Guidelines for manuscripts for the Journal of Mammalogy
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Supplemental feeding increased the number of red squirrels ( Tamiasciurus hudsonicus ) captured on two trapping grids, leading Sullivan (Journal of Mammalogy, 71:579–590, 1990) to infer that the population density of residents had increased. This conclusion is questionable because the residency status of captured squirrels was unknown.
Age is a basic demographic characteristic vital to studies of mammalian social organization, population dynamics, and behavior. To eliminate potentially confounding ontogenetic variation, morphological comparisons among populations of mammals typically are limited to mature individuals (i.e., those assumed to have ceased most somatic growth). In our morphometric studies of treeshrews (Scandentia), adult individuals are defined by the presence of fully erupted permanent dentition, a common criterion in specimen-based mammalogy. In a number of cases, however, we have had poorly sampled populations of interest in which there were potentially useful specimens that could not be included in samples because they lacked associated skulls. Such specimens typically are associated with external body and weight measurements recorded by the original collectors, and we sought to determine whether these data could be used successfully as a proxy for age or at least to establish maturity. We analyzed four traditional external dimensions (head-and-body length, tail length, hind foot length, and ear length) and weight associated with 103 specimens from two allopatric populations of the Lesser Treeshrew ( Tupaia minor Günther, 1876) from Peninsular Malaysia and from Borneo, which we treated as separate samples (populations). Individuals were assigned to one of eight age categories based on dental eruption stage, and measurements were compared among groups. In general, mean sizes of infants and subadults were smaller than those of adults, but the majority of subadults fell within the range of variation of adults. The large overlap among infants, subadults, and adults in external measurements and weight indicates that such measures are poor proxies for age in this species, probably for treeshrews in general, and possibly for other small mammals. This has significant implications for any investigation wherein relative age of individuals in a given population is an important consideration.
The kin structure of a species at relatively fine spatial scales impacts broad-scale patterns in genetic structure at the population level. However, kin structure rarely has been elucidated for migratory marine mammals. The Pacific walrus ( Odobenus rosmarus divergens ) exhibits migratory behavior linked to seasonal patterns in sea ice dynamics. Consequently, information on the spatial genetic structure of the subspecies, including kin structure, could aid wildlife managers in designing future studies to evaluate the impacts of sea ice loss on the subspecies. We sampled 8,303 individual walruses over a 5-year period and used 114 single-nucleotide polymorphisms to examine both broad-scale patterns in genetic structure and fine-scale patterns in relatedness. We did not detect any evidence of genetic structure at broad spatial scales, with low F ST values (≤ 0.001) across all pairs of putative aggregations. To evaluate kin structure at fine spatial scales, we defined a walrus group as a cluster of resting individuals that were less than one walrus body length apart. We found weak evidence of kin structure at fine spatial scales, with 3.72% of groups exhibiting mean relatedness values greater than expected by chance, and a significantly higher overall observed mean value of relatedness within groups than expected by chance. Thus, the high spatiotemporal variation in the distribution of resources in the Pacific Arctic environment likely has favored a gregarious social system in Pacific walruses, with unrelated animals forming temporary associations.
Morphometric analyses of the manus skeleton have proven useful in understanding species limits and morphological divergence among tupaiid treeshrews (Scandentia: Tupaiidae). Specimens in these studies are typically limited to mature individuals with fully erupted permanent dentition, which eliminates potentially confounding variation attributable to age, but can also exclude rare taxa and small island populations that are poorly represented in systematic collections. To determine the real limits associated with including immature animals in such studies, we used multivariate analyses to study sexual and age variation of the manus skeleton within two allopatric populations of the Lesser Treeshrew (Tupaia minor Günther, 1876) from the Malay Peninsula and from Borneo that we treated as separate samples. Individuals were aged using dental eruption of the permanent dentition. We also recorded the degree of epiphyseal fusion of the bones of the manus based on x-rays of study skins. We then tested our ability to distinguish the two populations using a series of discriminant function analyses of hand measurements from samples that included varying proportions of immature individuals and adults. We found no evidence of sexual dimorphism in hand proportions, permitting us to combine females and males in our samples. Epiphyseal fusion of the metacarpals and phalanges typically occurs by the time the third molars have completely erupted, and fusion of the distal epiphyses of the radius and ulna typically occurs by the time the permanent fourth premolars are in place. There is occasional asynchrony between dental age and epiphyseal fusion. In both populations, the hands of most infants and subadults provide morphometric values within the range of variation of adults, although they are typically distributed in the lower part of the adult range and have the potential to bias the sample toward lower mean size. The inclusion of infants and subadults when attempting to discriminate between two taxa generally results in lower rates of correct classifications, although the rates increase as the sample of immature individuals is limited to older subadults. As a general rule, we recommend that infants and subadults continue to be excluded from analyses when exploring taxonomic boundaries among treeshrews. In cases of extremely small sample sizes of adults, however, older subadults—in which the permanent third premolars are erupting or in place—can be used with appropriate caution.
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.
