Arvicolinae
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 or rank the taxon as a full family, the Arvicolidae.[3]
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 archeological sites in North America and Europe.[4]
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.[5]
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 size.
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 a larger body size and associated with an aquatic lifestyle.
Phylogeny
The phylogeny of Arvicolinae has been studied using morphological and molecular characters. Markers for the molecular phylogeny of arvicolines included the mitochondrial DNA cytochrome b (cyb) gene [6] and the exon 10 of the growth hormone receptor (ghr) nuclear gene.[7] The comparison of the cyb and ghr phylogenetic results seems to indicate that 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.[8] Sequencing complete mitochondrial genomes of voles [9] may help to distinguish between authentical genes and pseudogenes.
The complementary phylogenetic analysis of morphological and molecular characters [7][10] suggests that :
- Ellobius, Prometheomys and Lagurus are among the most basal arvicolines ;
- Dicrostonyx, Phenacomys and Arborimus may form a clade ;
- Core arvicolines include three subclades:
- Lemmini: Synaptomys, Lemmus, Myopus ;
- Clethrionomyini: Eothenomys, Myodes ;
- Arvicolini: Arvicola, Chionomys, Stenocranius and Microtus.
- Microtus sensu lato contains Alexandromys, ‘Neodon’, Mynomes, Lasiopodomys, Terricola, and Microtus sensu stricto ;
- 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.
Classification
Subfamily Arvicolinae - voles, lemmings, muskrats
The subfamily Arvicolinae contains ten tribes, seven of which are classified as voles, one as lemmings, and two as muskrats.
- Tribe Arvicolini
- Genus Arvicola - water voles
- Genus Blanfordimys
- Genus Chionomys - snow voles
- Genus Lasiopodomys
- Genus Lemmiscus
- Genus Microtus - voles
- Insular Vole, Microtus abbreviatus
- California Vole, Microtus californicus
- Rock Vole, Microtus chrotorrhinus
- Long-tailed Vole, Microtus longicaudus
- Mexican Vole, Microtus mexicanus
- Singing Vole, Microtus miurus
- Water Vole, Microtus richardsoni
- Zempoaltépec Vole, Microtus umbrosus
- Taiga Vole, Microtus xanthognathus
- Subgenus Microtus
- Field Vole, Microtus agrestis
- Anatolian Vole, Microtus anatolicus
- Common Vole, Microtus arvalis
- Cabrera's Vole, Microtus cabrerae
- Doğramaci's Vole, Microtus dogramacii
- Günther's Vole, Microtus guentheri
- Tien Shan Vole, Microtus ilaeus
- Persian Vole, Microtus irani
- Southern Vole, Microtus levis
- Paradox Vole, Microtus paradoxus
- Qazvin Vole, Microtus qazvinensis
- Schidlovsky's Vole, Microtus schidlovskii
- Social Vole, Microtus socialis
- European Pine Vole, Microtus subterraneus
- Transcaspian Vole, Microtus transcaspicus
- Subgenus Terricola
- Bavarian Pine Vole, Microtus bavaricus
- Calabria Pine Vole, Microtus brachycercus
- Daghestan Pine Vole, Microtus daghestanicus
- Mediterranean Pine Vole, Microtus duodecimcostatus
- Felten's Vole, Microtus felteni
- Liechtenstein's Pine Vole, Microtus liechtensteini
- Lusitanian Pine Vole, Microtus lusitanicus
- Major's Pine Vole, Microtus majori
- Alpine Pine Vole, Microtus multiplex
- Savi's Pine Vole, Microtus savii
- Tatra Pine Vole, Microtus tatricus
- Thomas's Pine Vole, Microtus thomasi
- Subgenus Mynomes
