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Issue: Vol. 4429 No. 1: 5 Jun. 2018
Type: Article
Published: 2018-06-05
DOI: 10.11646/zootaxa.4429.1.1
Page range: 1–52
Abstract views: 313
PDF downloaded: 8

Reassessment of the taxonomic status of Craseomys and three controversial species of Myodes and Alticola (Rodentia: Arvicolinae)

Key Laboratory of Bio-Resource and Eco-Environment of Ministry of Education, College of Life Sciences, Sichuan University, Chengdu 610065, Sichuan, PR China Sichuan Academy of Forestry, Chengdu 610066, Sichuan, PR China
Sichuan Academy of Forestry, Chengdu 610066, Sichuan, PR China
College of Tourism and Urban-rural Planning, Chengdu University of Technology, Chengdu 610058, Sichuan, PR China
Key Laboratory of Bio-Resource and Eco-Environment of Ministry of Education, College of Life Sciences, Sichuan University, Chengdu 610065, Sichuan, PR China
Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queen’s Park, Toronto, M5S 2C6, Ontario, Canada
Sichuan Academy of Forestry, Chengdu 610066, Sichuan, PR China
Key Laboratory of Bio-Resource and Eco-Environment of Ministry of Education, College of Life Sciences, Sichuan University, Chengdu 610065, Sichuan, PR China
Key Laboratory of Bio-Resource and Eco-Environment of Ministry of Education, College of Life Sciences, Sichuan University, Chengdu 610065, Sichuan, PR China
Sichuan Academy of Forestry, Chengdu 610066, Sichuan, PR China
Sichuan Academy of Forestry, Chengdu 610066, Sichuan, PR China
Key Laboratory of Bio-Resource and Eco-Environment of Ministry of Education, College of Life Sciences, Sichuan University, Chengdu 610065, Sichuan, PR China
Sichuan Academy of Forestry, Chengdu 610066, Sichuan, PR China
Mammalia Myodes shanseius Alticola stracheyi Alticola semicanus Phaulomys phylogeny geometric morphometrics

Abstract

The genera Myodes (red-backed voles) and Alticola (mountain voles) appear to be sister taxa based on morphological similarities, but molecular analyses fail to resolve them as monophyletic genera owing to the uncertain taxonomic status of Craseomys and Phaulomys. As a result of incomplete sampling of related specimens, ongoing controversies on the taxonomic positions of several generic and specific taxa necessitate further clarifications. Herein, we combined molecular, morphometric, and geometric morphometric approaches to analyze 217 specimens of 10 taxa of Myodes and Alticola systematically. We sequenced three genes (Cytb, COI, GHR) de novo from specimens with fresh tissues, and published sequences for M. shanseius and A. stoliczkanus for the first time. Based on this new molecular dataset, we produced phylogenetic trees using Bayesian inference, maximum likelihood, and maximum parsimony approaches. Our molecular and morphological analyses both identified three primary clades within Myodes and Alticola. The Craseomys–Phaulomys clade consistently separated from Myodes sensu stricto (s. str.) and Alticola s. str.–Platycranius. Our results support the resurrection of the genus Craseomys and the treatment of Phaulomys as its junior synonym. As Craseomys shanseius clustered with C. rufocanus in three gene phylogenies and this assessment was congruent with morphological results, we assigned C. shanseius to a subspecies of C. rufocanus. Specimens from one sampling site in Pulan County of Tibet possess M3 patterns typical of A. stoliczkanus and A. stracheyi, despite clustering together in matrilineal genealogy. Thus, we tentatively assigned A. stracheyi as a junior synonym of A. stoliczkanus. Our analyses confirmed the validity of A. semicanus and unambiguously distinguished it from A. argentatus by the ratio of tail length to head–body length, color of tail and feet, M3 pattern, and distribution.

 

References

  1. Abramson, N.I., Lebedev, V.S., Tesakov, A.S. & Bannikova, A.A. (2009) Supraspecies relationships in the subfamily Arvicolinae (Rodentia, Cricetidae): an unexpected result of nuclear gene analysis. Molecular Biology, 43, 834–846.
    https://doi.org/10.1134/S0026893309050148

    Abramson, N.I. & Lissovsky, A.A. (2012) Subfamily Arvicolinae Gray, 1821. In: Pavlinov, I.Y. & Lissovsky, A.A. (Eds.), The mammals of Russia: a taxonomic and geographic reference. KMK Scientific Press, Moscow, pp. 220–276.

