Evolutionary and ecological significance of non-shivering thermogenesis in the common vole

La production endogène à long terme de chaleur, même au repos, est une adaptation que l'on retrouve chez les animaux à sang chaud (les oiseaux et les mammifères). Cette production de chaleur a comme but le maintien d'une température constante du corps indépendamment de la température extérieure. A cette fin, les mammifères ont développé une forme de tissu spécialisé nommé tissu adipeux brun (BAT). Ce tissu est responsable de la conversion de nourriture en chaleur, procédé appelé thermogenèse sans frisson (NST = non-shivering thermogenesis). Durant ce procédé la uncoupling protein 1 (UCP1) convertit, au sein des mitochondries, la nourriture en chaleur au lieu de produire de l'ATP, molécule utilisée comme énergie cellulaire. On suppose que cette inefficacité de la conversion de la nourriture en ATP dans le BAT influence l'homéostasie de l'énergie, l'allocation des ressources ainsi que la régulation de processus gourmant en énergie comme la croissance et la reproduction. Afin de maintenir une température du corps constante, les mammifères doivent ajuster leur NST en fonction de la température ambiante. La NST devrait être donc plus importante que la croissance et la reproduction durant l'hiver que lors l'été/à haute altitude qu'à basse altitude. En effet, plusieurs études ont déjà démontré la dépendance de la NST aux divers paramètres environnementaux. Par contre, l'héritabilité de la NST ainsi que sa relation avec d'autres traits de caractère, ne sont que très peu connus, ceci malgré l'importance d'une telle information afin de pouvoir comprendre son potentiel évolutif. L'étude de l'importance évolutive et écologique sur la NST chez les campagnols des champs (Microtus arvalis) fut donc le but cette thèse de doctorat. Grâce aux informations collectées sur 4 générations de campagnols (chapitre 1), une dépendance saisonnière et journalière de la NST a été démontrée: elle augmente lors des périodes froides et diminue lors de la lactation. On a démontré que bien qu'étant plastique, la variation de la NST a une composante génétique significative. Elle est corrélée avec le taux d'activité métabolique au repos indiquant des contraintes intrinsèques. A l'aide d'une expérience de jardin commune, on a pu démontrer dans le chapitre 2 que les campagnols habitant en altitude ont une capacité génétique de thermogenèse sans frisson plus haute que celles de basse altitude. Ils produisent des portées plus petites et leur descendance grandit moins vite, surtout à partir du 10ème jour ce qui coïncide avec le début de la production de chaleur endogène. En choisissant artificiellement des campagnols avec une NST faible ou grande, on a pu démontrer une relation entre la NST et la développement de la masse corporelle. Les campagnols avec une haute NST grandissent plus lentement et sont plus légères à l'âge adulte que celles ayant une basse NST. A l'aide d'un croisement interligne entre les campagnols sélectionnés (avec basse et haute NST), on a pu montrer dans le chapitre 3 des effets « parent-of-origin >> du développement massique de la descendance, indiquant une empreinte génétique parentale. Cela veut dire que l'expression d'un allele dépend de l'origine parentale. De plus, des effets « parent-of-origin » des taux de base de norépinephrine et d'irisine ont pu être trouvés. Ces hormones sont connues pour influencer non seulement la TSF mais aussi d'autres caractéristiques. Ces influences ouvrent la voie à de nouvelles études sur la relation entre la TSF et l'histoire de vie. Dans le chapitre 4 on a démontré des effets à long terme de l'allocation des ressources en manipulant la taille des portées qui ont abouti à des différences dans l'investissement dans la reproduction et de la croissance de la descendance à la fois dans le cas de la reproduction manipulé et aussi dans le non - manipulée entre les femelles avec portées agrandies et réduites. Ensemble, ces résultats mettent en évidence le rôle central de la NST dans l'allocation des ressources sur la base d'un compromis entre le maintien et la croissance et ainsi transforme l'histoire de vie des mammifères. Ces études montrent comment les mammifères peuvent répondre rapidement à court et à long terme (c'est-à-dire par des réponses génétiques ou plastiques) à un changement rapide du climat. On montre aussi qu'il y a probablement une corrélation entre l'histoire de vie et des changements du comportement. Finalement mes résultats ont montré un lien étroit entre la NST et la croissance et les dimensions du corps. Ces résultats indiquent que le tissu adipeux brun et la NST pourraient être une cible thérapeutique intéressante pour traiter l'obésité.

