Agamid lizards (Agamidae, Acrodonta, Sauria, Reptilia) of Vietnam

June 12, 2017 | Autor: Nikolai Orlov | Categoria: Zoology, Vietnam, DISTRIBUTION, Distribution
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Mitt. Mus. Nat.kd. Berl., Zool. Reihe 83 (2007) zzz, 13–21 / DOI 10.1002/mmnz.200600021

Agamid lizards (Agamidae, Acrodonta, Sauria, Reptilia) of Vietnam Natalia B. Ananjeva *, 1, Nikolai L. Orlov1 & Nguyen Quang Truong**, 2 1 2

Zoological Institute, 199034, St. Petersburg, Universitetskaya nab., 1 Institute of Ecology and Biological Resources, Vietnamese Academy of Science and Technology (VAST), 18 Hoang Quoc Viet St., Hanoi, Vietnam

Received 16 October 2006, accepted 24 November 2006 Published online zzz With 3 figures Key words: Agamidae, Vietnam, taxonomic diversity, distribution.

Abstract Being a hotspot of plant and animal diversity and endemism, Vietnam is important for the conservation of biodiversity on a global scale. It is one of the most threatened areas in tropical Asia in terms of destruction of tropical rain forest as an environment with a unique Vietnamese fauna. Previous summaries on Vietnamese agamids, their taxonomy and distribution were published in the works of Bourret 1943, Nguyen & Ho 1996, Bobrov 1995, Ziegler 2002 and Ziegler et al. 2006. New, intensive study of the fauna of Vietnam as well as new opportunities offered by molecular methods make clear that taxonomic diversity in Southeast Asia is underestimated. The aim of this paper is to show and review the taxonomic diversity of different evolutionary lineages of agamids and their geographical distribution. This is based on new results of the authors and most recent evaluation of literature sources. # 2007 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

Introduction Study of mitochondrial DNA (Macey et al. 2000; Ananjeva 2004) allowed the construction of a phylogenetic tree of Acrodonta within Squamates and their main evolutionary lineages, which are associated in terms of their origin with Gondwana fragmenting into separate tectonic plates. There are 7 monophyletic lineages (Fig. 1). Such interpretations are based on the integration of morphological (Moody 1980) and molecular data (Macey et al. 2000). Six groups of agamids correspond to 6 subfamilies: 1) Uromastycinae Theobald, 1868 with the genus Uromastyx; 2) Leiolepidinae Fitzinger, 1843 with the genus Leiolepis; 3) Amphibolurinae Wagler, 1830 with all Australian and New Guinean species as well as Physignathus cocincinus; 4) Hydrosaurinae Kaup, 1828 with the genus Hydrosaurus; 5) South- and Southeast Asian Draconinae Fitzinger, 1826 with numerous Indian and Southeastern

genera (mostly arboreal or semi-arboreal) and 6) Afro-West-Asian Agaminae Spix, 1825. The Vietnamese agamid fauna has a complicated origin. It is represented by forms from 3 evolutionary lineages. Species in Vietnam belong to 3 subfamilies: Leiolepidinae, Amphibolurinae and Draconinae. The latter contains the maximum diversity of genera and species in South and Southeast Asia (Moody 1980; Macey et al. 2000; Ananjeva 2004). In the course of taxonomic revisions of the genera Acanthosaura (Kalyabina-Hauf et al. 2004; Orlov et al. 2006), Bronchocela (Hallermann 2005), Calotes (Hallermann 2000b; Ota & Hikida 1991, 1996; Vindum et al. 2003; Zug et al. 2006); Draco (Inger 1983; Musters 1983; McGuire & Kiew 2001), Gonocephalus (Manthey & Denzer 1991, 1991a, 1992, 1992a, 1993; Honda et al. 2002), Japalura (Ota 1989, 1991; Ota et al. 1998), Pseudocalotes (Hallermann & Bo¨hme 2000; Hallermann & McGuire), and Ptyctolaemus (Manthey & Nabhitabhata 1991;

* Corresponding author: e-mail: [email protected] ** e-mail: [email protected]

