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Taxonomic Diversity of Agamid Lizards (Reptilia, Sauria, Acrodonta, Agamidae) from China: a Comparative Analysis

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Taxonomic Diversity of Agamid Lizards (Reptilia, Sauria, Acrodonta, Agamidae) from China: a Comparative Analysis Asian Herpetological Research 2011, 2(3): 117-128 DOI: 10.3724/SP.J.1245.2011.00117 Taxonomic Diversity of Agamid Lizards (Reptilia, Sauria, Acrodonta, Agamidae) from China: A Comparative Analysis Natalia B. ANANJEVA1*, Xianguang GUO2 and Yuezhao WANG2 1 Zoological Institute, Russian Academy of Sciences, St. Petersburg 199034, Russia 2 Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, Sichuan, China Abstract Results of the analysis of taxonomic diversity and representation of different evolutionary lineages of agamid lizards (Reptilia, Sauria, Agamidae) are provided in this article. For comparison with the fauna of China, the following territories were selected: North Eurasia, Iran and different countries of South and Southeast Asia. There are 49 agamid species in China, comprising 12 genera and 4 subfamilies. Annotated check-list and identification keys to genera are provided. Among the 49 species of agamids present in China, the percent of endemic species is relatively high (22 species; 45%). Endemic species refer to 4 genera with the genus Japalura having the most, 10 of 14 species, proceeded by members of Tibetan Plateau lineage of Phrynocephalus, 8 of 12 species, and Calotes with 1 species and Laudakia with 3. Keywords Agamidae, China, taxonomy, biogeography, geographic distribution 1. Introduction well as Physignathus cocincinus; 4) Hydrosaurinae Каuр, 1828 with the genus Hydrosaurus; 5) South Studies of mitochondrial DNA for acrodont lizards Asian amd Southeast Asian Draconinae Fitzinger, 1826 (Honda et al., 2000; Macey et al., 2000; Ananjeva, with numerous Indian and Southeastern genera (mostly 2004) has allowed the phylogenetic reconstruction arbereal or semiarboreal); 6) Afro-west Asian Agaminae for squamate taxa comprising the clade Acrodonta, Spix, 1825. whose main evolutionary lineages were associated in We made an attempt to analyze the taxonomic their origin with the fragmentation of Gondwana into composition and representation of different evolutionary separate tectonic plates. Within Acrodonta, there are 7 lines (Amphibolurinae, Leiolepidinae, Draconinae monophyletic lineages: 1) Chamaeleo (Chamaeleonidae), and Agaminae) in the fauna of Asian agamid lizards. 2) Uromastyx, 3) Leiolepis, 4) Hydrosaurus, 5) Southeast For comparing agamid lizard species diversity from Asian group of genera, 6) Afro-southwest-Asian group of China, Iran, and North Eurasia (former Soviet Union genera, and 7) Australo-New Guinean group of genera + Mongolia), different regions of South and Southeast (including Physignathus cocincinus). Such interpretation Asia were selected. Differentiated diagnoses of agamid is based on the results of the integration of morphological lizards of China are provided which allow to identify (Moody, 1980) and molecular data (Macey et al., 2000). their taxonomic position at generic level. We use our Within the family Agamidae there are six major clades own data as well as published results from studies on (subfamilies): 1) Uromastycinae Theobald, 1868 with Oriental and Palearctic agamid lizards (Moody, 1980; the genus Uromastyx; 2) Leiolepidinae Fitzinger, 1843 Zhao and Adler, 1993) to develop a dichotomous key of with genus Leiolepis; 3) Amphibolurinae Wagler, the genera Acanthosaura (Orlov et al., 2006; Ananjeva et 1830 with all Australian and New Guinean species as al., 2008; Wood et al., 2009, 2010), Draco (Inger, 1983; Musters, 1983; McGuire and Heang, 2001), Japalura * Corresponding author: Prof. Natalia B. ANANJEVA, from the (Ota, 1989, 1991, 2000a, b; Ota et al., 1998), Calotes Zoological Institute, Russian Academy of Sciences, with her research mainly focusing on taxonomy, phylogeny and biogeography of widely (Hallermann, 2000; Ota and Hikida, 1991, 1996; Vindum distributed Eurasian amphibians and reptiles. et al., 2003; Zug et al., 2006), Pseudocalotes (Hallermann Tel: +7 (812) 3280811 Fax: +7 (812) 3282941 E-mail: [email protected] and Böhme, 2000; Hallermann and McGuire, 2001), Received: 13 May 2011 Accepted: 21 June 2011 Ptyctolaemus (Ananjeva and Stuart, 2001; Schulte II 118 Asian Herpetological Research Vol. 2 et al., 2004), Laudakia ( Ananjeva et al., 1990; Zhao Hallermann and Böhme, 2000; Hallermann and McGuire, et al., 1999), Phrynocephalus (Peters, 1984; Ananjeva 2001; Vindum et al., 2003; Zug et al., 2006; Orlov et et al., 2006; Barabanov and Ananjeva, 2007; Dunayev, al., 2006). Indexes of Syorensen-Chekanovsky were 2008), and Trapelus (Ananjeva et al., 2006; Macey and calculated as 2c/N1+N2 (Pesenko, 1982); program Excel Ananjeva, 2004). The identification key published in older 2003 was used to construct diagrams. monographs and field guides (Boulenger, 1885; Smith, Endemic species are noted by * in the text below. 