Microtus oeconomustundra vole

Geographic Range

Microtus oeconomus (pronounced e-KON-uh-mus), commonly known as the tundra or root vole, is one of only four Holarctic rodents and the only species of Microtus that is found on all northern continents. In the Nearctic, it is found from the extreme northern edge of British Columbia northward to the Arctic coast, and from western Northwest Territories, through Yukon Territory and nearly all of Alaska. It has the northernmost distribution of any Microtus species in North America, with occurrences to around 71°N in Alaska. In the Palearctic, tundra voles occupy a somewhat broader range, extending from Scandinavia and the Netherlands in the west, throughout northern Europe and Asia to Siberia in the east, and south as far as Mongolia. (Hoffmann and Koeppl, 1985; Musser and Carleton, 2005)

Habitat

Microtus oeconomus inhabits the tundra and taiga biomes and prefers mesic sedge meadows with abundant cover. It is commonly found along the edges of lakes and streams where this and similar habitats occur. Although this habitat usually occurs in relative lowlands, tundra voles are also found in wetter areas of mountainous regions at elevations of up to 2,500 meters. In the Palearctic, M. oeconomus is found in a broader range of habitats, commonly inhabiting mixed forest, taiga, and forest-steppe biomes with similar conditions. Habitat is most strongly selected on the basis of food quality and amount of cover provided. (Bergman and Krebs, 1993; Gromov and Polyakov, 1992; Hoffmann and Koeppl, 1985; Tast, 1966)

  • Range elevation
    0 to 2,500 m
    0.00 to ft

Physical Description

Microtus oeconomus is a medium-sized vole and varies in color dorsally from darker brown or grayish to lighter shades of cinnamon, rusty brown, or ocher. All color variations display a mixture of black-tipped hairs in the dorsal pelage. Sides are somewhat paler and the ventral surface is much lighter, from buff or ash gray to white. The tail is slightly to strongly dichromatic, being darker above and paler below. There is a fair amount of color variation in the species, which can often be related to habitat and geography. Tundra voles are generally lighter colored in more open habitats, and have yellow or grayish tones in tundra or forested habitats, respectfully. (Gromov and Polyakov, 1992; Hall, 1981; MacDonald, 2003)

Tundra voles display sexual size dimorphism, with adult males roughly 30% larger than females. The size and weight of the species is also variable relative to latitude and geography, especially so in the Palearctic. Individuals in populations at higher latitudes are generally larger and have proportionally smaller tails as a possible adaptation to colder temperatures. In addition, insular subspecies are generally larger than continental counterparts. Weight ranges from 25 to 80 g, with an average around 50 g. Total length ranges from 118 mm in the Old World, to 226 mm in the larger subspecies found in the New World. The tail is relatively short, generally being less than 30% of the total length. The name Microtus means “small ear” and refers to the short ears hidden in pelage that are a characteristic common to members of the genus. The dental formula for M. oeconomus follows the basic pattern for all Microtus: incisors 1/1, canines 0/0, premolars 0/0, molars 3/3. (Bondrup-Nielsen and Ims, 1990; Gromov and Polyakov, 1992; Hall, 1981; Lance and Cook, 1998; MacDonald, 2003; Nagorsen, 2002; Ringens, et al., 1977; Wilson and Ruff, 1999)

Microtus oeconomus maintains a relatively high basal metabolic rate (compared to most rodents) of 12.5 mL O2 per gram of body mass per hour. This is due primarily to the high thermoregulatory demands of small endothermic mammals in cold environments. (Bozinovic and Rosenmann, 1989; Ringens, et al., 1977)

  • Sexual Dimorphism
  • male larger
  • Range mass
    25 to 80 g
    0.88 to 2.82 oz
  • Average mass
    50 g
    1.76 oz
  • Range length
    118 to 226 mm
    4.65 to 8.90 in
  • Average basal metabolic rate
    12.5 cm3.O2/g/hr
  • Average basal metabolic rate
    0.566 W
    AnAge