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tories, type specimens and distribution. Recent data shows that Acanthosaura crucigera, Draco volans (Bobrov 1995; Nguyen & Ho 1996), Japalura yunnanensis (Bobrov 1995) do not occur in Vietnam. Material and methods Field study, samplings and observations were made in Vietnam (fig. 1) in 1987––2005 within the frame of Russian-Vietnamese projects studying biodiversity of amphibians and reptiles in mountain forests. Specimens were examined in the following museums: Zoological Institute, St. Petersburg (ZISP), Zoological Museum, Moscow State university (ZMMGU), Natural History Museum, [former British Museum (Natural History)], London (BMNH), Muse´um National d’Histoire Naturelle, Paris (MNHN); Zoological Museum Hamburg (ZMH), Zoological Museum of the Humboldt University, Berlin (ZMB), Zoological Museum A. Koenig (ZFMK), United States National Museum, Washington, D.C. (USNM); Field Museum of Natural History, Chicago (FMNH); Royal Ontario Museum, Toronto (ROM). Standard morphological characters were used (cf. Ota 1989, 1991; Ota et al. 1998; Hallermann 2000a; Hallermann & Bo¨hme 2000; Hallermann & McGuire 2001; Vindum et al. 2003; Zug et al. 2006; Orlov et al. 2006). Molecular methods are described elsewhere (Macey et al. 2000; Kalyabina-Hauf et al. 2004); this study was made in Department of Biology, Washington University, St. Louis, Missouri, USA and Laboratory of Molecular Biology, Heidelberg University. A complex of mt-DNA with 1434 bp (ND1, ND2, COI and 8 t-RNA genes, cytochrome b) was used.

Results. Annotated list of agamid lizards of Vietnam Subfamily Amphibolurinae Wagler, 1830

Fig. 1. Survey of the provinces of Vietnam: 1 –– Lai Chau; 2 –– Lao Cai; 3 –– Ha Giang; 4 –– Cao Bang; 5 –– Lang Son; 6 –– Quang Ninh; 7 –– Son La; 8 –– Yen Bai; 9 –– Tuyen Quang; 10 –– Bac Kan; 11 –– Thai Nguyen; 12 –– Bac Giang; 13  Phu Tho; 14 –– Vinh Phuc; 15 –– Than Pho; 16 –– Bac Ninh; 17  Hai Duong; 18 –– Hai Phong; 19 –– Hoa Binh; 20 –– Ha Tay; 21 –– Hung Yen; 22 ––Thai Binh; 23 –– Ninh Binh; 24  Ha Nam; 25 ––Nam Dinh; 26 –– Thanh Hoa; 27 –– Nghe An; 28 –– Ha Tinh; 29 –– Quang Binh; 30 –– Quang Tri; 31  Thua Thien-Hue; 32 –– Da Nang; 33 –– Quang Nam; 34  Kon Tum; 35 –– Quang Ngai; 36 –– Gia Lai; 37 –– Binh Dinh; 38 –– Phu Yen; 39 –– Dac Lac; 40 –– Khanh Hoa; 41  Binh Phuoc; 42 –– Lam Dong; 43 –– Ninh Thuan; 44 –– Tay Ninh; 45 –– Binh Duong; 46 –– Dong Nai; 47 –– Binh Thuan; 48 –– Long An; 49 –– Than Pho; 50 –– Ba Ria-Vung Tau; 51  Dong Thap; 52 –– Tien Giang; 53 –– Ben Tre; 54 –– An Giang; 55 –– Can Tho; 56 –– Vinh Long; 57 –– Tra Vinh; 58  Soc Trang; 59 –– Kien Giang; 60 –– Bac Lieu; 61 –– Ca Mau.

Ananjeva & Stuart 2001; Schulte et al. 2004) a number of new species were described. Recent results of faunistic, taxonomic and phylogenetic study of agamid lizards from Vietnam are summarized in an annotated list with type terri-

Physignathus cocincinus is the sister group to Australo-New Guinean agamids. At present the genus Physignathus includes 2 species: P. cocincinus and P. lessueri. Some authors provide evidence for polyphyletic origin of this genus (Moody 1993; Macey et al. 2000; Schulte et al. 2003) which represent additional support for ancient fragmentation of lizard taxa from both sides of Wallace’s line (Schulte et al. 2003). Agamids of this subfamily have femoral pores and lens-like receptors (Ananjeva 1997, 2004).