    Abramson, N.I., Petrova, T.V., Dokuchaev, N.E., Obolenskaya, E.V. & Lissovsky, A.A. (2012) Phylogeography of the gray red-backed vole Craseomys rufocanus (Rodentia: Cricetidae) across the distribution range inferred from nonrecombining molecular markers. Russian Journal of Theriology, 11, 137–156.
    https://doi.org/10.15298/rusjtheriol.11.2.04

    Adkins, R.M., Walton, A.H. & Honeycutt, R.L. (2003) Higher-level systematics of rodents and divergence time estimates based on two congruent nuclear genes. Molecular Phylogenetics and Evolution, 26, 409–420.
    https://doi.org/10.1016/S1055-7903(02)00304-4

    Aimi, M. (1980) A revised classification of the Japanese red-backed voles. Memoirs of the Faculty of Science, Kyoto University, Series of Biology, 8, 35–84.

    Allen, G.M. (1924) Microtinae collection by the American Museum Asian expeditions. 133, 6.

    Allen, G.M. (1940) The mammals of China and Mongolia. Natural History of Central Asia. Vol. XI. Part 2. American Museum of Natural History, New York, 730 pp. [oo, 621–1350]

    Anderson, M.P. (1909) XXXVII.—Description of a new Japanese vole. Annals and Magazine of Natural History, 4, 317–318.
    https://doi.org/10.1080/00222930908692678

    Blanford, W.T. (1875) List of Mammalia collected by the late Dr. Stoliczka when attached to the embassy under Sir D. Forsyth in Kashmir, Ladák, eastern Turkestan, and Wakhán, with descriptions of new species. Journal of the Asiatic Society of Bengal, 44, 105–112.

    Bodrov, S.Y., Kostygov, A.Y., Rudneva, L.V. & Abramson, N.I. (2016) Revision of the taxonomic position of the Olkhon mountain vole (Rodentia, Cricetidae). Biology Bulletin, 43, 136–145.
    https://doi.org/10.1134/S1062359016020035

    Bookstein, F.L. (1997) Morphometric tools for landmark data: geometry and biology. Cambridge University Press, Cambridge, 435 pp.

    Boratyński, Z., Alves, P.C., Berto, S., Koskela, E., Mappes, T. & Melo-Ferreira, J. (2011) Introgression of mitochondrial DNA among Myodes voles: consequences for energetics? BMC Evolutionary Biology, 11, 355.
    https://doi.org/10.1186/1471-2148-11-355

    Bradley, R.D. & Baker, R.J. (2001) A test of the genetic species concept: cytochrome-b sequences and mammals. Journal of Mammalogy, 82, 960–973.

    https://doi.org/10.1644/1545-1542(2001)082<0960:ATOTGS>2.0.CO;2

    Chaline, J. & Graf, J.D. (1988) Phylogeny of the Arvicolidae (Rodentia): biochemical and paleontological evidence. Journal of Mammalogy, 69, 22–33.
    https://doi.org/10.2307/1381744

    Chen, W.C., Hao, H.B., Sun, Z.Y., Liu, Y., Liu, S.Y. & Yue, B.S. (2012) Phylogenetic position of the genus Proedromys (Arvicolinae, Rodentia): evidence from nuclear and mitochondrial DNA. Biochemical Systematics and Ecology, 42, 59–68.
    https://doi.org/ 10.1016/j.bse.2012.01.002

    Conroy, C.J. & Cook, J.A. (1999) MtDNA evidence for repeated pulses of speciation within arvicoline and murid rodents. Journal of Mammalian Evolution, 6, 221–245.
    https://doi.org/10.1023/A:1020561623890

    Cook, J.A., Bidlack, A.L., Conroy, C.J., Demboski, J.R., Fleming, M.A., Runck, A.M., Stone, K.D. & MacDonald, S.O. (2001) A phylogeographic perspective on endemism in the Alexander Archipelago of southeast Alaska. Biological Conservation, 97, 215–227.
    https://doi.org/10.1016/S0006-3207(00)00114-2

    Cook, J.A., Runck, A.M. & Conroy, C.J. (2004) Historical biogeography at the crossroads of the northern continents: molecular phylogenetics of red-backed voles (Rodentia: Arvicolinae). Molecular Phylogenetics and Evolution, 30, 767–777.
    https://doi.org/10.1016/S1055-7903(03)00248-3

    Corbet, G.B. (1978) The mammals of the palaearctic region: a taxonomic review. British Museum (Natural History), London, 314 pp.