Reproduction

Various mating systems have been observed in Microtus oeconomus, including promiscuous, multi-partner groups, polygyny, and monogamy. The mating system adopted is determined primarily by the general spacing patterns of a specific population, which are in turn influenced by factors such as habitat, season, and population densities (discussed below). In general, a slightly biased female (57%) to male (43%) ratio favors a system of polygyny in tundra voles. Accordingly, single-male polygyny and multi-male promiscuity are the most common systems observed during normal breeding season. However, facultative monogamy is adopted in isolated territories or patchy habitats with low densities and can be relatively long-lasting. (Aars and Ims, 2002; Aars and Ims, 2002; Lambin, et al., 1992; Tast, 1966; Viitala, 1994)

Female tundra voles are polyestrous and often produce two to three litters per year. The breeding season generally lasts from late April to September during years with peak densities, and is around one to two months shorter in years of low densities. Winter breeding has been observed very rarely, and is attributed to unusually warm summers resulting in late fall re-growth. (Kaikusalo and Tast, 1984; Nowak, 1999; Whitney, 1976; Wilson and Ruff, 1999)

Litter sizes of Microtus oeconomus are relatively large for the genus and usually range from four to eight offspring, with an average of 6.9. Litter size increases with age of the female, and over-wintered females are usually much more productive than spring-born females. Litters produced by northern populations of tundra voles are more female biased than those of southern populations. This bias is thought to be influenced by several factors both before and after birth, but this is not well understood. The gestation period for M. oeconomus lasts 20 to 21 days. Neonates weigh about 3 g at birth, have no external hair (except vibrissae), and are essentially poikilothermic. The newborns are very altricial and depend heavily on the mother during development. After five days the young are covered in hair and their eyes open 11 to 13 days after birth. They develop quickly and are weaned after 18 days, at which point they are fairly independent. Maximum size is reached as early as two months after weaning. Sexual maturation is reached in three weeks by females, whereas males do not usually mature until six to eight weeks. This sex-based delay in maturation serves as a defense against inbreeding in that females are almost always fertilized before their male siblings are able to mate. (Aars, et al., 1995; Feldhamer, et al., 2003; Ims, 1994; Innes, 1978; Ru-Yung and Jing-Xiang, 1987; Wilson and Ruff, 1999)

  • Breeding interval
    Female tundra voles breed several times in a year.
  • Breeding season
    Breeding generally occurs from April to September. Very rarely, winter breeding has been observed
  • Range number of offspring
    4 to 8
  • Average number of offspring
    6.9
  • Average number of offspring
    7
    AnAge
  • Range gestation period
    20 to 21 days
  • Average weaning age
    18 days
  • Average time to independence
    18 days
  • Average age at sexual or reproductive maturity (female)
    3 weeks
  • Average age at sexual or reproductive maturity (female)
    Sex: female
    30 days
    AnAge
  • Range age at sexual or reproductive maturity (male)
    6 to 8 weeks

Females provide the greater investment in the offspring, including nest construction, protection, and care of the young. However, increases in paternal investment in southern populations of tundra voles have been observed. It has been suggested that this is a result of longer foraging times required by southern females during lactation due to increased resource limitation at lower latitudes. (Ims, 1997)

  • Parental Investment
  • altricial
  • pre-fertilization
    • provisioning
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female

Lifespan/Longevity

There is insufficient research on Microtus oeconomus for a definitive lifespan to be established. However, relatively few adults survive much longer than one year in the wild, and the maximum age for other species of Microtus is commonly around two years. (Feldhamer, et al., 2003)

Behavior

Microtus oeconomus exhibits a polyphasic activity pattern, with an average of seven bouts of activity per 24 hour time period. Although this type of ultradian rhythm is most prevalent, secondary crepuscular and circadian patterns have also been observed. In general, activity is higher during the day than at night and highest during the twilight hours. Activity levels also increase with decreases in temperature and are relatively unaffected by humidity and wind due to the protectiveness of preferred habitat types. Tundra voles are active throughout the year and do not hibernate or engage in bouts of torpor. (Feldhamer, et al., 2003; Halle, 1995; Feldhamer, et al., 2003; Halle, 1995)