Genus Physignathus Cuvier, 1829 Physignathus cocincinus Cuvier, 1829 Ty p e s p e c i m e n s : holotype MNHN 2537 (1975) (Guibe´ 1954). Ty p e l o c a l i t y : Cochinchina, south Vietnam [Cochinchine (Vietnam du Sud)]; coll. Diard. D i s t r i b u t i o n : Myanmar, south-eastern Thailand, Laos, Cambodia, Vietnam, and southern Yunnan and Guandon provinces, China (Smith 1935; Taylor 1963; Zhao & Adler 1993; Zhao et al. 1999). http://museum-zool.wiley-vch.de

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I n V i e t n a m : Lao Cai (Van Ban), Ha Giang (Vi Xuyen), Cao Bang (Nguyen Binh), Yen Bai (Phieng Ban, Van Yen), Bac Kan (Ba Be), Thai Nguyen (Vo Nhai), Lang Son (Huu Lung), Quang Ninh (Nam Son), Ninh Binh (Cuc Phuong), Bac Giang (An Lac), Hai Duong (Chi Linh), Son La (Ca Nang, Hua Trai, Muong Do, Xuan Nha), Hoa Binh (Thuong Tien, Binh Thanh, Mai Chau), Thanh Hoa (Nhu Xuan, Ben En), Nghe An (Pu Mat, Pu Huong), Ha Tinh (Huong Son, Vu Quang, Ky Anh, Cam Xuyen), Quang Binh (Minh Hoa: Cha Lo), Quang Tri (Dak Rong, Huong Hoa), Thua Thien-Hue (A Luoi, Huong Thuy, Huong Tra, Phu Loc, Nam Dong), Da Nang (Ba Na, Son Tra), Quang Nam (Tra My, Nam Giang, Tay Giang, Phuoc Son), Kon Tum (Ngoc Linh, Kon Plong), Gia Lai (K Bang: Son Lang, Dak Bang), Lam Dong (Loc Chau, Da Ban, Da Teh), Binh Phuoc (Nghia Trung), Dong Nai (Cat Tien), Kien Giang (Phu Quoc).

Subfamily Leiolepidinae Fitzinger, 1843 Study of mt-DNA (Macey et al. 2000) and morphological integument characters (Ananjeva et al. 2001) show the monophyly of this lineage, previously combined with Uromaxtyx into a subfamily (family) Uromastycinae (-dae) Theobald, 1868 (Moody 1980; Frost & Etheridge 1989). These lizards have femoral pores and lens-like receptors without hairs (Ananjeva 1997, 2004; Ananjeva et al. 2001). There are bisexual (L. belliana, L. guttata, L. reevesii, L. peguensis ) and parthenogenetic: triploid (L. triploida, L. guentherpetersi) or diploid (L. boehmei) (Darevsky & Kupriyanova 1993) examples.

I n V i e t n a m : records on Fukuok Island in the Siam Gulf (Nguyen & Ho 1996) and in Kien Giang Province are known (Darevsky & Kupriyanova 1993). Data on other localities must be confirmed because they can be referred to another species.

Leiolepis guentherpetersi Darevsky & Kupriyanova, 1993 Ty p e s p e c i m e n s : holotype L. guentherpetersi ZISP 20326, central Vietnam, Thuy Fu, about 17 km north-east of Hue. Coll. Tran Kien, February 1988, adult female. Paratypes ZISP 20327-29, 20331-33, the same data as holotype. Paratype 20330, Tiam Island, Da Nang Province, 29 April 1987. Coll. I.S. Darevsky, subadult female. Paratype 20334 (now ZFMK 52822), the same data as the holotype, adult female. Paratypes NHRM BJO 1939.099.3008/1-17 (Naturhistorika Riksmuseum, Stockholm), Chundu, about 50 km north-west from Hue. Coll. N. Berkengen, 4. February––2. April 1939. Ty p e l o c a l i t y : Thuy Fu, Thua Thien-Hue Province, in about 17 km north-east of Hue, Central Vietnam. D i s t r i b u t i o n : limited to the central provinces of Vietnam: Thua Thien-Hue (Thuy Phu, Thua Luu, Phu Loc), Da Nang (Son Tra), Quang Nam (Cu Lao Cham), Quang Ngai).

Leiolepis guttata Cuvier, 1829 Ty p e s p e c i m e n s : syntypes MNHN 2587 (2063), MNHN 2588 (2064), MNHN 6913 (2062). Ty p e l o c a l i t y : Cochinchina, southern Vietnam (Cochinchine, Viet-Nam du Sud); Diard (Guibe´ 1954).

Genus Leiolepis Cuvier, 1829 Genus includes 7 species, 4 of which are found in Vietnam.