    Ellerman, J.R. (1941) The families and genera of living rodents, Volume II Muridae. British Museum (Natural History), London, 642 pp.

    Ellerman, J.R. (1961) The fauna of India, including Pakistan, Burma and Ceylon: Mammalia (Second Edition), Vol 3, Rodentia. The Manager of Publications, Delhi, 884 pp.

    Ellerman, J.R. & Morrison-Scott, T.C.S. (1951) Checklist of palaearctic and Indian mammals, 1758 to 1946. British Museum (Natural History), London, 741 pp.
    https://doi.org/10.1086/399408

    Fan, L., Fan, Z., Yue, H., Zhang, X., Liu, Y., Sun, Z., Liu, S. & Yue, B. (2011) Complete mitochondrial genome sequence of the Chinese scrub vole (Neodon irene). Mitochondrial DNA, 22, 50–52.
    https://doi.org/10.3109/19401736.2011.603312

    Feng, Z.J., Cai, G.Q. & Zheng, C.L. (1986) The mammals of Xizang. The comprehensive scientific expedition to the Qinghai-Xizang Plateau. Science Press, Academia Sinica, Beijing, 423 pp. [in Chinese]

    Galewski, T., Tilak, M.K., Sanchez, S., Chevret, P., Paradis, E. & Douzery, E.J. (2006) The evolutionary radiation of Arvicolinae rodents (voles and lemmings): relative contribution of nuclear and mitochondrial DNA phylogenies. BMC Evolutionary Biology, 6, 80.
    https://doi.org/10.1186/1471-2148-6-80

    Gariepy, T.D., Lindsay, R., Ogden, N. & Gregory, T.R. (2012) Identifying the last supper: utility of the DNA barcode library for bloodmeal identification in ticks. Molecular Ecology Resources, 12, 646–652.
    https://doi.org/10.1111/j.1755-0998.2012.03140.x

    Gray, J.E. (1842) XXXVII.—Descriptions of some new genera and fifty unrecorded species of Mammalia. Annals and Magazine of Natural History, 10, 255–267.
    https://doi.org/10.1080/03745484209445232

    Gromov, I. & Polyakov, I.Y. (1977) Voles (Microtinae), Fauna of the USSR, Mammals. Vol. III. No. 8. Nauka Publisher, Leningrad. [in Russian]

    Guindon, S., Dufayard, J.F., Lefort, V., Anisimova, M., Hordijk, W. & Gascuel, O. (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology, 59, 307–321.
    https://dx.doi.org/10.1093/sysbio/syq010

    Heptner, V. & Rossolimo, O. (1968) Species composition and geographic variation of Asian mountain voles of the genus Alticola Blanford, 1881. Sbornik Trudy Zoologicheskogo Muzeya Moskovskogo Universiteta, 10, 53–93. [in Russian]

    Hillis, D.M. & Bull, J.J. (1993) An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis. Systematic Biology, 42, 182–192.
    https://doi.org/10.1093/sysbio/42.2.182

    Hinton, M.A.C. (1926) Monograph of the voles and lemmings (Microtinae) living and extinct. British Museum (Natural History), London, 475 pp.
    https://doi.org/10.5962/bhl.title.8319

    Howell, A.B. (1929) Mammals from China in the collections of the United States National MuseumProceedings of the United States National Museum, 75, 1–82.
    https://doi.org/10.2108/0289-0003(2000)17[477:GPOCBA]2.0.CO;2

    Huelsenbeck, J.P. & Rannala, B. (2004) Frequentist properties of Bayesian posterior probabilities of phylogenetic trees under simple and complex substitution models. Systematic Biology, 53, 904–913.
    https://doi.org/10.1080/10635150490522629

    Irwin, D.M., Kocher, T.D. & Wilson, A.C. (1991) Evolution of the cytochrome b gene of mammals. Journal of Molecular Evolution, 32, 128–144.
    https://doi.org/10.1007/bf02515385

