Crocodiles & Reptiles of Borneo

Indraneil Das and Ghazally Ismail
Institute of Biodiversity and Environmental Conservation,
Universiti Malaysia Sarawak,
94300 Kota Samarahan, Sarawak, Malaysia.
Email: [email protected]

This presentation covers all crocodiles and turtles known from the island of the island of Borneo. For each species is the current valid scientific name, the English name, maximum size recorded, synonymy, besides notes on identification, natural history and distribution.

Field work was supported by research grants (UNIMAS 120/98 [9] and 192/99 [4]) administered by the Universiti Malaysia Sarawak. We thank Fatimah Abang, Acting Director, Institute of Biodiversity and Environmental Conservation, for support and facilities and current and former colleagues at UNIMAS (Stuart James Davies, Stephen Oakley and Nicolas Pilcher) and at UBD (Joseph Kirbahkaran Charles, David Sydney Edwards, David Thomas Jones, Helen Pang Yoke Kiew and Samhan bin Nyawa) for various favours. Finally, we acknowledge Norzian binti Mohammed for scanning in the images depicted here.

Key to Agamidae.-

1.	Wing-like dermal expansion on sides of body ………………………………………………………… Draco No wing-like dermal expansion on sides of body …………………………………………………………… 2
2.	Tympanum concealed …………………………………………………………………………………………….. 3 Tympanum exposed ……………………………………………………………………………………………….. 4
3.	Dorsal crest present, enlarged unicarinate scales laterally ………………………………….. Aphaniotis Dorsal crest reduced or absent; enlarged tubercular scales laterally …………………… Phoxophrys
4.	Males with long compressed rostral appendage…………………………………………….. Harpesaurus Males without rostral appendage ………………………………………………………………………………. 5
5.	Throat fold present; dorsal scale unequal ………………………………………………….. Gonocephalus Throat fold absent; dorsal scale equal ………………………………………………………………………… 6
6.	Males without gular pouch; oblique fold in front of shoulder absent; > 65 rows of midbody scales; hindlimb > 80% SVL; tail > 290% SVL ……………………………………………………… Bronchocela Male with well developed gular pouch; oblique fold in front of shoulder; < 65 rows of midbody scales; hindlimb < 70% SVL; tail < 250% SVL ……………………………………………………………. 7
7.	Sides of head and throat with enlarged scales; adult SVL 130 mm ……………………. Hypsicalotes Sides of head and throat without enlarged scales; adult SVL < 100 mmm …………………………….8
8.	Gular sacs with lateral pockets; ground colour of body dark brownish gray, with a few distinct transverse white bands on dorsum; adult SVL 70 mm ……………………………………… Complicitus Gular sacs without lateral pockets; ground colour of body green, without transverse white bands on dorsum; adult SVL 82 mm ……………………………………………………………….. Pseudocalotes

Ophisaurus Daudin, 1803:

Ophisaurus F.-M. Daudin. 1803. Bull. Sci. Soc. Philomath ser. 2 3: 188. (type species: Anguis ventralis Linnaeus, 1766, by monotypy.)

Ophisaurus H. Wermuth. 1969. Das Tierreich (90): 29.

Derivation: Latin for snake-lizard, in allusion to its lack of limbs.

Aeluroscalabotes Boulenger, 1885:

Aeluroscalabotes G. A. Boulenger. 1885. Cat. Lizards British Mus. 1: 73. (type species: Pentadactylus felinus Günther, 1864, by monotypy and designation.)

Diagnosis: Postmentals enlarged; axillary pockets absent; claw between a dorsal and two lateral scutes; transverse enlarged lamellae restricted to base of digits.

Derivation: Greek for cat-like lizard.

