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A Review of the Softshell Turtles of the Genus Chitra, with the Description of New Taxa from Myanmar and Indonesia (Java)

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A Review of the Softshell Turtles of the Genus Chitra, with the Description of New Taxa from Myanmar and Indonesia (Java) Hamadryad Vol. 27, No. 1, pp. 11 – 56, 2002. Copyright 2002 Centre for Herpetology, Madras Crocodile Bank Trust. A REVIEW OF THE SOFTSHELL TURTLES OF THE GENUS CHITRA, WITH THE DESCRIPTION OF NEW TAXA FROM MYANMAR AND INDONESIA (JAVA) William P. McCord1 and Peter C. H. Pritchard2 1East Fishkill Animal Hospital, Hopewell Junction, NY 12533, U.S.A. (Fax: 845-221-2570; Email: [email protected]) 2Chelonian Research Institute, 401 South Central Ave, Oviedo, FL, 32765, U.S.A. (Fax: 407-977-5142; Email: [email protected]) ABSTRACT.– Taxonomic, distribution and morphological data are presented for the trionychid turtle genus Chitra (Testudines: Trionychidae). Many confusing taxonomic details are clarified. Types and type localities are described for both previously named Chitra species. Details of forelimb scalation, chromatic and pattern characteristics, original head drawings accentuating phenotypic characters, shell morphology, general skeletal details, and mitochondrial and nuclear gene sequencing results are presented. The overall range of the genus Chitra is discussed, with details of field collections confirming previously undescribed Indonesian populations. A new species of narrow-headed softshell, Chitra vandijki sp. nov. is described from the Ayeyarwaddy River drainage, Myanmar, distinguished from its congeners by its distribution, a unique combination of carapace, head and neck patterns, and genetic sequence divergence of a level corresponding to full species recognition. A new subspecies of narrow-headed softshell turtle, Chitra chitra javanensis ssp. nov., is described from eastern Java, Indonesia, distinguished from the nominate subspecies by its distribution, by details of carapace, dorsal head and chin patterns, and by a genetic sequence divergence level appropriate to subspecies status. Lastly, a first-use chresonymy is presented for the genus Chitra and included species. KEYWORDS.– Testudines, taxonomy, systematics, Trionychidae, Chitra, Ganges, Ayeyarwaddy, Mae Klong, Pasuruan. BACKGROUND CHITRA INDICA Thomas Hardwicke (1755-1835), who employed During much of the Nineteenth Century, the both Indian and British artists (Archer, 1962; spoils, trophies, and curiosities of the British Wheeler, 1998). These were completed while Empire flowed from the far corners of the globe Hardwicke was in the service of the East India into London, the hub of the entire colonial enter- Company, commanding the Bengal Artillery. prise. Among these items, little noticed at the They were brought to England when Hardwicke time, were both some original paintings and a retired in 1823, and he bequeathed them, along preserved specimen (having died en voyage)ofa with various other turtle paintings, and several truly extraordinary turtle from India. These mis- hundred drawings of other Indian reptiles and cellanea gave tangible evidence of the existence amphibians, to the British Museum of Natural of a giant Asiatic softshell turtle with a remark- History (= BMNH) upon his death in 1835 ably small, narrow head, and eyes so small and (Dawson, 1946). Another Englishman in service far forward that many biologists, seeing a skull, in India was Dr. Francis Buchanan-Hamilton are inclined to mistake the orbits for the nostrils. (1762-1829), who at different times, was super- The evidence for this strangely proportioned visor of both the “Calcutta Garden” (now the creature’s existence specifically included a pair Alipore Zoological Gardens, Kolkata) and the of water-colours painted for Major-General “Institute for Promotions of the Natural History 12 HAMADRYAD [Vol. 27, No. 1, of India” (“The Barrackpore Menagerie”) where and died on its passage.) Testudo membranacea. actual specimens were documented by “careful Nat. Hist. [specimen] No. 1238.B.” drawings” (Archer, 1962). Basically, the two The name Testudo membranacea was first men worked in a cooperative spirit, sometimes used by Blumenbach (1779) with the clearly er- having their staff copy paintings in each other’s roneous type locality “Guiana” (living collections. Today, these originals of trionychids are unknown in South America), and Hardwicke’s “coloured drawings” reside in was considered (e.g., by Wermuth and Mertens, bound volumes in the Zoological Library of the 1961, 1977) to be a questionable synonym of BMNH in London, and those of Buchanan-Ham- Testudo cartilaginea Boddaert (1770), which ilton in the India Office Library, also in London was the first trionychid turtle to be described. (Webb, 1980; Wheeler, 1998). Other artistic Both names describe general qualities of all works from Hardwicke and Buchanan-Hamilton trionychids, i.e. the