Nests are constructed out of coarse vegetation and lined with finer plant materials, such as sedges. Shallow burrows are dug by M. oeconomus in the soil and vegetation layers. These burrows are primarily used as traveling corridors along with well-developed runways created above ground and in natural features such as permafrost cracks. Tundra voles generally run in the open and walk under cover, lacking the bounding progression seen in most mammals. (Feldhamer, et al., 2003; Wilson and Ruff, 1999)

  • Range territory size
    3900 (high) m^2

Home Range

The home range, territory size, and spacing system of Microtus oeconomus vary widely depending on habitat, population density, season, and other factors. However, several trends have been observed. When food resources are of high quality and abundant, populations become increasingly clumped around these resources and males will display mutual territorial behavior. This behavior is also seen in response to periods of synchronous female breeding, where defense of mates becomes a higher priority. (Ims, 1987; Ostfeld, 1985)

Males generally occupy and utilize much larger territories than females, and this territory size is positively correlated with body size. Territories can range up to 3900 m^2 for males, but average 804 m^2 and have little overlap occurring between them. Females occupy much smaller territories, averaging 377 m^2, but are more philopatric than males and may share a territory with several other related females. The majority of male territories contain several female territories, and the resident male will often defend the breeding rights to all of the females contained within. Males travel approximately twice as far as females and may utilize an area as large as 12,000 m^2 in a single day. (Gliwicz, 1997; Lambin, et al., 1992; Litvin and Karaseva, 1968; Tast, 1966)

Communication and Perception

Olfaction is presumably the most important and well-developed sense in Microtus oeconomus. Scents are commonly used by the species to identify familiar or unfamiliar individuals and to determine their age, sex, reproductive condition, diet, or social status. Conspicuous hip and preputial glands used for scent production are located on both sexes of adult tundra voles. The glands are largest in males and older individuals, and are only present in females at higher latitudes. Scent marking behaviors such as scratching or rubbing are also displayed by M. oeconomus. Hearing is well-developed in tundra voles and they may occasionally use vocalizations for communication, although little is known about this behavior. (Feldhamer, et al., 2003; Hall, 1981)

Food Habits

Tundra voles are strictly vegetarian and preferentially feed on sedges (Carex sp. and Eupharium sp.) that grow in their favored habitat of wet, marshy tundra. Sedges make up around 70 to 80% of their diet, with the remainder comprising herbs, mosses, lichen, and small woody shrubs. These percentages vary seasonally, and a 30% decrease in sedge consumption is common in winter, with mosses and lichens becoming a relatively major component of the diet. Other plant foods that are preferred when available include Equisetum sp., Dryas integrifolia, Salix sp. and various grasses. (Batzli and Lesieutre, 1991; Tast, 1966)

In the fall, Microtus oeconomus creates large caches of stored seeds and rhizomes to supplement its winter diet. Native peoples of North America occasionally sought out these caches in order to obtain large quantities of desired foods, such as licorice root. (Wilson and Ruff, 1999)

  • Plant Foods
  • leaves
  • roots and tubers
  • wood, bark, or stems
  • bryophytes
  • lichens

Predation

Microtus oeconomus is an important prey source for many carnivores. During cycles of peak density, they have been known to comprise the majority of the diet for species such as the arctic fox (Vulpes lagopus) in Alaska. Other known terrestrial predators of M. oeconomus include weasels, martens, red foxes, and wolverines. In addition, numerous avian species prey on tundra voles, including owls, falcons, hawks, jaegers, gulls, and shrikes. (Anthony, et al., 2000; Sundell, et al., 2004; Wilson and Ruff, 1999)

