A review of the Cypriniform tribe Yunnanilini Prokofiev, 2010 from China, with an emphasis on five genera based on morphologies and complete mitochondrial genomes of some species
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摘要: 条鳅科鱼类是一类在生态学上具有重要性,在分类学上具有挑战性的淡水鱼类,广泛分布在亚洲大陆和临近岛屿,以及欧洲和非洲东北部。全世界记录条鳅科种类有47属、758种,其中,在中国有记录的有22属、268种。Prokofiev(2010)基于40个形态特征将条鳅科种类分为5个族,即Vaillantellini、北鳅族Lefuini、云南鳅族Yunnanilini、高原鳅族Triplophysini和条鳅族Nemacheilini。 利用条鳅科部分种的线粒体基因组,对云南鳅族的系统发育关系及其物种的分类地位进行研究。分子结果暗示:首先,云南鳅族不是单系的,云南鳅族种类被分为三支:黑斑云南鳅(Y. nigromaculatus)构成独立的一支,位于树的基部; Petruichthys brevis与条鳅族的南鳅属(Schistura)种类形成姐妹群;小条鳅属(Micronemacheilus)、异条鳅属(Paranemachilus)、沙猫鳅属(Traccatichthys)、云南鳅属(Yunnanilus)与北鳅族的北鳅属(Lefua)、岭鳅属(Oreonectes)和洞鳅属(Troglonectes)交互聚在一起;四川云南鳅(Y. sichuanensis)与高原鳅属种类聚为一支。其次,系统发育结果不支云南鳅属的单系性,黑斑云南鳅形成一个单系,与条鳅科其它种类形成姐妹群;靖西云南鳅(Y. jinxiensis)与颊鳞异条鳅(P. genelepis)聚成一个单系,而丽纹云南鳅(Y. pulcherrimus)与Micronemachelus cruciatus聚为一个单系。 在形态上,Eonemachilus(中文名:始条鳅属)、间条鳅属、小条鳅属、异条鳅属、原条鳅属、中条鳅属和云南鳅属可依据前、后鼻孔的位置、唇部结构、颊部是否被鳞,以及头部侧线管是否存在等联合性状,将这些属区分开来,如间条鳅属、异条鳅属、原条鳅属和沙猫鳅属的前、后鼻孔相邻,而始条鳅属、小条鳅属和云南鳅属的前、后鼻孔分开一小段距离。此外,异条鳅属颊部被鳞,而其它属种类颊部无鳞。小条鳅属和沙猫鳅属的上下唇均有较大的乳突。 在该研究中,依据分子和形态结果,将丽纹云南鳅移入小条鳅属;长须云南鳅和白莲云南鳅移到间条鳅属;靖西云南鳅移到异条鳅属。此外,原云南鳅属中黑斑云南鳅种组的种类被移到始条鳅属。 该研究尚存在一些不足,首先,因缺少间条鳅属模式种——郑氏间条鳅的组织样本,对于间条鳅属的分类地位及其系统发育关系尚未解决;其次,因缺少越南种类Micronemachelus cruciatus的标本,依据文献描述,该种与丽纹小条鳅在形态上不易区分,丽纹小条鳅与M. cruciatus的关系还有待进一步的研究;最后,后鳍云南鳅是目前仅记录于广西的云南鳅属的种类,虽然,蓝家湖等(2013)将其从岭鳅属分离出来,并移到云南鳅属,但是,后鳍云南鳅的前鼻孔形态与云南鳅属其它种类间的差别较大,该种是否为云南鳅属种类,还有待进一步的证实。
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关键词:
- 始条鳅属 /
- 间条鳅属 /
- 小条鳅属;异条鳅属;云南鳅属 /
- 系统分类
Abstract: The loach tribe Yunnanilini from China is reviewed here using morphological characters and complete mitochondrial genomes of select species. Molecular data suggest that the tribe Yunnanilini is not monophyletic and can be divided into three clades. Species of the Yunnanilus nigromaculatus group form an independent genus and are placed in Eonemachilus. In the phylogenetic tree, Y. jinxiensis clusters with Paranemachilus genilepis, and Y. pulcherrimus clusters with Micronemacheilus cruciatus, indicating that Y. jinxiensis and Y. pulcherrimus belong to Paranemachilus and Micronemacheilus, respectively. Based on morphological data, Y. bailianensis and Y. longibarbatus are placed in Heminoemacheilus, while Y. jinxiensis and Y. pulcherrimus, are placed in Paranemachilus and Micronemacheilus, respectively. Yunnanilus niulanensis and Y. qujinensis are treated as junior synonyms of Eonemachilus caohaiensis. Eonemachilus, Micronemacheilus, and Yunnanilus are show short separation between anterior and posterior nostrils. The genera can be distinguished from each other by mouth structure, lateral line and cephalic lateral-line canals, and papillae on median part of both lips. The anterior and posterior nostrils of Heminoemacheilus and Paranemachilus are closely set. Paranemachilus and Micronemacheilus are distinguished by cheeks covered with scales and lips with papillae, respectively. Our phylogenetic tree and morphological characters support Traccatichthys as a valid genus, which can be distinguished from Micronemacheilus by anterior and posterior nostrils closely set (vs. clearly separated). Four species are placed in Traccatichthys.-
Key words:
- Eonemachilus /
- Heminoemacheilus /
- Micronemacheilus, Paranemachilus, Yunnanilus /
- Taxonomy
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Figure 2. The types of lower lip, cephalic lateral-line pores and nostrils in Yunnanilini
