(Colubridae: Lycodon Boie, 1826) from Peninsular Malaysia
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Zootaxa 3815 (1): 051–067 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2014 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3815.1.3 http://zoobank.org/urn:lsid:zoobank.org:pub:FADC54BE-301E-40E7-9029-159082652822 A diminutive new species of cave-dwelling Wolf Snake (Colubridae: Lycodon Boie, 1826) from Peninsular Malaysia L. LEE GRISMER1, EVAN S.H. QUAH2, SHAHRUL ANUAR M.S2,3, MOHD ABDUL MUIN2, PERRY L. WOOD, JR4 & SITI AZIZAH MOHD NOR2 1Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California 92515 USA. E-mail: [email protected] 2School of Biological Sciences, Universiti Sains Malaysia, 11800 USM, Pulau Pinang, Penang, Malaysia. E-mails: [email protected], [email protected], [email protected] 3Center for Marine and Coastal Studies, Universiti Sains Malaysia, 11800 Minden, Pulau Pinang, Malaysia 4Department of Biology, Brigham Young University, 150 East Bulldog Boulevard, Provo, Utah 84602 USA. E-mail: [email protected] Abstract A newly discovered, diminutive, cave-dwelling, lowland species of the colubrid snake genus Lycodon Boie is described from a limestone cave along the Thai-Malaysian border in the state of Perlis, northwestern Peninsular Malaysia. Lycodon cavernicolus sp. nov. is most closely related to L. butleri Boulenger, an endemic, upland, forest-dwelling species from Peninsular Malaysia of the fasciatus group but is separated from L. butleri and all other species of the L. fasciatus group and the closely related L. ruhstrati group by having the combination of 245 (male) and 232 (female) ventral scales; 113 (male) and 92 (female) paired, subcaudal scales; a single precloacal plate; nine or 10 supralabials; 10 or 11 infralabials; a maximum total length of 508 mm (female); a relative tail length of 0.25–0.27; an immaculate venter in juveniles and dark- brown, posterior, ventral scale margins in adults; and dorsal and caudal bands in juveniles white. The discovery of L. cav- ernicolus sp. nov. adds to a rapidly growing list of newly discovered reptiles from karst regions and limestone forests of Peninsular Malaysia, underscoring the fact that these areas should be studied before they are quarried as they harbor a significant portion of the Peninsular Malaysia’s herpetological diversity. Key words: new species, Lycodon, karst, limestone, cave, conservation, endemic biodiversity, Peninsular Malaysia Introduction Karst formations are formed through the dissolution of layers of carbonate bedrock creating caves, sinkholes, and karst towers and compose some of Peninsular Malaysia’s most spectacular landscapes. Exposed karst surfaces are particularly subject to weathering and as such, karst towers and cliff faces are often marked by deep, corrugated surfaces resulting from years of erosion and fracturing. These erosive processes also create extensive networks of cave systems that permeate karst formations and create a unique microhabitat to which a number of organisms have become adapted (see Clements et al. 2006; Vermeulen & Whitten 1999). Despite the astonishing degree of floral endemism in karst habitats and their surrounding limestone forests (Kiew 1998) karst formation are generally not considered to harbor high numbers of endemic, terrestrial vertebrates (i.e. Alström et al. 2010; Jenkins et al. 2004; Woxvold et al. 2009) and as such, remain understudied by vertebrate systematists (see Grismer et al. 2014a). In contrast to this notion, however, our recent herpetological surveys of karst regions in Peninsular Malaysia have revealed 13 endemic, karst-adapted species of geckos (eight species of Cnemaspis—Grismer et al. 2008a,b, 2009; 2013; Wood et al. 2013 and five species of Cyrtodactylus—Grismer et al. 2012, 2014a,b,c) and a new species of limestone forest-adapted colubrid snake of the genus Dendrelaphis (Quah et al. in preparation). Remarkably, we have only explored approximately 2% of the known karst formations and associated limestone forests in Peninsular Malaysia (Price 2001) and anticipate that tens of additional new species will eventually be discovered as our explorations continue. Accepted by Z. Nagy: 21 May 2014; published: 12 Jun. 2014 51 Our surveys of the karst systems associated with the Banjaran Nakawan mountain range separating Peninsular Malaysia from southern Thailand resulted in the discovery of a new species of karst-adapted Rock Gecko (Cnemaspis biocellata Grismer, Chan, Nasir & Sumontha) and Bent-toed Gecko (Cyrtodactylus astrum Grismer, Wood, Quah, Anuar, Muin, Sumontha, Norhayati, Bauer, Wangkulangkul, Grismer & Pauwels). Continuing our surveys in this mountain range, we discovered two individuals of a small species of lowland snake from the karstic cave Gua Wang Burma in the Perlis State Park, Perlis (Fig. 1). We determined that these individuals belonged to the genus Lycodon because they have dorsoventrally flattened heads; vertically elliptical pupils; large