1935; Taylor, 1963) did not include diagnoses for the genera Bronchocoela and Pseudocalotes as they were at 3. Results and Discussion the time included in the collective genus Calotes prior to the phylogenetic work of Moody (1980). We used the 3.1 Identification key for the genera of agamid lizards recent data on the faunas of North Eurasia (Ananjeva of China et al., 2006), Iran (Anderson, 1999), China (Zhao and I Femoral pores present.................................................... Adler, 1963; Zhao et al., 1999), Vietnam (Ananjeva .................Subfamilies Amphibolurinae, Leiolepidinae et al., 2007), Myanmar (http://www/calacademy.org/ A. 4-8 femoral pores on each side, tail strongly research/herpetology/myanmar), Thailand (Taylor, compressed laterally....................................subfamily 1993, with addition), South Asia (Das, 1996), and the Amphibolurinae.............................genus Physignathus Sunda Archipelago (Manthey and Grossmann, 1997) B. 13-20 femoral pores on each side, tail roundish, for comparative analysis of taxonomic composition and covered with small scales of equal size ......................... presentation of different subfamilies in the fauna of Asian ....... subfamily Leiolepidinae.............. genus Leiolepis agamid lizards. II No true preanal or femoral pores, body mainly laterally This paper is an attempt to comparatively analyze the compressed, no callose scales............................... ........ agamid lizard fauna from China, and present a modified .................................................. subfamily Draconinae translation of the paper of Ananjeva and Wang (2008) with 1 Ribs much prolonged, supporting a wing-like dermal the additional data. An annotated list of agamid species expansions ............................................... genus Draco from China was prepared on the basis of the fundamental 2 No wing-like dermal expansions. Three parallel monographs on amphibians and reptiles of this country longitudinal folds on each side of the middle of the (Zhao and Adler, 1993; Zhao et al., 1999). throat forming a U-shaped figure................................. .......................................................genus Ptyctolaemus 2. Material and Methods 3 (4) Tympanum hidden............................genus Japalura 4 Tympanum exposed, visible from outside; spines Data for this study of agamid lizards were compounded present behind eye. Tail roundish, length of scales on from both field and laboratory work in different regions ventral side smaller than or equal to width ................... of Northern Eurasia, China, Vietnam, Sri Lanka and .......................................genus Oriocalotes Iran through a collaboration on several projects to study 5 (7) Dorsal scales normally have different sizes. Tail the biodiversity of amphibians and reptiles from these compressed, length of scales on ventral side larger countries. than width; spines present behind eye and in nape area Specimens deposited in the following museums were ..................................................... genus Acanthosaura examined: Zoological Institute, St. Petersburg (ZISP), 6 Dorsal scales of equal size, rombic, evenly spaced ....... Chengdu Institute of Biology, Chengdu (CIB), Zoological .................................................................................... 7 Museum, Moscow State University (ZMMGU), Natural 7 Scales on lateral sides oriented backwards and down- History Museum, [former British Museum (Natural wards........................................................................... 9 History)], London (NHM), Muséum National d’Histoire 8 Limbs moderately long; adpressed hind limbs do not Naturelle, Paris (MNHN); Zoological Museum Hamburg reaching shoulders, ventral scales not larger than (ZMH), Naturhistorisches Museum of the Humboldt dorsal scales; eggs oval ............... genus Pseudocalotes University, Berlin (ZMB), Zoological Forschungsinstitut 9 Scales on lateral sides oriented backwards and upwards and Museum, Koenig (ZFMK), United States National .................................................................genus Calotes Museum, Washington, (USNM), Field Museum of Natural III No true femoral pores, body mainly dorsoventrally History, Chicago (FMNH); Royal Ontario Museum, depressed ....................................subfamily Agaminae Toronto (ROM). Standard morphological characters were 1 Tympanum exposed; males with callose preeanal and used (Ota 1989, 1991; Ota et al., 1998; Hallermann, 2000; ventral scales; tail divided into distinct segments, No. 3 Natalia B. ANANJEVA et al. Taxonomic diversity of agamids from China 119 each composed of two or more whorls of scales rings, ed., Paris, 2: 37. tympanum superficial...........................genus Laudakia Study of mt DNA
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