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{{Short description|Subfamily of rodents}}
{{Taxobox
{{Automatic taxobox
| name = Arvicolines
| name = Arvicolinae
| fossil_range = Late [[Miocene]] - Recent
| fossil_range = Late [[Miocene]] – recent
| image = Meadvole.jpg
| image = Meadvole.jpg
| image_caption = [[Meadow Vole|Meadow vole]]
| image_caption = [[Meadow Vole|Meadow vole]] (''Microtus pennsylvanicus'')
| regnum = [[Animal]]ia
| taxon = Arvicolinae
| phylum = [[Chordate|Chordata]]
| authority = [[John Edward Gray|Gray]], 1821
| classis = [[Mammal]]ia
| ordo = [[Rodent]]ia
| superfamilia = [[Muroidea]]
| familia = [[Cricetidae]]
| subfamilia = '''Arvicolinae'''
| subfamilia_authority = [[John Edward Gray|Gray]], 1821
| subdivision_ranks = [[Genus|Genera]]
| subdivision_ranks = [[Genus|Genera]]
| subdivision =
| subdivision =
see text
see text
}}
}}

'''Arvicolinae''' is a [[subfamily]] of [[rodent]]s that includes the [[vole]]s, [[lemming]]s, and [[muskrat]]s. Its closest relatives are members of the other subfamilies in the [[Cricetidae]], the [[hamster]]s and [[New World rats and mice]]<ref>Steppan, S. J., R. A. Adkins, and J. Anderson. 2004. [[Phylogeny]] and [[divergence date]] estimates of rapid radiations in [[Muroidea|muroid]] [[rodent]]s based on multiple nuclear [[genes]]. Systematic Biology, 53:533-553.</ref>. Sometimes the subfamily Arvicolinae is placed in the family [[Muridae]] along with all other members of the superfamily [[Muroidea]] <ref>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.</ref>. It is also sometimes referred to as '''Microtinae''' or is recognized as a [[family (biology)|family]], '''Arvicolidae''' <ref>McKenna, M. C. and S. K. Bell. 1997. [[Taxonomic classification|Classification]] of [[Mammal]]s above the [[Species]] Level. Columbia University Press, New York.
The '''Arvicolinae''' are a [[subfamily]] of [[rodent]]s that includes the [[vole]]s, [[lemming]]s, and [[muskrat]]s. They are most closely related to the other subfamilies in the [[Cricetidae]] (comprising the [[hamster]]s and [[New World rats and mice]]<ref>{{cite journal |last1=Steppan |first1=S. J. |first2=R. A. |last2=Adkins |first3=J. |last3=Anderson |year=2004 |title=Phylogeny and divergence date estimates of rapid radiations in muroid rodents based on multiple nuclear genes |journal=Systematic Biology |volume=53 |issue=4 |pages=533–553 |doi=10.1080/10635150490468701|pmid=15371245 }}</ref>). Some authorities place the subfamily Arvicolinae in the family [[Muridae]] along with all other members of the superfamily [[Muroidea]].<ref>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.</ref> Some refer to the subfamily as the '''Microtinae''' (yielding the adjective "microtine")<ref name="Nakao-et-al-2010">{{cite journal | last1=Nakao | first1=Minoru | last2=Yanagida | first2=Tetsuya | last3=Okamoto | first3=Munehiro | last4=Knapp | first4=Jenny | last5=Nkouawa | first5=Agathe | last6=Sako | first6=Yasuhito | last7=Ito | first7=Akira | title=State-of-the-art ''Echinococcus'' and ''Taenia'': Phylogenetic taxonomy of human-pathogenic tapeworms and its application to molecular diagnosis | journal=[[Infection, Genetics and Evolution]] | publisher=[[Elsevier]] | volume=10 | issue=4 | year=2010 | issn=1567-1348 | doi=10.1016/j.meegid.2010.01.011 | pages=444–452| pmid=20132907 | bibcode=2010InfGE..10..444N }}</ref> or rank the taxon as a full [[family (biology)|family]], the '''Arvicolidae'''.<ref>McKenna, M. C. and S. K. Bell. 1997. [[Taxonomic classification|Classification]] of [[Mammal]]s above the [[Species]] Level. Columbia University Press, New York.
</ref>.
</ref>