Key to Gekkonidae.-

1.	Digits slender, not or slightly dilated …………………………………………………………………………… 2 Digits moderately to broadly dilated; terminal phalanges of all or digits II-V compressed ……….. 3
2.	Pupil vertical ………………………………………………………………………………………. Cyrtodactylus Pupil round …………………………………………………………………………………………….. Cnemaspis
3.	Terminal phalanges rising from within dilated portion; scansor broad or narrow …………………… 4 Terminal phalanges united with dilated portion to its distal end and extending beyond; scansor narrow ………………………………………………………………………………………………………………… 7
4.	Digits broadly dilated distally; scansor narrow; on toe IV covering approximately distal half; terminal scansors, distinctly wedge-shaped ………………………………………………………………….. 5 Digits uniformly dilated; scansors wide; on toe IV covering more than distal half; terminal scansors oval or squarish ………………………………………………………………………………………… 6
5.	First inner digit developed, broadly dilated …………………………………………………………… Gehyra First inner digit vestigial, not dilated ………………………………………………….. Hemiphyllodactylus
6.	Prominent cutaneous folds on lateral surfaces and posterior thigh; digits with prominent webs; dorsal without enlarged tubercle ………………………………………………………………… Cosymbotus Cutaneous folds absent; digits without webs or only at base between toes III and IV Hemidactylus
7.	Cutaneous folds present on limbs ………………………………………………………………………………. 8 Cutaneous folds on limbs absent ……………………………………………………………………………….. 9
8.	Exposed margin of scansor variably notched or usually some distal ones divided … Luperosaurus Scansor entire; outer margin nearly straight or with shallow notch for some distal one Ptychozoon
9.	Exposed margin of scansor variably notched or usually some distal ones divided . Lepidodactylus Scansor entire; distal ones with relatively straight, exposed margin or with shallow notch .. Gekko

Key to Scincidae.-

1.	Palatines and pterygoids separated in palate midline; limbs lacking ……………………. Brachymeles Palatines only, or palatine and pteryoids in contact; limbs present ……………………………………… 2
2.	Supranasals absent ………………………………………………………………………………………………… 3 Supranasals present ……………………………………………………………………………………………….. 5
3.	Lower eyelid with transparent disc; light vertebral stripe and dark dorsolateral stripe present ……. Lipinia Lower eyelid scaly; colouration various, but never as above ……………………………………………. 4
4.	Ear-opening superficial; a single large preanal …………………………………………… Tropidophorus Ear-opening sunk; several small preanals ……………………………………………….. Sphenomorphus
5.	Lower eyelid with transparent disc …………………………………………………………………….. Emoia Lower eyelid scaly ………………………………………………………………………………………………… 6
6.	Limbs reduced …………………………………………………………………………………………. Lygosoma Limbs well-developed …………………………………………………………………………………………….. 7
7.	Dorsal and lateral scales on body smooth ……………………………………………………. Lamprolepis Dorsal and lateral scales on body keeled …………………………………………………………………….. 8
8.	Dorsal scales moderately to strongly keeled; pterygoids separated; palatal notch reaching midorbital position ………………………………………………………………………………………… Mabuya Dorsal scales weakly keeled; pterygoid in contact anteriorly; palatal notch not reaching midorbital position ……………………………………………………………………………………………………………….. 9
9.	Linear pattern on forehead and body in juveniles and adults ……………………………. Apterygodon Banded pattern on forehead and body, especially in juveniles ……………………………………. Dasia