ABCtoolbox: a versatile toolkit for approximate Bayesian computations.

BACKGROUND: The estimation of demographic parameters from genetic data often requires the computation of likelihoods. However, the likelihood function is computationally intractable for many realistic evolutionary models, and the use of Bayesian inference has therefore been limited to very simple models. The situation changed recently with the advent of Approximate Bayesian Computation (ABC) algorithms allowing one to obtain parameter posterior distributions based on simulations not requiring likelihood computations. RESULTS: Here we present ABCtoolbox, a series of open source programs to perform Approximate Bayesian Computations (ABC). It implements various ABC algorithms including rejection sampling, MCMC without likelihood, a Particle-based sampler and ABC-GLM. ABCtoolbox is bundled with, but not limited to, a program that allows parameter inference in a population genetics context and the simultaneous use of different types of markers with different ploidy levels. In addition, ABCtoolbox can also interact with most simulation and summary statistics computation programs. The usability of the ABCtoolbox is demonstrated by inferring the evolutionary history of two evolutionary lineages of Microtus arvalis. Using nuclear microsatellites and mitochondrial sequence data in the same estimation procedure enabled us to infer sex-specific population sizes and migration rates and to find that males show smaller population sizes but much higher levels of migration than females. CONCLUSION: ABCtoolbox allows a user to perform all the necessary steps of a full ABC analysis, from parameter sampling from prior distributions, data simulations, computation of summary statistics, estimation of posterior distributions, model choice, validation of the estimation procedure, and visualization of the results.

Establishment success and resulting fitness consequences for vole dispersers

Dispersal is one of the most important, yet least understood phenomena of evolutionary ecology. Triggers and consequences of dispersal are difficult to study in natural populations since dispersers can typically only be identified a posteriori. Therefore, a lot of work on dispersal is either of a theoretical nature or based on anecdotal observation. This is especially true for cryptic species such as small mammals. We conducted an experiment on the common vole, Microtus arvalis, in semi-natural enclosures and investigated the spatial and genetic establishment success of residents and dispersers in their natal and new populations. Our study uses genetic data on the reproductive success of 1255 individuals to measure the fitness trajectories of the residents and dispersing individuals. In agreement with past studies, we found that dispersal was highly male-biased, and was most probably induced by the agonistic encounters with conspecifics, suggesting it could act as an inbreeding avoidance mechanism. There was low breeding success of dispersers into new populations. Although nearly 26% of identified dispersers reproduced in their natal populations, only seven percent reproduced in the new populations. Settlement appeared to be a pre-requisite for reproduction in both sexes, and animals that did not spatially settle into a new population dispersed again, usually on the same day of immigration. In the event that dispersers reproduced in the new population, they did so at relatively low population densities. We also found age-related differences between the sexes in breeding success, and male dispersers that subsequently established in the new population were young individuals that had not reproduced in their natal population, whereas successful females had already reproduced in their natal population. In conclusion, with our detailed field data on establishment and substantial parentage assignments to understand breeding success, we were able to gain an insight into the fitness of dispersers, and how the two sexes optimise their fitness. Taken together, our results help to further understand the relative advantages and costs of dispersal in the common vole.

Flea infestation reduces the life span of the common vole.

