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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.
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.
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.
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.
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.
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.
Mitochondrial phylogeography of moose (Alces alces) in North America
Nucleotide variation was assessed from the mitochondrial control region of North American moose ( Alces alces ) to test predictions of a model of range expansion by stepping-stone dispersal and to determine whether patterns of genetic variation support the current recognition of 4 subspecies. Haplotypes formed a star phylogeny indicative of a recent expansion of populations. Values of nucleotide and haplotype diversity were low continentwide but were greatest in the central part of the continent and lowest in peripheral populations. Despite low mitochondrial diversity, moose exhibited a high degree of differentiation regionally, which was not explained by isolation by distance. Our data indicate a pattern of colonization consistent with a large central population that supplied founders to peripheral populations (other than Alaska), perhaps through rare, long-distance dispersal events (leptokurtic dispersal) rather than mass dispersal by a stepping-stone model. The colonization scenario does not account for the low haplotype diversity observed in Alaska, which may be derived from a postcolonization bottleneck. Establishment of peripheral populations by leptokurtic dispersal and subsequent local adaptation may have been sufficient for development of morphological differentiation among extant subspecies.
Population estimates of Antillean manatees in Puerto Rico: An analytical framework for aerial surveys using multi-pass removal sampling
Effective management of the threatened Antillean manatee ( Trichechus manatus manatus ) in Puerto Rico requires reliable estimates of population size. Estimates are needed to assess population responses to management actions, and whether recovery objectives have been met. Aerial surveys have been conducted since 1976, but none adjusted for imperfect detection. We summarize surveys since 1976, report on current distribution, and provide population estimates after accounting for apparent detection probability for surveys between June 2010 and March 2014. Estimates in areas of high concentration (hotspots) averaged 317 ± 101, three times higher than unadjusted counts (104 ± 0.56). Adjusted estimates in three areas outside hotspots also differed markedly from counts (75 ± 9.89 versus 19.5 ± 3.5). Average minimum island-wide estimate was 386 ± 89, similar to the maximum estimate of 360 suggested in 2005, but fewer than the 700 recently suggested by the Puerto Rico Manatee Conservation Center. Manatees were more widespread than previously understood. Improving estimates, locally or island-wide, will require stratifying the island differently and greater knowledge about factors affecting detection probability. Sharing our protocol with partners in nearby islands (e.g., Cuba, Jamaica, Hispaniola), whose populations share genetic make-up, would contribute to enhanced regional conservation through better population estimates and tracking range expansion.
Taxonomic boundaries in Lesser Treeshrews (Scandentia, Tupaiidae: Tupaia minor)
The Lesser Treeshrew, Tupaia minor Günther, 1876 , is a small mammal from Southeast Asia with four currently recognized subspecies: T. m. minor from Borneo; T. m. malaccana from the Malay Peninsula; T. m. humeralis from Sumatra; and T. m. sincepis from Singkep Island and Lingga Island. A fifth subspecies, T. m. caedis , was previously synonymized with T. m. minor ; it was thought to occur in northern Borneo and on the nearby islands of Banggi and Balambangan. These subspecies were originally differentiated based on pelage color, a plastic feature that has proven to be an unreliable indicator of taxonomic boundaries in treeshrews and other mammals. To explore infraspecific variation among T. minor populations across the Malay Peninsula, Borneo, Sumatra, and smaller islands, we conducted multivariate analyses of morphometric data collected from the hands and skulls of museum specimens. Principal component and discriminant function analyses reveal limited differentiation in manus and skull proportions among populations of T. minor from different islands. We find no morphometric support for the recognition of the four allopatric subspecies and no support for the recognition of T. m. caedis as a separate subspecies on Borneo.
Disentangling genetic diversity of Myotis septentrionalis: population structure, demographic history, and effective population size
Myotis septentrionalis (Northern Long-eared Bat) has recently suffered a >90% decline in population size in North America due to white-nose syndrome (WNS). We assessed genetic diversity, population structure, current effective population size, and demographic history of M. septentrionalis distributed across the United States to determine baseline levels pre-WNS. We analyzed RADseq data from 81 individuals from Kentucky, Louisiana, Michigan, Minnesota, North Carolina, Oklahoma, and Wisconsin. Additionally, we examined population genetic structure using discriminant analysis of principal components, fastStructure, and STRUCTURE. We then estimated effective population size and demographic history using fastsimcoal2. Similar levels of genetic diversity were found across all samples. We found no population genetic structure in the varied analyses from these contemporary samples. The best model for demographic history estimated a rapid population expansion followed by a slower expansion approximately 340,000 years ago. The vagility of M. septentrionalis , along with male dispersal and random mating, may provide a buffer against serious bottleneck effects stemming from rapid population declines due to WNS. This research provides a baseline for tracking and monitoring the influence of WNS on genetic diversity such as potential reduced diversity or increased population structuring in the future.
Seasonal movements and Home-range use by female pronghorns in sagebrush-steppe communities of western South Dakota
Knowledge of seasonal movements by pronghorns ( Antilocapra americana ) within the easternmost extension of sagebrush-steppe communities is limited. Current hypotheses regarding movement patterns suggest that pronghorns initiate seasonal movements in response to severe winter weather, snowfall patterns, spatial and temporal variation in forage abundance, and availability of water. From January 2002 to August 2005, we monitored movements of 76 adult (≥1.5 years) female pronghorns on 2 study areas (Harding and Fall River counties) in western South Dakota. We collected 8,750 visual locations, calculated 204 home ranges, and documented 17 seasonal movements. Eighty-four percent ( n = 55) of pronghorns were nonmigratory and 10% ( n = 6) were conditional migrators. Mean distance between summer and winter range was 23.1 km ( SE = 2.8 km, n = 13). Five adult pronghorns (8%) dispersed a mean distance of 37.6 km ( SE = 12.4 km); of which 1 female moved a straight-line distance of 75.0 km. Winter and summer home-range size varied ( P < 0.0001) between study sites. Mean 95% adaptive kernel winter and summer home-range size of pronghorns was 55.5 and 19.7 km 2 , respectively, in Harding County and 127.2 and 65.9 km 2 , respectively, in Fall River County. Nonmigratory behavior exhibited by pronghorns was likely associated with minimal snow cover and moderate temperatures during winter 2002–2004. Variation in size of adult seasonal home ranges between sites was likely associated with differences in forage distribution and availability between regions.