    Iwasa, M.A. & Nakata, K. (2011) A note on the genetic status of the dark red-backed vole, Myodes rex, in Hokkaido, Japan. Mammal Study, 36, 99–103.
    https://doi.org/10.3106/041.036.0208

    Iwasa, M.A. & Suzuki, H. (2002) Evolutionary networks of maternal and paternal gene lineages in voles (Eothenomys) endemic to Japan. Journal of Mammalogy, 83, 852–865.
    https://doi.org/10.1644/1545-1542(2002)083<0852:ENOMAP>2.0.CO;2

    Iwasa, M.A., Utsumi, Y., Nakata, K., Kartavtseva, I.V., Nevedomskaya, I.A., Kondoh, N. & Suzuki, H. (2000) Geographic patterns of cytochrome b and Sry gene lineages in the gray red-backed vole Clethrionomys rufocanus from Far East Asia including Sakhalin and Hokkaido. Zoological Science, 17, 477–484.
    https://doi.org/10.2108/0289-0003(2000)17[477:GPOCBA]2.0.CO;2

    Jameson Junior, E.W. (1961) Relationships of the red-backed voles of Japan. Pacific Science, 15, 594–604.

    Jiang, J.Q. & Ma, Y. (1991) Study of taxonomic status of Craseomys shanseius Thomas in China Scientific treatise on systematic and evolutionary zoology, 1, 73–79. [in Chinese]

    Jiang, J.Q., Ma, Y. & Luo, Z.X. (1993) On the classification of the subspecies of Clethrionomys rufocanus in China (Rodentia: Cricetidae: Microtinae). Acta Zootaxonomica Sinica, 18, 114–122. [in Chinese]

    Jiang, X., Gao, J., Ni, L., Hu, J., Li, K., Sun, F., Xie, J., Bo, X., Gao, C., Xiao, J. & Zhou, Y. (2012) The complete mitochondrial genome of Microtus fortis calamorum (Arvicolinae, Rodentia) and its phylogenetic analysis. Gene, 498, 288–295.
    https://doi.org/10.1016/j.gene.2012.02.022

    Jiang, Z., Ma, Y., Wu, Y., Wang, Y., Feng, Z., Zhou, K., Liu, S., Luo, Z. & Li, C. (2015) China’s mammalian diversity. Biodiversity Science, 23, 351–364. [in Chinese]
    https://doi.org/10.17520/biods.2014202

    Kaneko, Y. (1990) Identification and some morphological characters of Clethrionomys rufocanus and Eothenomys regulus from USSR, Northeast China, and Korea in comparison with C. rufocanus from Finland. Journal of the Mammalogical Society of Japan, 14, 129–148.
    https://doi.org/10.11238/jmammsocjapan1987.14.129

    Kaneko, Y. (1992) Identification and morphological characteristics of Clethrionomys rufocanus, Eothenomys shanseius, E. inez and E. eva from the USSR, Mongolia, and northern and central China. Journal of the Mammalogical Society of Japan, 16, 71–95.
    https://doi.org/10.11238/jmammsocjapan.16.71

    Kaneko, Y., Nakashima, T. & Kimura, Y. (1992) Identification and vertical distribution of two species of Eothenomys on Ryo-Hakusan Mountains, central Honshyu, Japan. Bulletin of the Gifu Prefectural Museum, 11, 23–34. [in Japanese]

    Kimura, M. (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution, 16, 111–120.
    https://doi.org/10.1007/bf01731581

    Kimura, Y., Kaneko, Y. & Iwasa, M.A. (1999) Identification and vertical distribution of two species of Eothenomys in the Oze district, northeastern Honshu, Japan. Mammalian Science, 39, 257–268.

    Kimura, Y., Kaneko, Y. & Yoshida, T. (1994) Small mammalian fauna in Adatara mountain regions with special reference to genus Eothenomys. Fukushima Seibutsu, 37, 13–19.