Parasitism is often a source of variation in host's fitness components. Understanding and estimating its relative importance for fitness components of hosts is fundamental from physiological, ecological and evolutionary perspectives. Host-parasite studies have often reported parasite-induced reduction of host fecundity, whereas the effect of parasitism on host survival has been largely neglected. Here, we experimentally investigated the effect of infestation by rat fleas (Nosopsyllus fasciatus) on the life span of wild-derived male common voles (Microtus arvalis) bred in captivity. We found that the mean life span of parasitized voles was reduced by 36% compared to control voles. Parasitized voles had a smaller body size, but a relatively larger heart and spleen than control voles. These results indicate an effect of flea infestation on host life span and our findings strongly suggest that ectoparasites should be taken into account in the studies of host population dynamics.

Relationship between diet and reproductive success in the Israeli barn owl

The effect of diet on barn owl (Tyto alba) breeding biology has been well studied in the temperate regions but not in the more arid Middle East. In temperate regions, barn owls are darker colored and mainly prey upon Cricetidae rodents, whereas in arid regions, they are lighter colored and prey to a larger degree upon Muridae rodents. In this study we analyzed the diet and breeding success of 261 barn owl pairs nesting in Israel. The reproductive success of barn owls declined from March to August, and fledged more young when they consumed a larger proportion of social voles (Microtus socialis guentheri). Although the diet of the lighter colored barn owls in Israel comprises more Muridae than that of the darker morphs in temperate regions, in both regions the number of barn owl young increases with an increased proportion of voles in the diet.

The effects of spatial and temporal heterogeneity on the population dynamics of four animal species in a Danish landscape

Background: Variation in carrying capacity and population return rates is generally ignored in traditional studies of population dynamics. Variation is hard to study in the field because of difficulties controlling the environment in order to obtain statistical replicates, and because of the scale and expense of experimenting on populations. There may also be ethical issues. To circumvent these problems we used detailed simulations of the simultaneous behaviours of interacting animals in an accurate facsimile of a real Danish landscape. The models incorporate as much as possible of the behaviour and ecology of skylarks Alauda arvensis, voles Microtus agrestis, a ground beetle Bembidion lampros and a linyphiid spider Erigone atra. This allows us to quantify and evaluate the importance of spatial and temporal heterogeneity on the population dynamics of the four species. Results: Both spatial and temporal heterogeneity affected the relationship between population growth rate and population density in all four species. Spatial heterogeneity accounted for 23–30% of the variance in population growth rate after accounting for the effects of density, reflecting big differences in local carrying capacity associated with the landscape features important to individual species. Temporal heterogeneity accounted for 3–13% of the variance in vole, skylark and spider, but 43% in beetles. The associated temporal variation in carrying capacity would be problematic in traditional analyses of density dependence. Return rates were less than one in all species and essentially invariant in skylarks, spiders and beetles. Return rates varied over the landscape in voles, being slower where there were larger fluctuations in local population sizes. Conclusion: Our analyses estimated the traditional parameters of carrying capacities and return rates, but these are now seen as varying continuously over the landscape depending on habitat quality and the mechanisms of density dependence. The importance of our results lies in our demonstration that the effects of spatial and temporal heterogeneity must be accounted for if we are to have accurate predictive models for use in management and conservation. This is an area which until now has lacked an adequate theoretical framework and methodology.

Identification of Bartonella species in rodents, shrews and cats in Denmark: detection of two B-henselae variants, one in cats and the other in the long-tailed field mouse

