by the distance between habitats. When an organism relies on a central place for thermal refuge, the distance to food resources can potentially constrain foraging behavior. We investigated the effect of distance between thermal refuges and forage patches of the cold-intolerant marine mammal, the Florida manatee ( Trichechus manatus latirostris ), on foraging duration. We tested the alternative hypotheses of time minimization and energy maximization as a response to distance between habitats. We also determined if manatees mitigate foraging constraints with increased visits to closer thermal refuges. We used hidden Markov models to assign discrete behaviors from movement parameters as a function of water temperature and assessed the influence of distance on foraging duration in water temperatures above (> 20°C) and below (≤ 20°C) the lower critical limit of the thermoneutral zone of manatees. We found that with increased distance, manatees decreased foraging duration in cold water temperature and increased foraging duration in warmer temperatures. We also found that manatees returned to closer thermal refuges more often. Our results suggest that the spatial relationship of thermal and forage habitats can impact behavioral decisions regarding foraging. Addressing foraging behavior questions while considering thermoregulatory behavior implicates the importance of understanding changing environments on animal behavior, particularly in the face of current global change.
Genetic variation in black bears in Arkansas and Louisiana using microsatellite DNA markers
In the 1950s and 1960s, translocation projects reintroduced black bears ( Ursus americanus ) from Minnesota and Manitoba to Arkansas and Louisiana. Today, several geographically disconnected populations exist in Arkansas and Louisiana, but their origins are unclear. Some populations may represent a separate subspecies, U. a. luteolus , which is federally protected. We characterized 5 microsatellite loci in 5 isolated populations in Arkansas and Louisiana and compared them with genotypes from Minnesota. Our data indicate that bears of the Ozark and Ouachita mountains of Arkansas, an inland area of Louisiana, and those of Minnesota are similar in overall genetic diversity and allele frequencies, consistent with these populations being wholly or mostly descended from bears from the reintroduction programs. In contrast, bears from southeastern Arkansas and the coastal region of Louisiana genetically are more restricted and homogeneous. Because they exhibit a limited set of genotypes found in the other black bear populations, they represent isolated fragments of a single North American black bear population. Furthermore, genetic distance estimates indicate that the bears in southeastern Arkansas are more genetically distinct from bears in Louisiana, which are currently federally protected.
Selection of summer roosting sites by Indiana bats (Myotis sodalis) in Missouri
Summer roosting sites were studied at four maternity colonies of Indiana bats (Myotis sodalis) in northern Missouri. Colonies of Indiana bats used two types of roosts, primary and alternate, that differed in intensity of use, number, and probable function. Primary roosts were denned as roosts where use by >30 bats on more than one occasion was observed. The number of primary roosts per colony ranged from one to three. All primary roosts were in standing dead trees situated in trees exposed to direct sunlight. Alternate roosts were used by smaller numbers of bats. These roosts included both living and dead trees that typically were located within the shaded forest interior. Differences in patterns of use between types of roosts seemed to be influenced by weather conditions in that use of alternate roost trees increased during periods of elevated temperature and precipitation. Indiana bats have specific requirements for roost sites, but also must be able to relocate when loss of bark, tree fall, or other events render their current roost sites unusable. Practices of forest management within the summer range of Indiana bats should favor retention of large-diameter, mature, and senescent trees.
