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Long-term population dynamics are essentially unknown for the western heather vole (Phenacomys intermedius), a relatively rare microtine species. Our objectives were to provide a description of demographic changes in populations of P. intermedius and determine possible causes of these changes. We analyzed four detailed 12- to 22-year data sets that recorded changes in abundance of P. intermedius in a variety of managed forests in south-central British Columbia, Canada from 1993 to 2018. We evaluated three hypotheses (H) that may help explain population changes: heather voles will (H1) have higher populations in sites with (i) abundant dwarf shrubs or (ii) cover of woody debris; (H2) colonize new clear-cuts as dispersal sinks for young voles; and (H3) be at lower abundance when other Microtus species are present at equal or higher numbers. Populations of P. intermedius occurred commonly at ≤ 1 animal/ha (as a density index). However, in favorable habitats such as some new clear-cuts and in “open” pine stands, P. intermedius occurred at mean annual peak numbers of 5–6/ha (as a density index). There were two relatively high populations of P. intermedius in 1994–1995 and 1998, and possibly again in 2002 in the open pine stands. Mean numbers of lactating females and recruits per year tended to follow the pattern of changes in abundance. High populations of P. intermedius occurred in stands with abundant dwarf shrubs in open pine stands but not those with a cover of woody debris structures in clear-cuts, H1, therefore, was only partially supported. A relatively higher proportion of adult than juvenile heather voles on clear-cuts did not support H2: that these harvested sites would be dispersal sinks for young voles. Mean annual numbers of P. intermedius were positively correlated with those of M. longicaudus, at least on clear-cut sites, hence H3 was not supported. Phenacomys intermedius was essentially absent from our closed-canopy mature/old-growth forests but occurred in many managed forest habitats at low abundance (density index of ≤ 1/ha). In early successional habitats after clear-cutting and in open young pine stands this species occurred at moderate abundance and generally appeared stable in numbers across these forest landscapes.

Extensive habitat loss and degradation have transformed the grasslands of North America into one of the continent’s most vulnerable ecosystems and have greatly imperiled the plants and animals that depend on them. Small mammal communities play a pivotal role in maintaining the health and function of grasslands, and managing small mammal populations is an important, though overlooked, part of grassland restoration and management. We identified habitat variables in restored tallgrass prairie patches that were most strongly associated with variation in small mammal abundance and community composition, with the goal of aiding management efforts to manipulate small mammal populations. We found that small mammal abundance at the local vegetation scale was negatively related both to litter depth and plant diversity. At the landscape scale, small mammal abundance was positively associated with the amount of water surrounding a patch, and negatively associated with the amount of grassland surrounding a patch. Variation in small mammal community composition largely was governed by differences in habitat structure at the landscape scale, rather than differences in vegetation structure at the local scale. We suggested that managers interested in influencing small mammal abundance in grasslands employ tools such as prescribed fire to decrease litter depth if increased small mammal abundance is desired, or increase plant diversity by sowing a high diversity of seeds to depress small mammal abundance.

Urbanization is a selective force that is known to drive changes in the population dynamics of wildlife. How animals adapt to changing environmental conditions is crucial to their survival in these environments. Relative brain size, or cranial capacity, is a known proxy of behavioral flexibility, and can be used to assess how well a species has adapted to a particular environment. We examined changes in cranial capacity in a time series of small mammal skulls collected from urban and rural populations in southwestern Pennsylvania. Skulls from urban populations were collected from Allegheny County, an area that experienced rapid urbanization over the past century, and skulls of rural populations from the Powdermill Nature Reserve of the Carnegie Museum, which has remained relatively unchanged forest over the same period. Our results show that Peromyscus leucopus and Microtus pennsylvanicus from urban populations had significantly greater cranial capacity than their rural counterparts, but the opposite was true for Eptesicus fuscus. We found no difference in relative cranial capacity across time in any of the small mammal species. Our results suggest that a larger cranial capacity is selected for in an urban environment and reinforces the hypothesis that behavioral flexibility is important for animals to adapt to novel environments.

Sexual dimorphism evolves as a response to different selective pressures on males and females. In mammals, sexual selection on traits that improve a male’s ability to compete for access to mates is a common cause of sexual dimorphism. In addition to body mass, adaptations in specific components of the musculoskeletal system that increase strength, stability, and agility, may improve male fighting performance. Here we test the hypotheses that males, when compared to females, are more specialized for physical competition in their skeletal anatomy and that the degree of this sexual dimorphism increases with the intensity of male–male competition. In three species of voles (Cricetidae: Arvicolinae: Microtus), we found partial support for these hypotheses. Male-biased sexual dimorphism in a set of functional indices associated with improved fighting performance was identified in the postcranial anatomy. This dimorphism was greatest in the polygynous Microtus californicus, absent in the monogamous M. ochrogaster, and intermediate in the promiscuous or socially flexible M. oeconomus. However, in the skull, we found results opposite to our predictions. Females had larger skulls relative to overall skeletal size than did males. This may be associated with selection for increased food processing efficiency, which should be highly important because of the compounding effects of increased caloric requirements during gestation and lactation, and the generally low-quality diet of voles. In addition, larger heads in females may be associated with selection for greater digging ability or for defending offspring. These results suggest disparate selective pressures on the postcranial skeletons and skulls of male and female voles.

Individuals of either sex may display alternative behaviors to obtain copulations, but few studies have examined the breeding patterns of females and males in populations where individuals of both sexes exhibit alternative reproductive tactics (ARTs). In prairie voles (Microtus ochrogaster), most adults are territorial, residing at a single nest site either as male–female pairs or as solitary individuals. However, some adults adopt nonterritorial, wandering tactics. During two field seasons monitoring prairie vole populations maintained in seminatural enclosures, we found evidence that females exhibiting different ARTs bred differentially with resident and wandering males. Females residing at a nest with a male bred significantly more often with a paired resident male, primarily their social partner, and significantly less often with male wanderers compared to single resident females or wandering females. These patterns were not due to chance, because paired resident females produced offspring with paired resident males significantly more than expected based on the relative abundance of these males in the population, whereas single resident females produced offspring with male wanderers significantly more than expected based on the proportion of male wanderers in the population. We did not find any evidence that multiple paternity was greater in the litters of single resident females and wanderer females even though these females lacked a male social partner to limit mating access by multiple males. This suggests that mate guarding by a female’s male social partner was not the primary determinant of multiple paternity in the litters of females exhibiting different reproductive tactics. However, male ART did affect the likelihood of multiple paternity. Females that produced offspring with single resident or wanderer males had an increased likelihood of multiple paternity relative to females producing offspring with paired resident males. The results of this study show that female and male ARTs can affect breeding patterns.

Conservation efforts rely on robust taxonomic assessments that should be based on critical assessment of interspecific boundaries, infraspecific variation, and potentially distinctive peripheral populations. The meadow vole (Microtus pennsylvanicus) is widely distributed across North America, including 28 morphologically defined subspecies and numerous isolated populations. Because some subspecies are of high conservation concern, we examined geographic variation across the range of the species to test existing infraspecific taxonomy in terms of local and regional diversification. We sequenced mitochondrial DNA (mtDNA) from 20 subspecies of M. pennsylvanicus and contextualized infraspecific variation through comparison of pairwise genetic distances derived from an extended data set of 63 species of Microtus. We found strong support for at least three divergent clades within M. pennsylvanicus, with observed intraspecific clade divergence exceeding that between several pairwise comparisons of sister species within Microtus. Six nuclear genes were then sequenced to test the validity of mtDNA structure and to further evaluate the possibility of cryptic, species-level diversity using Bayes factor species delimitation (BFD) analyses. BFD consistently and decisively supported multiple species based on the multilocus approach. We propose that taxonomic revision of the meadow vole is required, with the eastern clade now identified as M. pennsylvanicus (Ord 1815), the western clade as M. drummondii (Audubon and Bachman 1853), and the coastal Florida clade as M. dukecampbelli (Woods, Post, and Kilpatrick 1982). We suggest that such an arrangement would more closely reflect evolutionary history and provide critical context for further examination of distinctive southern peripheral populations that harbor novel evolutionary legacies and adaptive potential.

