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The earliest undisputed arvicoline fossils are from the early Pliocene of northwest Asia, Europe, and North America. It is thought that arvicolines spread into southern Asia in the late Pliocene.

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Behavior

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Arvicolines are able to perceive tactile, visual, auditory, and chemical signals. Chemical signals are especially important for communication. Males, and sometimes females, mark their territories with secretions from their sebaceous flank glands. Also, some arvicolines are highly vocal and make a variety of chirping and chattering noises when disturbed or when engaged in a conflict with a conspecific.

Communication Channels: acoustic ; chemical

Other Communication Modes: scent marks

Perception Channels: visual ; tactile ; acoustic ; chemical

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Conservation Status

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In the subfamily Arvicolinae, the IUCN lists 23 lower risk species, 1 near threatened species (wood lemming, Myopus schisticolor), 4 vulnerable species (Central Kashmir vole, Alticola montosa, Mexican vole, Microtus mexicanus, Taiwan vole, Microtus kikuchii, and Japanese red-backed vole, Myodes andersoni), 2 endangered species (Alai mole vole, Ellobius alaicus and Baluchistan vole, Microtus transcaspicus), 3 critically endangered species (Wrangel lemming, Dicrostonyx vinogradovi, Evorsk vole, Microtus evoronensis, and Muisk vole, Microtus mujanensis), and 7 species lacking sufficient data to be ranked. Many arvicolines have restricted ranges, rendering them vulnerable to habitat loss.

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Comprehensive Description

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Arvicolinae is a large subfamily of cricetid rodents that are fairly uniform in appearance but diverse in their habits. There are 151 species in this family, in 28 genera. The genera are divided among 10 tribes.

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Benefits

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Some arvicolines carry diseases such as tularemia. Those that dwell in agricultural areas sometimes damage crops.

Negative Impacts: injures humans (carries human disease); crop pest

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Benefits

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Some arvicolines eat large quantities of insect larvae and therefore act as important controls on pest species. Others are hunted for their pelts or for food.

Positive Impacts: food ; body parts are source of valuable material; controls pest population

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Associations

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Arvicolines are primary and secondary consumers, and they provide a staple food source for many other species. The fossorial species turn over earth when they dig, and therefore may help to aerate the soil. Arvicolines are important for seed dispersal, and they impact forest regeneration by preying on tree seedlings (Manson et al. 2001). Because of their important role in forest dynamics, some are considered keystone species. Finally, because of their high reproductive output and cyclical boom and bust cycles in population numbers, the population dynamics of arvicolines often influences heavily the population dynamics of predators such as snowy owls and Canada lynx, and plant community composition through their grazing activity.

Ecosystem Impact: disperses seeds; soil aeration ; keystone species

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Trophic Strategy

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Arvicolines are primarily herbivorous, though some species are omnivorous. They consume leaves, grasses, forbs, roots, bulbs, bark, twigs, stems, pine needles, berries, nuts, seeds, lichen, fungi, insects, crayfish, mussels, and small fish. Some species cache food in their nests or burrows for use during times of shortage.

Foraging Behavior: stores or caches food

Primary Diet: carnivore (Piscivore , Insectivore , Eats non-insect arthropods, Molluscivore ); herbivore (Folivore , Frugivore , Granivore , Lignivore); omnivore ; mycophage

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Distribution

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The subfamily Arvicolinae has a Holarctic distribution. Arvicolines are found throughout North America from Guatemala northward, throughout Eurasia, in Japan, Taiwan, southwestern China, northern India, the Middle East including Asia Minor, and in Africa to Libya.

Biogeographic Regions: nearctic (Native ); palearctic (Native ); oriental (Native ); neotropical (Native )

Other Geographic Terms: holarctic

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Habitat

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Arvicolines inhabit a wide range of habitats within temperate, boreal, arctic, and montane biomes. These habitats include: dry and wet deciduous and coniferous forests, brushy or rocky mountain slopes, alpine meadows, prairies, steppes, agricultural fields, semidesert, cloud forests, tundra, riparian zones, lakes, marshes, and sphagnum bogs.

Habitat Regions: temperate ; tropical ; polar ; terrestrial

Terrestrial Biomes: tundra ; taiga ; savanna or grassland ; chaparral ; forest ; rainforest ; scrub forest ; mountains

Aquatic Biomes: lakes and ponds

Wetlands: marsh ; swamp ; bog

Other Habitat Features: agricultural

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Life Expectancy

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Most arvicolines only live for a few months in the wild. Captivity often extends the life span by several years.