D i s t r i b u t i o n : known only from Vietnam in Thua Thien Hue (Thuan An, Huong Phu, Phu Loc), Da Nang (Son Tra), Binh Dinh (Quy Nhon), Khanh Hoa (Nha Trang), Ninh Thuan (Thap Cham, Ninh Hai), Binh Thuan (Hoa Thang) provinces.

Leiolepis belliana (Gray, 1827) Leiolepis reevesii (Gray, 1831)

Ty p e s p e c i m e n s : not designated. Ty p e l o c a l i t y : Penang, Malaysia. D i s t r i b u t i o n : southern Myanmar, Vietnam, Laos, Cambodia, Thailand (north-eastern coast of Siam Gulf between Menam River and Mekong River Delta), coasts of southern Thailand and western Malaysia, as well as adjacent islands; the belt of the coasts of Sumatra and Banka Island (Indonesia) (Boulenger 1885; Smith 1935; Taylor 1963; Peters 1971; Schmitz et al. 2001).

Ty p e s p e c i m e n s : holotype BMNH 1946.8.14.59 (xxiv.78a), China, coll. J. Reeves. Ty p e l o c a l i t y : China (Boulenger 1885). In Vietnam 2 subspecies: Leiolepis reevesii reevesii (Gray, 1831) and L. reevesii rubritaeniata Mertens, 1961. D i s t r i b u t i o n : nominative subspecies is known from southern China including Hainan Island and from Vietnam. Leiolepis reevesii rubritaeniata is dis1http://museum-zool.wiley-vch.de

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tributed in Vietnam, the south-western part of Laos and north-eastern Thailand (Smith 1935; Taylor 1963; Peters 1971; Darevsky & Nguyen 2004). I n V i e t n a m : 1) Leiolepis reevesii reevesii (Gray, 1831) –– Thanh Hoa (Nga Son, Sam Son, Tinh Gia, Quang Xuong), Nghe An (Quynh Luu, Cua Lo), Ha Tinh (Nghi Xuan, Ky Anh), Quang Tri (Vinh Chap, Quang Tri), Thua Thien-Hue (Hue, Phu Loc) 2) Leiolepis reevesii rubritaeniata Mertens, 1961 –– Gia Lai Province (Chu Prong). (Darevsky & Nguyen 2004), not so far from the Vietnamese-Cambodian border. This record is about 200 km from the coast.

Ty p e l o c a l i t y : Cambodia. D i s t r i b u t i o n : Vietnam and Cambodia. Kalyabina-Hauf et al. 2004 and Orlov et al. 2006 revalidated this species name (Acanthosaura coronata) and Stuart et al. (2006) revalidated it in the course his study of the herpetofauna of Cambodia. I n V i e t n a m : southern provinces Lam Dong and Dong Nai.

Acanthosaura lepidogaster Cuvier, 1829 Ty p e s p e c i m e n s : lectotype MHNP 5076 and paralectotype MHNP 6895 (Guibe´ 1954).

Subfamily Draconinae Fitzinger, 1826 This subfamily is the most diverse group among those genera of arboreal and semi-arboreal agamids with a high percentage of endemics (Moody 1980). Among them there are monotypic genera (Lophocalotes, Mantheyus, Mictopholis) or those represented by 2––3 species (Aphaniotis, Cophotis, Otocryptis, Ptyctolaemus), as well as genera with high species diversity (Draco, Gonocephalus, Japalura). Draconin agamids exhibit very high degrees of diversification at the generic level (25 genera in South Asia and 24 genera in Southeast Asia). Phylogenetic studies conducted in recent decades have improved our knowledge about the composition and phylogeny of the subfamily. Some problems still exist relating to newly described or re-studied species in certain genera (Manthey & Grossmann 1997; Macey et al. 2000; Ananjeva & Stuart 2001; Schulte et al. 2004).

Ty p e l o c a l i t y : Cochin-China.

Genus Acanthosaura Gray, 1831

Ty p e s p e c i m e n s : holotype ZISP 23430, paratypes ZISP 23431-35, 23437-8, 23440-41, FMNH 252272, 252291-92, 252279, 252293-94, 252275, 252278, 252284-85, 252287, 252289, 262571, 258715, 258717-8 BMNH 1927.5.20.33, MNHN 1927-48, ZISP 23436, AMNH R-148545-6.

Genus includes 6 species, 4 of which are found in Vietnam.