Bartonella is a genus of gram-negative bacteria that includes a variety of human and veterinary pathogens. These pathogens are transmitted from reservoirs to secondary hosts through the bite of arthropod vectors including lice and fleas. Once in the secondary host, the bacteria cause a variety of pathologies including cat-scratch disease, endocarditis, and myocarditis. Reservoirs of these bacteria are numerous and include several species of large mammals, mesocarnivores, and small mammals. Research on reservoirs of these bacteria has focused on western North America, Europe, and Asia, with little focus on the eastern and central United States. We assessed the prevalence of zoonotic Bartonella species among prairie-dwelling rodent species in the midwestern United States. Tissue samples (n = 700) were collected between 2015 and 2017 from five rodent species and screened for the presence of Bartonella DNA via PCR and sequencing of two loci using Bartonella-specific primers. Bartonella were prevalent among all five species, with 13-lined ground squirrels (Ictidomys tridecemlineatus) serving as a likely reservoir of the pathogen B. washoensis, and other rodents serving as reservoirs of the pathogens B. grahamii and B. vinsonii subsp. arupensis. These results demonstrate the value of studies of disease ecology in grassland systems, particularly in the context of habitat restoration and human–vector interactions.

The Eurasian field vole (Microtus agrestis) comprises three evolutionarily significant units (ESUs). The northern ESU is found at higher latitudes across the western Palaearctic region and includes six, largely allopatric, mitochondrial DNA lineages that were derived from population bottlenecks. One of these lineages is found in southern Britain and nearby areas of continental Europe. A prominent sub-lineage is nested within, and therefore derived from, the part of this lineage occupying southern Britain. The sub-lineage consists of an abundant central haplotype together with a series of closely related haplotypes, a distribution that would result from either a recent population bottleneck or a selective sweep. To distinguish between these, we sequenced a Y-chromosome marker in 167 field voles from Britain and Europe, and analysed a panel of 13 autosomal microsatellite loci in 144 field voles from eight populations in Britain. The Y-chromosome marker showed a continental-scale pattern of variation that was not aligned with that of the mitochondrial marker, while microsatellite variation did not show any evidence for a bottleneck, tentatively favouring selection instead. This implies a role for both stochastic and selective processes in generating phylogeographical patterns at different scales in the field vole.

Voles are key mammals in understanding how social interactions can affect large-scale population processes. Previous studies have shown that at high population densities, meadow voles (Microtus pennsylvanicus) have a lower proportion of breeding animals, higher average corticosterone levels, and can be limited by female territorial spacing. Based on this, we compared corticosterone levels and spatial use between breeding and nonbreeding free-ranging adult meadow voles within populations. We measured intrasexual spatial overlap to examine if breeding females minimize occupying the same areas as other females, and noninvasively assessed corticosterone levels using fecal corticosterone metabolites (FCMs). We found that female meadow voles have much lower intrasexual spatial overlap than males, even though both sexes have similar range sizes, and that females have generally higher FCM levels than males. However, breeding and nonbreeding females did not differ from one another in spatial use or in FCM levels. Conversely, reproductive classes of males differed greatly in all measures: nonbreeding males had FCM levels that were two times higher than those of breeding males, occupied a smaller range, and had lower spatial overlap, indicating they were moving less widely than breeding males. We additionally validated an enzyme immunoassay for noninvasively measuring FCMs in meadow voles. The assay was successful in detecting an increase in corticosterone stimulated by adrenocorticotropic hormone injection; however, dexamethasone did not induce negative feedback. FCMs reflect circulating corticosterone levels approximately 5 h prior. These results highlight differences in FCMs and spacing in meadow voles related to sex and reproductive status, and reflect the respective strategies males and females employ during the breeding season.

The numbers of Ixodes ricinus (L.) and Dermacentor reticulatus (Fabricius) larvae and nymphs attached to small mammals are influenced by interspecific competition. The present study analyses data collected over several years in two study areas: Kosewo Górne (Mazurian District, N Poland; between July 1997 and July 2009) and Białowieża Primeval Forest (E Poland; in July 2007). In total, 975 ticks were collected from striped field mice (Apodemus agrarius), yellow-necked mice (A. flavicollis), bank voles (Myodes glareolus), and root voles (Microtus oeconomus). In total, of the 203 investigated rodents, 137 were infested with ticks and 39 demonstrated mixed infection. The numbers of the two tick species found on Apodemus mice were significantly negatively correlated with those on root voles; similarly, although bank voles were significantly more frequently infested by I. ricinus than by D. reticulatus, the reverse was observed in root voles. In addition, among the voles, each tick species was found in different locations on the host body, which could also result from competitive interactions; furthermore, competitive release regarding microhabitat selection was observed on hosts infested with one tick species. This competition may be driven by the limited area of host body available for foraging by ticks, i.e., safe areas of high vascularization covered by thin skin. However, the mechanisms of such competition require further investigation.

On the heterogametic sex chromosomes, the homology-driven processes of pairing, recombination, and segregation are restricted to a short region of X/Y homology known as the pseudoautosomal region (PAR). Although failure of X/Y associations in most mammals... In most mammals, the X and Y chromosomes synapse and recombine along a conserved region of homology known as the pseudoautosomal region (PAR). These homology-driven interactions are required for meiotic progression and are essential for male fertility. Although the PAR fulfills key meiotic functions in most mammals, several exceptional species lack PAR-mediated sex chromosome associations at meiosis. Here, we leveraged the natural variation in meiotic sex chromosome programs present in North American voles (Microtus) to investigate the relationship between meiotic sex chromosome dynamics and X/Y sequence homology. To this end, we developed a novel, reference-blind computational method to analyze sparse sequencing data from flow-sorted X and Y chromosomes isolated from vole species with sex chromosomes that always (Microtus montanus), never (Microtus mogollonensis), and occasionally synapse (Microtus ochrogaster) at meiosis. Unexpectedly, we find more shared X/Y homology in the two vole species with no and sporadic X/Y synapsis compared to the species with obligate synapsis. Sex chromosome homology in the asynaptic and occasionally synaptic species is interspersed along chromosomes and largely restricted to low-complexity sequences, including a striking enrichment for the telomeric repeat sequence, TTAGGG. In contrast, homology is concentrated in high complexity, and presumably euchromatic, sequence on the X and Y chromosomes of the synaptic vole species, M. montanus. Taken together, our findings suggest key conditions required to sustain the standard program of X/Y synapsis at meiosis and reveal an intriguing connection between heterochromatic repeat architecture and noncanonical, asynaptic mechanisms of sex chromosome segregation in voles.