  • Anti-predator Adaptations
  • cryptic

Ecosystem Roles

Populations of Microtus oeconomus experience cyclical fluctuations, reaching peak densities of 70 to 80 voles per hectare. As explained above, during these peak density years, tundra voles provide a large food base for many predator species. When present in such large numbers, they can also significantly alter the biological production of the tundra ecosystem through their foraging activities. Tundra voles share their habitat preferences with several similar species (e.g., M. longicaudus, M. pennsylvanicus, M. agrestis and M. xanthognathus) but generally either avoid competition through niche specialization, or out-compete the other species, as in the case of M. agrestis. (Tast, 1968; Whitney, 1976)

Tundra voles are host to a variety of internal and external parasites. Endoparasites include cestodes (Echinococcus sp., Paranoplocephala sp., and Taenia sp.), nematodes (Heligmosomoides sp.), and trematodes (Quinqueserialis nassalli). Ectoparasites include fleas (Siphonaptera sp.), lice (Polyplax sp.), and ticks (Ixodes angustus). (Timm, 1985)

Commensal/Parasitic Species

Economic Importance for Humans: Positive

Microtus oeconomus is commonly considered an ideal species for scientific research due to its adaptability to laboratory settings, quick development, and rapid population growth. Many important hypotheses have been tested using tundra voles as model species. As discussed above, tundra voles also provide an important food base for many carnivores (such as mink and foxes) that are highly valued for their fur. (Feldhamer, et al., 2003; Gromov and Polyakov, 1992)

  • Positive Impacts
  • research and education

Economic Importance for Humans: Negative

Tundra voles are known carriers of several diseases. Of most importance to humans is the role of Microtus oeconomus as a carrier of the bacterium (Francisella tularensis) that causes tularemia in humans. This disease is transmitted by direct contact and is fatal in about 7% of untreated human cases. Additionally, at peak densities they can compete with livestock for forage and cause damage to trees by gnawing on the roots. (Feldhamer, et al., 2003)

  • Negative Impacts
  • injures humans
    • carries human disease

Conservation Status

Overall, Microtus oeconomus is a relatively stable species and not currently listed as threatened or endangered on any listing. It is also categorized by the IUCN as a species of “least concern”. However, insufficient knowledge of several isolated subspecies has resulted in their categorization of “data deficient” by the IUCN. These subspecies include M. o. amakensis, M. o. elymocetes, M. o. innuitus, M. o. popofensis, M. o. punukensis, and M. o. sitkensis. Furthermore, two subspecies, M. o. mehelyi and M. o. arenicola, are categorized by the IUCN as “vulnerable” and “critically endangered”, respectively. Population declines in some subspecies have been attributed to habitat deterioration. In the case of M. o. arenicola, competition with M. arvalis has additionally contributed to population declines on some islands. (Gippoliti, 2006; Gippoliti, 2006)

Other Comments

Microtus oeconomus is most commonly described as comprising 10 subspecies in the Nearctic and 15 or more subspecies in the Palearctic. However, these numbers are subject to frequent debate and revision. New evidence suggests that Microtus oeconomus is a relatively recent colonizer of the Nearctic, and most likely crossed the Beringian land bridge during the Wisconsian (ca. 80,000-100,000 years ago) or Illinoian (ca. 130,000-300,000 years ago) glacial period. Recent research based on molecular evidence suggests that the species can be divided into four major clades. (Brunhoff, et al., 2003; Galbreath and Cook, 2004; Hall, 1981; Lance and Cook, 1998)

Contributors

Tanya Dewey (editor), Animal Diversity Web.

Chad Bieberich (author), University of Alaska Fairbanks, Link Olson (editor, instructor), University of Alaska Fairbanks.

Glossary

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

acoustic

uses sound to communicate

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

bog

a wetland area rich in accumulated plant material and with acidic soils surrounding a body of open water. Bogs have a flora dominated by sedges, heaths, and sphagnum.

chemical

uses smells or other chemicals to communicate

crepuscular

active at dawn and dusk

cryptic

having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.

diurnal
  1. active during the day, 2. lasting for one day.
endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

fertilization

union of egg and spermatozoan

folivore

an animal that mainly eats leaves.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

herbivore

An animal that eats mainly plants or parts of plants.

holarctic

a distribution that more or less circles the Arctic, so occurring in both the Nearctic and Palearctic biogeographic regions.