Small mammals and stray cats were trapped in two areas of North Zealand, Denmark, and their blood cultured for hemotrophic bacteria. Bacterial isolates were recovered in pure culture and subjected to 16S rDNA gene sequencing. Bartonella species were isolated from five mammalian species: B. grahamii from Microtus agrestis (field vole) and Apodemus flavicollis (yellow-necked field mouse); B. taylorii from M. agrestis, A. flavicollis and A. sylvaticus (long-tailed field mouse); B. tribocorum from A. flavicollis; R vinsonii subsp. vinsonii from M. agrestis and A. sylvaticus; and B. birtlesii from Sorex vulgaris (common shrew). In addition, two variant types of B. henselae were identified: variant I was recovered from three specimens of A. sylvaticus, and B. henselae variant 11 from I I cats; in each case this was the only B. henselae variant found. No Bartonella species was isolated from Clethrionomys glareolus (bank vole) or Micromys minutus (harvest mouse). These results suggest that B. henselae occurs in two animal reservoirs in this region, one of variant I in A. sylvaticus, which may be transmitted between mice by the tick Ixodes ricinus, and another of variant 11 in cats, which may be transmitted by the cat flea (Ctenocephalides felis). To our knowledge, this is the first report of the occurrence of B. henselae and B. tribocorum in Apodemus mice.

How predation and landscape fragmentation affect vole population dynamics

Background: Microtine species in Fennoscandia display a distinct north-south gradient from regular cycles to stable populations. The gradient has often been attributed to changes in the interactions between microtines and their predators. Although the spatial structure of the environment is known to influence predator-prey dynamics of a wide range of species, it has scarcely been considered in relation to the Fennoscandian gradient. Furthermore, the length of microtine breeding season also displays a north-south gradient. However, little consideration has been given to its role in shaping or generating population cycles. Because these factors covary along the gradient it is difficult to distinguish their effects experimentally in the field. The distinction is here attempted using realistic agent-based modelling. Methodology/Principal Findings: By using a spatially explicit computer simulation model based on behavioural and ecological data from the field vole (Microtus agrestis), we generated a number of repeated time series of vole densities whose mean population size and amplitude were measured. Subsequently, these time series were subjected to statistical autoregressive modelling, to investigate the effects on vole population dynamics of making predators more specialised, of altering the breeding season, and increasing the level of habitat fragmentation. We found that fragmentation as well as the presence of specialist predators are necessary for the occurrence of population cycles. Habitat fragmentation and predator assembly jointly determined cycle length and amplitude. Length of vole breeding season had little impact on the oscillations. Significance: There is good agreement between our results and the experimental work from Fennoscandia, but our results allow distinction of causation that is hard to unravel in field experiments. We hope our results will help understand the reasons for cycle gradients observed in other areas. Our results clearly demonstrate the importance of landscape fragmentation for population cycling and we recommend that the degree of fragmentation be more fully considered in future analyses of vole dynamics.

The effects of landscape modifications on the long-term persistence of animal populations

Background: The effects of landscape modifications on the long-term persistence of wild animal populations is of crucial importance to wildlife managers and conservation biologists, but obtaining experimental evidence using real landscapes is usually impossible. To circumvent this problem we used individual-based models (IBMs) of interacting animals in experimental modifications of a real Danish landscape. The models incorporate as much as possible of the behaviour and ecology of four species with contrasting life-history characteristics: skylark (Alauda arvensis), vole (Microtus agrestis), a ground beetle (Bembidion lampros) and a linyphiid spider (Erigone atra). This allows us to quantify the population implications of experimental modifications of landscape configuration and composition. Methodology/Principal Findings: Starting with a real agricultural landscape, we progressively reduced landscape complexity by (i) homogenizing habitat patch shapes, (ii) randomizing the locations of the patches, and (iii) randomizing the size of the patches. The first two steps increased landscape fragmentation. We assessed the effects of these manipulations on the long-term persistence of animal populations by measuring equilibrium population sizes and time to recovery after disturbance. Patch rearrangement and the presence of corridors had a large effect on the population dynamics of species whose local success depends on the surrounding terrain. Landscape modifications that reduced population sizes increased recovery times in the short-dispersing species, making small populations vulnerable to increasing disturbance. The species that were most strongly affected by large disturbances fluctuated little in population sizes in years when no perturbations took place. Significance: Traditional approaches to the management and conservation of populations use either classical methods of population analysis, which fail to adequately account for the spatial configurations of landscapes, or landscape ecology, which accounts for landscape structure but has difficulty predicting the dynamics of populations living in them. Here we show how realistic and replicable individual-based models can bridge the gap between non-spatial population theory and non-dynamic landscape ecology. A major strength of the approach is its ability to identify population vulnerabilities not detected by standard population viability analyses.