Spatial habitat use patterns of sea otters in coastal washington
Sea otter (Enhydra lutris kenyoni) movements, home range, and activity budgets were described from data collected during very-high-frequency radiotelemetry studies of 75 individuals on the outer coast of Washington State between 1992 and 1999. Sea otters were located at least once per week from 22 accessible sites along the coast. Over the 7-year study period, range expansion occurred from the core range north and east into the Strait of Juan de Fuca (SJF) as well as southward on the outer coast. Forty-three percent of the sea otters moved into the SJF at least once, most often in winter, using habitat that had not been occupied by sea otters since their extirpation 100 years ago. All sea otters spent portions of their time in the vicinity of Cape Alava, and many animals demonstrated consistent periodic seasonal shifts between specific portions of the coastline over several years. Ninety-five percent annual linear home ranges differed between sex and age classes. Adult males used the largest amount of coastline (50 km ?? 9 5D) and subadult females used the least (24 ?? 9 km). Both adult males and females demonstrated high seasonal periodicity in range use in summer and winter. Twenty-four-hour time budgets in the core portion of the range revealed on average sea otters spent 41% ?? 14% SD of the time foraging and 45% ?? 13% of the time resting (age and sex classes pooled). Adult and subadult female sea otters were most frequently found resting and foraging close to shore (< 1,000 m) and in shallow water (0-10 m), whereas adult and subadult males rested and foraged > 1,000 m offshore and at depths between 10 and 30 m. Given current rates of population growth and observed mobility, sea otters in Washington have high potential for range expansion into unoccupied habitat such as Grays Harbor, Willapa Bay, the SJF, or along Vancouver Island. ?? 2009 American Society of Mammalogists.
Mitochondrial-DNA variation among subspecies and populations of sea otters (Enhydra lutris)
We used restriction-enzyme analysis of polymerase-chain reaction-amplified, mitochondrial DNA (mtDNA) to assess genetic differentiation of subspecies and populations of sea otters, Enhydra lutris , throughout the range of the species. There were several haplotypes of mtDNA in each subspecies and geographically separate populations. MtDNA sequence divergence of haplotypes of sea otters was 0.0004–0.0041 base substitutions per nucleotide. E. L nereis appears to have monophyletic mitochondrial DNA, while E. I. lutris and E. I. kenyoni do not. Different frequencies of haplotypes of mtDNA among populations reflect current restriction of gene flow and the unique histories of different populations. There are two or three haplotypes of mtDNA and diversity of haplotypes is 0.1376–0.5854 in each population of otters. This is consistent with theoretical work, which suggests that population bottlenecks of sea otters probably did not result in major losses of genetic variation for individual populations, or the species as a whole.
Ecogeographic variation and taxonomic boundaries in Large Treeshrews (Scandentia, Tupaiidae: Tupaia tana Raffles, 1821) from Southeast Asia
The Large Treeshrew, Tupaia tana Raffles, 1821, is a small mammal (~205 g) from Southeast Asia with a complicated taxonomic history. Currently, 15 subspecies are recognized from Borneo, Sumatra, and smaller islands, and many were originally differentiated based on minor pelage differences and small sample sizes. We explored intraspecific variation in T. tana using quantitative osteological data obtained from the hands and skulls of museum specimens. Multivariate analyses reveal extensive overlap among T. tana populations in morphospace, indicating that the majority of currently recognized subspecies are not morphometrically distinct. In contrast, the separation between Bornean and Sumatran populations of T. tana is sufficient to recognize them as different subspecies. Comparisons of Bornean specimens to those on small, offshore islands reveal that the latter average smaller body size. This pattern is inconsistent with Foster’s island rule, which predicts that island populations of small mammals (< 5 kg) will average larger body size relative to mainland forms. A similar lack of support for ecogeographic rules has been noted in T. glis (Diard, 1820), suggesting that these “rules” are poor predictors of geographic variation in treeshrews.
