Small Mammal Mycophagy Response to Variations in Green-Tree Retention

Abstract: We studied the effects of 6 green-tree retention levels and patterns on the diets of northern flying squirrels (Glaucomys sabrinus), Townsend's chipmunks (Tamias townsendii), Siskiyou chipmunks (T. siskiyou), western red-backed voles (Myodes californicus), and southern red-backed voles (Myodes gapperi) using fecal pellet analysis. These rodents are truffle spore dispersers and prey for forest predators such as the northern spotted owl (Strix occidentalis caurina). Pretreatment diets showed differences in truffle and plant consumption among genera. Tree harvesting, especially in the 15% aggregated retention pattern, reduced frequency of Rhizopogon spores in the diet of voles, which may reflect a reduced ability of these animals to forage for Rhizopogon truffles, a decreased access to these truffles, or a reduction in Rhizopogon truffle abundance or frequency. Habitat island effects and edge effects provide conceptual frameworks for the reduction in consumption of Rhizopogon truffles by voles in green-tree aggregates. Overall, small mammal consumption of truffles showed little change in response to the treatments. Animals may be compensating for a locally declining food source by altering their foraging behavior. The long-term effect of this postulated behavioral compensation on small mammal energetics and population dynamics is unknown. Forest managers may reduce the impact of tree harvesting on these key forest ecosystem components by including green-tree aggregates within a dispersed retention matrix.     

Seasonal regulation in fluctuating small mammal populations: feedback structure and climate

We studied fluctuating populations of six small mammal species in the Appalachian Plateau of Pennsylvania, USA for 20 yr. We analyzed the feedback structure of these species using statistical time series models for spring and autumn abundances. All species showed a seasonal density-dependent structure, and in five of them first-order feedbacks were dominant in winter and summer. Instead, southern red-backed voles ( Clethrionomys gapperi ) showed a different feedback structure during winter and summer. In three species ( C. gapperi , Napaeozapus insignis and Peromyscus maniculatus ), environmental factors were more important during summer, while the opposite pattern was found in Blarina brevicauda and Tamias striatus . Snowfall influenced positively the winter population growth rates of southern red-backed voles, white-footed mice, woodland jumping mice and eastern chipmunks. We found seasonal differences in the effects of the small mammals assemblage on population growth rates of the two Peromyscus species. The common feedback structure between seasons observed in most of the species, particularly among voles and mice, points to a different feedback structure from northern cyclic small mammals. We conclude that a seasonal feedback structure dominated by intra- and inter-specific competitive interactions may be at the basis of the population dynamics of these species.     

The influence of weather on habitat use by small mammals

Summer habitat use by three species of forest small mammals was determined using tracking stations Nocturnal weather influenced habitat selection by deer mice and woodland jumping mice but not by red-backed voles Deer mice used all habitats equally on clear nights but were most active in mixed forest on cloudy, rainless nights and most active in a coniferous habitat on rainy nights Jumping mice were most active in mixed forest on clear and rainy nights but shifted to coniferous forest on cloudy dry nights Red-backed voles were most active in the coniferous habitat regardless of weather Microhabitat references within habitats reflected the same preferences as habitat selection Microhabitat selection by jumping mice also changes with weather The mechanism most likely responsible for the observed habitat selection changes is changing insect abundance associated with cloud cover and rainfall     

Effects of Logging Pattern and Intensity on Squirrel Demography

ABSTRACT We examined the effect of harvesting intensity and pattern on red squirrels (Tamiasciurus hudsonicus), northern flying squirrels (Glaucomys sabrinus), and yellow-pine chipmunks (Tamias amoenus) in mature inland Douglas-fir (Pseudotsuga menziesii glauca) forests in south-central British Columbia, Canada. We sampled squirrels 1 year before harvesting through 4 years after harvesting and estimated population parameters using open-population models. Relative to unharvested stands, each of the 3 species showed a strong response to tree removal. From 2 years to 4 years after logging, red squirrel density was 40% (SE = 7.1) lower in stands with 50% basal-area tree removal. From 1 year and up to 4 years after logging, northern flying squirrel density averaged 60% (SE = 5.2) lower in harvested treatments regardless of intensity or pattern of logging. In contrast, density of yellow-pine chipmunks increased markedly with increased logging intensity. Beginning 3 years after logging, yellow-pine chipmunk density was 734% (SE = 269) greater in stands with 50% basal-area tree removal. In the short term, harvesting intensity was a more important determinant of squirrel density than harvesting pattern. Retaining >10 m² per ha of live residual stand structure in mature inland Douglas-fir forests maintained habitat for forest-dependent species such as red squirrels and northern flying squirrels, albeit at lower densities.     

