中国竹鼠属(啮齿目:鼹形鼠科)分类与系统发育研究
收稿日期: 2025-07-16
录用日期: 2025-12-30
网络出版日期: 2026-06-04
基金资助
四川省基本科研项目(2023JBKY10);国家自然科学基金(31970399);国家公园动物多样性调查与生态系统敏感性研究(2025LCTD0402);四川省科技计划项目(2022YFS0483)
A taxonomic and phylogenetic study of the genus Rhizomys (Rodentia: Spalacidae) in China
Received date: 2025-07-16
Accepted date: 2025-12-30
Online published: 2026-06-04
鼹形鼠科(Spalacidae)物种专营地下生活,具有适应地下洞穴环境的系列表型特征,是适应进化研究的重要类群。竹鼠属(Rhizomys)位于该科系统演化的基部分支,近年来分子测序及适应进化研究逐步深入,但受新增标本量限制,属内物种及亚种形态度量比较和系统发育关系研究仍较薄弱、鉴别特征还需厘定。基于国内新采集竹鼠标本和查看的馆藏标本数据,利用形态学和分子系统学方法开展分类研究,结果表明:银星竹鼠(R. pruinosus)、中华竹鼠(R. sinensis)和大竹鼠(R. sumatrensis)可依据体背尖端白色的长柱状毛、脸颊颜色及尾长比例鉴定区分,尾长与头体长之比的均值分别为42.3%、27.9%和34.5%;中华竹鼠四川亚种(R. s. vestitus)个体显著大于福建亚种(R. s. davidi),前者颅全长大于70 mm,体背毛尖粉棕色段较长,脸颊灰白色,后者颅全长小于70 mm,体背灰棕色、毛尖棕色段较短;银星竹鼠云南亚种(R. p. senex)可依据牙齿较粗壮、体背柱状长毛明显稀疏且尖端白色区段较短的特征与华南亚种(R. p. latouchei)相区分;基于Cyt b基因构建的系统发育树显示银星竹鼠与大竹鼠亲缘关系较近,中华竹鼠为竹鼠属内较早分化的类群,种间遗传距离为9.3% ~ 15.9%,中华竹鼠四川亚种与福建亚种、银星竹鼠云南亚种与华南亚种间的遗传距离分别为2.3%和0.4%,揭示不同亚种间的分化程度存在差异。本研究为完善竹鼠属系统分类积累了基础数据,关于暗褐竹鼠(R. wardi)的分类地位有待更全面的采样研究来确定。
唐明坤 , 王旭明 , 刘莹洵 , 廖锐 , 彭步青 , 王新 , 刘少英 . 中国竹鼠属(啮齿目:鼹形鼠科)分类与系统发育研究[J]. 兽类学报, 2026 , 46(3) : 307 -319 . DOI: 10.16829/j.slxb.151126
Species of the family Spalacidae are exclusively subterranean, possessing a suite of phenotypic traits adapted to life in underground burrows, making them an important group for studying adaptive evolution. The genus Rhizomys represents a basal lineage within this family. While recent years have witnessed significant progress in molecular sequencing and adaptive evolution research on this genus, morphological comparisons and phylogenetic relationships among its constituent species and subspecies remain poorly understood due to limited availability of newly acquired specimens, and diagnostic characters still need to be clearly defined. Based on newly collected specimens and museum collections from China, we performed a systematic taxonomic study using morphological and molecular phylogenetic approaches. Our results demonstrate that Rhizomys pruinosus, R. sinensis, and R. sumatrensis can be distinguished by their long dorsal guard hairs with white tips, cheek coloration, and the ratio of tail length to head-body length (mean values: 42.3%, 27.9%, and 34.5%, respectively). The subspecies R. sinensis vestitus is significantly larger than R. s. davidi;it has a greatest skull length exceeding 70 mm, longer pinkish-brown tips on the dorsal fur, and greyish-white cheeks. In contrast, R. s. davidi has a greatest skull length less than 70 mm, greyish-brown dorsal fur with noticeably shorter brown tips. Rhizomys pruinosus senex can be distinguished from R. p. latouchei by its robust teeth and sparser dorsal long guard hairs with a shorter white tip section. Phylogenetic reconstruction based on the cytochrome b gene revealed a closer relationship between R. pruinosus and R. sumatrensis, with R. sinensis representing an early-diverging lineage within the genus. Interspecific genetic distances ranged from 9.3% to 15.9%. At the intraspecific level, the genetic distance was 2.3% between the subspecies R. s. vestitus and R. s. davidi, and 0.4% between R. p. senex and R. p. latouchei, indicating varying degrees of differentiation among subspecies. This study provides foundational data for the systematics of the genus Rhizomys; However, a definitive taxonomic assessment of R. wardi still awaits more comprehensive sampling.