A–D: Lower lip. A: Micronemacheilus pulcherrimus, with papillae; B: Yunnanilus pleurotaenia; C: Paranemachilus jinxiensis, with two strong furrows on each side, D: Y. chui, with slight furrows. E, F: Y. jiuchiensis, cephalic lateral-line pores. G: Eonemachilus caohaiensis, anterior and posterior nostrils well separated. H: Heminoemacheilus zhengbaoshani, anterior and posterior nostrils closely set. Abbreviations: A1+A2: Infraorbital canal; AN: Anterior nostril; B: Supraorbital canal; C: Supratemporal canal; D: Preoperculomandibular canal; F: Furrow; N: Nostril; P: Papillae; PN: Posterior nostril. Scale bar: 1 mm.
Figure 3. Map showing distribution of species of Eonemachilus, Heminoemacheilus, Paranemachilus, Micronemacheilus, and Yunnanilus
Red represents Eonemachilus, E1: E. altus; E2: E. bajiangensis; E3: E. caohaiensis; E4: E. longidorsalis; E5: E. niger; E6: E. nigromaculatus; E7: E. obtusirostris; E8: E. pachycephalus; E9: E. yangzonghaiensis. Purple represents Heminoemacheilus: H1: H. bailianensis; H2: H. longibarbatus; H3: H. zhengbaoshani; H4: H. hyalinus. Blue represents Micronemacheilus: M: M. pulcherrimus. Green represents Paranemachilus: P1: P. genilepis; P2: P. jinxiensis; P3: P. pingguoensis. Black represents Yunnanilus: Y1: Y. analis; Y2: Y. beipanjiangensis; Y3: Y. chui; Y4: Y. discoloris; Y5: Y. elakatis; Y6: Y. forkicaudalis; Y8: Y. jiuchiensis; Y9: Y. longibulla; Y10: Y. macrogaster; Y11: Y. macrositanus; Y12: Y. macrolepis; Y13: Y. nanpanjiangensis; Y14: Y. paludosus; Y15: Y. parvus; Y16: Y. pleurotaenia; Y17: Y. retrodorsalis; Y18: Y. sichuanensis; Y19: Y. spanisbripes; Y20: Y. chuanheensis.
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[1] An L, Liu BS, Li WX. 2009. Two new loaches of the genus Yunnanilus (Baliteridae) from Yunnan, China. Acta Zootaxonomica Sinica, 34(3): 630−638. (in Chinese) [2] Berg LS. 1938. On some South China loaches (Cobitidae, Pisces). Bulletin de la Société impériale des Naturalistes de Moscou, 47: 314−318. [3] Chaudhuri BL. 1911. Contributions to the fauna of Yunnan based on collections made by J. Coggin Brown, B. Sc. 1909-1910. Part II – Fishes. Records of the Indian Museum, 6(1): 13−24. [4] Chen WT, Yang JP, Li YF, Li XH. 2019. Exploring taxonomic diversity and biogeography of the family Nemacheilinae (Cypriniformes). Ecology and Evolution, 9(18): 10343−10353. doi: 10.1002/ece3.5553 [5] Chen XY. 2013. Checklist of fishes of Yunnan. Zoological Research, 34(4): 281−343. (in Chinese) [6] Chen ZM, Yang J, Yang JX. 2012. Description of a new species of the genus Yunnanilus Nichols, 1925 (Teleostei: Nemacheilidae) from Yunnan, China. Zootaxa, 3269(1): 57−64. doi: 10.11646/zootaxa.3269.1.5 [7] Ding RH. 1992. A new species of the Yunnanilus from Guizhou, China (Cypriniformes: Cobitidae). Acta Zootaxonomica Sinica, 17(4): 489−491. (in Chinese) [8] Ding RH. 1995. A new species of the genus Yunnanilus from western Sichuan, China (Cypriniformes: Cobitidae). Acta Zootaxonomica Sinica, 20(8): 253−256. (in Chinese) [9] Du CX, Zhang E, Chan BP. 2012. Traccatichthys tuberculum, a new species of nemacheiline loach from Guangdong Province, South China (Pisces: Balitoridae). Zootaxa, 3586(1): 304−312. doi: 10.11646/zootaxa.3586.1.28 [10] Du LN, Hou M, Chen XY, Yang JX. 2018. A new species of Yunnanilus (Cypriniformes: Nemacheilidae) from Sichuan, southwest China. Zootaxa, 4532(1): 137−144. doi: 10.11646/zootaxa.4532.1.9 [11] Du LN, Jiang YE, Chen XY, Yang JX, Aldridge D. 2017. A family-level macroinvertebrate biotic index for ecological assessment of lakes in Yunnan, China. Water Resources, 44(6): 864−874. doi: 10.1134/S0097807817090020 [12] Du LN, Lu YF, Chen XY. 2015. Yunnanilus qujinensis, a new species of loach from Yunnan, southwest China (Teleostei: Nemacheilidae). Ichthyological Exploration of Freshwaters, 26(3): 249−254. [13] Eagderi S, Mousavi-Sabet H, Freyhof J. 2019. Paraschistura makranensis, a new loach from the Jegin River drainage in southern Iran with comments on P. ilamensis and P. pasatigris (Teleostei: Nemacheilidae). Zootaxa, 4668(2): 258−270. doi: 10.11646/zootaxa.4668.2.6 [14] Eschmeyer WN. 2020. Catalog of fishes. Updated database version of April 2020. Catalog databases as made available to FishBase in April 2020. [15] Freyhof J, Serov DV. 2001. Nemacheiline loaches from Central Vietnam with descriptions of a new genus and 14 new species (Cypriniformes: Balitoridae). Ichthyological Exploration of Freshwaters, 12(2): 133−191. [16] Gransee A, Kaus A, Freyhof J, Borchardt D. 2019. New insights into the morphological and molecular variability of the genus Barbatula from across Mongolia. In: Frontiers in Marine Science 6 Conference Abstract: XVI European Congress of Ichthyology. [17] IUCN. 2018[2018-07-05]. The IUCN Red List of Threatened Species. Version 2018–1. http://www.iucnredlist.org. [18] Jiang WS, Zhao YP, Du LN, Wang M. 2021. Yunnanilus chuanheensis, a new loach species (Cypriniformes: Nemacheilidae) from the upper Lixianjiang River in Yunnan, China. Zoological Research, 42(2): 241−245. doi: 10.24272/j.issn.2095-8137.2020.287 [19] Kaya C, Turan D, Bayçelebi E, Kalayci G, Freyhof J. 2020a. Oxynoemacheilus cilicicus, a new nemacheilid loach from the Goksu River in southern Anatolia (Teleostei: Nemacheilidae). Zootaxa, 4808(2): 284−300. doi: 10.11646/zootaxa.4808.2.3 [20] Kaya C, Turan D, Kalayci G, Bayçelebi E, Freyhof J. 2020b. The westernmost known population of Paracobitis (Teleostei, Nemacheilidae), with the description of a new species from the Euphrates River in southern Anatolia. Zootaxa, 4838(4): 525−534. doi: 10.11646/zootaxa.4838.4.6 [21] Kottelat M. 1990. Indochinese Nemacheilines: A Revision of Nemacheiline Loaches (Pisces: Cypriniformes) of Thailand, Burma, Laos, Cambodia and Southern Viet Nam. München: 262 [22] Kottelat M. 2012. Conspectus cobitidum: an inventory of the loaches of the world (Teleostei: Cypriniformes: Cobitoidei). The Raffles Bulletin of Zoology, (S26): 1−199. [23] Kottelat M. 2013. The fishes of the inland waters of southeast Asia: a catalogue and core bibliography of the fishes known to occur in freshwaters, Mangroves and Estuaries. The Raffles Bulletin of Zoology Supplement, (S27): 1−663. [24] Kottelat M, Chu XL. 1988. Revision of Yunnanilus with descriptions of a miniature species flock and six new species from China (Cypriniformes: Homalopteridae). Environmental Biology of Fishes, 23(1-2): 65−94. doi: 10.1007/BF00000739 [25] Lan JH, Gan X, Wu TJ, Yang J. 2013. Cave Fishes of Guangxi, China. Beijing: Science Press, 104–139. (in Chinese) [26] Li WX, Wu DF, Xu K, Gao XM, Chen AL, Wu QL, et al. 1999. Fishes in the Heilongtan reservoir and its drainage. Sichuan Journal of Zoology, 18(1): 3−7. (in Chinese) [27] Liu SQ, Mayden RL, Zhang JB, Yu D, Tang QY, Deng X, et al. 2012. Phylogenetic relationships of the Cobitoidea (Teleostei: Cypriniformes) inferred from mitochondrial and nuclear genes with analyses of gene evolution. Gene, 508(1): 60−72. doi: 10.1016/j.gene.2012.07.040 [28] Luo FG, Huang J, Luo T, Yang J, Zhou HJ, Wen YH. 2019. Complete mitochondrial genome and phylogenetic analysis of Yunnanilus pulcherrimus (Cypriniformes, Nemacheilidae). Mitochondrial DNA Part B, 4(1): 1269−1270. doi: 