ACKNOWLEDGEMENTS

Field work and manuscript preparation were supported by research grants (UNIMAS 120/98 [9] and 192/99 [4]) administered by the Universiti Malaysia Sarawak. I thank Prof. Ghazally Ismail, Deputy Vice Chancellor, UNIMAS, for support and facilites and colleagues at UNIMAS (Fatimah Abang, Stuart Davies, Stephen Oakeley, Nicholas Pilcher and Ramlah Zainuddin) and at UBD (Joseph K. Charles, David Edwards, David Jones, Helen Pang and Samhan bin Nyawa) for various favours. Curatorial staff of the following institutions (institutional abbreviations follow Leviton et al., 1985, and are marked with asterisk; see Appendix I) provided facilities, data on specimens and/or loans of specimens: American Museum of Natural History, AMNH*, New York (Linda Ford, Darrel R. Frost, David Dickey and Margarete Arnold); the Australian Museum, AMS, Sydney (Allen E. Greer and Ross A. Sadlier); The Academy of Natural Sciences, ANSP, Philadelphia (Ned Gilmore); Brunei Museum, BM, Bandar Seri Begawan (Haji Jaya, Marina Wong and Haji Bakariah); Natural History Museum, BMNH*, London (E. Nicholas Arnold and Colin J. McCarthy); California Academy of Sciences, San Francisco, CAS* (Jens Vindum); Field Museum of Natural History, FMNH*, Chicago (Harold Knight Voris, Robert Frederick Inger, Alan Resetar and Cassandra Redhed); Museum of Comparative Zoology, Harvard University, MCZ*, Cambridge, MA (John E. Cadle, Ernest Edward Williams and José P. Rosado); Staatliches Museum für Tierkunde, Dresden, MTKD (Uwe Fritz); Swedish Museum of Natural History, NRM* (Erik Åhlander and Sven Kullander), Nationaal Natuurhistorisch Museum, RMNH*, Leiden (Marinus Hoogmoed); Sarawak Museum, SM*, Kuching (Peter Kedit and Chales Leh), Sabah Parks Zoological Museum, SP, Mount Kinabalu National Park Headquarters (Jamili Nais and Maklarim Lakim); Sabah State Museum, SSM, Kota Kinabalu (Anna Wong), Borneensis Collection, Universiti Malaysia Sabah, UMS (which also includes the collection of the erstwhile Universiti Kebangsaan Malaysia, Sabah Campus, UKMS), Kota Kinabalu (Maryati Mohammed, Axel Dalberg Poulsen, Ahmed Sudin and Lucy Kimsui); Universiti Brunei Darussalam UBD, Bandar Seri Begawan (Joseph K. Charles, David Edwards and Helen Pang); Florida Museum of Natural History, UF, Gainesville (Frederick Wayne King and David Auth); Raffles Museum for Biodiversity Research, University of Singapore, Department of Zoology, ZRC (USDZ*), Singapore (Peter Ng, C. M. Yang, Kelvin K. P. Lim and Tan Heok Hui); the United States National Museum, USNM*, Washington, D.C. (W. Ronald Heyer, Ronald I. Crombie and George R. Zug); Zoological Museum Amsterdam (ZMA*), Amsterdam (Leobertus van Tuijl); Zoologisches Institut und Museum, Universität Hamburg, Hamburg, ZMH (Jakob Hallermann); and Zoological Survey of India, ZSI, Calcutta (J. R. B. Alfred, Shyamal Kumar Chanda, Basudeb Dattagupta and Nimai Charan Gayen).

For providing literature and information to queries, I thank Kraig Ad.ler, Christopher C. Austin, Aaron Bauer, Chew Lun Chan, Patrick David, Itzchak Gilboa, Allen Greer, Wolfgang Grossmann, Jakob Hallermann, Tsutomu Hikida, Marinus Hoogmoed, Robert F. Inger, Werner Kästle, Tan Fui Lian, Colin McCarthy, Ulrich Manthey, Hidetoshi Ota, Alan Resetar, Robert Sprackland, Peter Uetz, Harold K. Voris and Van Wallach. Finally, I express my gratitude to Aaron M. Bauer and Bob Inger for reading a draft of the manuscript, offering numerous comments and suggestions. All errors that remain are mine.

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_____ and AARON MATTHEW BAUER. 1998. Systematics and biogeography of Bornean geckos of the genus Cnemaspis Strauch, 1887 (Sauria: Gekkonidae), with the description of a new species. Raffles Bull. Zool. 46(1): 11-28. (Cnemaspis dringi.)

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Reprinted around 1982 (fide Zhao and Adler, 1993), Oxford and IBH Publishing Co., New Delhi, Bombay and Calcutta.

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English translation: Observations on living earless monitors (Lanthonotus borneensis) by P. Gritis. Bull. Chicago Herpetol. Soc. 24(10): 185-188.

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______ and HIDETOSHI OTA. 1987. Thermal properties of humid tropical lizards in Sabah, Malaysia. In: 1987 Monbusho International Scientific Research Program. Research on behavioural strategy of small animals in the humid tropics 1987. pp: 72-86. Department of Zoology, Kyoto University, Kyoto.

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HONDA, MASANAO, MARI KOBAYASHI, HOI-SEN YONG, HIDETOSHI OTA and TSUTOMU HIKIDA. 1999. Taxonomic re-evaluation of the status of Draco cornutus Günther, 1864 (Reptilia: Agamidae). Amphibia-Reptilia 20(2): 195-210.

______, HIDETOSHI OTA, MARI KOBAYASHI, JARUJIN NABJITABHATA, HOI-SEN YONG & TSUTOMU HIKIDA. 1999. Phylogenetic relationships of the flying lizards, genus Draco (Reptilia: Agamidae). Zool. Sci. 16: 535-549.

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______. 1960. A review of the agamid lizards of the genus Phoxophrys Hubrecht. Copeia 1960(3): 221-225. (Phoxophrys borneensis.)

______. 1962. Rare lizard reaches Museum. Mus. News, Chicago March, 1962: 7.

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_____ and WALTER CREIGHTON BROWN. 1980. Species of the scincid genus Dasia Gray. Fieldiana Zool. n.s. (3): 1-11.