Key words: Rhizomyinae; Morphology; Phylogeny; Cyt b; Subspecies; Taxonomic key
| Allen G M,1940.The mammals of China and Mongolia[M]// Granger W ed. Natural history of central Asia,vol XI Part 2. New York:Central Asiatic Expeditions of the American Museum of Natural History,622?1289. | |
| Anderson J,1879.Anatomical and zoological researches in western Yunnan[M].London:Quaritch,1?984. | |
| Blyth E,1851. Notice of a collection of Mammalia,Birds,and Reptiles,procured at or near the station of Chérra Punji in the Khásia hills,north of Sylhet[J].Journal of the Asiatic Society of Bengal,20:519. | |
| Bradley R D, Baker R J,2001.A test of the genetic species concept:cytochrome?b sequences and mammals[J].Journal of Mammalogy,82(4):960?973.DOI:10.1644/1545-1542(2001)082<0960:ATOTGS>2.0.CO;2 . | |
| Brongersma L D,1936.On the subspecies of Rhizomys sumatrensis(Raffles)with some notes on related species[J].Zoologische Mededelingen,19(12):137?164. | |
| Dao V T, Cao V S,1990. Six new Vietnamese rodents[J].Mammalia,54(2):233?238. DOI:org/10.1515/mamm-1990-0207 . | |
| Ellerman J R, Morrison?Scott T C S,1951. Checklist of palaearctic and Indian mammals,1758 to 1946[M].London:Trustees of the British Museum(Natural History),550?553. | |
| Ellerman J R,1940.The Families and Genera of living rodents,Volume I. Rodents other than Muridae[M].London:Printed by Order of the Trustees of the British Museum,644?651. | |
| Emerling C A, Springer M S,2014. Eyes underground:regression of visual protein networks in subterranean mammals[J].Molecular Phylogenetics and Evolution,78:260?270. DOI:10.1016/j.ympev.2014.05.016 . | |
| Gray J E,1831. Characters of three new genera,including two new species of mammals from China[J].Proceedings of the Committee of Science and Correspondence of the Zoological Society of London,part I,95:1830?1831. | |
| Guo Y T, Zhang J, Xu D M, Tang L Z, Liu Z,2021. Phylogenomic relationships and molecular convergences to subterranean life in rodent family Spalacidae[J].Zoological Research,42(5):671?674. DOI:10.24272/j.issn.2095-8137.2021.240 . | |
| He Y, Hu S, Ge D, Connor T, Zhou C,2019. Evolutionary history of Spalacidae inferred from fossil occurrences and molecular phylogeny[J].Mammal Review,50(1):11?24.DOI:10.1111/mam.12170 . | |
| Hodgson B H,1842. Notice of the mammals of Xizang,with descriptions and plates of some new species[J].Journal of the Asiatic Society of Bengal,11:275?289. | |
| Holden M E, Cserkész T, Musser G G,2017. Family Sminthidae,Vol. 7: Rodents II [M]//Wilson D E,Lacher T E,Mittermeier R A eds. Handbook of the mammals of the world. Barecelona: Lynx Ediciones,134?136. | |
| Huang Xiaofu, Li Jialie,2004. Elementary study on the habitat selection of bamboo rats in Piankou Natural Reserve[J].Journal of China West Normal University(Natural Sciences),25(4):375?379.DOI:10.16246/j.issn.1673-5072.2004.04.010. (in Chinese with English abstract) | |