Distribution and Specificity of Helminths in Microtine Rodents: Evolutionary Implications

The taxonomic status of anoplocephaline cestodes of microtine rodents has been reviewed. Of the genus Andrya Railliet, 1883, five species are considered valid: A. macrocephala Douthitt, 1915; A. primordialis Douthitt, 1915; A. montana Kirshenblat, 1941 ; A. arctica Rausch, 1952; A. bairdi Schad, 1954. Of the genus Paranoplocephala Luehe, 1910, six species are regarded as valid: P. omphalodes (Hermann, 1783); P. blanchardi (Moniez, 1891); P. infrequens (Douthitt, 1915); P. variabilis (Douthitt, 1915); P. lemmi Rausch, 1952; P. neofibrinus Rausch, 1952. Andrya caucasica Kirshenblat, 1938, and A. bialowizensis Soltys, 1949, are regarded as synonyms of A. macrocephala. Paranoplocephala brevis Kirshenblat, 1938, is regarded as a synonym of P. infrequens. Three species, A. macrocephala, P. omphalodes, and P. infrequens, are holarctic in distribution, occurring mainly in species of Microtus. The uniformity of microtine rodents as hosts for various helminths has been discussed. It is concluded that Dicrostonyx is the most isolated genus from this standpoint, having two nematodes which have not been recorded from members of other genera, and harboring few helminths in common with others. This agrees with Hinton's conclusions, based on morphological characters of Dicrostonyx. From the present concept of Pleistocene glaciations, it is concluded that P. omphalodes and P. infrequens reached the St. Matthew Islands, in Bering Sea, as parasites of a vole from which Microtus abbreviatus has evolved. It appears that this vole arrived on these islands before North America was invaded, in the late Pleistocene, by the palearctic M. oeconomus and Clethrionomys rutilus,/i>. The present known distribution of P. omphalodes in North America corresponds about to that of M. oeconomus on the continent.

The Chromosomal Complement of the Yellow-Cheeked Vole

The karyotype of Microtus xanthognathus (Leach) is described, based on material from one female and one male vole. The diploid chromosomal number was found to be 54, and the fundamental number 62. The metacentric X-chromosome was of medium size and averaged 6.6% of the haploid complement. The designated Y-chromosome was near acrocentric. The specific distinction of M. xanthognathus and Microtus chrotorrhinus (Miller) was confirmed by the recognition of major differences in karyotype and differences in fundamental number. The distributional history of M. xanthognathus is briefly discussed.

Notes on Microtine Rodents from the Brooks Range, Arctic Alaska

In connection with parasitological studies carried on during 1949 for the U.S. Public Health Service, the writer collected a considerable number of mammals from the northern edge of the "Endicott" section of the Brooks Range, in Arctic Alaska. The mammalian fauna of this region is poorly known, since apparently no previous collecting has been done here. About 200 microtine rodents were collected, mostly near Tolugak Lake (latitude 68° 24' N, longitude 151° 26' W), near the head of the Anaktuvuk River Valley. A few specimens were also taken at Umiat, on the Colville River, about 80 miles north of Tolugak Lake (latitude 69° 23' N, longitude 152° 10' W). Five species are represented: Clethrionomys rutilis dawsoni (Merriam), Microtus oeconomus macfarlani Merriam, Microtus miurus paneaki, n. subsp., Lemmus trimucronatus alascensis Merriam and Dicrostonyx groenlandicus rubricatus (Richardson). More complete details concerning their ecology and reproduction will be presented in a later paper, at which time the other mammals obtained will also be considered. The specimens have been deposited in the U.S. National Museum.