10.1080/23802359.2019.1591222 [29] Min R, Chen XY, Yang JX, Winterbottom R, Mayden RL. 2012a. Phylogenetic relationships of the genus Homatula (Cypriniformes: Nemacheilidae), with special reference to the biogeographic history around the Yunnan-Guizhou Plateau. Zootaxa, 3586(1): 78−94. doi: 10.11646/zootaxa.3586.1.9 [30] Prokofiev AM. 2004. A new species of the genus Micronoemacheilus Rendahl, 1944 (Balitoridae: Nemacheinae) from Hainan (China) with notes on its status. Journal of Ichthyology, 44(3): 191−198. [31] Prokofiev AM. 2010. Morphological classification of loaches (Nemacheilinae). Journal of Ichthyology, 50(10): 827−913. doi: 10.1134/S0032945210100012 [32] Qiu BM. 2007. Molecular Phylogeny of Cyt B and D-loop Gene in Eleven Species of Yunnanilus, Yunnan University. Master Thesis, Yunnan University, Kunming. (in Chinese) [33] Regan CT. 1904. On a collection of fishes made by Mr. John Graham at Yunnan fu. Annals and Magazine of Natural History, Ser. 7, 13(75): 190–194. [34] Rendahl H. 1944. Einige Cobitiden von Annam und Tonkin. Göteborgs Kungliga Vetenskaps och Vitterhets Samhällas Handlingar, Series B, Matematiska och Naturvetenskapliga Skrifter, 3(3): 37. [35] Ronquist F, Teslenko M, Van Der Mark P, Ayres DL, Darling A, Höhna S, et al. 2012. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61(3): 539−542. doi: 10.1093/sysbio/sys029 [36] Sgouros K, Page LM, Orlofske SA, Jadin RC. 2019. A revised molecular phylogeny reveals polyphyly in Schistura (Teleostei: Cypriniformes: Nemacheilidae). Zootaxa, 4559(2): 349−362. doi: 10.11646/zootaxa.4559.2.8 [37] Shu SS, Jiang WS, Whitten T, Yang JX, Chen XY. 2013. Drought and China’s cave species. Science, 340(6130): 272. [38] Taggart JB, Hynes RA, Prodöuhl PA, Ferguson A. 1992. A simplified protocol for routine total DNA isolation from salmonid fishes. Journal of Fish Biology, 40(6): 963−965. doi: 10.1111/j.1095-8649.1992.tb02641.x [39] Tang QY, Liu HZ, Mayden R, Xiong BX. 2006. Comparison of evolutionary rates in the mitochondrial DNA cytochrome b gene and control region and their implications for phylogeny of the Cobitoidea (Teleostei: Cypriniformes). Molecular Phylogenetics and Evolution, 39(2): 347−357. doi: 10.1016/j.ympev.2005.08.007 [40] Yang JX. 1990. The Fishes of Yunnan, ChinaPart Ⅱ Cyprinidae. Beijing, China: Science Press. (in Chinese) [41] Yang JX. 1991. The fishes of Fuxian Lake, Yunnan, China, with description of two new species. Ichthyological Exploration of Freshwaters, 2(3): 193−202. [42] Yang JX, Chen YR. 1995. The Biology and Resource Utilization of the Fishes of Fuxian Lake, Yunnan. Kunming, China: Yunnan Science and Technology Press, 224. (in Chinese) [43] Yoğurtçuoğlu B, Kaya C, Geiger MF, Freyhof J. 2020. Revision of the genus Seminemacheilus, with the description of three new species (Teleostei: Nemacheilidae). Zootaxa, 4802(3): 477−501. doi: 10.11646/zootaxa.4802.3.5 [44] Zhang CG, Zhao YH. 2016. Species Diversity and Distribution of Inland Fishes in China. Beijing: Science Press, 127–151. (in Chinese) [45] Zheng CY. 1989. The Fishes of the Pearl River. Beijing: Science Press, 46–47. (in Chinese) [46] Zhu SQ. 1989. The Loaches of the Subfamily Nemacheilinae in China. Nanjing, China: Jiangsu Science and Technology Publishing House, 14–20. (in Chinese) [47] Zhu Y, Zhu DG. 2014. Description of a new species of the genus Heminoemachilus (Teleostei: Balitoridae) in Guangxi, China. Journal of Guangdong Ocean University, 34(6): 18−21. (in Chinese) -
ZR-2020-229 Supplementary Material.pdf
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