| Irwin D M, Kocher T D, Wilson A C,1991. Evolution of the cytochrome b gene of mammals[J].Journal of Molecular Evolution,32(2):128?144. DOI:10.1007/BF02515385 . | |
| Kimura M,1980. A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences[J].Journal of Molecular Evolution,16:111?120. DOI:10.1007/BF01731581 . | |
| Lan Xiaoguang,2023. Bamboo forest resources and utilization in Prehistoric in China[J].World Bamboo and Rattan,21(6):12?21.DOI:10.12168/sjzttx.2023.11.11.001. (in Chinese with English abstract) | |
| Li K X, Liu Y X, Zhang Y H, Fan Z X, Hu X X, Lai X L, Wei F W, Hu Y B,2025. Host genomic convergence,rather than gut microbiome convergence,underlies the convergent evolution of subterranean adaptation in mammals[J].Cell Reports,44(8):116110. DOI:10.1016/j.celrep.2025.116110 . | |
| Lin G H, Wang K, Deng X G, Nevo E, Zhao F, Su J P, Guo S C, Zhang T Z, Zhao H,2014. Transcriptome sequencing and phylogenomic resolution within Spalacidae(Rodentia)[J]. BMC Genomics,15:32. DOI:10.1186/1471-2164-15-32 . | |
| Liu Shaoying, Jin Wei, Liao Rui, Sun Zhiyu, Zeng Tao, Fu Jianrong, Liu Yang,Wangxin, Li Panfeng, Tang Mingkun, Chen Limin, Dong Li, Han Mingde, Gou Dan,2017. Phylogenetic study of Ochotona based on mitochondrial Cyt b and morphology with a description of one new subgenus and five new species[J].Acta Theriologica Sinica,37(1):1?43. DOI: 10.16829/j.slxb.201701001. (in Chinese with English abstract) | |
| Liu S Y, Liu Y, Guo P, Sun Z Y, Murphy R W, Fan Z X, Fu J R, Zhang Y P,2012. Phylogeny of oriental voles(Rodentia: Muridae:Arvicolinae):Molecular and morphological evidence[J].Zoological Science,29(9):610?622. DOI:10.2108/zsj.29.610 . | |
| Liu S Y, Sun Z Y, Zeng Z Y, Zhao E,2007. A new vole(Cricetidae:Arvicolinae:Proedromys)from the Liangshan Mountains of Sichuan Province,China[J].Journal of Mammalogy,88(5):1170?1178. DOI:10.1644/06-MAMM-A-141R2.1 . | |
| Milne?Edwards A,1868?1874.Recherches pour servir à l’histoire naturelle des mammifères[M].Paris:Masson,vol. I,1? 394. | |
| Musser G G, Carleton M D,1993. Family Muridae[M]//Wilson D E,Reeder D M eds. Mammal species of the world:a taxonomic and geographic reference,2nd Edition. Washington:Smithsonian Institution Press,685?687. | |
| Musser G G, Carleton M D. 2005. Superfamily Muroidea[M]//Wilson D E,Reeder D M eds. Mammal species of the world:a taxonomic and geographic reference,3rd Edition. Baltimore:Johns Hopkins University Press,894?1531. | |
| Raffles T S,1821. Descriptive catalogue of a zoological collection,made on account of the honorable East Indian Company,in the island of Sumatra and its vicinity,under the direction of Sir Thomas Stamford Raffles,Lieutenant?Governor of Fort Marlborough,with additional notices illurstrative of the natural history[J].Transactions of the Linnean Society of London,13:239?274. | |
| Ronquist F, Huelsenbeck J P,2003. MRBAYES 3: Bayesian phylogenetic inference under mixed models[J].Bioinformatics,19(12):1572?1574. DOI:10.1093/bioinformatics/btg180 . | |