Hibernation is an adaptation to survive periods of stress, from food limitation or harsh thermal conditions. A key question in contemporary ecology is whether rare, range-restricted species can change their behavior in response to climate change (i.e., through behavioral plasticity). The northern Idaho ground squirrel, Urocitellus brunneus (A. H. Howell, 1928), is a federally threatened species that hibernates for approximately 8 months per year within the bounds of its small range in central Idaho, USA. Changes in temperature, snow accumulation, and summer precipitation, all brought about as a result of climate change, may reduce survival or fecundity of northern Idaho ground squirrels if they cannot adapt to these climate changes. Hibernating species can respond to climate-change-induced thermal challenges in two ways: change their hibernation physiology and behavior (i.e., emergence date or number of torpor bouts) or alter their environment (i.e., change hibernacula depth or location). We explored a suite of intrinsic and extrinsic factors to document the extent to which they influenced hibernation behavior of northern Idaho ground squirrels. Emergence date was positively associated with snowpack and negatively associated with mean winter temperature. Mean minimum skin temperature was negatively associated with canopy closure and slope of a squirrel’s hibernaculum. Duration of the heterothermal period, number of euthermic bouts, and total time spent euthermic were positively associated with body mass. Immergence date and duration of the longest torpor bout were negatively associated with body mass. Warmer temperatures and less snow accumulation in the winter—caused by climate change—likely will cause altered emergence dates. Our results suggest that any future climate-induced changes in snowfall, ambient temperature, food availability, or habitat likely will impact survival of this rare ground squirrel, because such changes will cause changes in hibernation behavior, percent mass loss during hibernation, and duration of the active season when small mammals are more susceptible to predation.
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.
Habitat information for small mammals typically consists of anecdotal descriptions or infrequent analyses of habitat use, which often are reported erroneously as signifying habitat preference, requirements, or quality. Habitat preferences can be determined only by analysis of habitat selection, a behavioral process that results in the disproportionate use of one resource over other available resources and occurs in a hierarchical manner across different environmental scales. North American chipmunks ( Neotamias and Tamias ) are a prime example of the lack of studies on habitat selection for small mammal species. We used the Organ Mountains Colorado chipmunk ( N. quadrivittatus australis ) as a case study to determine whether previous descriptions of habitat in the literature were upheld in a multiscale habitat selection context. We tracked VHF radiocollared chipmunks and collected habitat information at used and available locations to analyze habitat selection at three scales: second order (i.e., home range), third order (i.e., within home range), and microhabitat scales. Mean home range was 2.55 ha ± 1.55 SD and did not differ between sexes. At the second and third order, N. q. australis avoided a coniferous forest land cover type and favored particular areas of arroyos (gullies) that were relatively steep-sided and greener and contained montane scrub land cover type. At the microhabitat scale, chipmunks selected areas that had greater woody plant diversity, rock ground cover, and ground cover of coarse woody debris. We concluded that habitat selection by N. q. australis fundamentally was different from descriptions of habitat in the literature that described N. quadrivittatus as primarily associated with coniferous forests. We suggest that arroyos, which are unique and rare on the landscape, function as climate refugia for these chipmunks because they create a cool, wet microclimate. Our findings demonstrate the importance of conducting multiscale habitat selection studies for small mammals to ensure that defensible and enduring habitat information is available to support appropriate conservation and management actions.
For more than 130 years, the type locality of the Plains Spotted Skunk, Spilogale putorius interrupta (Rafinesque, 1820) has been accepted to be along the upper Missouri River. The species’ description was based on a specimen observed by Constantine S. Rafinesque during his 1818 exploration of the Ohio River Valley, but Rafinesque never ventured into the animal’s geographic range west of the Mississippi River, calling into question the type locality and, therefore, the identity of the taxon. We reconstruct Rafinesque’s itinerary from his notes, publications, and correspondence and determine that Rafinesque probably observed the specimen on 20 September in Middletown, Kentucky, while traveling between Louisville and Lexington. He spent the day with John Bradbury, who participated in the 1811 Astor expedition up the Missouri River. On 1 April 1811, Bradbury collected the skin of a skunk, and evidence suggests that it was this skin that Rafinesque described. The type specimen of the Plains Spotted Skunk was obtained on the Missouri River flood plain in southern Chariton County or northern Saline County, Missouri, and this area should be considered the type locality for M. interrupta .