Resource pulses can generate cross-habitat dispersal of consumers, and therefore affect organisms even in areas where the resource pulses do not occur. We investigated this phenomenon at the elevational treeline in the Carpathian Mountains, where beech (Fagus sylvatica) masting caused an increase in abundance of forest rodents and intensified their use of alpine meadows. We tested 3 hypotheses concerning the impact of forest rodent spillover on the abundance of meadow-dwelling pine voles (Microtus subterraneus): 1) the competition hypothesis: if the spillover affects pine voles mostly through intensified competitive interactions, then pine voles should decline when forest rodents reach their peak abundance, i.e., 1 year after masting; 2) the apparent competition hypothesis: if predators switch to alternative prey when populations of forest rodents collapse, then pine voles should decline 2 years after masting; and 3) the apparent mutualism hypothesis: if the increase of forest rodents temporarily releases pine voles from predatory pressure, pine voles should increase in synchrony with forest rodents—1 year after masting. Our results, while correlative in nature, supported the apparent mutualism hypothesis: 1 year after masting, both forest rodents and pine voles strongly increased their abundance. Two years after masting, when populations of forest rodents crashed, abundance of pine voles returned to pre-masting levels rather than collapse. These findings suggest that pulsed spillover, known mostly from negative effects on organisms in recipient habitats, can also create indirect positive interactions. Furthermore, they illustrate how density-dependent spillover of animals might increase the spatial scale of masting effects beyond the habitats where seeds are released.

The needs of offspring change as they develop. Thus, parents should concomitantly change their investment based on the age-related needs of the offspring as they mature. Due to the high costs of parental care, it is optimal for parents to exhibit a shift from intense caregiving of young offspring to promoting independence in older offspring. Yet, the neural mechanisms that underlie shifts in parental behavior are poorly understood, and little is known about how the parental brain responds to offspring of different ages. To elucidate mechanisms that relate to shifts in parental behavior as offspring develop, we examined behavioral and neural responses of male and female prairie voles (Microtus ochrogaster), a biparental rodent, to interactions with offspring at different stages of development (ranging from neonatal to weaning age). Importantly, in biparental species, males and females may adjust their behavior differentially as offspring develop. Because the nonapeptides, vasopressin (VP) and oxytocin (OT), are well known for modulating aspects of parental care, we focused on functional activity of distinct VP and OT cell groups within the maternal and paternal brain in response to separation from, reunion (after a brief period of separation) with, or no separation from offspring of different ages. We found several differences in the neural responses of individual VP and OT cell groups that varied based on the age of pups and sex of the parent. Hypothalamic VP neurons exhibit similar functional responses in both mothers and fathers. However, hypothalamic and amygdalar OT neurons exhibit differential functional responses to being separated from pups based on the sex of the parent. Our results also reveal that the developmental stage of offspring significantly impacts neural function within OT, but not VP, cell groups of both mothers and fathers. These findings provide insight into the functional plastic capabilities of the nonapeptide system, specifically in relation to parental behavior. Identifying neural mechanisms that exhibit functional plasticity can elucidate one way in which animals are able to shift behavior on relatively short timescales in order to exhibit the most context-appropriate and adaptive behaviors.

Author(s): Patton, JL; Conroy, CJ | Abstract: We examined geographic trends in 4 morphological data sets (both craniodental and colorimetric measurements and scores of cranial foramina and qualitative craniodental variables) within and among 5 subspecies of the California vole, Microtus californicus, that occur within or adjacent to the Mojave Desert in eastern California. These analyses are corollary to those previously published on the same samples using both mitochondrial and nuclear gene sequences and microsatellite loci. The morphological and molecular data sets are generally concordant in supporting the existing infraspecific taxonomy, although important conflicts are evident. With the combination of data sets, we are able to assign previously unknown samples to these infraspecific taxa. We define and provide an emended diagnosis of each subspecies using the criteria established by Willi Hennig in his 1966 assessment of the qualities and characteristics of infraspecific taxa, which he understood to be fundamentally different than the hierarchical phylogenetic system he established as the conceptual framework for taxa at the species level and above. Hennig's views are contrary to how the subspecies concept has been applied in much of recent mammalian taxonomy.

Microtus agrestis (Linnaeus, 1761) is a relatively small microtine rodent commonly called the field vole. It has a compact body, blunt oval head, short round ears that barely protrude from the fur, and a short, bicolored, rather stiff tail. It is easily confused with sympatric or parapatric Microtus arvalis, M. oeconomus, and M. levis (M. rossiaermeridonalis) with indistinguishable juveniles. There is geographical variation of size and coloration ranging across Europe south from the Pyrenees to the Arctic coast and Alps eastward to the River Yenisei and Lake Baikal in Asia up to elevations of 2,100 m. It prefers wet meadow areas, riverside habitats, and forests with dense herbaceous cover. M. agrestis is considered to represent a trace of a 1st radiation of the genus Microtus in Europe.

The segregation of homologous chromosomes at the first meiotic division is dependent on the presence of at least one well-positioned crossover per chromosome. In some mammalian species, however, the genomic distribution of crossovers is consistent with a more stringent baseline requirement of one crossover per chromosome arm. Given that the meiotic requirement for crossing over defines the minimum frequency of recombination necessary for the production of viable gametes, determining the chromosomal scale of this constraint is essential for defining crossover profiles predisposed to aneuploidy and understanding the parameters that shape patterns of recombination rate evolution across species. Here, I use cytogenetic methods for in situ imaging of crossovers in karyotypically diverse house mice (Mus musculus domesticus) and voles (genus Microtus) to test how chromosome number and configuration constrain the distribution of crossovers in a genome. I show that the global distribution of crossovers in house mice is thresholded by a minimum of one crossover per chromosome arm, whereas the crossover landscape in voles is defined by a more relaxed requirement of one crossover per chromosome. I extend these findings in an evolutionary metaanalysis of published recombination and karyotype data for 112 mammalian species and demonstrate that the physical scale of the genomic crossover distribution has undergone multiple independent shifts from one crossover per chromosome arm to one per chromosome during mammalian evolution. Together, these results indicate that the chromosomal scale constraint on crossover rates is itself a trait that evolves among species, a finding that casts light on an important source of crossover rate variation in mammals.

We were able to substantially increase our knowledge of what is likely the least understood endangered terrestrial mammal in the United States, the Florida salt marsh vole (FSMV; Microtus pennsylvanicus dukecampbelli). We developed a predictive landscape model that estimated 264 ha of potential habitat for FSMVs. Evaluating our model, we found voles at 8 of the 36 sites sampled, yielding a model accuracy of 22% for a subspecies that previously was known from only 3 locations. Within areas of potential habitat, FSMVs selected patches of marsh vegetation > 0.49 ha with at least some (≥ 16.75% and ≤ 43.61%) smooth cordgrass (Spartina alterniflora) cover. Suggestive of a meta-population dynamic, FSMV activity decreased outside of patches of smooth cordgrass and saltgrass (Distichlis spicata) identified by the predictive landscape model. Our hierarchical approach to studying FSMVs allowed us to leverage a limited amount of data to ultimately produce important ecological information about an endangered species. This approach easily may be adapted to other mammals with similar information needs.