| Shih C M,1930. Preliminary report on the mammals from Yaoshan,Kwangsi,collected by the Yaoshan expedition,Sun Yatsen University,Canton,China[J].Bulletin of the Department of Biology,College of Science,Sun Yatsen University,4:10. | |
| Stamatakis A, Hoover P, Rougemont J,2008.A rapid bootstrap algorithm for the RAxML web servers[J].Systematic Biology,57(5):758?771. DOI:10.1080/10635150802429642 . | |
| Swinhoe R,1870.Catalogue of the mammals of China(south of the river Yangtsze)and of the island of Formosa[M]//Proceedings of the scientific meetings of the zoological society of London for the year 1870. London:Messrs.Longmans,Green,Reader and Dyer,637. | |
| Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S,2011.MEGA5:Molecular evolutionary genetics analysis using maximum likelihood,evolutionary distance,and maximum parsimony methods[J].Molecular Biology and Evolution,28(10):2731?2739. DOI:10.1093/molbev/msr121 . | |
| Tang Zhonghai, Peng Bo, You Zhangqiang, Liu Hao, Shi Hongyan,2009. Habitation selection and den structure characteristics of Rhizomys sinensis in Piankou Natural Reserve[J]. Chinese Journal of Zoology,44(6):36?40.DOI:10.13859/j.cjz.2009.06.005 (in Chinese with English abstract) | |
| Temminck C J,1832. Monographie over een nieuw geslacht van Knaagdier,onder den naam van Nyctocleptes [J].Bijdragen tot de Natuuren Sterrenkunde,7:1?8. | |
| Thomas O,1911. The Duke of Bedford’s zoological exploration of Eastern Asia. ⅩⅢ. On mammals from the provinces of Kan?su and Sze?chwan,western China[J].Proceedings of the Zoological Society of London,1911:158?180.DOI:10.1111/j.1469-7998.1912.tb07008.x . | |
| Thomas O,1915a. VII.-Notes on the Asiatic bamboo?rats(Rhizomys,etc.)[J]. Annals and Magazine of Natural History,16(91):56?61. | |
| Thomas O,1915b. XXXVII.-Further notes on Asiatic bamboo?rats [J]. Annals and Magazine of Natural History,16(94):313?317. | |
| Thomas O, 1916. L.-A new bamboo?rat from perak[J].Journal of Natural History,18(107):445?446.DOI:10.1080/00222931609486894 . | |
| Thomas O,1921.On small mammals from the Kachin Province,Northern Burma[J].Journal of the Bombay Natural History Society,27:499?505. | |
| Wan N, Chen J Q, Jiang M L, Wu M Z, Li H L, Wen M Z, Shen W Z, Liang X L, Yang X J, Liu Y J, Gao Y G, Nevo E, Li K X,2025. Genomic signatures of subterranean lifestyle and ecological adaptation in Myospalax [J].BMC Biology,23:332. DOI:10.1186/s12915-025-02431-4 . | |
| Wei Fuwen, Yang Qisen, Wu Yi, Jiang Xuelong, Liu Shaoying, Hu Yibo, Ge Deyan,Libaoguo, Yang Guang, Li Ming, Zhou Jiang, Li Song, Li Sheng, Yu Wenhua, Chen Bingyao, Zhang Zejun, Zhou Caiquan, Wu Shibao, Zhang Li, Chen Zhongzheng, Chen Shunde, Deng Huaiqing, Jiang Tinglei, Zhang Libiao, Shi Hongyan, Lu Xueli, Li Quan, Liu Zhu, Cui Yaqian, Li Yuchun, He Kai,2025. Catalogue of mammals in China(2024)[J].Acta Theriologica Sinica,45(1):1?16.DOI:10.16829/j.slxb.151039. (in Chinese with English abstract) | |
