The Herpetological Journal
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Volume 15, Number 1 January 2005 ISSN 0268-0130 THE HERPETOLOGICAL JOURNAL Published by the Indexed in BRITISH HERPETOLOGICAL SOCIETY Current Contents The Herpetological Journal is published quarterly by the British Herpetological Society and is issued free to members. Articles are listed in Current Awareness in Biological Sciences, Current Contents, Science Citation Index and Zoological Record. Applications to purchase copies and/or for details of membership should be made to the Hon. Secretary, British Herpetological Soc�ety, The Zoological Society of London, Regent's Park, London NWl 4RY, UK. Instructions to authors are printed inside the back cover. All contributions should be addressed to the Scientific Editor (address below). ScientificEd itor: Wolfgang Wuster, School of Biological Sciences, University of Wales, Bangor, Gwynedd, LL57 2UW, UK. E-mail: [email protected] Associate Scientific Editors: J. W. Arntzen (Leiden), R. Brown (Liverpool) Managing Editor: Richard A. Griffiths, The Durrell Institute of Conservation and Ecology, University of Kent, Canterbury, Kent CT2 7NS, UK. E-mail: [email protected] Associate Managing Editors: L. Gillett, M. Dos Santos, J. McKay Editorial Board: Donald Broadley (Zimbabwe) John Cooper (Uganda) John Davenport (Cork) Andrew Gardner (Abu Dhabi) Tim Halliday (Milton Keynes) Michael Klemens (New York) Colin McCarthy (London) Andrew Milner (London) Richard Tinsley (Bristol) BRITISH HERPETOLOGICAL SOCIETY Copyright It is a fundamental condition that submitted manuscripts have not been published and will not be simultaneously submitted or published elsewhere. By submitting a manu script, the authors agree that the copyright for their article is transferred to the publisher if and when the article is accepted for publication. The copyright covers the exclusive rights to reproduce and distribute the article, including reprints and photo graphic reproductions. Permission for any such activities must be sought in advance from the Editor. ADVERTISEMENTS The Herpetological Journal accepts advertisements subject to approval of contents by the Managing Editor, to whom enquiries should be addressed. Herpetological Journal website: http://biology.bangor.ac.uk/-bss 166/HJ FRONT COVER: Male Agama castroviejoi (J. M. Padial) HERPETOLOGICAL JOURNAL, Vol. 15, pp. 1-8 (2005) PHYLOGENETIC RELATIONSHIPS AMONG CHINESE RANIDS INFERRED FROM SEQUENCE DATA SET OF128 AND16S rDNA JIANPING JJANG 1•2 AND KAIY A ZHOU1 'lnstUute of Genetic Resources, Nanjing Normal University, Nanjing 210097, China 2Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 61004 I, China Phylogenetic relationships among twenty-nine species of Ranidae representing thirteen genera were investigated on the basis of I 005 base pairs of mitochondrial DNA sequences of I 2S and I 6S rRNA gene fragments. Sequence data were analyzed using maximum parsimony, likelihood maximum, and neighbour joining with all indel and missing/ambiguous sites deleted. Among the twenty-nine ran ids studied, two clades are well supported by the results of the three analyses, the first consists of twenty-one species in the genera Rana, Glandirana, Rugosa, Pelophylax, Amo/ops, Odorrana, and Hylarana; and the second includes eight species in the genera Fejervarya,Ho plobatrachus, Paa, Nanorana, A ltirana, and Limnonectes; the six genera with multi-species samples- including Amo/ops, Pelophylax, Rugosa, Rana, Odorrana, and Paa - are recognized as distinct lineages with higher bootstrap and quartet puzzling supports: the phylogenetic relationships between species within each lineage are resolved well. The results testify that the traditional genus Rana is heterogeneous. On the basis of the phylogenetic relationships of these tax a, it is suggested that the genera Paa, Nanorana, and Altirana should be removed from the subfamily Raninae and to be included in the subfamily Dicroglossinae. The torrent frog of the genus Amolops should be retained in the subfamily Raninae rather than in a distinct subfamily Amolopinae of its own. The inclusion of Fejervaryalimnocharis in the genus Limnonectes is not supported. Key words: China, molecular systematics, mtDNA, Ranidae INTRODUCTION era proposed by Fei et al. (1990), including na, Rugosa, Glandirana, and Odorrana, were With over 700 species, the family Ranidae is one of Pseudora treated as subgenera of the genus by Dubois the most species-rich amphibian families. It is distrib Rana so were the genera reintroduced by Fei uted throughout the world, except southern South (1992), et al. including and con America and most of Australia(Frost, 1985, Duellman, ( 1990), Pelophylax, Hylarana, Rana sisting of only brown frogs. Zhao and 1993 ). There are a few reports on the