The California vole, Microtus californicus, restricted to habitat patches where water is available nearly year-round, is a remnant of the mesic history of the southern Great Basin and Mojave deserts of eastern California. The history of voles in this region is a model for species-edge population dynamics through periods of climatic change. We sampled voles from the eastern deserts of California and examined variation in the mitochondrial cytb gene, three nuclear intron regions, and across 12 nuclear microsatellite markers. Samples are allocated to two mitochondrial clades: one associated with southern California and the other with central and northern California. The limited mtDNA structure largely recovers the geographical distribution, replicated by both nuclear introns and microsatellites. The most remote population, Microtus californicus scirpensis at Tecopa near Death Valley, was the most distinct. This population shares microsatellite alleles with both mtDNA clades, and both its northern clade nuclear introns and southern clade mtDNA sequences support a hybrid origin for this endangered population. The overall patterns support two major invasions into the desert through an ancient system of riparian corridors along streams and lake margins during the latter part of the Pleistocene followed by local in situ divergence subsequent to late Pleistocene and Holocene drying events. Changes in current water resource use could easily remove California voles from parts of the desert landscape.

Arvicoline rodents serve as model organisms for investigating the intricacies of ecological/evolutionary feedbacks that characterize multiannual fluctuations in abundance. A population of California voles (Microtus californicus) in central California was followed for 3.5 years using 4 small grids located 50 m apart. Blood samples were scored for 4 polymorphic allozyme loci by starch gel electrophoresis. These were carefully chosen for their potential to reveal details of genetic structuring among demes. A low density of voles developed into a peak followed by a rapid decline to low density. Heterozygosity averaged 20.2% (n = 709) with loci-specific shifts associated with density, season, and cyclic phase. Statistically significant heterogeneity in genotypic and allelic frequencies among the grids was found for 3 of the loci. Pairwise comparisons among the grids for each locus separately (n = 24) yielded 11 significantly disparate pairs for genotypic frequencies and 10 for alleles. Significant correlations...

There is now considerable evidence for the survival of temperate species within glacial refugia that were situated at relatively high latitudes, notably the Carpathian Basin and Dordogne region in Europe. However, the prevalence of fossil remains in such locations is rarely matched by molecular evidence for their contribution to subsequent geographical and demographic expansion of the species in question. One obstacle to this has been insufficient analysis of modern samples from the relevant areas, in particular the parts of eastern Europe that surround the Carpathian refugium. In the present study, we examine the patterns of variation in mitochondrial DNA of the common vole (Microtus arvalis), obtained from existing museum specimens and from newly-collected samples obtained in this area. We show that common voles from one of six extant mitochondrial DNA lineages have colonized most of the species' range in eastern Europe. We contend that the post-glacial dispersal of this lineage most likely originated from the Carpathian refugium, adding support to the argument that such northern refugia made an important contribution to existing genetic diversity in Europe.

Although the biological significance of individual variation in physiological traits is widely recognized, studies of their association with fitness in wild populations are surprisingly scarce. We investigated the effect of individual phenotypic variation in body mass, resting (RMR) and peak metabolic rates (PMR) on mortality of the root vole Microtus oeconomus. Body mass and metabolic rates varied significantly among consecutive years and were also age dependent, as individuals born in late summer and autumn were characterized by significantly lower body mass and metabolic rates than animals born earlier. At the beginning of winter voles born in spring and early summer exhibited reduced body mass and metabolic rates, whereas animals born later maintained lower body mass and RMR, which may be interpreted as phenotypic plasticity enhancing the probability of survival. Body mass had no significant effect on vole survival during summer. In contrast, smaller individuals were characterized by lower mortality during early winter, whereas higher body mass was positively associated with survival later in the season. High body-mass-corrected RMR positively affected survival in both summer and winter. The effect of PMR was apparent only during winter, though its direction (and correlation with RMR) varied among years. Deep snow cover negatively affected the survival of voles in both early and late winter. Ambient temperature was positively associated with winter survival, except for late winter, when rising temperature caused flooding of vole habitat. We conclude that the lack of consistency in the directionality and strength of the effects of body mass and metabolic rates on winter survival does not undermine their importance, but rather demonstrates the ability of individuals to adjust metabolic rate to changing environmental conditions. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113, 297–309.

Extreme weather variation on the northern Great Plains of North America can potentially influence the abundance of grassland rodents across vast areas. We used the remains of 33,697 small mammals collected from owl pellets in central and western Canada over 15 years to determine the influence of weather on the annual abundance of deer mice (Peromyscus maniculatus), sagebrush voles (Lemmiscus curtatus), and meadow voles (Microtus pennsylvanicus). Weather variation affected the annual abundances of all 3 species; however, influence on deer mouse and sagebrush vole annual abundances was relatively small compared to that on meadow voles. This finding may indicate that factors other than weather (i.e., habitat availability) are more important for the abundance of deer mice and sagebrush voles at the landscape scale. In contrast, meadow voles were positively associated with the duration of snow cover above the hiemal threshold (20 cm), exhibiting up to 5-fold increases (i.e., irruptions) in abundance f...

Prey often react to predation risk by foraging preferentially in the safety of cover rather than in more risky open patches. Yet this pattern of patch use also can be caused by dominant interspecific competitors. We develop a simple theory of this form of apparent predation risk that describes the patch use of an optimal forager confronted with dominant individuals. The theory predicts that subordinate animals should increase their use of safe foraging patches as the density of nearby dominants increases. We tested the theory with meadow voles (Microtus pennsylvanicus) and southern red-backed voles (Myodes gapperi). We used dyadic encounters to confirm that meadow voles are dominant over red-backed voles. We then evaluated their respective foraging patterns in pairs of covered and open patches in 4 adjacent subgrids in an old-field enclosure. Subordinate redbacked voles foraged indifferently between covered and open patches when few meadow voles were present. Red-backed voles increased their use of both patches as the number of nearby meadow voles increased. Givingup densities were lowest, and harvesting efficiency highest, in covered patches when the number of nearby meadow voles was high. These results document competition between the 2 species and suggest that vigilance toward dominant meadow voles magnifies the risk experienced by red-backed voles in open patches. Investigators assessing foraging behavior between ‘‘safe’’ and ‘‘risky’’ patches might misinterpret the competitive effect as predation risk unless they 1st account for competition among foraging individuals.

Little is known about the macrohabitat associations of rodents and shrews in prairie landscapes because of the logistic constraints of conventional trapping. We used the remains of 60,972 small mammals in owl pellets to assess factors affecting small mammal composition across 4.3 million hectares of the northern Great Plains of North America. Cropland with clay soils was dominated by deer mice (Peromyscus maniculatus), whereas areas with higher proportions of native grassland and moderately sandy soils supported communities with more sagebrush voles (Lemmiscus curtatus). Areas with clay soils and higher annual precipitation were associated with higher proportions of house mice (Mus musculus), meadow voles (Microtus pennsylvanicus), and shrews (Blarina brevicauda and Sorex species), whereas drier areas with sandier soils and lower annual precipitation were dominated by olive-backed pocket mice (Perognathus fasciatus) and northern grasshopper mice (Onychomys leucogaster). Contrary to extrapolations...

Many female small mammals face limited food availability during lactation. These dams may have to choose between altering the amount of maternal behavior they provide to their young, reducing the size of their litter, or not adjusting their behavior or litter size. How females allocate energy to maternal investment may depend on the energy costs of different lactation stages. We hypothesized that the amount of time female voles provided maternal behavior would differ if they were deprived of food during early, middle, or late lactation. We tested this hypothesis by placing lactating female meadow voles (Microtus pennsylvanicus) into 1 of 4 groups: dams that underwent a 30% caloric restriction during days 1–7 of lactation, those that were food restricted on days 8– 14, those that were food restricted on days 15–21, and dams that did not undergo food restriction during lactation. Dams that were not food restricted spent more time engaged in maternal behavior than dams that were food restricted during lactation. Dams that were food restricted during days 8–14 of lactation displayed the most pronounced decline in maternal behavior relative to dams that were restricted during days 1–7 or days 15–21 of lactation. This effect was most dramatic in the amount of time that dams spent licking their pups. Reduced licking of pups may affect the mother–pup bond, inducing pups to possibly develop deficits in their social and sexual behavior as adults. The results also suggest that when they are faced with a food shortage, particularly during the 1st week of lactation, lactating female meadow voles do not reduce the size of their litter but do show a decrease in maternal behavior toward pups.