Adaptive intraspecific divergence: an example using the animal cytochrome b gene

The topologies of phylogenetic trees characterized by a high level of intraspecific divergence between the phylogenetic DNA groups (clades) are often explained in terms of the theory of Pleistocene refugia. To elucidate the issue of the adaptive role of intraspecific divergence, the changes in the physicochemical properties of amino acids in the course ofcladogenesis (MM01 model of the TreeSAAP 3.2 package) were analyzed in this work using as an example the nucleotide sequences of the cytochrome b gene in some species of northern animals (lemmings, redbacked voles, chipmunks, flying squirrels, ermines). It was shown that the process of intraspecific divergence was rarely accompanied by radical amino acid substitutions in cytochrome b caused by adaptation (directional selection). In connection with this, the hypothesis is discussed according to which the adaptive variants formed in the species at the peak of cold were lost with climatic warming due to the drift or selection against individuals adapted to cold.     

Adaptive intraspecific divergence: An example using the animal cytochrome <Emphasis Type="Italic">b</Emphasis> gene

The topologies of phylogenetic trees characterized by a high level of intraspecific divergence between the phylogenetic DNA groups (clades) are often explained in terms of the theory of Pleistocene refugia. To elucidate the issue of the adaptive role of intraspecific divergence, the changes in the physicochemical properties of amino acids in the course of cladogenesis (MM01 model of the TreeSAAP 3.2 package) were analyzed in this work using as an example the nucleotide sequences of the cytochrome b gene in some species of northern animals (lemmings, redbacked voles, chipmunks, flying squirrels, ermines). It was shown that the process of intraspecific divergence was rarely accompanied by radical amino acid substitutions in cytochrome b caused by adaptation (directional selection). In connection with this, the hypothesis is discussed according to which the adaptive variants formed in the species at the peak of cold were lost with climatic warming due to the drift or selection against individuals adapted to cold.     

Comparative phylogeography of eastern chipmunks and white-footed mice in relation to the individualistic nature of species

Palaeoecological studies have demonstrated that ecological communities as a whole did not remain stable throughout the climatic fluctuations of the Quaternary. The result is that long-term associations of species cannot be inferred by contemporary associations in ecological communities. Therefore, the evolutionary significance of any contemporary ecological interactions among species and of the biotic community within which species have evolved also cannot be assumed from contemporary conditions. Comparative phylogeographic data provide a method to identify species within ecological communities that have shared biogeographic histories. We present an example of a long-term association between populations of two mammalian species, eastern chipmunks (Tamias striatus) and white-footed mice (Peromyscus leucopus), which are commonly associated with deciduous forest habitats. The distribution of mitochondrial DNA variation in T. striatus and P. leucopus from previously glaciated regions of the eastern United States support the hypothesis that, in at least part of their range, genetic lineages of the two species have expanded from similar population sources since the Last Glacial Maximum. In addition, the spatial concordance of genetic lineages of T. striatus and P. leucopus with the oak-savannah forest formations of Wisconsin and Illinois, suggest that populations associated with this community colonized the area in association with a set of arboreal species that comprise their deciduous forest habitat.     

Coexistence of white-footed mice and deer mice may be mediated by fluctuating environmental conditions

White-footed mice, Peromyscus leucopus noveboracensis, and deer mice, P. maniculatus nubiterrae, occur sympatically throughout much of the Appalachian Mountains of the eastern United States. Previous studies have shown that these two species are behaviorally and ecologically similar and do not partition food or microhabitat. In this paper I use a 14-year data set to demonstrate that the two species have differential population growth and survival rates in response to food abundance and season, which may mediate their coexistence. The ratio of white-footed mice to deer mice ranged from 0.5:1 to 6:1. During summer and times of tood abundance, white-footed mice gained a numerical advantage over deer mice in 10 of 14 years, whereas following winter and poor food production, deer mice had higher survival in 9 of 13 years. The major decline in white-foote mice, but not deer mice, was associated with poor mast (acorn) production in autumn. Differential survival of deer mice may be due to their physiological adaptations to cold temperature and efficient use of torpor during food shortage. Inter-year variance in food production and climatic conditions appear to change the competitive advantage of each species often enough to permit coexistence. Coexistence of these two ecologically similar species may be maintained by different physiological limitations in a fluctuating environment.     