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Morphology

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Arvicolines are medium to large muroid rodents, ranging in head and body length from 70 mm to over 300 mm, and ranging in tail length from 5 to 295 mm. The tail is always shorter than the head and body. Arvicolines weigh anywhere from 15 grams to over 1.8 kg. They have stout bodies with small, rounded ears, blunt snouts, and short legs. The eyes are relatively large. Adult males, and sometimes females, have large sebaceous glands on the rump, hips, flanks, or tail region. Most arvicolines have cursorially adapted feet, and some have long claws for digging. Arvicoline fur is usually thick and ranges from long to short and from smooth to harsh. In some species the texture changes with the seasons, becoming shorter and thinning out in the summer. The tail is covered with fur in most species, and sometimes bears a terminal tuft. The fur on the dorsal surface of arvicolines can be various shades of brown or gray, and in some species it has a distinct red or yellow cast. The fur on the ventral surface is pale brown, white, cream, buff, yellowish, or gray. Some species have bicolored tails that are darker above than below. There are polymorphic arvicoline populations, with two or more color morphs living in the same area.

The dental formula of arvicolines is 1/1, 0/0, 0/0, 3/3 = 16. The incisors may be orthodont, opisthodont, or proodont, and the molars may either be rooted or evergrowing. The molars bear a prismatic enamel pattern. Arvicolines have relatively large skulls. The squamosomastoid foramen is always present, and most have a stapedial foramen as well. The palatine process of the maxillary and the palatine are thickened dorsoventrally, in conjunction with the molars' large alveolar capsules. There are usually longitudinal furrows and ridges, as well as tiny perforations, in the bony palate. There are 13 thoracic vertebrae and six lumber vertebrae in the arvicoline vertebral column.

Arvicolines have stomachs that are either one- or two-chambered, and their large intestines and ceca are extremely complex. However, the small intestine is quite short. There is no supraorbital branch of the stapedial artery; instead, the infraorbital branch supplies blood to the orbits. Arvicolines have a diploid chromosome number between 18 and 62.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry ; polymorphic

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Associations

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Hawks, owls, snakes, and small mammalian carnivores are the main predators of arvicolines. Arvicolines can be quite vicious, gnashing their teeth and biting when threatened. Also, their neutral-colored fur probably keeps them somewhat camouflaged.

Known Predators:

  • hawks Accipitridae
  • owls Strigiformes
  • snakes Serpentes
  • mammalian carnivores Carnivora

Anti-predator Adaptations: cryptic

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Reproduction

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Most arvicolines are promiscuous, with males and females both having multiple mates. In some species, a copulatory plug forms and seals the female's reproductive tract, preventing subsequent males from successfully fertilizing the female's eggs. However, a few species, such as muskrats and prairie voles, live in monogamous pairs and share the responsibility of raising young. In fact, in captivity it has been shown that prairie voles stay with their parents for more than one breeding period and help raise their younger siblings (Gruder-Adams and Getz 1985).

Mating System: monogamous ; polygynandrous (promiscuous) ; cooperative breeder

Many arvicolines are capable of breeding year round, and some species even give birth to litters under snow cover. Others concentrate their reproductive efforts during the warmer months and breed from spring to autumn, with a peak in breeding occurring from late spring to early summer. In some species, ovulation is not spontaneous; rather, it is induced by the act of mating. Females are polyestrus, giving birth to anywhere from 1 to 7 litters per year. They often become impregnated again as soon as they give birth, due to a postpartum estrus. However, implantation of the embryo is delayed in some species while the female is lactating. Gestation from the time the embryo implants is 16 to 30 days. Litter sizes average 3 to 7 young, but some females have as few as one and as many as 13 young in a litter. The young are relatively precocial and develop rapidly, opening their eyes at 8 to 16 days and becoming weaned and independent at 12 to 35 days. Females often breed in the year that they are born, becoming sexually mature as early as 14 days. Males mature somewhat later than females.

Key Reproductive Features: iteroparous ; seasonal breeding ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; induced ovulation ; viviparous ; delayed implantation ; post-partum estrous

Most female arvicolines invest little in each individual offspring, instead employing a strategy of high reproductive output. They build nests in which they rear their litters and nurse their relatively precocial young for 12 to 35 days. Often females become highly aggressive when nursing, fiercely defending their litters against intruding males. Male parental care occurs in this group, with males of some species brooding the young in the nest or retrieving them when they wander away. Also, care of youngsters by older siblings has been reported for arvicolines kept in captivity (Gruder-Adams and Getz 1985).