Arvicoline voles do not possess sophisticated mechanisms to tolerate prolonged fasting and can starve to death in less than 24 h. Two experiments were performed to determine if the common vole (Microtus arvalis) and the tundra vole (M. oeconomus) have thermoregulatory adaptations to enhance survival during food deprivation or in winter, and to reveal the possible presence of ultradian body temperature (Tb) rhythms. Laboratory-bred voles (n = 5–6 for both species) were equipped with intra-abdominal data loggers that registered Tb every 10 min and were subjected to 12-h fasting periods during photophase and scotophase. In addition, 10 animals of both species were placed in outdoor cages at natural ambient temperature (Ta) and their Tb was registered at 3-h intervals between October and June. Fasted voles did not enter torpor, but the average 24-h Tb decreased by 0.2–0.5°C in winter, yielding modest energy savings. The decrease in the Tb correlated with Ta but not photoperiod and could, thus, be int...

Male prairie voles become more responsive to infants following cohabitation with a female. Exposure to female sensory cues prior to offspring birth may influence male paternal tendencies by modifying his response to infant odors in particular or to odors in general. To test these hypotheses, males were housed with an unfamiliar female or a same-sex sibling for 13 days then examined for their response towards either live infants or infant-like inanimate objects covered with one of three odors: water, infant, sub-adult. We recorded the number of males that retrieved and manipulated the infants or odor-covered objects and measured the frequency and duration of time males spent attending to them or engaged in other non-social activities. Female-Cohabited males approached the container holding infant-odor covered objects faster than Male-Cohabited males, but showed no differences in time spent manipulating those objects. Males in both groups spent more time manipulating live infants than odor-covered objects. However, Female-Cohabited subjects were more likely to manipulate odor-covered objects as well as live infants than Male-Cohabited subjects. Additionally, the frequency of self-grooming in Female-Cohabited males was higher for water-covered objects compared to Male-Cohabited males. In presence of water and live infants, Female-Cohabited males groomed themselves with greater frequency than in presence of infant odor or sub-adult odor. The data suggest that female cues increase the male’s sensitivity to infant odors and enhance the salience of non-social odors.

Understanding the impact of small mammals on the development of grasslands requires information on their food preferences. We conducted feeding trials with 4 species of common small mammals, prairie voles (Microtus ochrogaster), meadow voles (M. pennsylvanicus), white-footed mice (Peromyscus leucopus), and eastern cottontails (Sylvilagus floridanus), to determine the relative palatability of plant shoots and seeds in introduced grasslands of east-central Illinois. Because we needed to conduct many feeding trials we developed a rapid method for comparison of palatability of plants to multiple species of small mammals in the field. We evaluated the method by comparison with previous data on food preference and palatability for voles at nearby study sites. Shoots of herbaceous legumes (Medicago sativa and Trifolium pratense) and dandelions (Taraxacum officianale) were among the most palatable foods for voles, and bluegrass (Poa pratensis) was among the least palatable. Meadow voles found palatable more species of graminoids than did prairie voles, and meadow voles, but not prairie voles, found palatable a greater percentage of graminoids than of forbs. A greater percentage of introduced plants than of native plants were palatable to prairie voles, but this likely reflected a high proportion of introduced graminoids among tested plants. White-footed mice found palatable a few forbs and a greater percentage of annual plants compared with perennial plants, whereas cottontails found palatable many forbs and woody plants and a greater percentage of perennial plants. White-footed mice tended to eat larger amounts and more kinds of seeds than voles, and only white-footed mice tended to eat more of large-seeded species. Plant type did not appear to significantly affect palatability of seeds for any mammal. We used these results to predict the likely impact of small mammals on the development of nonnative grasslands.

Following dispersal from 1 group, individuals may join other established social groups. Such intergroup transfer may increase access to potential mates and decrease mate competition. We used data from 402 individuals to examine patterns of intergroup transfer in prairie voles (Microtus ochrogaster). Nearly 32% of established social groups (single female units, male–female pairs, or communal groups of at least 2 adults of the same sex) were joined by 1 or more individuals. Most individuals (76%) that joined social groups were wanderers that were either unmarked, recently marked during grid trapping, or marked transients; 70% were males. Joining a group was not contingent upon recent disappearance of residents. Total number of residents positively affected the probability of a female joining a social group, whereas number of adult female residents and population density negatively affected it. Some individuals (24%) moved directly from one group to another without an intervening wandering stage; we refer to these instances of intergroup transfer as direct transfers. Most direct transferers moved into nearby groups, but not the closest group. Males were more likely than females to directly transfer into groups with potential mates and without potential competitors. Thus, males directly transferred in a manner consistent with maximizing reproductive opportunities. In contrast, 25% of females directly transferred into groups without potential mates and 96% into groups with at least 1 adult female. Females may be less constrained by group composition with respect to potential mates because wandering males, with which females can mate, are prevalent. All-male groups almost never occur in our population, so females probably cannot avoid joining groups with competitors.

Natural hybridization between species is not a rare event. In arvicoline rodents, hybridization is known to occur in the wild and/or in captivity. In the Microtus arvalis group, cytogenetic studies revealed that there were two distinct chromosomal forms (2n = 46 but a different fundamental number of autosomes). These forms have been attributed to two cryptic species: the common (arvalis) and Altai (obscurus) voles. Recently, individuals with intermediate karyotypes (F1 and backcrosses) were discovered in central European Russia, and, for this reason, other studies have regarded obscurus and arvalis as conspecific. In the present study, to address the question of the species limits in the Altai vole and to infer its evolutionary history, a phylogeographical analysis combined with multivariate morphometric methods and original chromosome data was performed. Two obscurus lineages were identified: the Sino-Russian and South Caucasian lineages. Both lineages are characterized by low genetic diversity, resulting, in the former, from a past bottleneck event caused by encroaching periglacial areas and, in the latter, from recent rapid population divergence. Introgressive hybridization between the Altai and common voles appears to be the result of a secondary contact following the Last Glacial Maximum in central European Russia. Despite the fact that speciation is an ongoing process in most arvicoline species, the common and Altai voles are genetically divergent, morphologically and karyologically distinct, and exhibit contrasting evolutionary histories. For all these reasons, they should be ranked as species: M. arvalis and M. obscurus.