    Koh, H.S., Yang, B.K., Heo, S.W., Jang, K.H. & In, S.T. (2011) Genetic distinctiveness of the Korean red-backed vole (Myodes regulus) from Korea, revealed by mitochondrial cytochrome b gene sequences. Biochemical Genetics, 49, 153–160.
    https://doi.org/10.1007/s10528-010-9395-3

    Kohli, B.A., Speer, K.A., Kilpatrick, C.W., Batsaikhan, N., Damdinbazar, D. & Cook, J.A. (2014) Multilocus systematics and non-punctuated evolution of Holarctic Myodini (Rodentia: Arvicolinae). Molecular Phylogenetics and Evolution, 76, 18–29.
    https://doi.org/10.1016/j.ympev.2014.02.019

    Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: molecular evolutionary genetics analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33, 1870–1874.
    https://doi.org/10.1093/molbev/msw054

    Lebedev, V.S., Bannikova, A.A., Tesakov, A.S. & Abramson, N.I. (2007) Molecular phylogeny of the genus Alticola (Cricetidae, Rodentia) as inferred from the sequence of the cytochrome b gene. Zoologica Scripta, 36, 547–563.
    https://doi.org/10.1111/j.1463-6409.2007.00300.x

    Lin, Y.H., Waddell, P.J. & Penny, D. (2002) Pika and vole mitochondrial genomes increase support for both rodent monophyly and glires. Gene, 294, 119–129.
    https://doi.org/10.1016/S0378-1119(02)00695-9

    Liu, S., Jin, W., Liu, Y., Murphy, R.W., Lv, B., Hao, H., Liao, R., Sun, Z., Tang, M., Chen, W. & Fu, J. (2017) Taxonomic position of Chinese voles of the tribe Arvicolini and the description of 2 new species from Xizang, China. Journal of Mammalogy, 98, 166–182.
    https://doi.org/10.1093/jmammal/gyw170

    Liu, S., Liu, Y., Guo, P., Sun, Z., Murphy, R.W., Fan, Z., Fu, J. & Zhang, Y. (2012) Phylogeny of oriental voles (Rodentia: Muridae: Arvicolinae): molecular and morphological evidence. Zoological Science, 29, 610–622.
    https://doi.org/10.2108/zsj.29.610

    Liu, S., Sun, Z., Zeng, Z. & Zhao, E. (2007) A new vole (Cricetidae: Arvicolinae: Proedromys) from the Liangshan Mountains of Sichuan Province, China. Journal of Mammalogy, 88, 1170–1178.
    https://doi.org/10.1644/06-mamm-a-141r2.1

    Luo, J., Yang, D., Suzuki, H., Wang, Y., Chen, W.J., Campbell, K.L. & Zhang, Y.P. (2004) Molecular phylogeny and biogeography of Oriental voles: genus Eothenomys (Muridae, Mammalia). Molecular Phylogenetics and Evolution, 33, 349–362.
    https://doi.org/10.1016/j.ympev.2004.06.005

    Luo, Z.X., Chen, W. & Gao, W. (2000) Mammalia, Vol.6, Rodentia, Part III: Cricetidae. In: Luo, Z.X., Chen, W. & Gao, W. (Eds.), Fauna Sinica. Science Press, Beijing, pp. 333–388. [in Chinese]

    Mead, J.I. & Nadachowski, A. (1999) Alticola stoliczkanus. Mammalian Species, 624, 1–4.

    Mezhzherin, S.V. & Serbenyuk, M.A. (1992) Biochemical variability and genetic divergence of palearctic Arvicolidae–the genus Clethrionomys Tilesius, 1850. Genetika, 28, 143–153.

    Miller, G.S. (1900) Preliminary revision of the European redbacked mice. Proceedings of the Washington Academy of Sciences, 2, 83–109.

    Miller, G.S. (1906) Some voles from the Tian Shan region. Annals and Magazine of Natural History, 7, 371–375.
    https://doi.org/10.1080/00222930608562539

    Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Gateway Computing Environments Workshop (GCE), 2010, 1–7.
    https://doi.org/10.1109/GCE.2010.5676129

    Musser, G.G. & Carleton, M.D. (1993) Muridae: Arvicolinae. In: Wilson, D.E. & Reeder, D.M. (Eds.), Mammal species of the world: a taxonomic and geographic reference. 2nd Edition. Smithsonian Institution Press, Washington, D.C., pp. 501–536.

    Musser, G.G. & Carleton, M.D. (2005) Superfamily Muroidea. In: Wilson, D.E. & Reeder, D.M. (Eds.), Mammal species of the world: a taxonomic and geographic reference. 3rd Edition. Johns Hopkins University Press, Baltimore, Maryland, pp. 894–1531.