Utah prairie dogs (UPDs, Cynomys parvidens ) are colonial, herbivorous rodents listed under the Endangered Species Act as threatened. Little is known about UPD population dynamics at higher elevations in the species’ range. From 2013 through 2016, we studied UPDs on five colonies at 2,645 to 2,873 m elevation on the Awapa Plateau, Utah, USA. Primary production increases with precipitation and precipitation increases with elevation on the plateau. We hypothesized that UPD body condition, reproduction, survival, and population growth all would vary directly with precipitation and elevation. Each year, we live-trapped UPDs from late-Jun through Aug, weighing each UPD, aging it as adult or pup, measuring its right hind foot, marking it for unique identification, and releasing it at point of capture. Fleas from live-trapped UPDs and opportunistically collected rodent carcasses, and rodent carcasses themselves, were tested for the agent of sylvatic plague ( Yersinia pestis ), a lethal invasive pathogen. Adult UPD body condition (mass:foot) increased with elevation. In addition, UPD reproduction (pups:adults) and population growth (λ) increased with precipitation. Annual survival declined from 0.49 in 2013–2014 to 0.24 in 2015–2016. We captured 421 UPDs in 2013 but only 149 in 2016. Sylvatic plague may have contributed to population declines. Notwithstanding, plague detection (yes/no by colony and year) had no statistical effect on population growth or annual survival, raising suspicion about the predictive value of binary plague detection variables. Generally speaking, efforts to conserve UPDs may benefit from the restoration and preservation of large colonies at mesic sites.
Pelt color serves many functions from signaling to crypsis to thermoregulation and its purpose has been a lively source of debate in biology for over a century. Determining the effects of both habitat and human influences on pelt color patterns can be difficult. We made novel use of a multispecies occupancy model by defining “pelt color” as “species.” We then used this model to test predictions and estimate pelt color patterns concurrently for three carnivore species in Idaho, United States. We predicted pelt patterns of all three carnivores would be affected by environmental variables as well as human disturbance. Areas of Idaho where baiting was allowed and preferential harvest possible did not explain pelt patterns in black bears and neither did forest cover. Road density was positively associated with detection probability but negatively associated with occupancy of both black and brown pelt bears, however. Gray pelt wolves were found more often in areas with higher road densities than black wolves. As predicted, black, but not gray, wolves were positively associated with forest cover. Both red and black pelt foxes were positively associated with increasing elevation and road density. Black pelt foxes were negatively associated with forest cover, mirroring the habitat use described for native black pelt foxes. We demonstrate how using noninvasively collected data and extending multispecies occupancy models can allow biologists to study the distribution of different pelt colors in wild populations.
Movement patterns of lions ( Panthera leo ) reveal how they hunt large herbivores in heterogeneous landscapes such as the Kruger National Park in South Africa. Large herbivores are distributed differently on the landscape and therefore have different vulnerabilities as prey for lions. For instance, blue wildebeest ( Connochaetes taurinus ) occupy small grazing lawns at night but are difficult for lions to capture because open areas lack cover for stalking. African buffalo ( Syncerus caffer ) aggregate in large herds but are less available because these herds only intermittently enter the home ranges of individual lion prides. Unlike large herds of wildebeest and buffalo, plains zebra ( Equus quagga ) move widely in small herds while browsing greater kudus ( Tragelaphus strepsiceros ) and giraffes ( Giraffa camelopardalis giraffa ) generally occur in lower densities. We used spatial data derived from GPS collars to investigate several hypotheses regarding the movements of three lion prides in response to their prey. We found that lions were most active and moved longer distances during nighttime than during daytime. Lions remained within their core home ranges on 87% of nights and wandered to the outlying areas of the home ranges every second night. Lions visited grazing lawns, that is, area of short grass, where wildebeest herds resided every second night, and moved toward the direction of buffalo herds within 2 km of vicinity. Lions spent more time near riverbanks that provided dense woody cover at night than expected but concentrated only weakly near sites with surface water where herbivores drank in the dry season. Our study contributes to understanding how lions vary their movements in response to the spatial and temporal heterogeneity in the relative availability and vulnerability of multiple prey species.
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.
Space use by mammals can differ among age-classes, sexes, or seasons, and these processes are recognized as adaptive behavioral strategies. Semi-fossorial ground squirrels, in particular, have shown age- and sex-specific patterns in their aboveground movement behaviors. We studied space use of Mohave ground squirrels ( Xerospermophilus mohavensis ) at the Freeman Gulch study site in the central region of their range in the Mojave Desert, California. We documented the timing of their full annual cycle, investigated correlates of size of home ranges of adults and distance of long-distance movements by juveniles, and evaluated whether juvenile body masses and movements were related to interannual climatic variation. Adult males emerged from burrows and entered hibernation sooner than did adult females. Home ranges were larger for males ( x ¯ "> x ¯ x¯ ± SD = 0.50 ± 0.40 km 2 ) than females (0.05 ± 0.03 km 2 ), especially during the mating season. Maximum distances moved by juveniles did not differ significantly between males (1.6 ± 2.3 km) and females (1.0 ± 1.8 km), and both sexes were equally likely to travel long distances from natal sites. The longest-distance movement we documented was 7.7 km by a juvenile male. Juveniles born in a year of low vegetation productivity gained significantly less mass (97 ± 7 g) and traveled shorter distances than those born in a year of higher vegetation productivity (177 ± 20 g). Our findings were similar to those documented in the Coso Range study site in the northern region of the geographic range of the species, indicating that movement patterns are similar range-wide. Mohave ground squirrels responded strongly to climate variation, and global climate change will likely result in a northward shift in suitable habitats. Informed conservation planning, therefore, will be essential for this rare, declining species.