seemingly non-bony, carti- are thought to have been deposited with the East laginous or membrane-like shell. It is uncertain India Company, or with the Library of the Asi- what actual species was represented by the name atic Society of Bengal, but cannot now be traced T. membranacea. No type was designated. Gray (S. Biswas, pers. comm. to Webb, 1980). (1831b) did not clarify the matter when he placed Hardwicke’s surviving turtle drawings, thir- it (with question) in the synonymy of the African teen in number, bear BMNH identification num- Trionyx Niloticus, which in itself was an unjusti- bers 28 to 40. The illustrations of the new, giant fied replacement name for the older name Tes- softshell turtle, numbers 31 (dorsal) and 40 tudo triunguis Forskål (1775). We hereby (ventral), bear the handwritten inscription declare the name Testudo membranacea a nomen “Sewteree – Country Name Found in the Ganges dubium for the reasons stated above (see also – grows to the size of 240 lbs. Futtehghur May”. chresonymy). The illustrations themselves are well done and A later entry, by Richard Owen in an clearly show the four plastral callosities; the 1859:104 RCS catalogue, reads: “No. 685 [led- small, very narrow head with short nasal tube; ger #]. The Indian Mud-Tortoise (Trionyx the extraordinarily small, anteriorly located indicus, Gray; Testudo membranacea, eyes; the longitudinal lines on the neck; the Henderson). Fig. Gray, Illust. Ind. Zool. vol. i. p. paramedian neck lines forming a “V”; the com- 80. Cat. of Tortoises in Brit. Mus. p. 49. Hab. plex, vermiform marbling of the leathery shell; Penang. [= same basic locality as RCS “Nat. Hist. and the concave posterior border of the bony car- No. 1238.B” (= RCS 1238.B)] Presented by Dr. apace that together characterize the genus Henderson.” (Farkas, 1994: 117). This specimen Chitra. is also presently unaccounted for and is likely the Hardwicke also collected specimens, but all same (RCS 1238.B) specimen as is mentioned in three collections that he shipped to London were the 1817 Royal College of Surgeons ledger “No. lost by shipwreck (Gray, 1872). Nevertheless, it 903”. appears that one narrow-headed softshell speci- J. E. Gray (1800-1875), the prolific chelonian men, reportedly from the East Indies, did reach (among other topics) specialist at the British Mu- London safely, although it cannot be located to- seum (BMNH) for much of the 19th century, re- day (Farkas, 1994: 117). This was received in a produced some of Hardwicke’s and fluid-preserved state by the Royal College of Buchanan-Hamilton’s paintings (see Archer, Surgeons (= RCS), where in 1817 (Farkas, 1994: 1962) in his oversized folio “Illustrations of In- 118) it was listed in the “Rough Minute Book” of dian Zoology,” a two volume work, comprised of the College as “No. 903 [= ledger or entry #]. Mr. 202 colored plates, originally produced as 16 is- Henderson. East Indies. Sept. 4th 1818. A speci- sues in 20 parts (Wheeler, 1998) between 1830 men of the soft shell’d tortoise, called ‘mud-tor- and 1835. Although Hardwicke provided most of toise.’ (Brought by the General Kyd Indiaman, the illustrations and financed the work, Gray ed- October, 2002] NEW CHITRA FROM MYANMAR AND JAVA 13 ited and described the new species illustrated thor’s Illustrations of Indian Zoology and clearly [with a few exceptions credited to Bell refers (although without quoting catalogue num- (1829-1842) by Bourret (1941: 158) and ber) to an actual specimen deposited in the RCS Wheeler (1998: 349), plus see “T. Bell” in collection. This would be the Henderson speci- Iverson (1992: 166), but we agree with men (RCS 1238.B) alluded to above, which is Boulenger (1889), Wermuth and Mertens (1961) presumed destroyed by wartime bombing in and Iverson (1992), crediting the contested spe- 1941 (E. Allen, pers. comm. to Farkas, 1994). cies to Gray (1831b: 20, 21, 23) being first pub- The actual wording of Gray’s two brief de- lished], so this work is attributed solely to Gray scriptions is as follows: (Wheeler, 1998: 348). The accompanying text, Gray 1831a, p. 18: “Indian Trionyx. Trionyx the “Prodromus Faunae Indicae” was never Indicus, Gray, Illust. Ind. Zool. t. Olive green, published, as explained by Dawson (1946: 63). with black-edged, irregular pale tortuous and Plate 80 (Fig. 1), copied as a composite of forked streaks; sternal callosities four, the hinder Hardwicke’s paintings (BMNH 31 and 40) of the ones rounded triangular. India.” narrow-headed softshell species, is thought to Gray 1831b, p. 47: “Trionyx Indicus,(Indian have been published in October of 1831 Trionyx.) – Testa supra subconvexa olivaceo (Kinnear, 1925; Sawyer, 1971; Webb, 1980; viridi, lineis irregularibus tortuosis vel furcatis Wheeler,1998), and is identified in the second of nigro marginatis ornata, sterno 4-calloso, callis two unnumbered introductory pages as “Egyp- lateralibus quadrangularibus, posticis longe
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