    Nadachowski, A. & Mead, J.I. (1999) Alticola argentatus. Mammalian Species, 625, 1–4.
    https://doi.org/10.2307/3504506

    Pääbo, S. & Wilson, A.C. (1988) Polymerase chain reaction reveals cloning artefacts. Nature, 334, 387–388.
    https://doi.org/10.1038/334387b0

    Pavlinov, I.Y. (2003) Systematics of modern mammals. Moscow State University, Moscow, 537 pp. [in Russian]

    Pavlinov, I.Y. & Rossolimo, O.L. (1987) Systematics of mammals of the USSR. Moscow University Press, Moscow, 282 pp. [in Russian]

    Posada, D. & Buckley, T.R. (2004) Model selection and model averaging in phylogenetics: advantages of Akaike information criterion and Bayesian approaches over likelihood ratio tests. Systematic Biology, 53, 793–808.
    https://doi.org/10.1080/10635150490522304

    Posada, D. & Crandall, K.A. (1998) Modeltest: testing the model of DNA substitution. Bioinformatics, 14, 817–818.
    https://doi.org/10.1093/bioinformatics/14.9.817

    R Core Team (2017) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from: https://www.r-project.org/ (accessed 13 December 2017)

    Rohlf, F.J. (2015) The tps series of software. Hystrix, the Italian Journal of Mammalogy, 26, 9–12.
    https://doi.org/10.4404/hystrix-26.1-11264

    Ronquist, F., Teslenko, M., Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542.
    https://doi.org/10.1093/sysbio/sys029

    Rossolimo, O. & Pavlinov, I.J. (1992) Species and subspecies of Alticola s. str. (Rodentia: Arvicolidae). In: Horacek, I. & Vohralik, V. (Eds.), Prague studies in mammalogy. Charles University Press, Prague, pp. 149–176.

    Rossolimo, O., Pavlinov, I.Y. & Hoffmann, R. (1994) Systematics and distribution of the rock voles of the subgenus Alticola s. str. in the People’s Republic of China (Rodentia, Arvicolinae). Acta Theriologica Sinica, 14, 86–99.

    Rossolimo, O.L. (1989) Revision of Royle’s high-mountain vole Alticola (A.) argentatus (Mammalia: Cricetidae). Zoologicheskii Zhumal, 68, 104–114. [in Russian]

    Sansalone, G., Kotsakis, T. & Piras, P. (2015) Talpa fossilis or Talpa europaea? using geometric morphometrics and allometric trajectories of humeral moles remains from Hungary to answer a taxonomic debate. Palaeontologia Electronica, 18, 1–17.
    https://doi.org/10.26879/560

    Severtzov, N.A. (1879) Remarks on vertebrate fauna of Pamir. Zapiski Turkestanskogo Otdela Obshchestva Lubitelei Estetvoznanya, 1, 63–64. [in Russian]

    Sheets, H.D. (2014) Morphometric software IMP 8.0. Department of Geology, SUNY at Buffalo, New York. Available from: http://www3.canisius.edu/~sheets/IMP%208.htm (accessed 2 April 2017)

    Sheng, H.L., Ohtaishi, N. & Lu, H.J. (1999) Wild mammals of China. China Forestry Publishing House, Beijing, 297 pp. [in Chinese]

    Sikes, R.S., Gannon, W.L. & the Animal Care and Use Committee of the American Society of Mammalogists (2011) Guidelines of the American society of mammalogists for the use of wild mammals in research. Journal of Mammalogy, 92, 235–253.
    https://doi.org/10.1644/10-mamm-f-355.1

    Smith, A.T., Xie, Y., Hoffmann, R.S., Lunde, D., MacKinnon, J., Wilson, D.E., Wozencraft, W.C. & Gemma, F. (2010) A guide to the mammals of China. Princeton University Press, New Jersey, 576 pp.

    https://doi.org/10.1515/9781400834112

    Stöver, B.C. & Müller, K.F. (2010) TreeGraph 2: combining and visualizing evidence from different phylogenetic analyses. BMC Bioinformatics, 11, 7.
    https://doi.org/10.1186/1471-2105-11-7

    Swofford, D.L. (2003) PAUP*: phylogenetic analysis using parsimony (* and other methods). Version 4.0b10. Sinauer Associates, Sunderland, Mass. [software]

    Tamura, K. & Nei, M. (1993) Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees. Molecular Biology and Evolution, 10, 512–526.
    https://doi.org/10.1093/oxfordjournals.molbev.a040023

    Tanaka, R. (1971) A research into variation in molar and external features among a population of Smith’s red-backed vole for elucidation of its systematic rank. Japanese Journal of Zoology, 16, 163–176.