Acanthosaura capra Gu¨nther, 1861 Ty p e s p e c i m e n s : lectotype BMNH 1946.8.13.90, paralectotype BMNH 1946.8.13.89. Type locality Cambodia. D i s t r i b u t i o n : Vietnam and Cambodia (Mondolikiri Province) (Orlov et al. 2006: Fig. 20; Stuart et al. 2006). I n V i e t n a m : Dong Nai, Khanh Hoa and Lam Dong provinces in southern Vietnam.

D i s t r i b u t i o n : Acanthosaura lepidogaster is distributed in southern China, Thailand, Laos, Vietnam and Myanmar. The most widely-distributed species of the genus. I n V i e t n a m : reliable records confirmed by morphological identification as well as mt-DNA are known from the northern provinces southwards to Ha Tinh Province in central Vietnam (Lao Kai, Ha Giang, Tuyen Quang, Cao Bang, Bak Kan, Lang Son, Son La, Vinh Phuk, Ha Tay, Hoa Binh, Quang Ninh, Thai Ngyen, Ha Tay, Ninh Binh, Nghe An, Ha Tinh, Quang Binh, Quang Tri, Thua Thien-Hue, Da Nang, Quang Nam and Kon Tum).

Acanthosaura nataliae Orlov, Nguyen & Nguyen, 2006

Ty p e l o c a l i t y : Tram Lap Village, 40 km northwest of Kannack Town, An Khe District, Gia Lai Province, Vietnam. D i s t r i b u t i o n : Central Vietnam and southern Laos (Saravan and Xekong provinces) (Orlov et al. 2006 fig. 20). I n V i e t n a m : Gia Lai, Kon Tum, Quang Nam, Da Nang, Thua Thien-Hue and Quang Tri provinces.

Acanthosaura coronata Gu¨nther, 1861

Genus Bronchocela Kaup, 1827

Ty p e s p e c i m e n s : syntypes BMNH 1946.8.13.81, BMNH 1946.8.13.82, BMNH 1946.8.13.83.

Genus includes 8 species, 3 of which are found in Vietnam. http://museum-zool.wiley-vch.de

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Bronchocela orlovi Hallermann, 2004 Ty p e s p e c i m e n : holotype ZISP 22827. Ty p e l o c a l i t y : Buon Luoi, An Khe district, Gia Lai Province, Vietnam (14 200 N, 108 360 E), altitude 750 m, coll. N.L. Orlov, May 1993. D i s t r i b u t i o n : Vietnam, north part of Tay Nguyen Plateau, Annam mountain. I n V i e t n a m : Gia Lai Province, only known from type locality

I n V i e t n a m : Cao Bang (Nguyen Binh), Bac Kan (Ba Be), Thai Nguyen (Vo Nhai), Vinh Phuc (Tam Dao), Ninh Binh (Cuc Phuong), Thanh Hoa (Ben En), Nghe An (Pu Mat, Vinh), Ha Tinh (Ky Anh, Tan Ap), Quang Binh (Phong Nha-Ke Bang), Quang Tri (Vinh Linh, Huong Hoa), Thua ThienHue (Huong Thuy), Quang Nam (Tra My, Nam Giang, Tay Giang), Kon Tum (Kon Plong), Dong Nai (Cat Tien), Ba Ria-Vung Tau (Con Dao) provinces.

Calotes mystaceus Dume´ril & Bibron, 1837 Bronchocela smaragdina Gu¨nther, 1864 Ty p e s p e c i m e n s : holotype MNHN 2557 (2005). Ty p e s p e c i m e n s : syntypes BMNH 1946.8.11.3536 (adult females). Ty p e l o c a l i t y : Cambodia. D i s t r i b u t i o n : south of Annamit Mountains in Vietnam (Lam Dong Province) and in Cambodia (Mondolkiri Province). I n V i e t n a m : Lam Dong Province.

Bronchocela vietnamensis Hallermann & Orlov, 2005 Ty p e s p e c i m e n s : holotype FMNH 25295 (adult male), coll. Nikolai Orlov and Ilya Darevsky, Vicinity of Tram Lap village (14 200 N, 108 360 E), Annamit mountains, An Khe district, Gia Lai Province, Vietnam, 3th April, 1995, paratype ZISP 22845 (adult female), coll. Nikolai Orlov, August 1993, the same data as for holotype. Ty p e l o c a l i t y : Tram Lap village, 14 20 N.L., 108 36 E.L., Annamit mountains, 40 km from Kannack settlement, An Khe, Gial Lai Province, Vietnam. D i s t r i b u t i o n : species is known only from type locality.