Parental Investment: precocial ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Male, Female); pre-independence (Provisioning: Female, Protecting: Male, Female)

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Arvicolinae

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The Arvicolinae are a subfamily of rodents that includes the voles, lemmings, and muskrats. They are most closely related to the other subfamilies in the Cricetidae (comprising the hamsters and New World rats and mice[1]). Some authorities place the subfamily Arvicolinae in the family Muridae along with all other members of the superfamily Muroidea.[2] Some refer to the subfamily as the Microtinae (yielding the adjective "microtine")[3] or rank the taxon as a full family, the Arvicolidae.[4]

The Arvicolinae are the most populous group of Rodentia in the Northern Hemisphere. They often are found in fossil occlusions of bones cached by past predators such as owls and other birds of prey. Fossils of this group are often used for biostratigraphic dating of paleontological and archeological sites in North America and Europe.[5]

Description

The most convenient distinguishing feature of the Arvicolinae is the nature of their molar teeth, which have prismatic cusps in the shape of alternating triangles. These molars are an adaptation to a herbivorous diet in which the major food plants include a large proportion of abrasive materials such as phytoliths; the teeth get worn down by abrasion throughout the adult life of the animal and they grow continuously in compensation.[6]

Arvicolinae are Holarctic in distribution and represent one of only a few major muroid radiations to reach the New World via Beringia. (The others are the three subfamilies of New World rats and mice.) Arvicolines do very well in the subnival zone beneath the winter snowpack, and persist throughout winter without needing to hibernate. They are also characterized by extreme fluctuations in population numbers.

Most arvicolines are small, furry, short-tailed voles or lemmings, but some, such as Ellobius and Hyperacrius, are well adapted to a fossorial lifestyle. Others, such as Ondatra, Neofiber, and Arvicola, have evolved larger body sizes and are associated with an aquatic lifestyle.

Phylogeny

The phylogeny of the Arvicolinae has been studied using morphological and molecular characters. Markers for the molecular phylogeny of arvicolines included the mitochondrial DNA cytochrome b (cyb) gene [7] and the exon 10 of the growth hormone receptor (ghr) nuclear gene.[8] The comparison of the cyb and ghr phylogenetic results seems to indicate nuclear genes are useful for resolving relationships of recently evolved animals. As compared to mitochondrial genes, nuclear genes display several informative sites in third codon positions that evolve rapidly enough to accumulate synapomorphies, but slow enough to avoid evolutionary noise. Of note, mitochondrial pseudogenes translocated within the nuclear genome complicate the assessment of the mitochondrial DNA orthology, but they can also be used as phylogenetic markers.[9] Sequencing complete mitochondrial genomes of voles [10] may help to distinguish between authentic genes and pseudogenes.

The complementary phylogenetic analysis of morphological and molecular characters [8][11] suggests:

Some authorities have placed the zokors within the Arvicolinae, but they have been shown to be unrelated.

A 2021 study found Lemmini to be the most basal group of Arvicolinae. The study also found Arvicola to actually fall outside the tribe Arvicolini, and to be sister to the tribe Lagurini.[12]

Classification

Skull of a bank vole: Note the distinctive molar pattern characteristic of arvicolines.

Subfamily Arvicolinae - voles, lemmings, muskrats

The subfamily Arvicolinae contains eleven tribes, eight of which are classified as voles, two as lemmings, and one as muskrats.[13] Recent changes to the subfamily include disbanding genus Myodes in favor of genera Clethrionomys and Craseomys (and disbanding Myodini in favor of Clethrionomyini), moving most of the genera from Arvicolini to Microtini, and renaming Phenacomyini as Pliophenacomyini.[13]