relationships of (1994, 1995) Inger ( criticised the new taxonomy proposed by groups from lineages or regions, based on morphologi 1996) this, cal and (or) molecular data sets, such as Boulenger Fei et al. (1990) and Dubois (1992). Despite Dubois' s classificationis already being adopted in influ (1920), Liu Hu (1961), Wallace et al. (1973), & ential works, forexample, Emerson et al. (1993), Marmayou et al. (2000), Additions and Corrections to Amphibian Sp ecies of the Wo rld (Duellman, 1993) and Emerson et al. (2000a), Sumida et al. (2003). Neverthe less, the taxonomy of the Ranidae is still very Amphibian Sp ecies of the World (ver. 2.21 online, Frost, problematic (Dubois, 1999) because the phylogenetic 2002). relationships within the family are still poorly known The purpose of the present work was therefore three fold: (1) to clarify the phylogenetic relationships (Duelhnan Trueb, 1985). & some groups of the family Ranidae; to test On the basis of a phenetic analysis, Dubois (1992) between (2) heterogeneity of the traditional genus Rana; (3) to placed the species in the family into seven subfamilies, fur ther discuss the systematic issues of some ranid groups 88 genera and subgenera including some new genera, (e.g. Paa) and reintroduced some genera previously proposed by Dubois ( 1992) and Fei et al. ( 1990). All the topics will be addressed here using the sequence data set named (e.g. Limnonectes, Hoplobatrachus). According to the classification of Dubois ( 1992), Chinese ranids of 16S and 12S mitochondrial DNA. should be grouped into two subfamilies MATERIALS AND METHODS Dicroglossinae and Raninae - while Fei et al. (1990) classed them into three subfamilies - Raninae, SPECIES STUDIED AmolopinaeYang, 1989, and Occidozyginae Fei, Ye et Twenty-nine species (Table 1) representing 13 gen Huang, 1990. In the subfamilyRaninae, all the new gen- era were examined. Tissue samples were all derived from thighmuscle preserved in either ethanol (95%) or Correspondence: K. Zhou, Institute of Genetic Resources, in refrigerator (-20°C). Specimens were kept in the College of Life Sciences, Nanjing Normal University, herpetological collection of the Institute of Genetic Re Nanjing 210097, China. E-mail: kyzhounj@l 26.com sources, Nanjing Normal University (NJNU) and 2 J. JIANG AND K. ZHOU TABLE l. Species studied in the present study. Family Subfamily Subfamily Genus Genus Traditional Species Locality of collection (Fei et al., (Dubois, (Fei et al., (Dubois, Genus 1990) 1992) 1990) 1992) (Frost, (Fei, 1999) 1985) OUTGROUP Bufonidae Bufo Bufo gargarizans Nanjing, Jiangsu Microhylidae Ka/oula Ka/oula pulchra Zhaoan,Fujian Hylidae Hy la Hy la chinensis Zhaoan, Fujian INGROUP Ranidae Raninae Raninae Rana Rana Rana chensinensis Yuzhong, Gansu Raninae Raninae Rana Rana Rana amurensis Mudanj iang, Heilongjiang Raninae Raninae Rana Rana Rana zhenhaiensis Zhenhai, Zhejiang Raninae Raninae Rana Rana Rana omeimontis Emei Mt., Sichuan Raninae Raninae Rana Rana Rana chaochiaoensis Muli, Sichuan Raninae Raninae Pelophylax 'Rana Rana 11igromaculata Huoqiu, Anhui Raninae Raninae Pelophylax Rana Rana hubeiensis Huoqiu, Anhui Raninae Raninae Hylarana Rana Rana adenopleura Nanjing, Fujian Raninae Raninae Hylarana Rana Rana guentheri Guangzhou, Guangdong. Raninae Raninae Rugosa Rana Rana emeljanovi Dahan, Liaoning Raninae Raninae Rugosa Rana Rana tientaiensis Tianmushan,Zhejiang Raninae Raninae Glandirana Rana Rana minima Fuzhou, Fuj ian Raninae Raninae Odorrana Rana Rana margaretae Wawushan, Sichuan Raninae Raninae Odorrana Rana Rana livida Hejiang, Sichuan Raninae Raninae Odorrana Rana Rana schmackeri Emei Mt., Sichuan Raninae Raninae Odorrana Rana Rana hejiangensis Hejiang, Sichuan Raninae Raninae Odorrana Rana Rana grahami Muli, Sichuan Raninae Raninae Paa Paa Rana boulengeri Wawushan, Sichuan Raninae Raninae Paa Paa Rana sp inosa Tianmushan, Zhejiang Raninae Raninae Paa Paa Rana robertingeri Hejiang, Sichuan Raninae Dicroglossinae Fejervarya Limnonectes Rana /imnocharis Nanj ing, Jiangsu Raninae Dicroglossinae Limnonectes Limnonectes Rana fujianensis Nanjing, Fujian Raninae Dicroglossinae Hoplobatrachus Hoplobatrachus Rana rugulosus Haikou, Hainan Raninae Raninae Nanorana Na norana Nanoran a pleskei Songpan, Sichuan Raninae Raninae Altirana Na norana Altirana parkeri Lasha, Xizang Amolopinae Raninae Amo/ops Amo/ops Amo/ops daiyunensis Nanjing, Fujian Amolopinae Raninae Amo/ops Amo/ops Amo/ops ricketti Hejiang, Sichuan Amolopinae Raninae Amolop s Amo/ops Amo/ops wuyiensis Huangshan, Anhui Amolopinae Raninae Amo/ops Amo/ops Amo/ops mantzorum Wawushan, Sichuan Chengdu Institute of Biology, Chinese Academy of Sci age of the Ranidae. Since the relationships among the ences (CIB). Taxonomic assignment of examined families mentioned above have not been resolved so