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Cladus: Eukaryota
Supergroup: Opisthokonta
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Classis: Mammalia
Subclassis: Theria
Infraclassis: Placentalia
Ordo: Rodentia
Subordo: Myomorpha
Superfamilia: Muroidea
Familia: Cricetidae
Subfamilia: Arvicolinae
Genera: †Altaiomys - Alticola - Arborimus - Arvicola - Blanfordimys - Caryomys - Chionomys - Clethrionomys - Dictrostonyx - Dinaromys - Ellobius - Eolagurus - Eothenomys - Hyperacrius - Lagurus - Lasiopodomys - Lemmiscus - Lemmus - Microtus - Myodes - Myopus - Neodon - Neofiber - Ondatra - Phaiomys - Phenacomys - Proedromys - Prometheomys - Synaptomys - Volemys

Name

Arvicolinae Gray, 1821

Vernacular names
Internationalization
日本語: ハタネズミ亜科


References

* Arvicolinae on Mammal species of the World.
* Mammal Species of the World: A Taxonomic and Geographic Reference, 2 Volume Set edited by Don E. Wilson, DeeAnn M. Reeder

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Arvicolinae is a subfamily of rodents that includes the voles, lemmings, and muskrats. Its closest relatives are members of the other subfamilies in the Cricetidae, the hamsters and New World rats and mice[1]. Sometimes the subfamily Arvicolinae is placed in the family Muridae along with all other members of the superfamily Muroidea[2]. It is also sometimes referred to as Microtinae or is recognized as a family, Arvicolidae [3].

They are the most populous group of Rodentia in the Northern Hemisphere. They are often 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 arvicolines are most easily identified based on their molar teeth, which show prismatic cusps consisting of alternating triangles. These molars are ever-growing and are well adapted to a herbivorous lifestyle.

Arvicolines are Holarctic in distribution and represent one of the only major muroid radiations to reach the New World via Beringia. The other 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.

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

The subfamily Arvicolinae contains ten tribes, seven of which are classified as voles, one as lemmings, and two as muskrats.