The Arvicolinae are the most populous group of [[Rodent]]ia 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 [[biostratigraphy|biostratigraphic]] dating of paleontological and archeological sites in North America and Europe.<ref>{{Cite book
| last = Klein
| first = Richard
| title = The Human Career: Human Biological and Cultural Origins
| publisher = The University of Chicago Press
| location = London
| year = 2009
| page = 25
| isbn = 978-0-226-43965-5}}</ref>


== Description ==
== Description ==
The most convenient distinguishing feature of the Arvicolinae is the nature of their [[Molar (tooth)|molar]] teeth, which have prismatic [[Cusp (dentistry)|cusps]] in the shape of alternating triangles. These molars are an adaptation to a [[herbivore|herbivorous]] diet in which the major food plants include a large proportion of abrasive materials such as [[phytolith]]s; the teeth get worn down by abrasion throughout the adult life of the animal and they grow continuously in compensation.<ref>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.</ref>


Arvicolinae are [[Holarctic]] in distribution and represent one of only a few major [[Muroidea|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 [[hibernation|hibernate]]. They are also characterized by extreme fluctuations in population numbers.
The arvicolines are most easily identified based on their [[Molar (tooth)|molar]] teeth, which show prismatic cusps consisting of alternating triangles. These molars are ever-growing and are well adapted to a [[herbivore|herbivorous]] lifestyle.


Most arvicolines are small, furry, short-tailed [[vole]]s or [[lemming]]s, 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 animal|aquatic]] 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 [[hibernation|hibernate]]. They are also characterized by extreme fluctuations in population size.


== Phylogeny ==
Most arvicolines are small, furry, short tailed [[vole]]s or [[lemming]]s, 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 animal|aquatic]] lifestyle.
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 <ref>Conroy CJ, Cook JA. 1999. MtDNA evidence for repeated pulses of speciation within arvicoline and murid rodents. J. Mammal. Evol. 6:221-245.</ref>
and the exon 10 of the [[growth hormone receptor]] (''ghr'') nuclear gene.<ref name="Galewski2006">Galewski T, Tilak M, Sanchez S, Chevret P, Paradis E, Douzery EJP. 2006. [http://www.biomedcentral.com/1471-2148/6/80 The evolutionary radiation of Arvicolinae rodents (voles and lemmings): relative contribution of nuclear and mitochondrial DNA phylogenies. BMC Evol. Biol. 6:80].</ref> 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 gene]]s 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 [[Homology (biology)#Orthology|orthology]], but they can also be used as phylogenetic markers.<ref>Triant DA, DeWoody JA. 2008. Molecular analyses of mitochondrial pseudogenes within the nuclear genome of arvicoline rodents. Genetica 132:21-33.</ref>
Sequencing complete mitochondrial genomes of voles <ref>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.</ref> may help to distinguish between authentic genes and pseudogenes.


The complementary phylogenetic analysis of morphological and molecular characters <ref name="Galewski2006"/><ref>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.</ref>
Some authorities have placed the [[zokor]]s within the Arvicolinae, but they have been shown to be unrelated.
suggests:
* ''[[Ellobius]]'', ''[[Prometheomys]]'', and ''[[Lagurus (rodent)|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 [[zokor]]s within the Arvicolinae, but they have been shown {{by whom|date=January 2017}} to be unrelated.
The subfamily Arvicolinae contains ten tribes, seven of which are classified as voles, one as lemmings, and two as muskrats.

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]].<ref>{{Cite journal|last1=Abramson|first1=Natalia I.|last2=Bodrov|first2=Semyon Yu|last3=Bondareva|first3=Olga V.|last4=Genelt-Yanovskiy|first4=Evgeny A.|last5=Petrova|first5=Tatyana V.|date=2021-11-19|title=A mitochondrial genome phylogeny of voles and lemmings (Rodentia: Arvicolinae): Evolutionary and taxonomic implications|journal=PLOS ONE|language=en|volume=16|issue=11|pages=e0248198|doi=10.1371/journal.pone.0248198|issn=1932-6203|pmc=8604340|pmid=34797834|bibcode=2021PLoSO..1648198A |doi-access=free }}</ref>


