A catalogue and systematic overview of the shield-tailed snakes (Serpentes: Uropeltidae) Robert Alexander PYRON Department of Biological Sciences, The George Washington University, 2029 G St. NW, Washington, D.C. 20052 (United States) [email protected] Sumaithangi Rajagopalan GANESH Chennai Snake Park, Rajbhavan Post, Chennai, Tamil Nadu 600 022 (India) Amit SAYYED Wildlife Protection and Research Society, Satara, Maharashtra 415 001 (India) Vivek SHARMA 393 Sanjeevni Nagar, Jabalpur, Madhya Pradesh 482 003 (India) Van WALLACH 4 Potter Park, Cambridge, Massachusetts 02138 (United States) Ruchira SOMAWEERA CSIRO Land and Water, Floreat WA 6014 (Australia) Published on 30 December 2016 urn:lsid:zoobank.org:pub:BFFD82EF-50C9-42BF-8493-DF57591EA4FF Pyron R. A., Ganesh S. R., Sayyed A., Sharma V., Wallach V. & Somaweera R. 2016. — A catalogue and systematic over- view of the shield-tailed snakes (Serpentes: Uropeltidae). Zoosystema 38 (4): 453-506. https://doi.org/10.5252/z2016n4a2 ABSTRACT We present a catalogue and systematic overview of Uropeltidae Müller, 1832 based on both new and previously published molecular and morphological data, and a new molecular phylogenetic analysis. We support the monophyly and distinctiveness of Brachyophidium Wall, 1921, Melanophidium Günther, 1864, Platyplectrurus Günther, 1868, Pseudoplectrurus Boulenger, 1890, and Teretrurus Beddome, 1886. We move Uropeltis melanogaster (Gray, 1858), U. phillipsi (Nicholls, 1929), and Pseudotyphlops Schlegel, 1839 to Rhinophis Hemprich, 1820, and re-name Pseudotyphlops philippinus (Müller, 1832) as R. saf- fragamus (Kelaart, 1853), and U. smithi Gans, 1966 as U. grandis (Beddome, 1867). Thanks to these changes, the taxonomy of all these genera is based on monophyletic entities. Diagnoses based on meristic and mensural characters for external and internal anatomy are provided for the family and all genera, and accounts are given for all currently recognized species, summarizing known morphological variation. We note several taxa that continue to be of uncertain phylogenetic affinity, and outline necessary future studies of variation in systematically valuable characters such as rostral and tail morphology. Cryptic KEY WORDS variation is likely present in many species, and additional collection of specimens and DNA-sequence Taxonomic revision, India, data will likely be needed to provide conclusive resolution for remaining taxonomic issues. Numerous Sri Lanka, questions remain for the systematics of Uropeltidae, and we hope that this study will provide a platform phylogenetics, for ongoing research into the group, including the description of cryptic species, clarifying the phyloge- shield-tailed snakes, new synonyms, netic placement of some remaining taxa, and quantifying the range of intra- and inter-specific variation new combinations. in crucial morphological characters. ZOOSYSTEMA • 2016 • 38 (4) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com 453 Pyron R. A. et al. RÉSUMÉ Catalogue et révision systématique des serpents à queue armée (Serpentes: Uropeltidae). Nous présentons un catalogue et une révision systématique des Uropeltidae Müller, 1832 basés sur des données moléculaires et morphologiques déjà publiées et nouvelles, et sur une nouvelle analyse phylogénétique moléculaire. Nous confirmons la monophylie et la validité de Brachyophidium Wall, 1921, Melanophidium Günther, 1864, Platyplectrurus Günther, 1868, Pseudoplectrurus Boulen- ger, 1890, et Teretrurus Beddome, 1886. Nous transférons Uropeltis melanogaster (Gray, 1858), U. phillipsi (Nicholls, 1929), et Pseudotyphlops Schlegel, 1839 dans Rhinophis Hemprich, 1820 et nous renommons Pseudotyphlops philippinus (Müller, 1832) en R. saffragamus (Kelaart, 1853) et U. smithi Gans, 1966 en U. grandis (Beddome, 1867). Grâce à ces changements, la taxonomie de tous ces genres semble basée sur des entités monophylétiques. Des diagnoses fondées sur des carac- tères méristiques et métriques de l’anatomie externe et interne sont fournies pour la famille et pour tous les genres, et des descriptions sont données pour toutes les espèces actuellement reconnues, résumant la variation morphologique connue. Nous indiquons plusieurs taxons dont les relations phylogénétiques restent incertaines et mettons en avant les études qui seront nécessaires dans le futur sur la variation de caractères significatifs en systématique, comme la morphologie du rostre et de la queue. Une variation cryptique est probablement présente chez de nombreuses espèces et la collecte MOTS CLÉS supplémentaire de spécimens et de données sur les séquences d’ADN sera certainement nécessaire Révision taxonomique, Inde, pour résoudre les problèmes taxonomiques restants. De nombreuses questions subsistent concernant