- Subgenus Alexandromys
- Clarke's Vole, Microtus clarkei
- Evorsk Vole, Microtus evoronensis
- Reed Vole, Microtus fortis
- Gerbe's Vole, Microtus gerbei
- Taiwan Vole, Microtus kikuchii
- Lacustrine Vole, Microtus limnophilus
- Maximowicz's Vole, Microtus maximowiczii
- Middendorf's Vole, Microtus middendorffi
- Mongolian Vole, Microtus mongolicus
- Japanese Grass Vole, Microtus montebelli
- Muisk Vole, Microtus mujanensis
- Tundra Vole (Root Vole), Microtus oeconomus
- Sakhalin Vole, Microtus sachalinensis
- Subgenus Stenocranius
- Subgenus Pitymys
- Subgenus Pedomys
- Subgenus Hyrcanicola
- Genus Neodon - mountain voles
- Genus Phaiomys
- Genus Proedromys
- Genus Volemys
- Tribe Dicrostonychini - collared lemmings
- Genus Dicrostonyx
- St. Lawrence Island Collared Lemming, Dicrostonyx exsul (Dicrostonyx groenlandicus exsul)
- Northern Collared Lemming, Dicrostonyx groenlandicus
- Ungava Collared Lemming, Dicrostonyx hudsonius
- Victoria Collared Lemming, Dicrostonyx kilangmiutak (Dicrostonyx groenlandicus kilangmiutak)
- Nelson's Collared Lemming, Dicrostonyx nelsoni (Dicrostonyx groenlandicus nelsoni)
- Ogilvie Mountains Collared Lemming, Dicrostonyx nunatakensis
- Richardson's Collared Lemming, Dicrostonyx richardsoni
- Bering Collared Lemming, Dicrostonyx rubricatus (Dicrostonyx groenlandicus rubricatus)
- Arctic Lemming, Dicrostonyx torquatus
- Unalaska Collared Lemming, Dicrostonyx unalascensis
- Wrangel Lemming, Dicrostonyx vinogradovi
- Tribe Ellobiusini - mole voles
- Tribe Lagurini
- Tribe Lemmini - lemmings
- Tribe Myodini
- Genus Alticola - voles from Central Asia
- Subgenus Alticola
- Subgenus Aschizomys
- Subgenus Platycranius
- Genus Caryomys
- Genus Eothenomys - voles from East Asia
- Genus Hyperacrius - voles from Pakistan
- Genus Myodes - red-backed voles
- Anderson's Red-backed Vole, Myodes andersoni
- Western Red-backed Vole, Myodes californicus
- Tien Shan Red-backed Vole, Myodes centralis
- Southern Red-backed Vole, Myodes gapperi
- Bank Vole, Myodes glareolus
- Imaizumi's Red-backed Vole, Myodes imaizumii
- Korean Red-backed Vole, Myodes regulus
- Hokkaido Red-backed Vole, Myodes rex
- Grey Red-backed Vole, Myodes rufocanus
- Northern Red-backed Vole, Myodes rutilus
- Shansei Vole, Myodes shanseius
- Smith's Vole, Myodes smithii
- Tribe Neofibrini
- Tribe Ondatrini - muskrat
- Tribe Pliomyini
- Tribe Prometheomyini
- incertae sedis
Fossil species
References
- ^ 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.
- ^ 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.
- ^ McKenna, M. C. and S. K. Bell. 1997. Classification of Mammals above the Species Level. Columbia University Press, New York.
- ^ Klein, Richard (2009). The Human Career: Human Biological and Cultural Origins. London: The University of Chicago Press. p. 25. ISBN 9780226439655.
- ^ 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.
- ^ Conroy CJ, Cook JA. 1999. MtDNA evidence for repeated pulses of speciation within arvicoline and murid rodents. J. Mammal. Evol. 6:221-245.
- ^ 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.
- ^ Triant DA, DeWoody JA. 2008. Molecular analyses of mitochondrial pseudogenes within the nuclear genome of arvicoline rodents. Genetica 132:21-33.
- ^ 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.
- ^ 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.
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