    Tang, Y., Yukawa, T., Bateman, R.M., Jiang, H. & Peng, H. (2015) Phylogeny and classification of the East Asian Amitostigma alliance (Orchidaceae: Orchideae) based on six DNA markers. BMC evolutionary biology, 15, 96.
    https://doi.org/10.1186/s12862-015-0376-3

    Thomas, O. (1880) XL.—Description of a new species of Arvicola from northern India. Annals and Magazine of Natural History, 6, 322–323.
    https://doi.org/10.1080/00222938009458945

    Thomas, O. (1905) On some new Japanese mammals presented to the British Museum by Mr. R. Gordon Smith. Annals and Magazine of Natural History, 15, 487–495.
    https://doi.org/10.1080/03745480509442838

    Thomas, O. (1907) The Duke of Bedford’s zoological exploration in eastern Asia.—IV. List of small mammals from the islands of Saghalien and Hokakaido, Proceedings of the Zoological Society of London, 77, 404–414.

    Thomas, O. (1908) The Duke of Bedford’s zoological exploration in eastern Asia.—X. List of mammals from the Provinces of Chih‐li and Shan‐si, N. China. Proceedings of the Zoological Society of London, 78, 635–646.
    https://doi.org/10.1111/j.1469-7998.1908.tb07397.x

    Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The CLUSTAL_X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 25, 4876–4882.
    https://doi.org/10.1093/nar/25.24.4876

    Triant, D.A. & DeWoody, J.A. (2006) Accelerated molecular evolution in Microtus (Rodentia) as assessed via complete mitochondrial genome sequences. Genetica, 128, 95–108.
    https://doi.org/10.1007/s10709-005-5538-6

    Triant, D.A. & DeWoody, J.A. (2009) Demography and phylogenetic utility of numt pseudogenes in the southern red-backed vole (Myodes gapperi). Journal of Mammalogy, 90, 561–570.
    https://doi.org/10.1644/08-mamm-a-149r1.1

    Venables, W.N. & Ripley, B.D. (2002) Modern applied statistics with S. 4th Edition. Springer, New York, 495 pp.
    https://doi.org/10.1007/978-0-387-21706-2

    Wójcik, J.M., Kawałko, A., Marková, S., Searle, J.B. & Kotlík, P. (2010) Phylogeographic signatures of northward post-glacial colonization from high-latitude refugia: a case study of bank voles using museum specimens. Journal of Zoology, 281, 249–262.
    https://doi.org/10.1111/j.1469-7998.2010.00699.x

    Wang, Y.X. (2003) A complete checklist of mammal species and subspecies in China: a taxonomic and geographical reference. China Forestry Publishing House, Beijing, 394 pp. [in Chinese]

    Xia, L., Yang, Q.S., Ma, Y., Feng, Z.J. & Zhou, L.Z. (2006) A guide to the measurement of mammal skull III: Rodentia and Lagomorpha. Chinese Journal of Zoology, 41, 68–71. [in Chinese]

    Yang, Q.S., Xia, L., Ma, Y., Feng, Z.J. & Quan, G.Q. (2005) A guide to the measurement of mammal skull I: basic measurement. Chinese Journal of Zoology, 40, 50–56. [in Chinese]

    Zelditch, M.L., Swiderski, D.L. & Sheets, H.D. (2012) Geometric morphometrics for biologists: a primer second edition. Elsevier Academic Press, London, 488 pp.

    Zeng, T., Jin, W., Sun, Z.Y., Liu, Y., Murphy, R.W., Fu, J.R., Wang, X., Hou, Q.F., Tu, F.Y., Liao, R., Liu, S.Y. & Yue, B.S. (2013) Taxonomic position of Eothenomys wardi (Arvicolinae: Cricetidae) based on morphological and molecular analyses with a detailed description of the species. Zootaxa, 3682, 85–104.
    https://doi.org/10.11646/zootaxa.3682.1.3