We tested the hypothesis that domestic sheep (Ovis aries) in alpine meadows of central Norway negatively affect populations of field voles (Microtus agrestis) because grazing reduces cover and increases the concentration of secondary compounds in forage plants. We maintained 3 treatments for 3–4 years on 11 experimental plots. Treatments included fencing to remove sheep; fertilization to improve forage quality; and grazed, unfertilized controls. Within experimental plots we collected data on cover of vascular plants, forage quality (nitrogen, fiber, and secondary compounds), and indicators of performance for vole populations (summer density, winter activity, and body size). Sheep appeared to reduce the cover of vascular plants, but quality of 4 species of forage plants showed little response to grazing. Concentrations of nitrogen in forage increased on fertilized plots, as did the ratio of nitrogen/(fiber plus phenolic compounds). Voles occurred sporadically on the experimental plots, however, and indicators of vole performance differed little among treatments. Furthermore, the quality of preferred food (Carex bigelowii) of voles showed no clear response to fertilization, and the activity of voles on plots showed no significant correlation with quality of C. bigelowii irrespective of treatment. We concluded that removal of grazing by sheep in these meadows did not have a substantial short-term impact on forage quality or on vole populations, but longer-term studies are needed because the composition of vegetation may change slowly in response to exclusion of herbivores.

The Preble’s meadow jumping mouse (Zapus hudsonius preblei [PMJM]) is a rare rodent of southeastern Wyoming and central Colorado that has been the center of debates regarding subspecies’ genetic identity and the application of the Endangered Species Act. I analyzed a 7-year PMJM mark–recapture data set using a temporal symmetry model (Pradel model) to estimate apparent survival (u), recruitment ( f ), population change (k), and vital rate influence on k. Deer mouse (Peromyscus maniculatus) captures depressed u, suggesting that competition for resources may decrease PMJM survival. Vole-mediated habitat changes or voles’ affinity for quality riparian habitats may explain why PMJM u and f increased with meadow vole (Microtus pennsylvanicus) captures. Based on early-summer and late-summer sampling from 2000 to 2006, k estimates were 0.87 6 0.06 SE and 0.87 6 0.11 SE, respectively, and f had a greater influence on k than did u. This PMJM population is losing connectivity to eastern, northern, and southern tributaries from habitat degradation and storm-water and municipal runoff erosion. The loss of the adjoining habitat and the PMJMs that were supported by this habitat prevents new recruitment via immigration. Because of the importance of recruitment to PMJM population stability, tributaries and the riparian habitat along these tributaries are vital to PMJM conservation. Scaleappropriate habitat sampling, assessments of reproductive success, and detailed demographic studies to estimate vital demographic parameters will help identify how particular habitat components impact fecundity and immigration.

When isolated, pups of muroid rodents emit ultrasonic vocalizations, an indication that they are stressed by being alone and exposed to cooling temperatures. Rate of vocalizations is greatest at the end of their 1st week and beginning of the 2nd week, declines in the 2nd week as eyes open and thermoregulation becomes fully established, and ceases in the 3rd week. Young of 1 vole species, the prairie vole (Microtus ochrogaster), vocalize significantly more than those of other species, which has been attributed to the social structure of the species, because they have monogamous mating behavior and high level of biparental care. To determine whether this vocalizing behavior is typical of vole species with monogamous social systems, I examined calling behavior in another monogamous species with biparental care, the pine (or woodland) vole (M. pinetorum), and I compared it to that of a polygynous species that has only maternal care, the meadow vole (M. pennsylvanicus). Pups were isolated for 20 min at 228C 6 28C and the number of ultrasonic calls they emitted was monitored. Pine vole pups vocalized at high rates, similar to prairie voles, emitting a mean of 47 calls/min at the ages when calling was greatest (8–16 days). In contrast, meadow vole pups vocalized significantly less, emitting a mean of 9 calls/min at the age of greatest calling (5–9 days). Pine voles continued vocalizing to 23 days, like prairie voles but different from other vole species, including meadow voles, which cease calling earlier. I conclude that young voles of monogamous species, which are highly affiliative and have paternal as well as maternal care, respond to the stress of isolation more strongly than do other vole species and emit more ultrasonic vocalizations than species that are less social and have less parental care.

The intervening landscape between patches of forest (i.e., matrix) has enormous potential to mitigate the negative effects of forest fragmentation. However, to release this potential requires understanding of how individual species perceive matrix. Here we investigated use of matrix by pine martens (Martes martes) in a region with low forest cover (Scotland). We radiotracked 11 martens to determine their habitat preferences, then combined our data with those published from 5 additional Scottish landscapes to examine how home-range size (i.e., population density) and diet of martens varied with forest edge density (i.e., fragmentation). Our tracking showed that although mature forest was the most preferred habitat, certain matrix habitats (scrub and tussock grassland) also were consistently selected. These 2 habitats provided martens with fundamental resources that are of limited availability within intensively managed plantation forests: den sites and primary prey (Microtus agrestis). Our synthesis of data across landscapes indicated martens benefit from supplemental resources in matrix habitats; consumption of small mammals increased with fragmentation and coincided with an initial increase in marten population densities. However, population densities of martens decreased once fragmentation passed a threshold level. Our results demonstrate that habitat complementation at the landscape-scale is essential for some forest-associated species. Resource supplementation from matrix habitats may be particularly important in regions with a long history of low-forest cover or where forest cover is now dominated by afforested plantations, which may lack essential resources.

Many mammal species can distinguish between opposite-sex conspecifics that differ in a certain trait. In that coat coloration is associated with differences in physiological and behavioral traits, coat color may affect the attractiveness of odor cues produced by conspecifics. Individuals may be able to respond preferentially to conspecifics with a particular coat color. In the present study, we test the hypothesis that scent marks of brown and blond voles differ in their attractiveness to male and female conspecifics. Male voles and brown females did not discriminate between blond- and brown-coated opposite-sex conspecifics suggesting that they are neither selecting potential mates dissociatively nor associatively. However, blond females behaved as if the scent marks of blond males were more attractive than were the scent marks of brown males. Our data suggest that blond females who are already conspicuous to predators, may select blond males as mates because they do not appreciably increase the risk of detection to predators, particularly avian predators. Moreover, because these conspicuous males have survived to mate they may have good genes that reflect their relatively higher quality [Current Zoology 58 (2): 221–227 , 2012].

Lemmings construct nests of grass and moss under the snow during winter, and counting these nests in spring is 1 method of obtaining an index of winter density and habitat use. We counted winter nests after snow melt on fixed grids on 5 areas scattered across the Canadian Arctic and compared these nest counts to population density estimated by mark–recapture on the same areas in spring and during the previous autumn. Collared lemmings were a common species in most areas, some sites had an abundance of brown lemmings, and only 2 sites had tundra voles. Winter nest counts were correlated with lemming densities estimated in the following spring (rs 5 0.80, P , 0.001), but less well correlated with densities the previous autumn (rs 5 0.55, P , 0.001). Winter nest counts can be used to predict spring lemming densities with a log-log regression that explains 64% of the observed variation. Winter nest counts are best treated as an approximate index and should not be used when precise, quantitative lemming density estimates are required. Nest counts also can be used to provide general information about habitat-use in winter, predation rates by weasels, and the extent of winter breeding.

Within socially monogamous species, a male's reproductive success depends on his success at obtaining a social partner with which he mates, as well as gaining extra-pair matings. We investigated the impacts of body mass and parasitism on 2 measures of a male's success at obtaining a social partner (number of female social associations and relative strength of a social bond to a single female) and 2 measures of reproduction (number of females with which a male sires offspring and the number of offspring sired) in natural populations of prairie voles (Microtus ochrogaster) located in Kansas and Indiana. Kansas males with greater endoparasite loads were less likely to have one social partner. Instead, they associated with multiple females although they did not sire offspring with more females than males with lower levels of endoparasitism. We did not find any relationship between endoparasite infestation and the number of females with which males associated in Indiana. There was no association betwe...