Strongyloides robustus and the northern sympatric populations of northern (Glaucomys sabrinus) and southern (G. volans) flying squirrels

Within North America, northern (Glaucomys sabrinus) and southern (Glaucomys volans) flying squirrels occupy distinct ranges with limited overlap. Sympatry in northern latitudes coincides with northern hardwood vegetation from Minnesota to New England. Strongyloides robustus is an intestinal parasite that infects both species but appears to be deleterious only to northern flying squirrels. As a result, S. robustus could be a critical determinant of flying squirrel population characteristics in at least some areas of sympatry. However, cold weather could potentially limit the distribution of S. robustus in northern climates. Therefore, we assessed fecal samples from both flying squirrel species to determine the presence of the nematode in Wisconsin. Strongyloides robustus was found in 12 flying squirrel scat samples and infected 52% of southern flying squirrels and 11% of northern flying squirrels. Prevalence of S. robustus infection for northern flying squirrels was substantially lower than previously reported from more southern regions. This is the northernmost documentation of S. robustus in flying squirrels and the first documentation of S. robustus parasitizing flying squirrels in Wisconsin.     

Climate change induced hybridization in flying squirrels

There is now unequivocal evidence for global climate change; however, its potential impacts on evolutionary processes remain unclear. Many species have responded to contemporary climate change through shifts in their geographic range. This could lead to increased sympatry between recently diverged species; likely increasing the potential for hybridization. Recently, following a series of warm winters, southern flying squirrels ( Glaucomys volans ) in Ontario, Canada rapidly expanded their northern range limit resulting in increased sympatry with the closely related northern flying squirrel ( Glaucomys sabrinus ). This provided the opportunity to test the prediction that contemporary climate change can act as a catalyst creating conditions for the formation of hybrid zones. Following extensive sampling and molecular analyses (nuclear and mitochondrial DNA), we identified the occurrence of hybridization between sympatric G. sabrinus and G. volans . There was evidence of backcrossing but not of extensive introgession, consistent with the hypothesis of recent rather than historic hybridization. To our knowledge, this is the first report of hybrid zone formation following a range expansion induced by contemporary climate change. This is also the first report of hybridization between North American flying squirrel species.     

An Experimental Test of Competition among Mice,Chipmunks, and Squirrels in Deciduous Forest Fragments

Mixed hardwood forests of the northeast United States support a guild of granivorous/omnivorous rodents including gray squirrels (Sciurus carolinensis), eastern chipmunks (Tamias striatus), and white-footed mice (Peromyscus leucopus). These species coincide geographically, co-occur locally, and consume similar food resources. Despite their idiosyncratic responses to landscape and patch variables, patch occupancy models suggest that competition may influence their respective distributions and abundances, and accordingly their influence on the rest of the forest community. Experimental studies, however, are wanting. We present the result of a large-scale experiment in which we removed white-footed mice or gray squirrels from small, isolated forest fragments in Dutchess County, New York, and added these mammals to other fragments in order to alter the abundance of these two species. We then used mark–recapture analyses to quantify the population-level and individual-level effects on resident mice, squirrels, and chipmunks. Overall, we found little evidence of competition. There were essentially no within-season numerical responses to changes in the abundance of putative competitors. Moreover, while individual-level responses (apparent survival and capture probability) did vary with competitor densities in some models, these effects were often better explained by site-specific parameters and were restricted to few of the 19 sites we studied. With only weak or nonexistent competition among these three common rodent species, we expect their patterns of habitat occupancy and population dynamics to be largely independent of one another.     