Classification


Subfamily Arvicolinae - voles, lemmings, muskrats

* Tribe Arvicolini
o Genus Arvicola - water voles
+ European (or Northern) Water Vole, Arvicola amphibius (Arvicola terrestris)
+ Southwestern (or Southern) Water Vole, Arvicola sapidus
+ Montane Water Vole, Arvicola scherman
o Genus Blanfordimys
+ Afghan Vole, Blanfordimys afghanus
+ Bucharian Vole, Blanfordimys bucharicus
o Genus Chionomys - snow voles
+ Caucasian Snow Vole, Chionomys gud
+ European Snow Vole, Chionomys nivalis
+ Robert's Snow Vole, Chionomys roberti
o Genus Lasiopodomys
+ Brandt's Vole, Lasiopodomys brandtii
+ Plateau Vole, Lasiopodomys fuscus
+ Mandarin Vole, Lasiopodomys mandarinus
o Genus Lemmiscus
+ Sagebrush Vole, Lemmiscus curtatus
o 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
# Beach Vole, Microtus breweri
# Gray-tailed Vole, Microtus canicaudus
# Montane Vole, Microtus montanus
# Creeping Vole, Microtus oregoni
# Meadow Vole, Microtus pennsylvanicus
# Townsend's Vole, Microtus townsendii
+ 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
# Narrow-headed Vole, Microtus gregalis
+ Subgenus Pitymys
# Guatemalan Vole, Microtus guatemalensis
# Tarabundí Vole, Microtus oaxacensis
# Woodland Vole, Microtus pinetorum
# Jalapan Pine Vole, Microtus quasiater
+ Subgenus Pedomys
# Prairie Vole, Microtus ochrogaster
+ Subgenus Hyrcanicola
# Schelkovnikov's Pine Vole, Microtus schelkovnikovi
o Genus Neodon - mountain voles
+ Juniper Vole, Neodon juldaschi
+ Chinese Scrub Vole, Neodon irene
+ Sikkim Vole, Neodon sikimensis
+ Forrest's Vole, Neodon forresti
o Genus Phaiomys
+ Blyth's Vole, Phaiomys leucurus
o Genus Proedromys
+ Duke of Bedford's Vole, Proedromys bedfordi
+ Proedromys liangshanensis
o Genus Volemys
+ Szechuan Vole, Volemys millicens
+ Marie's Vole, Volemys musseri
* Tribe Dicrostonychini - collared lemmings
o 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
o Genus Ellobius - mole voles
+ Alai Mole Vole, Ellobius alaicus
+ Southern Mole Vole, Ellobius fuscocapillus
+ Transcaucasian Mole Vole, Ellobius lutescens
+ Northern Mole Vole, Ellobius talpinus
+ Zaisan Mole Vole, Ellobius tancrei
* Tribe Lagurini
o Genus Eolagurus
+ Yellow Steppe Lemming, Eolagurus luteus
+ Przewalski's Steppe Lemming, Eolagurus przewalskii
o Genus Lagurus
+ Steppe Lemming, Lagurus lagurus
* Tribe Lemmini - lemmings
o Genus Lemmus - true lemmings
+ Amur Lemming, Lemmus amurensis
+ Norway lemming, Lemmus lemmus
+ Siberian Brown Lemming, Lemmus sibiricus
+ North American Brown Lemming, Lemmus trimucronatus
+ Wrangel Island Lemming, Lemmus portenkoi
o Genus Myopus
+ Wood Lemming, Myopus schisticolor
o Genus Synaptomys - bog lemmings
+ Northern Bog Lemming, Synaptomys borealis
+ Southern Bog Lemming, Synaptomys cooperi
* Tribe Myodini
o Genus Alticola - voles from Central Asia
+ Subgenus Alticola
# White-tailed Mountain Vole, Alticola albicauda
# Silver Mountain Vole, Alticola argentatus
# Gobi Altai Mountain Vole, Alticola barakshin
# Central Kashmir Vole, Alticola montosa
# Royle's Mountain Vole, Alticola roylei
# Mongolian Silver Vole, Alticola semicanus
# Stoliczka's Mountain Vole, Alticola stoliczkanus
# Tuva Silver Vole, Alticola tuvinicus
+ Subgenus Aschizomys
# Lemming Vole, Alticola lemminus
# Large-eared Vole, Alticola macrotis
# Lake Baikal Mountain Vole, Alticola olchonensis
+ Subgenus Platycranius
# Flat-headed Vole, Alticola strelzowi
o Genus Caryomys
+ Ganzu Vole, Caryomys eva
+ Kolan Vole, Caryomys inez
o Genus Eothenomys - voles from East Asia
+ Kachin Red-backed Vole, Eothenomys cachinus
+ Pratt's Vole, Eothenomys chinensis
+ Southwest China Vole, Eothenomys custos
+ Pére David's Vole, Eothenomys melanogaster
+ Yunnan Red-backed Vole, Eothenomys miletus
+ Chaotung Vole, Eothenomys olitor
+ Yulungshan Vole, Eothenomys proditor
+ Ward's Red-backed Vole, Eothenomys wardi
o Genus Hyperacrius - voles from Pakistan
+ True's Vole, Hyperacrius fertilis
+ Murree Vole, Hyperacrius wynnei
o 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
o Genus Neofiber
+ Round-tailed Muskrat, Neofiber alleni
* Tribe Ondatrini - muskrat
o Genus Ondatra
+ Muskrat, Ondatra zibethicus
* Tribe Pliomyini
o Genus Dinaromys
+ Balkan Snow Vole, Dinaromys bogdanovi
* Tribe Prometheomyini
o Genus Prometheomys
+ Long-clawed Mole Vole, Prometheomys schaposchnikowi
* incertae sedis
o Genus Arborimus - tree voles
+ White-footed Vole, Arborimus albipes
+ Red Tree Vole, Arborimus longicaudus
+ Sonoma Tree Vole, Arborimus pomo
o Genus Phenacomys - heather voles
+ Western Heather Vole, Phenacomys intermedius
+ Eastern Heather Vole, Phenacomys ungava

Fossil species

* Tribe Dicrostonychini - collared lemmings
o Genus Predicrostonyx †
+ Hopkins's Lemming, Predicrostonyx hopkinsi †
* Tribe Clethrionomyini
o Genus Altaiomys

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 [5] and the exon 10 of the growth hormone receptor (ghr) nuclear gene [6]. 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 [7]. Sequencing complete mitochondrial genomes of voles [8] may help to distinguish between authentical genes and pseudogenes.

The complementary phylogenetic analysis of morphological and molecular characters [6][9] 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:
o Lemmini: Synaptomys, Lemmus, Myopus ;
o Clethrionomyini: Eothenomys, Myodes ;
o 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.

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. ^ McKenna, M. C. and S. K. Bell. 1997. Classification of Mammals above the Species Level. Columbia University Press, New York.
4. ^ Klein, Richard (2009). The Human Career: Human Biological and Cultural Origins. London: The University of Chicago Press. p. 25. ISBN 9780226439655.
5. ^ Conroy CJ, Cook JA. 1999. MtDNA evidence for repeated pulses of speciation within arvicoline and murid rodents. J. Mammal. Evol. 6:221-245.
6. ^ 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.
7. ^ Triant DA, DeWoody JA. 2008. Molecular analyses of mitochondrial pseudogenes within the nuclear genome of arvicoline rodents. Genetica 132:21-33.
8. ^ 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.
9. ^ 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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