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2008, Schleip, Revision of the Genus Leiopython.Pdf Journal of Herpetology, Vol. 42, No. 4, pp. 645–667, 2008 Copyright 2008 Society for the Study of Amphibians and Reptiles Revision of the Genus Leiopython Hubrecht 1879 (Serpentes: Pythonidae) with the Redescription of Taxa Recently Described by Hoser (2000) and the Description of New Species WULF D. SCHLEIP Fichtenweg 11, 53340 Meckenheim, Germany; E-mail: [email protected] ABSTRACT.—This study is the first comprehensive systematic study on the python genus Leiopython Hubrecht 1879 native to New Guinea. The taxonomic arrangement recently made is critically reviewed, and proper descriptions for taxa herein recognized as valid are provided. Twenty external morphological characters were recorded from 90 preserved specimens from throughout most of the distribution of the genus. Thirteen characters were used with principal coordinate analysis to test the diversity of populations from different distributions. Additional evidence for some species was obtained by maximum parsimony and maximum likelihood analysis of mitochondrial DNA sequences (cytochrome b gene) taken from GenBank. Besides three conventional taxa, two new species from the mainland, and one new island species were recognized in accordance with the evolutionary species concept. Additionally, a new locality record is provided. The island of New Guinea is known for its thon on the northern Torres Strait Islands of extremely diverse flora and fauna and for its Australia. A detailed description of such spec- high level of species endemism (e.g., McDowell, imens can be found in Barker and Barker (1994), 1984; Heads, 2002; Kraus and Allison, 2002; stating that populations would occur on the Austin, 2006). This megadiversity may result islands of Dauan, Saibai, and ‘‘possibly Boigu.’’ from a variety of climates (McApline et al., 1983; Nevertheless, reports of the occurrence on the Prentice and Hope, 2007) and habitats (Austin, mainland of Australia around Pascoe River, 2006) such as tropical savannahs, mixed low- Cape York Peninsula (Ehmann, 1992) remain land forests, rain forests, montane forests, and unconfirmed (Barker and Barker, 1994), as does even alpine vegetation at higher altitude. The the occurrence on Aru Island, mentioned by herpetofauna of New Guinea currently consists Hoser (2000). Some authors assumed that, in the of 109 snake species (Allison, 2007), including former case, specimens may have been con- several endemic python species (Rawlings et al., fused with Liasis fuscus Peters (Kend, 1997; 2004; Allison, 2007). However, some areas are O’Shea, 2007). However, the occurrence of difficult to access, and it is likely that many Leiopython on Normanby Island (McDowell, species remain undiscovered (e.g., Austin, 1975; McDiarmid et al., 1999) appears dubious 2006). Just recently, O’Shea et al. (2004) reported because McDowell (1975) had erroneously the occurrence of Antaresia maculosa in the assigned Bara Bara to the Normanby Island, Western Province, previously unknown from rather than to the mainland of Papua New New Guinea. Guinea (PNG) at the Milne Bay Province as The genus Leiopython (Serpentes: Pythonidae) stated by Boulenger (1898) and Koopman represents medium-sized, unpatterned, terres- (1982). trial, and nocturnal snakes (Barker and Barker, Although specimens of the genus Leiopython 1994; O’Shea, 1996; Kend, 1997) that are widely Hubrecht 1879 are well represented in natural distributed throughout tropical and subtropical history museum collections around the world, New Guinea and closely associated with vari- this genus remains taxonomically rather under- ous wetland habitats. Several populations occur studied. Even though the genus was considered on offshore islands (McDowell, 1975; Barker monotypic until recently, two distinct popula- and Barker, 1994; O’Shea, 1996), such as tions, separated by the Central Mountain Range, Salawati Island (terra typical of Leiopython were recognized in the international pet trade gracilis Hubrecht, 1879), Biak Island (Bron- for over the last 30 years (McDowell, 1975; gersma, 1956), and the remote Mussau Island Parker, 1982; Barker and Barker, 1994; O’Shea, of the Bismarck Archipelago (McDowell, 1975). 1996), called the ‘‘golden’’ or ‘‘northern race’’ Various authors (e.g., McDowell, 1975; Cogger and the ‘‘black’’ or ‘‘southern race.’’ Neverthe- et al., 1983; Barker and Barker, 1994; O’Shea, less, published locality data of specimens and 1996) have mentioned the occurrence of Leiopy- distributional ranges were always assigned to 646 WULF D. SCHLEIP Leiopython albertisii in the literature (e.g., Parker, eny of pythonid snakes (Rawlings et al., 2008) 1982; McDowell, 1984; Barker and Barker, 1994; identified Leiopython Hubrecht and Bothrochilus A. Allison, Reptiles and amphibians of the Fitzinger as sister species. Rawlings et al. (2008) trans-Fly region, New Guinea, report sub- suggest their placement into a single genus, mitted to World Wildlife Fund, South Pacific namely Bothrochilus Fitzinger, as the oldest Program, WWF PNG Madang Office, Madang, available synonym. However, this placement Papua New Guinea, available at http://www. will require further study. wwfpacific.org.fj/publications/png/Transfly_ Aim of the Study.—In this paper, I critically reptiles_report.pdf [February 2008], unpubl. evaluate the current taxonomic arrangement data. 2006). and provide detailed descriptions of the taxa Taxonomic History.—Hubrecht (1879), obvi- recognized as valid. I also provide new mor- ously unaware of the prior description of Liasis phological and ecological data on the genus that albertisii by Peters and Doria 1878, described a may serve for future research. The examination new species based on a single specimen from of a larger number of specimens from through- Salawati Island. He introduced it to his new out the distribution of the genus makes this the genus Leiopython, an intermediate genus be- first comprehensive systematic study of the tween Liasis Gray 1842 and Nardoa Gray 1842 genus. It is hypothesized that the genus forms (generic name was preoccupied for a starfish, a species complex comprising several species Nardoa Gray 1840) as Leiopython gracilis, charac- under the evolutionary species concept (ESC; terized by a pitted rostral and the presence of sensu Frost and Kluge, 1994). The hypothesis of pits in the supralabials. However, Boulenger species diversity was tested using principal (1893) did not follow Hubrecht’s proposal and coordinate analysis of morphological characters. synonymized Leiopython with Liasis. Later, Stull Additionally, the analysis of available mito- (1935) placed the taxon albertisii as a subspecies chondrial DNA (cytochrome b) sequences pro- of Liasis fuscus Peters 1873 and, although she vides evidence for the separation of the so- had not provided reasoning for her action, called northern and southern races. Stull’s proposal was widely followed by subse- quent workers (e.g., Loveridge, 1948; De Haas, 1950; Brongersma, 1953, 1956). Nevertheless, MATERIALS AND METHODS after examining the skulls of the two taxa, Museum Abbreviations.—Museum abbrevia- Worrell (1961) reassigned specific level to tions used in this paper are those of Leviton et albertisii. The ambiguous taxonomic status of al. (1985) with the following additions of Liasis the type species of (Gray had not International Commission on Zoological No- designated a type species for Liasis) prompted menclature (ICZN); School of Biological Scienc- Cogger et al. (1983) to synonymize Liasis with es, Bangor University, Wales, UK (UWB); ‘‘Vida Bothrochilus Fitzinger 1843 (for discussion, see Preciosa’’ International, Boerne, Texas, USA Stimson and McDowell, 1986). Later, Under- (VPI); and Southeastern Louisiana University, wood and Stimson (1990) conducted a phyloge- Hammond, Louisiana, USA (SELU). netic study and synonymized Bothrochilus and Liasis with Morelia Gray 1842. Kluge (1993) Specimens Examined.—A total of 90 preserved contradicted this study in parts, examining 121 specimens from natural history museum collec- external and internal morphological and behav- tions was examined (Appendix 1). Additional- ioral characters in a phylogenetic study, finding ly, data was obtained from literature sources evidence for the distinction of the taxon albertisii (Peters and Doria, 1878; Hubrecht, 1879; Bron- from other python species and, therefore, gersma, 1953, 1956; McDowell, 1975; Under- resurrected the oldest available synonym Leio- wood and Stimson, 1990; Kluge, 1993). Further- python Hubrecht. More recently, Hoser (2000) more, curatorial staff members of natural introduced two new subspecies and one species history museums provided scale counts along to the genus, but subsequent workers did not with pictures of some specimens (see Acknowl- follow this taxonomic arrangement because of edgments). The numbers of available specimens inadequate descriptions and the lack of evi- from some localities (Salawati, Biak, Mussau dence for the taxa described (see Wu¨ ster et al., and Emirau Islands, Enga Province, PNG, the 2001; Williams et al., 2006). Besides the weakly upper Fly-River region of the Western Province, defined diagnoses and erroneous authorship PNG, and Merauke, Indonesia) were small, and (i.e., assigning Leiopython albertisii to Gray 1842 specimens from other localities (e.g., Torres instead of Peters and Doria 1878), emendation Strait Islands, Australia and southwestern Pa- of the subspecific names is required (Wu¨ ster et pua) were unavailable for this study. Data and al., 2001) because of erroneous latinization pictures
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