Mycophagy by small mammals in the coniferous forests of North America: nutritional value of sporocarps of Rhizopogon vinicolor, a common hypogeous fungus

We evaluated the nutritional value of sporocarps of Rhizopogon vinicolor, a common hypogeous fungus in the coniferous forests of North America, for two small mammal species: the Californian red-backed vole (Clethrionomys californicus) and the northern flying squirrel (Glaucomys sabrinus). Although the nitrogen concentration of sporocarps was high, much of it was in non-protein form or associated with cell walls, suggesting that it may be of low nutritional value or protected from mammalian digestive enzymes. Sporocarps also had high concentrations of cell wall constituents, indicating low availability of digestible energy. When fed a diet of this fungus alone in a controlled feeding experiment both mammal species lost a small amount of body mass. Digestibilities of dry matter, nitrogen, cell wall constituents and energy from sporocarps by both species were lower than the digestibilities of other food types by other similarly sized small mammals. Red-backed voles digested the various components of sporocarps at least as well as the flying squirrels, even though they were almost six-fold smaller in body mass. This observation supports the notion that red-backed voles, like other microtine rodents, have morphological and physiological adaptations of the digestive system that are postulated to permit greater digestion of fibrous diets than predicted on the basis of body size. Despite this, our results re-affirm previous conclusions that hypogeous fungi are only of moderate nutritional value for most small, hindgut-fermenting mammals. Future studies should focus on the importance of mixed-species of fungi in the diet of small mammalian mycophagists. Accepted: 4 December 1998     

A meta-analysis of forest age and structure effects on northern flying squirrel densities

Research on the impact of clearcut logging and partial harvesting practices on northern flying squirrels (Glaucomys sabrinus) has shown inconsistent and contrary results, limiting the use of this species as a management indicator species. Much of this variability in study results is due to the labor intensive nature of studying flying squirrels, resulting in small sampling sizes (average = 5.2 sites, n = 14) and high variation (CV = 0.59) across studies. We conducted a meta-analysis of relevant studies from North America to determine how forestry practices affect flying squirrel abundance. Mean effect size was −1.18 (P < 0.001; n = 14) for all studies, indicating a strong difference between control stands and those regenerating postclearcut or following partial harvesting. Our results support the association of northern flying squirrels with mature, uncut forest and their suitability as ecological indicators of these vegetation types. © 2011 The Wildlife Society.     

Emigration effects on estimates of age‐ and sex‐specific survival of two sciurids

Age‐ and sex‐specific survival estimates are crucial to understanding important life history characteristics, and variation in these estimates can be a key driver of population dynamics. When estimating survival using Cormack–Jolly–Seber (CJS) models, emigration is typically unknown but confounded with apparent survival. Consequently, especially for populations or age classes with high dispersal rates, apparent survival estimates are often biased low and temporal patterns in survival might be masked when site fidelity varies temporally. We used 9 years of annual mark–recapture data to estimate age‐, sex‐, and time‐specific apparent survival of Humboldt''s flying squirrels (Glaucomys oregonensis) and Townsend''s chipmunks (Neotamias townsendii). For Humboldt''s flying squirrels, these estimates support a small body of research investigating potential variation in survival among age and sex classes, but age‐ and sex‐specific survival has not been evaluated for Townsend''s chipmunks. We also quantified the effects of age‐ and sex‐specific emigration on confounded estimates of apparent survival. Our estimates of juvenile flying squirrel survival were high relative to other small mammal species and estimates for both species were variable among years. We found survival differed moderately among age and sex classes for Humboldt''s flying squirrels, but little among age and sex classes for Townsend''s chipmunks, and that the degree to which emigration confounded apparent survival estimates varied substantially among years. Our results demonstrate that emigration can influence commonly used estimates of apparent survival. Unadjusted estimates confounded the interpretation of differences in survival between age and sex classes and masked potential temporal patterns in survival because the magnitude of adjustment varied among years. We conclude that apparent survival estimators are robust during some time periods; however, when emigration rates vary in time, the effects of emigration should be carefully considered and accounted for.     

