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.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. Some refer to the subfamily as the Microtinae (yielding the adjective "microtine") or rank the taxon as a full family, the Arvicolidae.McKenna, M. C. and S. K. Bell. 1997. Classification of Mammals above the Species Level. Columbia University Press, New York.
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.
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.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 https://animaldiversity.org.
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 Conroy CJ, Cook JA. 1999. MtDNA evidence for repeated pulses of speciation within arvicoline and murid rodents. J. Mammal. Evol. 6:221-245. and the exon 10 of the growth hormone receptor (ghr) nuclear gene.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. 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.Triant DA, DeWoody JA. 2008. Molecular analyses of mitochondrial pseudogenes within the nuclear genome of arvicoline rodents. Genetica 132:21-33. Sequencing complete mitochondrial genomes of voles 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. may help to distinguish between authentic genes and pseudogenes.
The complementary phylogenetic analysis of morphological and molecular characters 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. suggests: Ellobius, Prometheomys, and Lagurus are among the most basal arvicolines. Dicrostonyx, Phenacomys, and Arborimus may form a clade. Core arvicolines include four subclades: Lemmini: Synaptomys, Lemmus, Myopus Myodini: Eothenomys, Myodes Arvicolini: Arvicola Microtini: Alexandromys, Chionomys, Hyperacrius, Lasiopodomys, Lemmiscus, Microtus, Mictomicrotus, Neodon, Proedromys, Stenocranius, Volemys Ondatra and Dinaromys positions are uncertain, probably compromised by the convergent evolution of morphological characters.
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.
Classification
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. 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.
Tribe Arvicolini Genus Arvicola - water voles European (or Northern) water vole (Arvicola amphibius or Arvicola terrestris) Southwestern (or southern) water vole (Arvicola sapidus) Montane water vole (Arvicola scherman) Tribe Microtini Genus Alexandromys Alpine vole (Alexandromys alpinus) Evorsk vole (Alexandromys evoronensis) Reed vole (Alexandromys fortis) Taiwan vole (Alexandromys kikuchii) Lacustrine vole (Alexandromys limnophilus) Maximowicz's vole (Alexandromys maximowiczii) Middendorff's vole (Alexandromys middendorffi) Mongolian vole (Alexandromys mongolicus) Japanese grass vole (Alexandromys montebelli) Muya Valley vole (Alexandromys mujanensis) Tundra vole or root vole (Alexandromys oeconomus) Sakhalin vole (Alexandromys sachalinensis) Shantar vole (Alexandromys shantaricus) Genus Chionomys - snow voles Caucasian snow vole, Chionomys gud European snow vole, Chionomys nivalis Robert's snow vole, Chionomys roberti Lazistan snow vole (Chionomys lasistanius) Stekolnikov's snow vole (Chionomys stekolnikovi) Genus Hyperacrius - voles from Pakistan True's vole, Hyperacrius fertilis Murree vole, Hyperacrius wynnei Genus Lasiopodomys Brandt's vole, Lasiopodomys brandtii Mandarin vole, Lasiopodomys mandarinus Genus Lemmiscus Sagebrush vole, Lemmiscus curtatus Genus Microtus - voles Insular vole, M. abbreviatus California vole, M. californicus Rock vole, M. chrotorrhinus Long-tailed vole, M. longicaudus Mexican vole, M. mexicanus Singing vole, M. miurus North American water vole, M. richardsoni Zempoaltépec vole, M. umbrosus Taiga vole, M. xanthognathus Subgenus Microtus Field vole, M. agrestis Anatolian vole, M. anatolicus Common vole, M. arvalis Cabrera's vole, M. cabrerae Doğramaci's vole, M. dogramacii Elbeyli vole, M. elbeyli Günther's vole, M. guentheri Harting's vole, M. hartingii Tien Shan vole, M. ilaeus Persian vole, M. irani Mediterranean field vole, M. lavernedii Turkish vole, M. lydius Kerman vole, M. kermanensis Southern vole, M. levis