Genus Calotes Rafinesque, 1815 Genus includes 20 species, 3 of which are found in Vietnam.

Calotes emma Gray, 1845 Ty p e s p e c i m e n s : lectotype BMNH 1946.8.11.26 (xxii.31a). Ty p e l o c a l i t y : Afghanistan [in error] East Indian Company). D i s t r i b u t i o n : India (Assam), Myanmar, southern China, Laos, Thailand, Cambodia, Vietnam, northern provinces of West-Malaysia.

Ty p e l o c a l i t y : Myanmar (Birmanie). D i s t r i b u t i o n : China (Yunnan), India (Andaman and Nicobar islands), Myanmar, Thailand, Vietnam, Cambodia, Laos and Vietnam. I n V i e t n a m : Vietnam: Lang Son (Mau Son), Son La (Ca Nang, Hua Trai, Chieng Hac, Ta Sua), Nghe An (Vinh, Pu Mat), Ha Tinh (Huong Son), Quang Tri (Vinh Linh), Thua Thien-Hue (A Luoi, Phu Loc, Loc Hai), Quang Nam (Tay Giang, Phuoc Son), Kon Tum (Kon Plong), Gia Lai (K Bang), Dak Lak (Ea Kao, Krong Pak), Dak Nong (Nam Da), Lam Dong (Di Linh, Lang Bian), Binh Phuoc (Nghia Trung), Khanh Hoa (Nha Trang), Ninh Thuan (Nha Ho, Ninh Hai), Dong Nai (Cat Tien), Binh Duong (Thu Dau Mot), Tay Ninh (Lo Go-Sa Mat), Ho Chi Minh City (Thu Duc), Kien Giang (Ha Tien)

Calotes versicolor (Daudin, 1802) Ty p e s p e c i m e n s : unknown (not designated) (Guibe´ 1954). Ty p e l o c a l i t y : not designated by author. Smith (1935) designated it as Pondicherry, India. D i s t r i b u t i o n : Iran, Afghanistan, Pakistan, Nepal, Bhutan, India, Sri Lanka, Myanmar, southern China (Yunnan, Hainan Province, Hong Kong), Laos, Thailand, Vietnam, Singapore, West-Malaysia, and Indonesia (Sumatra Island). I n V i e t n a m : Vietnam: Lao Cai (Van Ban), Yen Bai (Van Yen: Na Hau, Phieng Ban), Lang Son (Lang Son, Huu Lien), Vinh Phuc (Tam Dao), Hai Duong (Chi Linh), Son La (Co Ma, Chieng Hac, Muong Do), Ninh Binh (Cuc Phuong), Thanh Hoa (Ben En), Nghe An (Hoang Mai, Pu Mat, Pu Huong), Ha Tinh (Huong Son), Quang Tri (Dak Rong, Huong Hoa, Vinh Linh), Thua Thien-Hue (A Luoi, Hue, Phu Loc, Thuan An), Da Nang (Son Tra, Ba Na), Quang Nam (Tra My, Cu Lao Cham), Kon Tum (Kon Plong, Dien Binh, Mom Ray, Tan Lap), Gia Lai (Song Ba, Dak Bang), Dak Lak 1http://museum-zool.wiley-vch.de

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(Buon Ma Thuot, Ban Don), Dak Nong (Nam Da), Lam Dong (Loc Nam, Loc Chau), Ninh Thuan (Nha Ho, Ninh Hai), Dong Nai (Cat Tien), Tay Ninh (Lo Go-Sa Mat, Ba Den Mountain), Ba Ria-Vung Tau (Con Dao), Can Tho (Can Tho), Hau Giang (Phung Hiep), Kien Giang (Ha Tien, Phu Quoc), Ca Mau (U Minh).

D. m. haasei. Hainan Island (China) and North Vietnam (Tonkin). In Vietnam –– Bac Kan, Tuyen Quang, Quang Ninh, Son La, Vinh Phuc and Ha Tay. D. m. whiteheadi. Eastern Thailand, Cambodia, central and southern Vietnam (Ha Tinh, Thua Thien-Hue, Da Nang, Quang Nam, Lan Dong and Ba Ria-Vung Tau). Subspecies of Draco maculatus from Central Vietnam (Ha Tinh, Thua Thien-Hue need to be confirmed (Ziegler 2002)).