World Map

Found in northern North America and northern Europe or Asia.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

monogamous

Having one mate at a time.

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

polar

the regions of the earth that surround the north and south poles, from the north pole to 60 degrees north and from the south pole to 60 degrees south.

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

polygynous

having more than one female as a mate at one time

scent marks

communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

solitary

lives alone

stores or caches food

places a food item in a special place to be eaten later. Also called "hoarding"

tactile

uses touch to communicate

taiga

Coniferous or boreal forest, located in a band across northern North America, Europe, and Asia. This terrestrial biome also occurs at high elevations. Long, cold winters and short, wet summers. Few species of trees are present; these are primarily conifers that grow in dense stands with little undergrowth. Some deciduous trees also may be present.

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

tundra

A terrestrial biome with low, shrubby or mat-like vegetation found at extremely high latitudes or elevations, near the limit of plant growth. Soils usually subject to permafrost. Plant diversity is typically low and the growing season is short.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

References

Aars, J., H. Andreassen, R. Ims. 1995. Root voles: litter sex ratio variation in fragmented habitat. Journal of Animal Ecology, 64(4): 459-472.

Aars, J., R. Ims. 2002. Intrinsic and climatic determinants of population demography: the winter dynamics of tundra voles. Ecology, 83(12): 3449–3456.

Anthony, R., N. Barten, P. Seiser. 2000. Foods of arctic foxes (Vulpes lagopus) during winter and spring in western Alaska. Journal of Mammalogy, 81(3): 820-828.

Batzli, G., C. Lesieutre. 1991. The influence of high quality food on habitat use by arctic microtine rodents. Oikos, 60(3): 299-306.

Bergman, C., C. Krebs. 1993. Diet overlap of collared lemmings and tundra voles at Pearce-Point, Northwest-Territories. Canadian Journal of Zoology, 71(9): 1703-1709.

Bondrup-Nielsen, S., R. Ims. 1990. Reversed sexual size dimorphism in microtines: are females larger than males or are males smaller than females?. Evolutionary Ecology, 4: 261-272.

Bozinovic, F., M. Rosenmann. 1989. Maximum metabolic rate of rodents: physiological and ecological consequences on distributional limits. Functional Ecology, 3(2): 173-181.

Brunhoff, C., K. Galbreath, V. Fedorov, J. Cook, M. Jaarola. 2003. Holarctic phylogeography of the root vole (Microtus oeconomus): implications for late Quaternary biogeography of high latitudes. Molecular Ecology, 12: 957-968.

Feldhamer, G., B. Thompson, J. Chapman. 2003. Wild mammals of North America : biology, management, and conservation. 2nd ed. Baltimore, Maryland: Johns Hopkins University Press.

Galbreath, K., J. Cook. 2004. Genetic consequences of Pleistocene glaciations for the tundra vole (Microtus oeconomus) in Beringia. Molecular Ecology, 13: 135-148.

Gippoliti, S. 2006. "2006 IUCN Red List of Threatened Species" (On-line). Microtus oeconomus ssp. arenicola. Accessed December 04, 2006 at http://www.iucnredlist.org/search/details.php/40769/summ.

Gliwicz, J. 1997. Space use in the root vole: basic patterns and variability. Ecography, 20(4): 383-389.

Gromov, I., I. Polyakov. 1992. Voles (Microtinae). Washington D.C: Smithsonian Institution Libraries and the National Science Foundation.

Hall, R. 1981. The mammals of North America. 2nd ed. New York, NY: Wiley and Sons.

Halle, S. 1995. Diel pattern of locomotor activity in populations of root voles Microtus oeconomus . Journal of Biological Rhythms, 10(3): 211-224.