Sri Lanka, la systématique des Uropeltidae telles que la description d’espèces cryptiques, la clarification du pla- phylogénie, cement phylogénétique des quelques taxons restants, et la quantification de l’extension de la variation serpents à queue armée, synonymies nouvelles, intra- et interspécifique des caractères morphologiques déterminants. Nous espérons que ce travail combinaisons nouvelles. servira de base pour poursuivre la recherche sur ce groupe. INTRODUCTION Fitzinger (1843) had fixedUropeltis ceylanica Cuvier, 1829 as the type species of Uropeltis Cuvier, 1829, indicating its prior- The shield-tailed snakes, family Uropeltidae Müller, 1832 ity over Silybura Peters, 1861, which was in widespread use (sensu Pyron et al. 2013a), are a small group (54 species) of in the 19th century (see Beddome 1886; Boulenger 1893a). poorly studied, fossorial snakes, endemic to peninsular India However, the characters that Smith (1943) used to charac- and Sri Lanka (see Bossuyt et al. 2004). Most species were terize individual species were often subjective and potentially described in a burst of activity in the 19th century. Numerous ambiguous (e.g., general descriptions of color patterns), and junior synonyms resulted from this early period of discovery, genera such as Uropeltis give the appearance of being non- including Coloburus Duméril & Bibron, 1851; Crealia Gray, monophyletic. In particular, some species (e.g., U. melanogaster 1858; Dapatnaya Kelaart, 1853; Maudia Gray, 1858; Mitylia [Gray, 1858]; U. phillipsi [Nicholls, 1929]) have characters Gray, 1858; Morina Gray, 1858; Siluboura Gray, 1858 (later such as the rostral dividing the nasals and enlarged terminal unjustifiably emended as Silybura Peters, 1861); and Wallia scutes on the tail, that are otherwise more common in spe- Werner, 1925 (see Gans 1966; McDiarmid et al. 1999 and cies of Rhinophis Hemprich, 1820. Smith also did not usually Wallach et al. 2014). mention or discuss type material, a convention that was not Of the 54 known, extant species, 23 were described by in wide use at the time. Colonel Richard Henry Beddome (see Wallach et al. 2014), Following the implementation of the Code, taxonomic a British officer in India who published extensively on South checklists were later provided by Gans (1966), McDiarmid Asian herpetofauna (Ganesh 2010). Beddome (1886) later et al. (1999), and Wallach et al. (2014), clearing up some provided an overview of the known species and numerous nomenclatural issues and consolidating lists of the type ma- subspecies. Beddome’s accounts use generic and specific names terial and localities, though they did not otherwise discuss (e.g., Silybura Peters, 1861; Rhinophis trevelyanus Kelaart, distributions or diagnostic characters at or above the species 1853) that are not in use today, as they were later designated level. Other research into the biology and interrelationships as junior synonyms (see McDiarmid et al. 1999). within the group has been continuous (e.g., Gans et al. 1978; Building on Beddome (1886), the last comprehensive Rajendran 1985; Gower 2003; Comeaux et al. 2010), but little systematic account of the family that included diagnostic progress in higher-level taxonomy has been made, other than characteristics and distributional data for individual species noting a general disarray in taxonomy (Gans 1966; Rieppel & was that of Malcolm Smith in 1943. Smith was a British Zaher 2002; Olori & Bell 2012). herpetologist and physician working primarily in Thailand Molecular phylogenies have shown for at least 26 years who published a volume on the serpent fauna of British India, that the genera Pseudotyphlops Schlegel, 1839, Rhinophis, and including Sri Lanka and Burma (Smith 1943). This included Uropeltis are either non-monophyletic, or render other genera a key and summary descriptions of the known species, and paraphyletic (e.g., Cadle et al. 1990; Bossuyt et al. 2004; Py- established the taxonomic framework at the genus level used ron et al. 2013a, b). However, a lack of sufficient taxonomic by most subsequent researchers. In particular, Smith noted that and character sampling precluded confident phylogenetic 454 ZOOSYSTEMA • 2016 • 38 (4) Systematics of Uropeltidae resolution and comprehensive taxonomic revision addressing ertheless, informed by our new data, we find a compelling all species and genera in a unified framework. Such a revision, need to consolidate and synthesize current knowledge of the including synonymies, lists of type material, character-based phylogeny, taxonomy, and nomenclature of the currently descriptions, observations of potential cryptic diversity, and recognized species of shield-tailed snakes. phylogenetic evidence, would be inordinately beneficial to Here,