Ambiguities in using telomere length for age determination in two North American bat species
The age of an animal, determined by time (chronological age) as well as genetic and environmental factors (biological age), influences the likelihood of mortality and reproduction and thus the animal’s contribution to population growth. For many long-lived species, such as bats, a lack of external and morphological indicators has made determining age a challenge, leading researchers to examine genetic markers of age for application to demographic studies. One widely studied biomarker of age is telomere length, which has been related both to chronological and biological age across taxa, but only recently has begun to be studied in bats. We assessed telomere length from the DNA of known-age and minimum known-age individuals of two bat species using a quantitative PCR assay. We determined that telomere length was quadratically related to chronological age in big brown bats ( Eptesicus fuscus ), although it had little predictive power for accurate age determination of unknown-age individuals. The relationship was different in little brown bats ( Myotis lucifugus ), where telomere length instead was correlated with biological age, apparently due to infection and wing damage associated with white-nose syndrome. Furthermore, we showed that wing biopsies currently are a better tissue source for studying telomere length in bats than guano and buccal swabs; the results from the latter group were more variable and potentially influenced by storage time. Refinement of collection and assessment methods for different non-lethally collected tissues will be important for longitudinal sampling to better understand telomere dynamics in these long-lived species. Although further work is needed to develop a biomarker capable of determining chronological age in bats, our results suggest that biological age, as reflected in telomere length, may be influenced by extrinsic stressors such as disease.
Taxonomic status of Myotis occultus
The taxonomic status of the Arizona myotis ( Myotis occultus ) is uncertain. Although the taxon was described as a distinct species and currently is regarded as such by some authors, others have noted what they interpreted as intergradation with the little brown bat ( M. lucifugus carissima ) near the Colorado-New Mexico state line. In this study, we used protein electrophoresis to compare bats of these nominal taxa. We examined 20 loci from 142 specimens referable to M. occultus and M. lucifugus from New Mexico, Colorado, and Wyoming. Nine of the 20 loci were polymorphic. Results show that there were high similarities among samples, no fixed alleles, and minor divergence from Hardy-Weinberg equilibrium. Our results suggest that the two nominal taxa represent only one species and that M. occultus should be regarded as a subspecies of M. lucifugus .
Temporal-spatial distribution of American bison (Bison bison) in a tallgrass prairie fire mosaic
Fire and bison (Bison bison) are thought to be historically responsible for shaping prairie vegetation in North America. Interactions between temporal-spatial distributions of bison and prescribed burning protocols are important in current restoration of tallgrass prairies. We examined dynamics of bison distribution in a patch-burned tallgrass prairie in the south-central United States relative to bison group size and composition, and burn age and temporal distribution. Bison formed larger mixed groups during summer and smaller sexually segregated groups the rest of the year, and bison selected dormant-season burn patches in the 1st posture growing season most often during spring and summer. Large bison herds selecting recently burned areas resulted in seasonally variable and concentrated grazing pressure that may substantially alter site-specific vegetation. These dynamics must be considered when reintroducing bison and fire into tallgrass prairie because variable outcomes of floral richness and structural complexity are likely depending on temporal-spatial distribution of bison. ?? 2006 American Society of Mammalogists.
Killer whales and marine mammal trends in the North Pacific - A re-examination of evidence for sequential megafauna collapse and the prey-switching hypothesis
Springer et al. (2003) contend that sequential declines occurred in North Pacific populations of harbor and fur seals, Steller sea lions, and sea otters. They hypothesize that these were due to increased predation by killer whales, when industrial whaling's removal of large whales as a supposed primary food source precipitated a prey switch. Using a regional approach, we reexamined whale catch data, killer whale predation observations, and the current biomass and trends of potential prey, and found little support for the prey-switching hypothesis. Large whale biomass in the Bering Sea did not decline as much as suggested by Springer et al., and much of the reduction occurred 50-100 yr ago, well before the declines of pinnipeds and sea otters began; thus, the need to switch prey starting in the 1970s is doubtful. With the sole exception that the sea otter decline followed the decline of pinnipeds, the reported declines were not in fact sequential. Given this, it is unlikely that a sequential megafaunal collapse from whales to sea otters occurred. The spatial and temporal patterns of pinniped and sea otter population trends are more complex than Springer et al. suggest, and are often inconsistent with their hypothesis. Populations remained stable or increased in many areas, despite extensive historical whaling and high killer whale abundance. Furthermore, observed killer whale predation has largely involved pinnipeds and small cetaceans; there is little evidence that large whales were ever a major prey item in high latitudes. Small cetaceans (ignored by Springer et al.) were likely abundant throughout the period. Overall, we suggest that the Springer et al. hypothesis represents a misleading and simplistic view of events and trophic relationships within this complex marine ecosystem. ?? 2007 by the Society for Marine Mammalogy.