Dynamics of the Arctic Fox Population on St. Lawrence Island, Bering Sea

English abstract: We hypothesized that the arctic fox, Alopex lagopus (Linnaeus), population on St. Lawrence Island was cyclic and that its fluctuations in size. structure, and productivity were correlated with the relative size of the population of northern voles, Microtus oeconomus Pallas, the primary prey. Based on a nine-year study, we determined that the variations in size of the fox and vole populations were similar, but they both were of low amplitude and not closely correlated. The high pregnancy rate (mean, 86%/yr) and numbers of young conceived (mean, 11.5/pregnancy) did not vary significantly among years, probably because of the consistently abundant and diverse food supply available to the foxes. The age composition of the trappers' catch of foxes each winter also was comparatively stable, but it was closely correlated with the size of the vole population in the previous summer. The survival of the young foxes during the summer probably was dependent on the availability of the voles, The composition of the catch also appeared to be influenced by immigration of faxes from the adjacent continents via the pack ice. French abstract: Nous avons émis I'hypothèse que la population du renard arctique, Alopex lagopus (Linnaeus), sur I'île Saint Lawrence était cyclique el que les fluctuations concernant sa tailIe, sa structure et sa productivité étaient corrélées à la taille relative de la population du campagnol nordique. Microtus oeconomus Pallas, sa principale proie. En nous appuyant sur une étude menée sur neuf ans, nous avons déterminé que les variations dans la taille des populations du renard et du campagnol étaient semblables. mais que toutes deux avaient une faible amplitude et n'étaient pas corrélées de façon étroite. Le taux de grossesse élevé (moyenne 86 p. cent/an) et Ie nombre dc petits conçus (moyenne 11,5/grossesse) ne variaient pas de façon significative au cours des ans, probablement à cause de I'abondance et de la variété de sources de nourriture pour les renards. La composition d'âge des prises des trappeurs était également stable d'un hiver à I'autre, mais elle était corrélée de façon étroite avec la taille de la population dc campagnols au cours de I'été précédent. La survie des renardeaux au cours de I'été dépendait probablement de la disponibilité des campagnols. La composition des prises semblait aussi être influencée par I'immigration des renards venant des terres continentales adjacentes par la voie de la banquise.

The Specific Status of the Narrow-Skulled Vole (Subgenus Stenocranius Kashchenko) in North America