== Classification ==
== Classification ==


[[Image:Rötelmausschädel.jpg|thumb|right|200px|Skull of a [[Bank Vole]]. Note the distinctive molar pattern characteristic of arvicolines.]]
[[Image:Rötelmausschädel.jpg|thumb|right|200px|Skull of a [[bank vole]]: Note the distinctive molar pattern characteristic of arvicolines.]]
*Subfamily '''Arvicolinae''' - [[vole]]s, [[lemming]]s, [[muskrat]]s
'''Subfamily Arvicolinae''' - [[vole]]s, [[lemming]]s, [[muskrat]]s

**Tribe [[Arvicolini]]
The subfamily Arvicolinae contains eleven tribes, eight of which are classified as voles, two as lemmings, and one as muskrats.<ref name=MDD>{{cite journal|author=Mammal Diversity Database |year=2023|title=Mammal Diversity Database (Version 1.11) [Data set] |journal=Zenodo |doi=10.5281/zenodo.7830771}}</ref> Recent changes to the subfamily include disbanding genus ''[[Red-backed vole (disambiguation)|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]].<ref name=MDD/>
***Genus ''[[Arvicola]]'' - water voles

***Genus ''[[Blanfordimys]]'' - Afghan vole and Bucharian vole
* Tribe [[Arvicolini]]
***Genus ''[[Chionomys]]'' - snow voles
***Genus ''[[Lasiopodomys]]''
** Genus ''[[Arvicola]]'' - water voles
*** [[European water vole|European (or Northern) water vole]] (''Arvicola amphibius'' or ''Arvicola terrestris'')
***Genus ''[[Lemmiscus]]'' - sagebrush vole
*** [[Southwestern water vole|Southwestern (or southern) water vole]] (''Arvicola sapidus'')
***Genus ''[[Microtus]]'' - voles
*** [[Montane water vole]] (''Arvicola scherman'')
***Genus ''[[Neodon]]'' - mountain voles
* Tribe [[Microtini]]
***Genus ''[[Phaiomys]]''
***Genus ''[[Proedromys]]'' - Duke of Bedford's vole
** Genus ''[[Alexandromys]]''
***[[Alpine vole]] (''Alexandromys alpinus'') {{small|split off from ''A. mongolicus''}}<ref name=alpinus>{{BioRef|asm|title=''Alexandromys alpinus''|id=1006515}}</ref>
***Genus ''[[Volemys]]''
***[[Evorsk vole]] (''Alexandromys evoronensis'')
**Tribe [[Dicrostonychini]] - collared lemmings
***[[Reed vole]] (''Alexandromys fortis'')
***Genus ''[[Dicrostonyx]]'' - collared lemmings
***[[Taiwan vole]] (''Alexandromys kikuchii'')
***Genus ''[[Predicrostonyx]]'' †
***[[Lacustrine vole]] (''Alexandromys limnophilus'')
**Tribe [[Ellobiusini]] - mole voles
***[[Maximowicz's vole]] (''Alexandromys maximowiczii'')
***Genus ''[[Ellobius]]'' - mole voles
***[[Middendorff's vole]] (''Alexandromys middendorffi'')
**Tribe [[Lagurini]]
***[[Mongolian vole]] (''Alexandromys mongolicus'')
***Genus ''[[Eolagurus]]'' - steppe lemmings
***[[Japanese grass vole]] (''Alexandromys montebelli'')
***Genus ''[[Lagurus]]'' - steppe vole
***[[Muya Valley vole]] (''Alexandromys mujanensis'')
**Tribe [[Lemmini]] - lemmings
***[[Tundra vole]] or root vole (''Alexandromys oeconomus'')
***Genus ''[[Lemmus]]'' - true lemmings
***[[Sakhalin vole]] (''Alexandromys sachalinensis'')
***Genus ''[[Myopus]]''
***[[Shantar vole]] (''Alexandromys shantaricus'') {{small|split off from ''A. maximowiczii''}}<ref name=shantaricus>{{BioRef|asm|title=''Alexandromys shantaricus''|id=1002027}}</ref>
***Genus ''[[Synaptomys]]'' - bog lemmings
** Genus ''[[Chionomys]]'' - snow voles
**Tribe [[Myodini]]
*** [[Caucasian snow vole]], ''Chionomys gud''
***Genus ''[[Alticola]]'' - voles from [[Central Asia]]
*** [[European snow vole]], ''Chionomys nivalis''
***Genus ''[[Caryomys]]''
*** [[Robert's snow vole]], ''Chionomys roberti''
***Genus ''[[Eothenomys]]'' - voles from [[East Asia]]
*** [[Lazistan snow vole]] (''Chionomys lasistanius'') {{small|split from ''C. gud''}}<ref name=lasistanius>{{BioRef|asm|title=''Chionomys lasistanius''|id=1002043}}</ref>
***Genus ''[[Hyperacrius]]'' - voles from [[Pakistan]]
*** [[Stekolnikov's snow vole]] (''Chionomys stekolnikovi'') {{small|recently described}}<ref name=stekolnikovi>{{BioRef|asm|title=''Chionomys stekolnikovi''|id=1006652}}</ref><ref>{{cite journal |last1=Golenishchev |first1=F. N. |last2=Malikov |first2=V. G. |last3=Bannikova |first3=A. A. |last4=Zykov |first4=A. E. |last5=Yiğit |first5=N. |last6=Çolak |first6=E. |year=2022 |title=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 |journal=Russian Journal of Theriology |volume=21 |issue=1 |pages=1–12|url=https://www.researchgate.net/publication/361085614|doi=10.15298/rusjtheriol.21.1.01|s2cid=250649779 |doi-access=free }}</ref>
***Genus ''[[Myodes]]'' - red-backed voles
** Genus ''[[Hyperacrius]]'' - voles from [[Pakistan]]
**Tribe [[Neofibrini]]
*** [[True's vole]], ''Hyperacrius fertilis''
***Genus ''[[Neofiber]]'' - round-tailed muskrat
*** [[Murree vole]], ''Hyperacrius wynnei''
**Tribe [[Ondatrini]] - muskrat
***Genus ''[[Ondatra]]'' - muskrat
** Genus ''[[Lasiopodomys]]''
*** [[Brandt's vole]], ''Lasiopodomys brandtii''
**Tribe [[Pliomyini]]
*** [[Mandarin vole]], ''Lasiopodomys mandarinus''
***Genus ''[[Dinaromys]]'' - voles from the [[Dinaric Alps]]
** Genus ''[[Lemmiscus]]''
**Tribe [[Prometheomyini]] - Prometheus mouse
*** [[Sagebrush vole]], ''Lemmiscus curtatus''
***Genus ''[[Prometheomys]]'' - Prometheus mouse
**''[[incertae sedis]]''
** Genus ''[[Microtus]]'' - voles
***Genus ''[[Arborimus]]'' - tree voles
*** [[Insular vole]], ''M. abbreviatus''
***Genus ''[[Phenacomys]]'' - heather voles
*** [[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 (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 (rodent)|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''
*** [[North American brown lemming|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''
*** [[Père David's vole]], ''E. melanogaster''
*** [[Yunnan red-backed vole]], ''E. miletus''
*** [[Chaotung vole]], ''E. olitor''
*** [[Yulungshan vole]], ''E. proditor''
*** [[Ward's red-backed vole]], ''E. wardi''
** Genus ''[[Craseomys]]'' - red-backed voles
*** [[Anderson's red-backed vole]], ''C. andersoni''
*** [[Imaizumi's red-backed vole]], ''C. imaizumii''
*** [[Korean red-backed vole]], ''C. regulus''
*** [[Hokkaido red-backed vole]], ''C. rex''
*** [[Grey red-backed vole]], ''C. rufocanus''
*** [[Shansei vole]], ''C. shanseius''
*** [[Smith's vole]], ''C. smithii''
** Genus ''[[Clethrionomys ]]'' - red-backed voles
*** [[Western red-backed vole]], ''C. californicus''
*** [[Tien Shan red-backed vole]], ''C. centralis''
*** [[Southern red-backed vole]], ''C. gapperi''
*** [[Bank vole]], ''C. glareolus''
*** [[Northern red-backed vole]], ''C. rutilus''
* Tribe [[Ondatrini]] - muskrats
** Genus ''[[Neofiber]]''
*** [[Round-tailed muskrat]], ''N. alleni''
**Genus ''[[Ondatra]]''
*** [[Muskrat]], ''O. zibethicus''
* Tribe [[Pliophenacomyini]]
**Genus ''[[Arborimus]]'' - tree voles
*** [[White-footed vole]], ''A. albipes''
*** [[Red tree vole]], ''A. longicaudus''
*** [[Sonoma tree vole]] or California red tree mouse, ''A. pomo''
** Genus ''[[Phenacomys]]'' - heather voles
*** [[Western heather vole]], ''P. intermedius''
*** [[Eastern heather vole]], ''P. ungava''
*Tribe [[Pliomyini]]
** Genus ''[[Dinaromys]]''
*** [[Balkan snow vole]], ''D. bogdanovi''
* Tribe [[Prometheomyini]]
** Genus ''[[Prometheomys]]''
*** [[Long-clawed mole vole]], ''P. schaposchnikowi''