Experimental infection of nontarget species of rodents and birds with Brucella abortus strain RB51 vaccine

The Brucella abortus vaccine strain RB51 (SRB51) is being considered for use in the management of bnucellosis in wild bison (Bison bison) and elk (Cervus elaphus) populations in the Greater Yellowstone Area (USA). Evaluation of the vaccines safety in non-target species was considered necessary prior to field use. Between June 1998 and December 1999, ground squirrels (Spermophilus richardsonii, n = 21), deer mice (Peromyscus maniculatus, n = 14), prairie voles (Microtus ochrogaster, n = 21), and ravens (Corvus corax, n = 13) were orally inoculated with SRB51 or physiologic saline. Oral and rectal swabs and blood samples were collected for bacteriologic evaluation. Rodents were necropsied at 8 to 10 wk and 12 to 21 wk post inoculation (PI), and ravens at 7 and 11 wk PI. Spleen, liver and reproductive tissues were collected for bacteriologic and histopathologic evaluation. No differences in clinical signs, appetite, weight loss or gain, or activity were observed between saline- and SRB51-inoculated animals in all four species. Oral and rectal swabs from all species were negative throughout the study. In tissues obtained from SRB51-inoculated animals, the organism was isolated from six of seven (86%) ground squirrels, one of six (17%) deer mice, none of seven voles, and one of five (20%) ravens necropsied at 8, 8, 10, and 7 wk PI, respectively. Tissues from four of seven (57%) SRB51-inoculated ground squirrels were culture positive for the organism 12 wk PI; SRB51 was not recovered from deer mice, voles. or ravens necropsied 12, 21, or 11 wk, respectively, PI. SRB51 was not recovered from saline-inoculated ground squirrels, deer mice, or voles at any time but was recovered from one saline-inoculated raven at necropsy, 7 wk PI, likely attributable to contact with SRB51-inoculated ravens in an adjacent aviary room. Spleen was time primary tissue site of colonization in ground squirrels, followed by the liver and reproductive organs. The results indicate oral exposure to SRB51 does not produce morbidity or mortality in ravens, ground squirrels, deer mice, or prairie voles.     

Site occupancy and cavity use by the northern flying squirrel in the boreal forest

Tree cavities are used as shelter and breeding nests by numerous avian and mammalian species. In cold environments, tree cavities are often proposed as the best winter nest choice because of the superior protection they offer from precipitation, wind, cold temperatures, and predators. As such, they represent a critical resource, which has the potential to limit population size of non-excavating species. We assessed factors affecting site occupancy in the boreal forest by northern flying squirrels, a secondary user of tree cavities, and to identify which nest type is preferred during the colder days of the autumn–winter period. We trapped flying squirrels twice in 59 aspen-dominated stands in the autumn period using low- (1.5 m above ground-level) and high-mounted (4 m) traps to determine site occupancy. A total of 85 individuals were captured on 2,880 trap-nights. During the winter period, we radio-tracked 26 individuals to 87 diurnal nests in 220 locations. None of the habitat variables considered (cavity availability, woody debris, and lateral cover) explained site occupancy. Detectability decreased with precipitation, and was lower using high traps than low traps. Both females and males used tree cavities (26%), external nests (39%), and ground nests (35%). In cold weather, females preferred ground nests, whereas males preferred external nests. Our results do not support the hypothesis that tree cavities represent a limiting factor to northern flying squirrels in cold environments. Instead, this species seems to be a generalist and is opportunistic, using a variety of nest types. Nevertheless, practices ensuring the persistence of large diameter live cavity trees, providing better insulative properties, are likely to increase the relative use of tree cavities as nest sites by northern flying squirrels. © 2011 The Wildlife Society.     

Soluble blood proteins and hemoglobin from red-backed and large- toothed voles

Electrophoretic pattern of serum proteins of northern red-backed and large-toothed voles was examined. Seven main protein zones were distinguished. In four of them variability was observed, possibly genetically determined. Polymorphism of transferrins was thoroughly studied. 6 alleles and only 7 phenotypes of this protein were found in the northern red-backed vole, while 3 alleles and 4 phenotypes were revealed in the large-toothed red-backed vole, two alleles being common. Distribution of phenotype frequencies observed in both species differs significantly from the expected value. The portion of heterozygous phenotypes is quite high in both species, being 0.304 and 0.400 in the northern red-backed and large-toothed red-backed voles, respectively. Hemoglobins of the species studied were identical in electrophoretic mobility and monomorphic.     