On morphological and zoogeographical grounds, discussed in the present paper, it is concluded that the narrow-skulled vole in North America, previously designated Microtus (Stenocranius) miurus Osgood, is conspecific with the Eurasian M. (Stenocranius) gregalis Pallas. Fourteen subspecies in Eurasia and 5 in North America are now recognized, but it is probable that the number in Eurasia will be reduced through future investigation. The Eurasian subspecies of this vole comprise two major groups, of which one occupies the tundra zone and the other occurs across central Asia below latitude 60° N; their geographic ranges are largely separate but evidently become confluent in northeastern Siberia. The members of the northern group of Eurasian subspecies and the North American forms are closely related; the present distribution of the latter indicates post-glacial dispersal from the Amphiberingian Refugium. It is believed that the tundra-inhabiting voles in Eurasia likewise survived the Pleistocene glaciations in northern refugia, while the members of the southern group of subspecies probably represent populations that survived south of the limits of the continental glaciers. The ranges of the two Eurasian groups probably have become confluent during post-glacial time in northeastern Siberia as a result of the southward spread of the northern forms. At least, the subspecies having the intervening range closely resembles members of the northern group. Some of the ecological and ethological characteristics of these voles are briefly discussed. The chromosome number of one of the North American subspecies of narrow-skulled vole was determined to be 54; this is the first time that the chromosomes of a member of the subgenus Stenocranius have been investigated. A karyogram has been included. German abstract: Auf morphologischen und tiergeographischen Grundlagen, die in dieser Arbeit besprochen wurden, ist festgestellt worden, daß die schmalschädlige Wiihlmaus in Nordamerika, friiher Microtus (Stenocranius) miurus Osgood bezeichnet, mit der palaearktischen Art M. (Stenocranius) gregalis Pallas identisch ist. Zur Zeit gelten 14 Unterarten in Eurasien und 5 in Nordamerika als unterscheidbar; vermutlich aber wird die Zahl der palaearktischen Unterarten durch eingehendere Untersuchungen künftig vermindert werden. Auf Grund ihrer Verbreitung bilden die palaearktischen Unterarten zwei beinahe vollständig getrennte Gruppen. Die Wühlmäuse der nördlichen Gruppe bewohnen die Tundrazone, während die Vertreter der zweiten Gruppe über Mittelasien südlicher als 60° N.B. verbreitet sind. Die Verbreitungsgebiete der zwei Gruppen verbinden sich anscheinend. Die nordamerikanischen schmalschädligen Wühlmäuse sind mit den in der Tundrazone vorkommenden palaearktischen Formen nahe verwandt; sie haben sich wahrscheinlich während der Postglazialzeit aus dem Amphiberingschen Refugium verbreitet. Möglicherweise überlebten die tundrabewohnenden Wühlmäuse Eurasiens die Eiszeit ebenfalls in vereinzelten Refugien in Nordostsibirien, während die Formen der südlichen Gruppe sie jenseits der Grenzen des Festlandsgletschers überlebten. Wahrscheinlich wurden die zwei Verbreitungsgebiete dieser Art in Eurasien erst während der Postglazialzeit durch das Vordringen der nordischen Formen verbunden, da eine nähere Verwandtschaft zwischen den nördlichen und der dazwischenliegenden Unterart besteht. Einige ökologische und ethologische Eigentümlichkeiten dieser Wühlmäuse werden kurz besprochen. Es wurde festgestellt, daß eine der nordamerikanischen Unterarten der schmalschädligen Wühlmaus 54 Chromosomen hat; sie ist der einzige Vertreter der Untergattung Stenocranius, dessen Chromosomen untersucht worden sind.

BURROW-BUILDING STRATEGIES AND HABITAT USE OF VOLES IN PACIFIC NORTHWEST ORCHARDS

Seidel and Booth (1960) wrote that the "life histories of the genus Microtus are not numerous in the literature." In support of his observation he cited 6 publications, all dated between 1891 and 1953. Since then the literature has exploded with a proliferation of publications. An international literature review recently revealed over 3,500 citations for the genus. When Pitymys and Clethrionomys are included another 350 and 1,880, respectively, were found. Over the last 10 years approximately 3 new publications on voles appeared every 4 days; a significant output for what some would consider such an insignificant species. Most of the publications were the result of graduate research projects on population dynamics and species ecology. As such, many do not explore more than the rudimentary ecological relationships between the animal and their environments. Unfortunate, as well, is that all but one confined their observations to only a small part of their total environment. For many of these animals, their life underground may be more important for their survival than that above ground. Trapping studies conducted by Godfrey and Askham (1988) with permanently placed pitfall live traps in orchards revealed a significant inverse population fluctuation during the year. During the winter, when populations are expected to decrease, as many as 6 to 8 mature Microtus montanus were collected at any 1 time in the traps after several centimeters of snow accumulation. During the summer, when populations are expected to increase, virtually no animals were collected in the traps. According to current population dynamics theory, greater numbers of animals, including increasingly larger numbers of immature members of the community, should appear in any sample between the onset of the breeding period, generally in the spring, taper off during the latter part of the production season, usually late summer, and then decline as the limiting factors begin to take effect. For us, we trapped more animals in the fall and early winter than we did during the spring and summer. A review of the above literature did little to answer our question. Where are the animals going during the summer and why?