| Xia Lin, Yang Qisen, Ma Yong, Feng Zuojian, Zhou Lizhi,2006. A guide to the measurement of mammal skull Ⅲ:Rodentia and Lagomorpha[J].Chinese Journal of Zoology,41(5):68?71.DOI:10.13859/j.cjz.2006.05.013. (in Chinese with English abstract) | |
| Xu Longhui,1984.Studies on the biology of the hoary bamboo rat(Rhizomys pruinosus Blyth)[J].Acta Theriologica Sinica,4(2):99?105.DOI:10.16829/j.slxb.1984.02.005. (in Chinese with English abstract) | |
| Xu Y, Liu X H, Tu F Y,2014. Complete mitochondrial genome of Chinese bamboo rat,Rhizomys sinensis and species divergence comparison[J].Mitochondrial DNA,27(3):1773?1774. DOI:10.3109/19401736.2014.963806 . | |
| Xu Y, Wen B, Zhu K L, Du Z H, Bai X J,2018. The complete mitochondrial genome of Rhizomys sumatrensis(Rodentia: Spalacidae)and its phylogenetic implications[J].Conservation Genetics Resources,10:685?688.DOI:10.1007/s12686-017-0901-3 . | |
| Yang Qisen, Xia Lin, Ma Yong, Feng Zuojian, Quan Guoqiang,2005. A guide to the measurement of mammal skull Ⅰ:basic measurement[J].Chinese Journal of Zoology,40(3):50?56. DOI:10.13859/j.cjz.2005.03.011. (in Chinese with English abstract) | |
| Smith A T,解焱,2009. 中国兽类野外手册[M].长沙:湖南教育出版社,87?95. | |
| 王酉之,胡锦矗,1999. 四川兽类原色图鉴[M].北京:中国林业出版社,245?246. | |
| 王应祥,2003.中国哺乳动物种和亚种分类名录与分布大全[M].北京:中国林业出版社,212?214. | |
| 刘少英,石红艳,李松,陈顺德,徐怀亮,胡杰,刘洋,余文华,孙治宇,李晟,2023. 四川兽类志[M].北京:中国林业出版社,803?824. | |
| 刘少英,靳伟,廖锐,孙治宇,曾涛,符建荣,刘洋,王新,李盼峰,唐明坤,谌利民,董立,韩明德,苟丹,2017. 基于Cyt b基因和形态学的鼠兔属系统发育研究及鼠兔属1新亚属5新种描述[J].兽类学报,37(1):1?43. DOI:10.16829/j.slxb.201701001 . | |
| 杨奇森,夏霖,马勇,冯祚建,全国强,2005.兽类头骨测量标准Ⅰ:基本量度[J].动物学杂志,40(3):50?56.DOI:10.13859/j.cjz.2005.03.011 . | |
| 何业恒,1980.中国竹鼠分布的变迁[J].湘潭大学学报(哲学社会科学版),(3):53?58.DOI:10.13715/j.cnki.jxupss. 1980.03.009 . | |
| 欧阳新举,1960.中华竹鼠(Rhizomys sinensis vestitus Milne?Edwards)的初步观察[J].动物学杂志,(5):200?202. DOI:10.13859/j.cjz.1960.05.004 . | |
| 罗蓉,1993.贵州兽类志[M].贵阳:贵州科技出版社,238?246. | |
| 夏霖,杨奇森,马勇,冯祚建,周立志,2006.兽类头骨测量标准Ⅲ:啮齿目,兔形目[J].动物学杂志,41(5):68?71.DOI:10.13859/j.cjz.2006.05.013 . | |
| 徐龙辉,1984.花白竹鼠(Rhizomys pruinosus)的生物学研究[J].兽类学报,4(2):99?105.DOI:10.16829/j.slxb. 1984.02.005 . | |
| 唐中海,彭波,游章强,刘昊,石红艳,2009.中华竹鼠的洞穴结构及其生境利用特征[J].动物学杂志,44(6):36?40.DOI:10.13859/j.cjz.2009.06.005 . | |
| 黄小富,李家烈,2004.四川片口自然保护区中华竹鼠对生境选择的初步研究[J].西华师范大学学报(自然科学版),25(4):375?379. DOI:10.16246/j.issn.1673-5072.2004.04.010 . | |
| 蒋海军,杨耀华,周言言,田军东,路纪琪,2024. 河南发现银星竹鼠[J].四川动物,43(3):345?346. | |
| 蓝晓光,2023.史前中国竹林资源与利用[J].世界竹藤通讯,21(6):12?21. DOI:10.12168/sjzttx.2023.11.11.001 . | |
| 潘清华,王应祥,岩崑,2007.中国哺乳动物彩色图鉴[M].北京:中国林业出版社,311?314. | |
| 魏辅文,杨奇森,吴毅,蒋学龙,刘少英,胡义波,葛德燕,李保国,杨光,李明,周江,李松,李晟,余文华,陈炳耀,张泽钧,周材权,吴诗宝,张立,陈中正,陈顺德,邓怀庆,江廷磊,张礼标,石红艳,卢学理,李权,刘铸,崔雅倩,李玉春,何锴,2025.中国兽类名录(2024版)[J].兽类学报,45(1):1?16. DOI:10.16829/j.slxb.151039 . | |
| 魏辅文,杨奇森,吴毅,蒋学龙,刘少英,2022. 中国兽类分类与分布[M].北京:科学出版社,78?87. |
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