Paleontology can provide a deep-time dimension to observations about recent reactions of small mammals to climate change. Obtaining this perspective for voles (Microtus), a common and important constituent of North American mammal communities, has been difficult because species identification based on their dental remains is problematic. Here I demonstrate that geometric morphometrics and discriminant analyses can use commonly fossilized dental features to identify the 5 extant species of Microtus in California: M. californicus (California vole), M. longicaudus (long-tailed vole), M. montanus (montane vole), M. oregoni (Oregon vole), and M. townsendii (Townsend’s vole). Analyses of landmarks on the lower 1st molar (m1) provide more accurate identification than those of the 3rd upper molar (M3), and it is important to use jackknife misidentification metrics to assess the precision of discriminant analyses. Addition of semilandmark curves on m1 does not improve accuracy. The utility of these techniques is demonstrated by identifying Microtus specimens from 2 California fossil localities, Pacheco 2 and Prune Avenue, which provides the first evidence for extralimital presence of M. longicaudus at both localities. The presence of M. longicaudus at these low-elevation sites indicates that pronounced geographic range shifts in this species that have been observed in California over the last 100 years also occurred during previous climate changes. Eventually it might be possible to ascertain whether current range shifts are exceeding those that typified responses to past climate changes.

Management agencies and quantitative ecologists need robust estimates of population density. The best way of converting population estimates of livetrapped small mammals to population density is not clear. We estimated population density on livetrapping grids with 4 estimators applied to 3 species of boreal forest and 3 species of tundra rodents to test for relative differences in density estimators. We used 2 spatial estimators proposed by Efford (2009) and 2 traditional boundary-strip estimators designed for grid livetrapping. We analyzed mark– recapture data from 104 trapping sessions from the boreal forest at Kluane, Yukon (n 5 4,818 individuals), and 56 trapping sessions from tundra areas of Herschel Island and Komakuk Beach in northern Yukon (n 5 1,327 individuals). For boreal forest rodents on average both boundary-strip methods produced density estimates larger than Efford’s maximum-likelihood (ML) estimator by as much as 50% at all population densities up to 25 animals/ha. For tundra rodents both boundary-strip methods produced density estimates smaller than Efford’s ML at low density (,1.5/ha) and larger than Efford’s ML density by 36–63% at high density (25/ha). Efford’s inverse prediction estimator produced larger density estimates than the ML estimator by 4% for the boreal forest and 32% for the tundra rodents. Relationships were high between all the estimators, such that trends in density could be inferred from all methods. Determining the bias in population density estimators in small mammals will require data from populations spatially closed and completely enumerated. For our small mammals Efford’s ML estimator typically provided density estimates smaller than those produced by conventional boundary-strip estimators.

Zones of contact between divergent biological forms within or between species are critical to the study of speciation. How characters flow across contact zones can be informative of the speciation process. To better understand this phenomenon in a mammal, we investigated cranial shape change in a contact zone between northern and southern phylogeographical groups of California voles (Microtus californicus). We took 12 linear measurements of skulls, one measurement of the mandible, and coded the presence and absence of two skull foramina for 427 specimens. In multivariate analyses, skulls within parental regions were correctly assigned more than 90% of the time. In the contact zone, 49% were classified as northern and 51% as southern, with a bimodal distribution of posterior probability values. Foraminal patterns in the contact zone were intermediate between northern and southern regions. A cline analysis for coastal populations suggested a similar centre for mitochondrial and nuclear markers, although a centre for the morphological data was offset. Cranial morphology indicates an intermediate area with overlap between the two regions, as suggested by the molecular data, with a pattern distinct from mitochondrial DNA or nuclear DNA markers. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104, 264–283.

Previous studies have shown that individuals responded preferentially to the mark of the top-scent donor relative to that of the bottom-scent donor of an over-mark. However, terrestrial mammals are likely to encounter over-marks consisting of the scent marks of more than two same-sex conspecifics in the intersections of runways, near the nests of sexually receptive female conspecifics, and inside and along the borders of the territories of conspecifics. We determined how meadow voles, Microtus pennsylvanicus, respond to the marks of the top-, middle-, and bottom-scent donors of an over-mark. We tested the hypothesis that voles exposed to an over-mark will respond preferentially to the scent marks that were deposited more recently, the scent marks that were on top or near the top of the over-mark, compared to the scent marks that were deposited earlier or near the bottom of the over-mark. Voles spent more time investigating the mark of the top-scent donor than that of the either the middle- or bottom-scent donor. However, males but not female voles spent more time investigating the middle-scent mark than the bottom-scent mark. We also tested the hypothesis that voles evaluate and respond to over-marks differently from single scent marks. Voles spent more time investigating the marks of the top-, middle-, and bottom-scent donors compared to scent marks that were not part of the over-mark. Voles can distinguish among the overlapping scent marks of three scent donors and sex differences exist in the values they appear to attach to each of these scent marks.

Summary Our study used ‘ stand structures ’ and ‘ ecological indicators ’ as inferences to managing dry Douglas-fi r ( Pseudotsuga menziesii (Mirbel) Franco) forests for timber, biodiversity and range habitat attributes at 10 – 12 years after partial harvesting. Forest structures included densities and sizes of Douglas-fi r and species and structural diversity of total conifers. Ecological indicators included cattle forage species in the understor e y vegetation and three small mammal species: a closed-canopy specialist, the southern red-backed vole ( Myodes gapperi Vigors) and two early successional species, the long-tailed vole ( Microtus longicaudus Merriam) and heather vole ( Phenacomys intermedius Merriam). Harvesting treatments were across a gradient of historical cut, individual tree selection and patch cut sites in the upper Interior Douglas-fi r (IDF dk ) biogeoclimatic zone near Summerland, British Columbia, Canada. Small mammal populations were sampled from 2006 to 2008. Closed-canopy forest structure seemed to maintain populations of red-backed voles in the harvested sites. Development of understor e y vegetation in harvest-origin openings was similar across the levels of partial cutting and provided at least fi ve plant species for cattle forage. Patches of grassland and shrubland in all treatment sites maintained viable populations of long-tailed voles and heather voles. These three small mammal indicators represent both late and early successional conditions in these IDF forests, which need to be managed for multiple objectives. Long-term monitoring of indicators will verify relationships with the components of biodiversity that they represent.

The behaviors that surround copulation are characterized as sociosexual behaviors. These behaviors displayed by males that are directed at females may include allogrooming, wrestling, chasing, approach, and time spent together. The data supported the hypothesis that the duration of sociosexual behaviors differs during the pre-copulatory, peri-copulatory, and post-copulatory phases of the mating bout in meadow voles. Voles spent more time approaching conspecifics during the pre- and peri-copulatory phases than during the post-copulatory phase. Voles spent more time allogrooming, wrestling, and chasing during the pre-copulatory phase than during the peri- and post-copulatory phases. Voles spent similar amounts of time together during the pre-, peri-, and post-copulatory phases. The data suggest that sociosexual behaviors displayed by males may be involved in setting the pace and temporal components of the mating bout. During the pre-copulatory phase particular behaviors by male voles may attract females, during the peri-copulatory phase some of these behaviors may stimulate or motivate the female to mate, and during the post-copulatory phase certain behaviors may prepare the male to mate again.