Individual variation in space use by female spotted hyenas
Large carnivores range more widely than many other terrestrial mammals, and this behavior tends to bring them into frequent conflict with humans. Within any carnivore population, individual variation in patterns of space use should be expected to make some animals more vulnerable than others to risks of mortality from humans and other sources. In this study, our goal was to document variation among individuals in space use by female spotted hyenas ( Crocuta crocuta ). We examined predictions of hypotheses suggesting that space use by female hyenas is affected by reproductive state, social rank, and local prey abundance. Home-range size, distance at which females were found from the current communal den, and distance at which they were found from the nearest territorial boundary all varied significantly with the 3 independent variables. Females with den-dwelling cubs had smaller home ranges, were found closer to the communal den, and were found farther from the territorial boundary than were females with no den-dwelling cubs. Neither social rank nor prey availability significantly influenced the space-use patterns of females with den-dwelling cubs. Among females with no den-dwelling cubs, high-ranking females had smaller home ranges, were closer to the communal den, and were farther from the territorial boundary than were low-ranking females. The females ranging most widely were low-ranking individuals with no den-dwelling cubs when they were observed during periods of prey scarcity.
Climatic and density influences on recruitment in an irruptive population of Roosevelt elk
Current paradigms of ungulate population ecology recognize that density-dependent and independent mechanisms are not always mutually exclusive. Long-term data sets are necessary to assess the relative strength of each mechanism, especially when populations display irruptive dynamics. Using an 18-year time series of population abundances of Roosevelt elk ( Cervus elaphus roosevelti ) inhabiting Redwood National Park in northwestern California we assessed the influence of population size and climatic variation on elk recruitment and whether irruptive dynamics occurred. An information-theoretic model selection analysis indicated that abundance lagged 2 years and neither climatic factors nor a mix of abundance and climatic factors influenced elk recruitment. However, density-dependent recruitment differed between when the population was declining and when the population increased and then stabilized at an abundance lower than at the start of the decline. The population displayed irruptive dynamics.
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.
Predicting niche spaces of expanding Evening Bat (Nycticeius humeralis) populations following white-nose syndrome establishment
Observations of the Evening Bat ( Nycticeius humeralis ), a species historically distributed abundantly throughout the southeastern United States, are speculated to have increased in the Midwest. One hypothesis for this expansion in geographic distribution is that local extirpations of other bat species resulted in the expanded realized niche spaces for evening bats. In Indiana, such niche spaces may have been created by declines in populations of the Northern Long-eared Bat ( Myotis septentrionalis ), Indiana Bat ( M. sodalis ), Little Brown Bat ( M. lucifugus ), Big Brown Bat ( Eptesicus fuscus ), and the Tri-colored Bat ( Perimyotis subflavus ) due to white-nose syndrome (WNS). Our goal was to estimate the occupancy of Evening Bat in Indiana post-WNS establishment relative to the occupancy of other bat species before significant population declines caused by WNS. We expected that indices of occupancy of nearly extirpated species pre-WNS establishment would best predict current Evening Bat observations, and this would elucidate the niche space evening bats are now filling. We hypothesized that Evening Bat populations may be expanding their geographic range due to compensatory community dynamics, and that their realized niche space may be expanding in part due to losses of other bat species from WNS. We constructed multi-season Bayesian occupancy models using informative priors and integrative prior knowledge to test our predictions. We found that evening bats are occupying the niche space they were already occupying pre-WNS establishment. Furthermore, our results indicate that evening bats may be filling the niche space left behind by Myotis spp. ( M. sodalis and M. lucifigus ). These results can help us understand the dynamics of bat communities in a post-WNS establishment landscape and may also help to inform conservation of imperiled Myotis species.