Family AGAMIDAE 11-19
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HAROLD F. DE LISLE, LYNN R. G. RAW, DANIEL A. MELNIKOV AGAMIDAE A CATALOG OF RECENT SPECIES HAROLD F. DE LISLE, LYNN R. G. RAW, DANIEL A. MELNIKOV © Harold F. De Lisle, Daniel Melnikov, Lynn R.G. Raw, 2016 Email: [email protected] Cover photo: Psammophilus dorsalis, male in breeding colours, Hyderabad, India, by JM Garg Back cover: Chlamydosaurus kingie, Hann River area, Queensland, Australia, photo by Stephen Zozaya CONTENTS INTRODUCTION …............................................................................. 1 CATALOG …............................................................................................... 13 ACKNOWLEGEMENTS …........................................................... 99 REFERENCES . …................................................................................. 105 TAXON INDEX …................................................................................... 143 INTRODUCTION Family Agamidae Iguaniens (part), Cuvier, 1817, Le règne animal distribué d’après son organisation, pour servir de base a l’historire naturelle des animaux et d’introduction a l’anatomie comparée . Vol. 2. Les reptiles, les poissons, les mollusques et les annélides. Déterville, Paris Ascalabotae (part), Merrem, 1820, Versuch eines Systems der Amphibien I (Tentamen Systematis Amphibiorum). J. C. Krieger, Marburg, 191 pp. Stellionidae (part), Gray 1825, A synopsis of the genera of reptiles and Amphibia, with a description of some new species. Annals of Philosophy , 10: 196 Agamidae Gray, 1827 , A Synopsis of the Genera of Saurian Reptiles in which some new Genera are indicated, and the others reviewed by actual Examination. Philosophical Magazine, London, 2 (2): 57 Iguaniens acrodontes Duméril & Bibron, 1837, Erpetologie Générale ou Histoire Naturelle Complete des Reptiles. Vol.4. Libr. Encyclopédique Roret, Paris Uromasticidae Theobald, 1868, Catalogue of the reptiles of British Burma, embracing the provinces of Pegu, Martaban, and Tenasserim; with descriptions of new or little-known species. Journal of the Linnean Society , London, Zool., 10: 34 The large family Agamidae contains Some Agamid is adapted to almost some 450 species, which are mostly every type of terrestrial habitat. From sea confined to Asia, Australia, and Africa. level to the top of tropical mountain; from Before the recent split of the family hot desert (Sahara) to cold desert (Gobi). Iguanidae, the two generas were often Some are arenicolous, many are saxicolous , compared, with the Iguanidae largely found many more are arboreal. A few are almost in the Western Hemisphere. semiaquatic, but none are semifossial . Perhaps the major anatomical Most Agamids are insectivores, but a difference between the two families is in few are omnivorous. A very few are dentition. Agamids have acrodont and herbivores. heterdont dentition, a character they only All Agamids are oviparous. Most share with the Chamaeleonidae. Dermal species lay leathery eggs with the oval shape appendages such as crests, dewlaps, and that are typical of most reptile eggs, while a even wing-like mebranes ( Draco ) are often few genera ( Calotes ) lay spindle-shaped present. Sexual dimorphism is the rule, with eggs. There are few enigmatic and the males often having brilliant colors and chrestomatic examples of viviparous more pronounced dermal appendages. Agamids. They are extraordinary The eye of Agamids is provided with representatives of different subfamilies. complete lids and the pupil is always round. Live-bearing Agaminae are found in Some species have no tympanum while Phrynocephalus from high elevations of others have large exposed tympanum. Only Tibet and Himalaya ( Ph. vlangalii , Ph. a few genera have femoral pores typical of theobaldi , Ph. putjatai and others) , and lizards. The tongue is rather short and broad. lowland desert of Takla-Makan ( Ph. Limbs are well developed. Tail is usually forsyhtii ). Viviparous Draconinae are long and tough Cophotis ceyloniaca and Harpesaurus borneensis . another with G . robinsonii, and a third with Recent phylogenetic studies have Pseudocalotes ). found that the genera Moloch and At present (2019) there are 53 genera Chelosania render Hypsilurus paraphyletic . in the Family Agamidae. This includes Lophognathus is non-monophyletic. Also, several monotype genera, many of which are Gonocephalus (G. robinsonii is only part of the morbid thirst for naming new distantly related to other Gonocephalus ) and genera in the last two decades (Kaiser et al., Japalura (with species distributed 2013). There are also ditype genera among three distantly related clades, including one allied with Ptyctolaemus, . Acanthocercus Fitzinger, 1843 (13 species) Physignathus Cuvier, 1829 (1 species) Acanthosaura Gray, 1831 (11 species) Pogona Storr, 1982 (8 species) Agama Daudin, 1802 (47 species) Psammophilus Fitzinger, 1843 (2 species) Amphibolurus Wagler, 1830 (4 species) Pseudocalotes Fitzinger, 1843 (23 species) Aphaniotis Peters, 1864 (2 species) Pseudocophotis Manthey and Grossmann. 