Microtus pennsylvanicus is represented in Mexico only by the Chihuahuan meadow vole (M. p. chihuahuensis), known from only 1 disjunct population in a small and isolated marsh in the arid lands of northern Chihuahua. Livetrapping conducted between 2000 and 2004 provided no specimens of M. p. chihuahuensis, nor was any sign of this vole observed. By the end of this study the marsh providing water had been drained, thereby destroying the vole's habitat. Surveys of other marshes in northern Chihuahua also failed to produce evidence of the species. We therefore conclude that M. p. chihuahuensis has been extirpated from its only known locality. Using “ancient” DNA from museum specimens we evaluated genetic divergence between museum specimens of M. p. chihuahuensis and 46 extant Microtus species and subspecies. Our results support the subspecific status of M. p. chihuahuensis. The loss of this subspecies is an example of population extinction, a very severe form of biodiversity loss. Until recently such...

The phylogeographic architecture of the common vole, Microtus arvalis, has been well-studied using mitochondrial DNA and used to test hypotheses relating to glacial refugia. The distribution of the five described cytochrome b (cyt b) lineages in Europe west of Russia has been interpreted as a consequence of postglacial expansion from both southern and central European refugia. A recently proposed competing model suggests that the 'cradle' of the M. arvalis lineages is in western central Europe from where they dispersed in different directions after the Last Glacial Maximum. In the present study, we report a new cyt b lineage of the common vole from the Balkans that is not closely related to any other lineage and whose presence might help resolve these issues of glacial refugia. The Balkan phylogroup occurs along the southern distributional border of M. arvalis in central and eastern Bosnia and Herzegovina, Montenegro, and eastern Serbia. Further north and west in Slovenia, Bosnia and Herzegovina, and Serbia, common voles belong to the previously-described Eastern lineage, whereas both lineages are sympatric in one site in Bosnia and Herzegovina. The Balkan phylogroup most reasonably occupied a glacial refugium already known for various Balkan endemic species, in contrast to the recently proposed model. South-east Europe is an absolutely crucial area for understanding the postglacial colonization history of small mammals in Europe and the present study adds to the very few previous detailed phylogeographic studies of this region.

Small mammals in boreal forest ecosystems fluctuate dramatically in abundance and 1 possible mechanism to explain these changes is the bottom-up hypothesis of variation in food supplies. Here we ask if variation in berry crops produced by 6 major species of dwarf shrubs and herbs, epigeous mushroom crops, and white spruce seeds allow us to predict changes in the abundance of the red-backed vole (Myodes [5 Clethrionomys] rutilus), the deer mouse (Peromyscus maniculatus), and field voles (Microtus oeconomus and M. pennsylvanicus combined) over 13 years (1997–2009) in the Kluane Lake region of the southwestern Yukon, Canada. M. rutilus is the dominant rodent in these forests, comprising 64% of the catch. Overwinter survival is a key demographic variable in all these rodents, and the winter food supply—principally berries produced the previous summer— may be 1 key to overwinter survival. We predicted that berry, mushroom, and tree seed crops in year t would produce changes in rodent density in year t + 1. We could explain statistically 78–98% of the variation in May and August abundance of all 3 rodent species with indices of berry crops and mushrooms in the previous summer. For M. rutilus the critical predictor was berry crops of Empetrum nigrum. For P. maniculatus, the critical species were Arctostaphylos uva-ursi, A. rubra, and mushrooms. Spruce seed crops were not significantly correlated with rodent densities or changes in density. A large fraction of the variation in rodent numbers in this ecosystem is explained by a simple bottom-up model of population limitation. DOI: 10.1644/ 09-MAMM-A-005.1.

To examine phylogenetic relationships within the Asian lineage of voles (Microtus) belonging to subgenus Alexandromys, the mitochondrial cytochrome b gene (cytb) was sequenced for its representatives, and the results were compared with the cytogenetic, morphological, and paleontological data. In all the trees inferred from maximum likelihood, parsimony, and Bayesian phylogenetic analyses, the Asian clade is subdivided into highly supported Alexandromys s.s. and moderately supported Pallasiinus lineages. Four subclades are recovered within Alexandromys: (1) Microtus maximowiczii and Microtus sachalinensis; (2) Microtus miiddendorffii s.l., Microtus mongolicus and Microtus gromovi; (3) Microtus fortis; and (4) Microtus limnophilus. Thus, M. limnophilus demonstrates clear affinities to Alexandromys s.s. but not to Microtus oeconomus (subgenus Pallasiinus), which was always regarded as its sibling species. The results obtained indicate M. mongolicus as a member of Alexandromys but not of the Microtus arvalis group, thus being concordant with the cytogenetic data. The mitochondrial data support the species status of M. gromovi; moreover, its placement as a part of a trichotomy with M. miiddendorffii s.l. and M. mongolicus contradicts the traditional affiliation of M. gromovi with M. maximowiczii. The divergence rate of cytb third position transversions in Microtus is estimated at approximately 8% per Myr, which corresponds to approximately 30% per Myr for all substitution types at all codon positions. The maximum likelihood distance based on complete sequence showed a tendency for a progressive underestimation of divergence and time for older splits. According to our molecular clock analysis employing nonlinear estimation methods, the split between Alexandromys and Pallasiinus and basal radiation within Alexandromys date back to approximately 1.2 Mya and 800 Kya, respectively. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99, 595–613.

Seventeen extant populations of Microtus (Terricola) savii have been investigated for correlations of first lower molar shape to climatic variables by means of geometric morphometrics, and controlling for phylogenetic inheritance. Comparative methods revealed that climatic variables and phylogeography provide a very similar contribution to variation in first lower molars morphology, whereas tooth size does not appear to be affected by climatic conditions. Climate-related changes have been recognized in the anteroconid portion of the tooth. This indicates that molar tooth variation is strongly influenced by climatic conditions, although in a complex way. Calabrian populations, often ascribed to Microtus (Terricola) brachycercus, form a distinct cluster, in agreement with the most recent genetic analyses.

Microtus californicus (Peale, 1848) is a cricetid commonly called the California vole or California meadow mouse. A sexually dimorphic, medium-sized vole, M. californicus is 1 of 62 species in the genus Microtus. It is found in the interior valleys of southwestern Oregon, most of California, and northern Baja California, Mexico. M. californicus is found in a wide range of habitats from arid uplands to wet meadows and salt marshes. Several subspecies are listed as of conservation concern; the Amargosa vole, M. c. scirpensis, is listed as federally endangered.

The study of alternative model organisms has yielded tremendous insights into the regulation of behavioral and physiological traits not displayed by more widely used animal models, such as laboratory rats and mice. In particular, comparative approaches often exploit species ideally suited for investigating specific phenomenon. For instance, comparative studies of socially monogamous prairie voles and polygamous meadow voles have been instrumental toward gaining an understanding of the genetic and neurobiological basis of social bonding. However, laboratory studies of less commonly used organisms, such as prairie voles, have been limited by a lack of genetic tools, including the ability to manipulate the genome. Here, we show that lentiviral vector-mediated transgenesis is a rapid and efficient approach for creating germline transgenics in alternative laboratory rodents. Injection of a green fluorescent protein (GFP)-expressing lentiviral vector into the perivitelline space of 23 single-cell embryos yielded three live offspring (13 %), one of which (33%) contained germline integration of a GFP transgene driven by the human ubiquitin-C promoter. In comparison, transfer of 23 uninjected embryos yielded six live offspring (26%). Green fluorescent protein is present in all tissues examined and is expressed widely in the brain. The GFP transgene is heritable and stably expressed until at least the F(2) generation. This technology has the potential to allow investigation of specific gene candidates in prairie voles and provides a general protocol to pursue germline transgenic manipulation in many different rodent species.