Genus Draco Linnaeus, 1758 Genus includes about 20 species 2 of which are found in Vietnam.

Genus Gonocephalus Kaup, 1825 Genus includes 16 species, 1 of which is founds in Vietnam.

Draco indochinensis Smith, 1928 Ty p e specimens: BMNH 1946.8.26.77 (1926.6.29) –– holotype, adult female; BMNH 1946.8.26.78 (1927.5.20.19) –– paratype, coll. Delacoure et Lowe (Musters 1983). Ty p e l o c a l i t y : Bockor, Cambodia. D i s t r i b u t i o n : Kamche Mountain, south-east Cambodia (border with Vietnam) and Vietnam. I n V i e t n a m : southern provinces Kon Tum (Kon Tum, Kon Plong), Khanh Hoa (Nha Trang), Lam Dong (Da Teh), Dong Nai (Cat Tien), Tay Ninh (Tay Ninh, Ba Den Mountain). This was previously considered to be Draco blanfordii indochinensis Musters, 1983 and Draco taeniopterus indochinensis. McGuire & Kiew (2001) confirmed the independent species status of this agamid.

Gonocephalus grandis (Gray, 1845) Ty p e s p e c i m e n s : syntypes BMNH 1946.8.14.83 (xxiii.16a) Rangoon. (J. E. Gray), BMNH 1946.9.4.26 (xxiv.16b), Rangoon. Coll. J. E. Gray. Ty p e l o c a l i t y : Rangoon = Yangon, southern Myanmar. D i s t r i b u t i o n : southern Thailand, lau Tioman, Sunda region, Laos and recent record in south-east Laos 2004) changes the border to more along the coast of Siam Gulf.

Malaysia, PuVietnam. The (Teynie´ et al. than 1000 km

I n V i e t n a m : first record of Gonocephalus grandis Plei Manu, in 25 km to the north-west from Plei Ku, Gia Lai Province, Vietnam.

Genus Japalura Gray, 1853 Draco maculatus (Gray, 1845 ) Ty p e s p e c i m e n s : holotype BMNH 1946.8.27.22 (xxiii.9a), Penang, coll. Hay, paratypes: BMNH 1946.8.27.5––6 (xxiii.9 b––c), Asia. Coll. J. MacGregor (Musters 1983). Ty p e l o c a l i t y : Penang, Malaysia. D i s t r i b u t i o n : southern China, Myanmar, Laos, Thailand, Vietnam and West-Malaysia. I n V i e t n a m : general distribution Vietnam: Bac Kan (Ngan Son, Ba Be), Tuyen Quang (Na Hang), Vinh Phuc (Tam Dao), Quang Ninh (Ba Mun, Cai Bau), Hai Duong (Chi Linh), Ha Tay (Ba Vi), Son La (Hua Trai, Muong Do, Xuan Nha), Ninh Binh (Cuc Phuong), Thanh Hoa (Ben En, Tho Xuan), Nghe An (Pu Mat, Pu Huong), Ha Tinh (Vu Quang, Ky Anh), Quang Binh (Phong Nha-Ke Bang), Thua Thien-Hue (A Luoi, Bach Ma), Da Nang (Ba Na), Quang Nam (Tra My, Nam Giang), Lam Dong (Di Linh), Ba Ria-Vung Tau (Con Dao).

Genus includes 24 species, 2 of which are found in Vietnam.

Japalura chapaensis Bourret, 1937 Ty p e s p e c i m e n s : holotype: MHNP 1948.45, Sa Pa (Chapa), Lao Cai Province, Vietnam (Guibe´ 1954). Ty p e l o c a l i t y : Sa Pa (Chapa), Lao Cai Province, Vietnam. D i s t r i b u t i o n : limited by territory of Vietnam (Northern provinces Cao Bang and Lao Cai). Japalura fasciata Mertens, 1926 Ty p e s p e c i m e n s : holotype SMF 9871 (4310, 1), adult male, coll. H. Fruhstorfer 1904. Ty p e l o c a l i t y : Tonkin (north Vietnam). D i s t r i b u t i o n : northern Vietnam, Lang Son Province (Mauson Mt.), as well as central and southhttp://museum-zool.wiley-vch.de

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ern China. J. szechwanensis Hu & Zhao, 1966 was synonymized with J. fasciata (Ota 2000). Further records indicate a much wider distribution range within China.