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_____ and WILLIAM HOSMER. 1965. New species of scincid lizards of the genus Sphenomorphus from Sarawak. Israel J. Zool. 14: 134-140. (Sphenomorphus cyanolaemus, Sphenomorphis haasi.)

_____ and FREDERIC WAYNE KING. 1961. A new cave-dwelling lizard of the genus Cyrtodactylus. Sarawak Mus. J. n.s. 10(17-18): 274-276. (Cyrtodactylus cavernicolus.)

_____ and MAKLARIN LAKIM. 1998. Rediscovery of the agamid lizard Calotes kinabaluensis de Grijs (Lacertilia: Agamidae) in Borneo, with notes on its habitat. Herpetol. Rev. 29(3): 143-144.

_______ and ROBERT BUTLER STUEBING. 1989. Frogs of Sabah. Sabah Parks Publication No. 10. Sabah Parks Trustees, Kota Kinabalu. iv + 133 pp.

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______ and ______. 1997. A field guide to the frogs of Borneo. Natural History Publications/Science and Technology Unit, Kota Kinabalu. x + 205 pp.

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_______ and MICHAEL J. ECKARDT. 1969. Hemidactylus garnotii Dumeril and Bibron, a triploid all-female species of gekkonid lizard. Copeia 1969: 651-664.

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_____. “1829-1831” 1830. Observations générales sur les reptiles recuellis dans le voyage. In: Voyage autor du monde, exécute par ordre du Roi, sur la corvette de sa Majesté, La Coquille, pendant les annés 1822, 1823, 1824 et 1825. L. I. Duperrey (Ed). Zoologie Tome Second, 1ére Partie.. Arthus Bertrand, Libraire-Éditeur, Paris. 65 pp. (Scincus cyanurus, Scincus atrocostatus.)

See Zhao and Adler (1993: 209) for dates of publication of the various chapters.

LEVITON, ALAN EDWARD, STEVEN CLEMENT ANDERSON, and RICHARD H. GIBBS. 1988. Standards in herpetology and ichthyology. Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Supplement no. 1: Additions and corrections. Copeia 1988(1): 280-282.

_______, _______, _______, ELIZABETH HEAL and CHARLES ERIC DAWSON. 1985. Standards in herpetology and ichthyology. Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985(3): 802-832.

LIDTH DE JEUDE, THEODORUS WILLEM VAN. 1893. On reptiles from North Borneo. Notes Leyden Mus. 15: 250-257. (Japalura ornata.)

_____. 1904. Reptilia from the Malay Archipelago. In: Zoologische Ergebnisse einer Reise in Niederländisch ost-Indien 2: 178-192. M. Weber (Ed). E. J. Brill, Leiden.

______. 1905. Zoological results of the Dutch scientific expedition to central Borneo. The reptiles. Part I. Lizards. Notes Leyden Mus. 25(16): 187-202. (Lygosoma büttikoferi, Lygosoma hallieri, Lygosoma Nieuwenhuisii, Ophisaurus büttikoferi, Tropidophorus iniquus, Tropidophorus micropus.)

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______ and ______. 1982b. Exkursion am Mt. Kinabalu 4100 m, Nordborneo. Teil 3. Checklist der Herpetofauna oberhalb 600m ü. NN. Herpetofauna, Weinstadt 5(23): 20-31.

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_______ and ______. 1989a. A new triploid Hemidactylus (Gekkonidae: Sauria) from Taiwan, with comments on morphological and karyological variation on the H. garnotii-vietnamensis complex. J. Herpetol. 23(1): 50-60.

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______, SHOWICHI SENGOKU and TSUTOMU HIKIDA. 1996. Two new species of Luperosaurus (Reptilia: Gekkonidae) from Borneo. Copeia 1996(2); 433-439. (Luperosaurus yasumai.)

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______. 1864. Über einige Säugethiere (Mormops, Macrotus, Vesperus, Molossus, Capromys), Amphibien (Platydactylus, Otocryptis, Euprepes, Ungalia, Dromicus, Tropidonotus, Xenodon, Hylodes) und Fische (Sillago, Sebastes, Channa, Myctophum, Carassius, Barbus, Capoëta, Poecilia, Saurenchelys, Leptocephalus). Monatsber. Königl. Preuss. Akad. Wissensch. Berlin 1864: 381-399.

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