1997 (2 Bronchocela Kaup, 1827 (12 species) species) Bufoniceps Arnold, 1992 (1 species) Pseudotrapelus Fitzinger, 1843 (6 species) Calotes Cuvier, 1817 (23 species) Ptyctolaemus Peters, 1865 (2 species) Ceratophora Gray, 1835 (4 species) Rankinia Wells & Wellington, 1984 (1 species) Chelosania Gray, 1845 ( 1 species) Salea Gray, 1845 (3 species) Chlamydosaurus Gray, 1825 (1 species) Sarada Deepak, Karanth and Giri, 2016 (3 species) Complicitus Manthey & Grossmann, 1997 (1 species) Sitana Cuvier, 1829 (13 species) Cophotis Peters, 1861 (2 species) Thaumatorhynchus Parker, 1924 (1 species) Coryphophylax Blyth, 1860 (2 species) Trapelus Cuvier, 1817 (13 species) Cryptagama Witten, 1984 (2 species) Tropicagama Melville et al., 2019 (1 species) Cristidorsa Wang et al., 2018 (2 species) Tympanocryptis Peters, 1863 (15 species) Ctenophorus Fitzinger, 1843 (29 species) Uromastyx Merrem, 1820 (18 species) Dendragama Doria, 1888 (3 species) Xenagama Boulenger, 1895 (4 species) Diploderma Hallowell, 1861 (24 species) Diporiphora Gray, 1842 (22 species) Draco Linnaeus, 1758 (39 species) Gonocephalus Kaup, 1825 (16 species) Gowidon Wells & Wellington. 1984 (2 species). Harpesaurus Boulenger, 1885 (5 species) Hydrosaurus Kaup, 1828 (3 species) Hypsicalotes Manthey & Denzer, 2000 (1 species) Hypsilurus Peters, 1867 (16 species) Intellagama Wells & Wellington, 1985 (1 species) Japalura Gray. 1853 (7 species) Laudakia Gray, 1845 (20 species) Leiolepis Cuvier, 1829 (9 species) Lophocalotes Günther, 1872 (2 species) Lophognathus Gray. 1842 (2 species) Lophosaurus Günther, 1872 (3 species) Lyriocephalus Merrem, 1820 (1 species) Malayodracon Denzer, 2015 (1 species) Mantheyus Ananjeva and Stuart, 2001 (1 species) Moloch Gray, 1841 (1 species) Monilesarus Pal et al, 2018 (4 species) Otocryptis Wagler, 1830 (3 species) Phoxophrys Hubrecht, 1881(5 species) Phrynocephalus Kaup, 1825 (31 species) Many authors (Anajeva, 2011) subdivide the Southeast Asian Draconinae Fitzinger, 1826 agamids into six subfamilies: (1) with numerous Indian and Southeastern Uromastycinae Theobald, 1868 with the genera (mostly arboreal or semi-arboreal) genera Uromastyx and Saara ; (2) and (6) Afro-West-Asian Agaminae Spix, Leiolepidinae Fitzinger, 1843 with the genus 1825. Leiolepis ; (3) Amphibolurinae Wagler, 1830 The exact position of many genera is with all Australian and New Guinean still debated among taxonomists. species; (4) Hydrosaurinae K аuр, 1828 with the genus Hydrosaurus ; (5) South- and This is the phylogeny according to Pyron, Burbrink and Wiens, 2013( Pyron et al. BMC Evolutionary Biology 2013) Abbreviations of museums holding the types are as follows AMNH - American Museum of Natural History, Division of Vertebrate Zoology (Herpetology), 79th Street and Central Park West, New York, New York 10024, USA. ANSP - Academy of Natural Sciences, Department of Herpetology, 19th and the Benjamin Franklin Parkway, Philadelphia, Pennsylvania 19103, USA. ANWC – Australian National Wildlife Collection, Canberra, Australia BA – Insitute of Rare Animals and Plants, Sichuan Teachers College, Nanchong, Sichuan Province 637702, China BMNH – The Natural History Museum, Department of Zoology, Cromwell Road, London SW7 5BD, United Kingdom. BPBM - Bernice P. Bishop Museum, Department of Zoology, 1355 Kalihi Street, Honolulu, Hawaii 96818, USA. BSP - Bureau Science Philippines (destroyed in WWII). CAS - California Academy of Sciences, Department of Herpetology, Golden Gate Park, San Francisco, California 94118, USA. CIB - Chengdu Institute of Biology, Academia Sinica, P.O. Box 416, Chengdu, Sichuan, P.R. China. DB.ULPGC - Departamento de Biología, Universidad de Las Palmas de Gran Canaria, Calle Juan de Quesada, 30, 35001 Las Palmas de Gran Canaria, Las Palmas, Spain FMNH - Field Museum, Division of Amphibians and Reptiles, Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605, USA. GNHM - Göteborgs Naturhistoriska Museum, Museivägen 10, 413 11, Göteborg, Sweden HLMD - Hessisches Landesmuseum, Darmstadt, Germany. HUJ - Hebrew University, Department of Zoology, Jerusalem, Israel. IEBR - Institute of Ecology and Biological Resources, No 18 Hoang Quoc Viet Street, Hanoi, Vietnam IZANU - Zoological Institute. Ukrainian National Academy of Sciences, Kiev, Ukraine KIZ - Kunming Institute of Zoology, Chinese Academy of Sciences, 32 East Jaio Chang Road; 650223, Kunming, Yunnan, China KUZ - Department of Zoology, Graduate School of Science, Kyoto University, Sakyo, Kyoto 606-8502, Japan. HNM - Hungarian National Museum, Budapest,