Hoffmann, R., J. Koeppl. 1985. Zoogeography. Pp. 84–115 in R Tamarin, ed. Biology of New World Microtus . Boston, MA: The American Society of Mammalogists.

Ims, R. 1997. Determinants of geographic variation in growth and reproductive traits in the root vole. Ecology, 78(2): 461-470.

Ims, R. 1994. Litter sex ratio variation in laboratory colonies of two geographically distinct strains of the root vole Microtus oeconomus . Ecography, 17(2): 141-146.

Ims, R. 1987. Male spacing systems in microtine rodents. The American Naturalist, 130(4): 475-484.

Innes, D. 1978. A reexamination of litter sizes in some North American microtines.. Canadian Journal of Zoology, 56(7): 1488-1496.

Kaikusalo, A., J. Tast. 1984. Winter breeding of microtine rodents at Kilpisjaervi, Finnish Lapland. Carnegie Mus. Nat. Hist. Spec. Publ., 10: 243-252.

Lambin, X., C. Krebs, B. Scott. 1992. Spacing system of the tundra vole (Microtus oeconomus) during the breeding season in Canada’s western arctic. Canadian Journal of Zoology, 70(10): 2068-2072.

Lance, E., J. Cook. 1998. Biogeography of tundra voles (Microtus oeconomus) of Beringia and the southern coast of Alaska. Journal of Mammalogy, 79(1): 53-65.

Litvin, V., E. Karaseva. 1968. A study of the daily migrations of Microtus oeconomus Pall. Voles by radioactive tagging. Zoologicheskii Zhurnal, 47(11): 1701-1706.

MacDonald, S. 2003. The small mammals of Alaska: A field handbook of the shrews and small rodents. Unpublished Draft.

Musser, G., M. Carleton. 2005. Superfamily Muroidea. Pp. 894-1522 in D Wilson, D Reeder, eds. Mammal Species of the World. Baltimore, Maryland: Johns Hopkins University Press.

Nagorsen, D. 2002. An identification manual to the small mammals of British Columbia. British Columbia, Canada: Ministry of Sustainable Resource Management.

Nowak, R. 1999. Walker's Mammals of the World. Baltimore, Maryland: Johns Hopkins University Press.

Ostfeld, R. 1985. Limiting resources and territoriality in microtine rodents. The American Naturalist, 126(1): 1-15.

Ringens, P., G. Folk, J. Berberich. 1977. Cold acclimation in the tundra vole. Acta Theriologica, 22(3): 67-74.

Ru-Yung, S., Z. Jing-Xiang. 1987. Postnatal development of thermoregulation in root vole and some comments on the index of homeothermy ability. Journal of Thermal Biology, 12(4): 267-272.

Sundell, J., O. Huitu, H. Henttonen, A. Kaikusalo, E. Korpimäki, H. Pietiäinen, P. Saurola, I. Hanski, E. Korpimaeki, H. Pietiaeinen. 2004. Large-scale spatial dynamics of vole populations in Finland revealed by the breeding success of vole-eating avian predators. Journal of Animal Ecology, 73(1): 167-178.

Tast, J. 1968. Influence of the root vole, Microtus oeconomus (Pallas), upon the habitat selection of the field vole, Microtus agrestis (L.), in northern Finland. Helsinki, Finland: Suomalainen Tiedeakatemia.

Tast, J. 1966. The root vole, Microtus oeconomus (Pallas), as an inhabitant of seasonally flooded land. Annales Zoologici Fennici, 3: 127-171.

Timm, R. 1985. Parasites. Pp. 455-528 in R Tamarin, ed. Biology of New World Microtus . Boston, MA: American Society of Mammalogists.

Viitala, J. 1994. Monogamy in free-living Microtus oeconomus . Annales Zoologici Fennici, 31(3): 343-345.

Whitney, P. 1976. Population ecology of two sympatric species of subarctic microtine rodents. Ecological Monographs, 46(1): 85-104.

Wilson, D., S. Ruff. 1999. The Smithsonian book of North American mammals. Washington, D.C.: Smithsonian Institution Press.