Paradox vole, M. paradoxus Qazvin vole, M. qazvinensis Portuguese field vole, M. rosianus Schidlovsky's vole, M. schidlovskii Social vole, M. socialis European pine vole, M. subterraneus Transcaspian vole, M. transcaspicus Subgenus Blanfordimys Afghan vole, M. afghanus Bucharian vole, M. bucharicus Juniper vole, M. juldaschi Subgenus Terricola Bavarian pine vole, M. bavaricus Calabria pine vole, M. brachycercus Daghestan pine vole, M. daghestanicus Mediterranean pine vole, M. duodecimcostatus Felten's vole, M. felteni Liechtenstein's pine vole, M. liechtensteini Lusitanian pine vole, M. lusitanicus Major's pine vole, M. majori Alpine pine vole, M. multiplex Sicilian pine vole, M. nebrodensis Savi's pine vole, M. savii Tatra pine vole, M. tatricus Thomas's pine vole, M. thomasi Subgenus Mynomes Gray-tailed vole, M. canicaudus Western meadow vole, M. drummondi Florida salt marsh vole, M. dukecampbelli Montane vole, M. montanus Creeping vole, M. oregoni Eastern meadow vole, M. pennsylvanicus Townsend's vole, M. townsendii Subgenus Pitymys Guatemalan vole, M. guatemalensis Tarabundí vole, M. oaxacensis Woodland vole, M. pinetorum Jalapan pine vole, M. quasiater Subgenus Pedomys Prairie vole, M. ochrogaster Subgenus Hyrcanicola Schelkovnikov's pine vole, M. schelkovnikovi Genus Mictomicrotus Liangshan vole, Mictomicrotus liangshanensis Genus Neodon - mountain voles Bershula mountain vole (Neodon bershulaensis) Bomi mountain vole (Neodon bomiensis) Chayu mountain vole (Neodon chayuensis) Clarke's vole (Neodon clarkei) Forrest's mountain vole (Neodon forresti) Plateau vole (Neodon fuscus) Chinese scrub vole (Neodon irene) Blyth's vole (Neodon leucurus) Liao Rui's mountain vole (Neodon liaoruii) Linzhi mountain vole (Neodon linzhiensis) Medog mountain vole (Neodon medogensis) Namchabarwa mountain vole (Neodon namchabarwaensis) Nepalese mountain vole (Neodon nepalensis) Nyalam mountain vole (Neodon nyalamensis) Shergyla mountain vole (Neodon shergylaensis) Sikkim mountain vole (Neodon sikimensis) Genus Proedromys Duke of Bedford's vole (Proedromys bedfordi) Genus Stenocranius Narrow-headed vole (Stenocranius gregalis) Radde's vole (Stenocranius raddei) Genus Volemys Szechuan vole (Volemys millicens) Marie's vole (Volemys musseri) Tribe Dicrostonychini - collared lemmings Genus Dicrostonyx Northern collared lemming, D. groenlandicus Nelson's collared lemming, D. nelsoni Ungava collared lemming, D. hudsonius Ogilvie Mountains collared lemming, D. nunatakensis Richardson's collared lemming, D. richardsoni Arctic lemming, D. torquatus Unalaska collared lemming, D. unalascensis Tribe Ellobiusini - mole voles Genus Ellobius - mole voles Alai mole vole, E. alaicus Southern mole vole, E. fuscocapillus Transcaucasian mole vole, E. lutescens Northern mole vole, E. talpinus Zaisan mole vole, E. tancrei Tribe Lagurini Genus Eolagurus Yellow steppe lemming, E. luteus Przewalski's steppe lemming, E. przewalskii Genus Lagurus Steppe lemming, L. lagurus Tribe Lemmini Genus Lemmus - true lemmings Amur lemming, L. amurensis Norway lemming, L. lemmus Beringian lemming, L. nigripes East Siberian lemming, L. paulus West Siberian lemming, L. sibiricus Canadian lemming, L. trimucronatus Genus Myopus Wood lemming, M. schisticolor Genus Synaptomys - bog lemmings Northern bog lemming, S. borealis Southern bog lemming, S. cooperi Tribe Clethrionomyini Genus Alticola - voles from Central Asia Subgenus Alticola White-tailed mountain vole, A. albicauda Silver mountain vole, A. argentatus Gobi Altai mountain vole, A. barakshin Central Kashmir vole, A. montosa Royle's mountain vole, A. roylei Mongolian silver vole, A. semicanus Stolička's mountain vole, A. stoliczkanus Tuva silver vole, A. tuvinicus Subgenus Aschizomys Lemming vole, A. lemminus Large-eared vole, A. macrotis Lake Baikal mountain vole, A. olchonensis Subgenus Platycranius Flat-headed vole, A. strelzowi Genus Caryomys Ganzu vole, C. eva Kolan vole, C. inez Genus Eothenomys - voles from East Asia Kachin red-backed vole, E. cachinus Pratt's vole, E. chinensis Southwest China vole, E. custos