=== Fossil species ===
=== Fossil species ===
**Tribe [[Clethrionomyini]]
* Tribe [[Arvicolini]]
***Genus ''[[Altaiomys]]''
** Genus ''[[Mimomys]]'' †
* Tribe [[Dicrostonychini]] - collared lemmings
** Genus ''[[Predicrostonyx]]'' †
* Tribe [[Clethrionomyini]]
** Genus ''[[Altaiomys]]'' †
** Genus ''[[Pitymimomys]]'' †
** Genus ''[[Borsodia]]'' †
** Genus ''[[Allophaiomys]]'' †
** Genus ''[[Prolagurus]]'' †
* Tribe [[Pliomyini]]
** Genus ''[[Pliomys]]'' † (extinct c. 12,000 years ago)


== Phylogeny ==
== See also ==


* [[Hantavirus]]
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 <ref>Conroy CJ, Cook JA. 1999. MtDNA evidence for repeated pulses of speciation within arvicoline and murid rodents. J. Mammal. Evol. 6:221-245.</ref>
* [[Isla Vista virus]]
and the exon 10 of the [[growth hormon receptor]] (''ghr'') nuclear gene <ref>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.</ref>.


== References ==
== References ==
{{reflist}}
{{Reflist}}


== External links ==
== External links ==
{{Wikispecies|Arvicolinae}}
{{Wikispecies|Arvicolinae}}
{{Cricetidae}}
{{Arvicolinae}}
{{Taxonbar|from=Q840750}}
{{Authority control}}


[[Category:Muroid rodents]]
[[Category:Voles and lemmings| ]]
[[Category:Voles and lemmings| ]]
[[Category:Mammal subfamilies]]

[[Category:Taxa named by John Edward Gray]]
[[br:Arvicolinae]]
[[bg:Полевки]]
[[ca:Arvicolí]]
[[de:Wühlmäuse]]
[[es:Arvicolinae]]
[[eo:Arvikolenoj]]
[[fr:Arvicolinae]]
[[it:Arvicolinae]]
[[ka:მემინდვრიები]]
[[lb:Wullmais]]
[[hu:Pocokformák]]
[[nl:Woelmuizen]]
[[ja:ハタネズミ亜科]]
[[pl:Nornikowate]]
[[pt:Arvicolinae]]
[[ru:Полёвковые]]
[[sl:Voluhar]]
[[fi:Muridae]]
[[sv:Sorkar]]
[[uk:Полівкові]]
[[zh:田鼠]]

Latest revision as of 02:04, 2 November 2024

Arvicolinae
Temporal range: Late Miocene – recent
Meadow vole (Microtus pennsylvanicus)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Rodentia
Family: Cricetidae
Subfamily: Arvicolinae
Gray, 1821
Genera

see text

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

[edit]

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

[edit]

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 [by whom?] 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

[edit]
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

[edit]

See also

[edit]

References

[edit]
  1. ^ Steppan, S. J.; Adkins, R. A.; Anderson, J. (2004). "Phylogeny and divergence date estimates of rapid radiations in muroid rodents based on multiple nuclear genes". Systematic Biology. 53 (4): 533–553. doi:10.1080/10635150490468701. PMID 15371245.
  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. 10 (4). Elsevier: 444–452. Bibcode:2010InfGE..10..444N. 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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