Small Mammal Responses to Fine Woody Debris and Forest Fuel Reduction in Southwest Oregon

Abstract: Despite its importance for wildlife, most forests in the Pacific Northwest contain low volumes of large downed wood compared to fine woody debris (FWD). We used a replicated experiment to compare short-term responses of deer mice (Peromyscus maniculatus) and western red-backed voles (Clethrionomys californicus) among 3 arrangements of FWD: piled, lopped and scattered, and pile burning, a commonly used method of fuel reduction in commercial Douglas fir (Pseudotsuga menziesii) forests in southwest Oregon, USA. We assessed habitat use, density, and survival of mice and voles during 2 consecutive summers (Jun—Aug 1999 and 2000). Both mice and voles used FWD cover disproportionately from its availability, and they differed in their responses to specific FWD arrangements. Mice used piled FWD (proportional use = 37.0%, 90% CI = 33.0–44.0) 43% more than expected (26.0). Number of mice captured (x̄ = 1.9 mice, 90% CI = 1.5–2.5) and index of home range size (x̄ = 4.8 m, 90% CI = 0.7–8.9) at individual FWD piles decreased up to 16% and increased up to 50%, respectively, for each 1-m increase in distance from piles. Voles used all FWD cover classes in proportion to availability, but number of voles captured increased slightly (x̄ = 0.016 voles/m, 90% CI = 0.001–0.031) for each 1-m increase in distance from piles. Piled FWD had no discernable effect on population density and apparent survival of mice, but analyses had low power (0.25, 0.67). Our results suggest that piling FWD would benefit deer mice, whereas lopped and scattered FWD might benefit voles. Thus, a combination of methods to reduce fire risk should be considered to accommodate multiple small mammal species. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):625–632; 2008)     

Implications of raccoon latrines in the epizootiology of baylisascariasis.

Raccoons (Procyon lotor) frequently establish defecation sites, termed latrines, on large logs, stumps, rocks, and other horizontally oriented structures/surfaces. These latrines are important foci of infective eggs of Baylisascaris procyonis, a nematode parasite of raccoons which is pathogenic to numerous species of mammals and birds. To examine the role of raccoon latrines in this animal-parasite interaction, we documented animal visitations to raccoon latrines in two large forested tracts and two woodlots in Indiana (USA) during 1994 and 1995. Species richness of vertebrate visitors did not differ between sites or years, but species composition differed by site and year. Fourteen mammal and 15 bird species were documented visiting raccoon latrines. Small granivorous mammals, including white-footed mice (Peromyscus leucopus), eastern chipmunks (Tamias striatus), and tree squirrels (Sciurus carolinensis, S. niger, Tamiasciurus hudsonicus) were the most common visitors to latrine sites. White-footed mice, chipmunks, white-breasted nuthatches (Sitta carolinensis), and opossums (Didelphis virginiana) were photographed foraging on undigested seeds in raccoon feces. Active foraging at latrines also was shown experimentally; seeds embedded in raccoon feces were removed at a greater rate at latrine sites than at nonlatrines. We conclude that raccoon latrines are visited routinely by a variety of vertebrates, especially small granivorous rodents and birds which forage for seeds in raccoon feces, and that raccoon latrines are probable sites of transmission of B. procyonis to susceptible mammals and birds.     

Habitat and landscape correlates of southern flying squirrel use of red-cockaded woodpecker clusters

Southern flying squirrels (Glaucomys volans) can have significant negative impacts on red-cockaded woodpecker (Picoides borealis) reproductive success and group size. Although direct control of southern flying squirrels may be necessary in small red-cockaded woodpecker populations (<30 groups), creation of high quality habitat through landscape management is the preferred method for managing larger woodpecker populations. Thus, we determined the habitat and landscape factors within 100 m, 400 m, and 800 m of cluster centers that were related to southern flying squirrel use of red-cockaded woodpecker cavities at the Carolina Sandhills National Wildlife Refuge, South Carolina. At all spatial scales, the number of cavities in the cluster was the most influential variable determining use by southern flying squirrels. At the 400-m and 800-m scales, the amount of stream length was also positively associated with the presence of flying squirrels. The proximity and amount of hardwoods surrounding clusters were not related to southern flying squirrel use at any spatial scale; thus, removal or conversion of hardwood stands surrounding red-cockaded woodpeckers may not be necessary for reducing cavity kleptoparasitism by flying squirrels. However, when establishing recruitment clusters, areas with streams should be avoided and addition of artificial cavities to existing clusters should be done judiciously to minimize the number of excess cavities. Published 2012. This article is a U.S. Government work and is in the public domain in the USA.