Fossil species

See also

References

  1. ^ Steppan, S. J., R. A. Adkins, and J. Anderson. 2004. Phylogeny and divergence date estimates of rapid radiations in muroid rodents based on multiple nuclear genes. Systematic Biology, 53:533-553.
  2. ^ Musser, G. G. and M. D. Carleton. 2005. Superfamily Muroidea. Pp. 894-1531 in Mammal Species of the World a Taxonomic and Geographic Reference. D. E. Wilson and D. M. Reeder eds. Johns Hopkins University Press, Baltimore.
  3. ^ Nakao, Minoru; Yanagida, Tetsuya; Okamoto, Munehiro; Knapp, Jenny; Nkouawa, Agathe; Sako, Yasuhito; Ito, Akira (2010). "State-of-the-art Echinococcus and Taenia: Phylogenetic taxonomy of human-pathogenic tapeworms and its application to molecular diagnosis". Infection, Genetics and Evolution. Elsevier. 10 (4): 444–452. doi:10.1016/j.meegid.2010.01.011. ISSN 1567-1348. PMID 20132907.
  4. ^ McKenna, M. C. and S. K. Bell. 1997. Classification of Mammals above the Species Level. Columbia University Press, New York.
  5. ^ Klein, Richard (2009). The Human Career: Human Biological and Cultural Origins. London: The University of Chicago Press. p. 25. ISBN 978-0-226-43965-5.
  6. ^ Myers, P., R. Espinosa, C. S. Parr, T. Jones, G. S. Hammond, and T. A. Dewey. 2006.; "The Diversity of Cheek Teeth" ; The Animal Diversity Web (online). Accessed November 26, 2011 at http://animaldiversity.org.
  7. ^ Conroy CJ, Cook JA. 1999. MtDNA evidence for repeated pulses of speciation within arvicoline and murid rodents. J. Mammal. Evol. 6:221-245.
  8. ^ a b Galewski T, Tilak M, Sanchez S, Chevret P, Paradis E, Douzery EJP. 2006. The evolutionary radiation of Arvicolinae rodents (voles and lemmings): relative contribution of nuclear and mitochondrial DNA phylogenies. BMC Evol. Biol. 6:80.
  9. ^ Triant DA, DeWoody JA. 2008. Molecular analyses of mitochondrial pseudogenes within the nuclear genome of arvicoline rodents. Genetica 132:21-33.
  10. ^ Lin Y-H, Waddell PJ, Penny D. 2002. Pika and vole mitochondrial genomes increase support for both rodent monophyly and glires. Gene 294:119-129.
  11. ^ Robovsky J, Ricánková V, Zrzavy J. 2008. Phylogeny of Arvicolinae (Mammalia, Cricetidae): utility of morphological and molecular data sets in a recently radiating clade. Zool. Scripta 37:571–590.
  12. ^ Abramson, Natalia I.; Bodrov, Semyon Yu; Bondareva, Olga V.; Genelt-Yanovskiy, Evgeny A.; Petrova, Tatyana V. (2021-11-19). "A mitochondrial genome phylogeny of voles and lemmings (Rodentia: Arvicolinae): Evolutionary and taxonomic implications". PLOS ONE. 16 (11): e0248198. Bibcode:2021PLoSO..1648198A. doi:10.1371/journal.pone.0248198. ISSN 1932-6203. PMC 8604340. PMID 34797834.
  13. ^ a b Mammal Diversity Database (2023). "Mammal Diversity Database (Version 1.11) [Data set]". Zenodo. doi:10.5281/zenodo.7830771.
  14. ^ "Alexandromys alpinus". ASM Mammal Diversity Database. American Society of Mammalogists.
  15. ^ "Alexandromys shantaricus". ASM Mammal Diversity Database. American Society of Mammalogists.
  16. ^ "Chionomys lasistanius". ASM Mammal Diversity Database. American Society of Mammalogists.
  17. ^ "Chionomys stekolnikovi". ASM Mammal Diversity Database. American Society of Mammalogists.
  18. ^ Golenishchev, F. N.; Malikov, V. G.; Bannikova, A. A.; Zykov, A. E.; Yiğit, N.; Çolak, E. (2022). "Diversity of snow voles of the "nivalis" group (Chionomys, Arvicolinae, Rodentia) in the eastern part of the range with a description of a new species". Russian Journal of Theriology. 21 (1): 1–12. doi:10.15298/rusjtheriol.21.1.01. S2CID 250649779.

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Arvicolinae: Brief Summary

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The Arvicolinae are a subfamily of rodents that includes the voles, lemmings, and muskrats. They are most closely related to the other subfamilies in the Cricetidae (comprising the hamsters and New World rats and mice). Some authorities place the subfamily Arvicolinae in the family Muridae along with all other members of the superfamily Muroidea. Some refer to the subfamily as the Microtinae (yielding the adjective "microtine") or rank the taxon as a full family, the Arvicolidae.

The Arvicolinae are the most populous group of Rodentia in the Northern Hemisphere. They often are found in fossil occlusions of bones cached by past predators such as owls and other birds of prey. Fossils of this group are often used for biostratigraphic dating of paleontological and archeological sites in North America and Europe.

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