I n V i e t n a m : northern, central and southern provinces (Thua Thien-Hue, Da Nang, Quang,Nam, Lam Dong, Kon Tum, Gia Lai, Phu Yen and Dac Lac).

I n V i e t n a m : species only known from type locality in Lang Son Province. Discussion Genus Pseudocalotes Fitzinger, 1843 Genus includes about 10 species, 2 of which are found in Vietnam

Pseudocalotes brevipes (Werner, 1904 ) Ty p e s p e c i m e n : (1928 alpha).

lectotype

MNHN 1907.164

Ty p e l o c a l i t y : Tonkin ¼ North Vietnam. D i s t r i b u t i o n : Vietnam and southern China. I n V i e t n a m : Lang Son (Mau Son), Vinh Phuc (Tam Dao), Ha Tay (Ba Vi), Cao Bang (Nguyen Binh), Hai Duong (Chi Linh).

Pseudocalotes microlepis (Boulenger, 1887) Ty p e s p e c i m e n : BMNH 1946.8.11.21 (formerly 89.3.25.14) Pla-Pu, Tennasserim Coll. L. Fea. Ty p e l o c a l i t y : Pla-Pu, Tennasserim, 1200 m. D i s t r i b u t i o n : Vietnam, India (Assam), Myanmar, southern China.

A biogeographic analysis of the lizard fauna of Vietnam was first attempted by Bourret (1943) and later by Bobrov (1993). The list of specified Vietnamese agamids was compared with those of several regions of South-East Asia (Myanmar, Thailand, Sunda region and non-Palearctic China (Manthey & Grossmann 1997; Zhao & Adler 1993; Zhao et al. 1999; www.http://www.calacademy.org/research/herpetology). Comparison shows (Fig. 2) species diversity range from 22 (Vietnam) to 29 (China) species for the continental regions Vietnam, China, Myanmar and Thailand. The Sunda region has a much higher species diversity (60 spp.). There are 10 species shared between Vietnam and China. The index of faunal similarity (IFS) is ––0.39; 8 –– between Vietnam and Myanmar, 9 –– between Vietnam and Thailand and 5 –– between Vietnam and the Sunda region (IFS 0.12). Among the 22 species in Vietnam, 10 species (45%) are endemics (fig. 3). Distribution of 9 of the 10 endemics is delimited by the southern part of the Annamit Mountains. This is additional evidence for designating an Annam faunistic territory with tropical mountain forests. Inger (1999) wrote that this region is one of the centers of active cryptic speciation in Asia for Anura. Only one endemic species, L. guentherpetersi, is described from Cen-

Fig. 2. Comparison of the number of subfamilies, genera and species of agamids in different regions of South-East Asia.

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Ananjeva, N. B. et al., Agamid lizards of Vietnam

Fig. 3. Groups of agamids of Vietnam.

tral Vietnam, one more from the Tonkin region (north Vietnam) –– Japalura chapaensis. The other 8 species are known only from the Central Highland within Vietnam and adjacent regions of Laos and Cambodia (3 species of Acanthosaura, 3 species of Bronchocela, 1 species of Draco, 1 species of Leiolepis). In general the agamid fauna in Vietnam is characterized by high level of endemism and cryptic diversity. The data on cryptic diversity in genera such as Acanthosaura and Bronchocela confirm ideas about diversification of the fauna of tropical forests (Moritz et al. 2000), which is illustrated by numerous examples from tropical faunas of Central Africa, Australia and South America.

Acknowledgements We would like to thank Alain Dubois and Annemarie Ohler (MNHN), Wolfgang Bo¨hme (ZFMK), Rainer Gu¨nther (ZMB), Jakob Hallermann (ZMH), Robert Inger, Harold Voris, and Alan Resetar (FMNH), V.F. Orlova, A. Dunaev (ZMMGU), Colin McCarthy (BMNH), Robert Murphy (ROM) and George Zug (USNM) providing us museum specimens for this study. We are grateful to Ilya Darevsky for critical reading earlier draft of the manuscript; to Vladimir Bobrov and Dmitry Semenov for providing of their data. Our sincerest thanks are also due to two reviewers, Ulrich Manthey, Berlin and Jakob Hallermann, Hamburg for corrections and recommendations that considerably improved the manuscript. We acknowledge the support of Le Xuan Canh, Ho Thu Cuc and Nguyen Xuan Dang (IEBR) for field work in Vietnam. This work was supported by grants RFFI 05-0448156 and program for support of scientific schools NSH 4212.2006.4.

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