nostrils; strongly arched upper maxillary bones whose anterior sections angle inwards; curved, enlarged, anterior maxillary teeth followed by a diastema; 17 anterior and mid-body and 15 posterior rows of dorsal scales bearing eight weakly keeled, medial rows; and rounded, weakly notched, ventral scales (Lanza 1999). We added these specimens to the molecular phylogenetic data set of Siler et al. (2013) and determined they were conspecific and most closely related to L. butleri Boulenger, an upland species endemic to Peninsular Malaysia. However, these specimens differ from L. butleri and all other species of Lycodon in having a unique suite of morphological and color pattern characters. We therefore consider it a new species and describe it below. FIGURE 1. Distribution of Lycodon cavernicolus sp. nov. and L. butleri in Peninsular Malaysia. Stars designate type localities. 52 · Zootaxa 3815 (1) © 2014 Magnolia Press GRISMER ET AL. Material and methods Morphological analysis. Morphological and color pattern data were taken from two specimens from Gua Wang Burma cave, Perlis and 12 specimens of Lycodon butleri from throughout its range. Data evaluated from species of the ruhstrati group were acquired from Vogel et al. (2009). Data presented for species of the fasciatus group were obtained from Zhang et al. (2011). Scale counts and scale nomenclature follow Vogel et al. (2009). All body measurements were made to the nearest millimeter. The number of ventral scales was counted according to Dowling (1951). Half ventrals were not counted, except if they were present on both sides (i.e. divided ventrals). The terminal scute (precloacal plate) was not included in the number of ventrals. The dorsal scale row counts are given at one head length behind the head, at midbody (i.e., at the level of the ventral scale corresponding to a one- half of the total number of ventrals), and at one head length before the vent. We considered sublabials as those scales that had more than one-half their length below a supralabial. The values for paired head characters are listed in Tables 2 and 3 in left/right order. The light-colored bands on the body (dorsal bands) and tail (caudal bands) were counted on one side. Indiscernible or incomplete bands were counted as a single band and bands that were partly fused were counted as two. The collar (i.e. the band on the back of the head) was not counted as a band and bands covering the precloacal plate were added to the dorsal band count. Ventral or belly pattern was considered either banded, spotted, banded anteriorly and spotted posteriorly, immaculate, or with dark-brown posterior edges on the ventral scales. Character abbreviations are SVL–Snout-vent length (mm), TaL–tail length (mm), TL–total length (mm), Rel–TL relative tail length TaL/TL, ASR–dorsal scale rows at neck, MSR–dorsal scale rows at midbody, PSR–dorsal scale rows before vent, Keel–number of keeled dorsal scale rows, VEN–number ventral plates, PreVEN–number of preventrals, VEN not–ventrals notched or not, VEN keel–ventrals keeled or not, SC–number of subcaudal scales, ANA–anal plate single or divided, Lor–number of loreal scales, Lo touch–loreal scale entering the orbit, SL–number of supralabials, SL/Eye–numbers of the supralabials entering orbit, Larg SL–largest supralabial, IL–number of infralabials, IL-tot–total number of infralabials, IL/1st child–number of infralabials in contact with the anterior chin shield, PreOc–number of preoculars, PostOc–number of postoculars. Museum abbreviations are BM–The Natural History Museum, London, UK; BNHS: Bombay Natural History; ZMB–Museum für Naturkunde, Berlin; LSUHC–La Sierra University Herpetological Collection, La Sierra University, Riverside, California, USA. USMHC–Universiti Sains Malaysia Herpetological Collection Molecular analysis. Sequence data from a 1,110 base pair fragment of the cytochrome b gene (cyt b) was obtained from six specimens of Lycodon butleri and the two specimens from the Gua Wang Burma which in total comprised the ingroup. Outgroup samples included 43 specimens from Siler et al. (2013) comprising 18 species. An additional outgroup sequence was obtained for L. albofuscus (USMHC 1457). New sequences used in this study are deposited in GeneBank (Table 1). Genomic DNA was isolated from liver tissue stored in 95% ethanol and extracted using the animal tissue protocol provided by the iNtRON Biotechnology G-spinTM total DNA Extraction Kit (Korea). Cyt b was amplified using a double-stranded Polymerase Chain Reaction (PCR) under the following conditions:1.0 µl genomic DNA (10–30 µg of DNA), 0.5 µl light strand primer (5μM) HI4910 5’–GACCTGTGATMTGAAAAACCAYC–3’ (Burbrink et al. 2000), 0.5 µl heavy strand primer (5μM) THRSN2 5’–CTTTGGTTTACAAGAACAATGCT–3’ (Burbrink et al. 2000), 2.5 µl of iNtRON MgCl2-free PCR Buffer, 2.5 µl of iNtRON 15 mM MgCl2, 0.2 µl iNtRON Taq DNA Polymerase, 0.5 µl dNTP Mix 10 mM each dNTP, and 17.3 µl nuclease free water. PCR reactions were completed using a G-Storm GS482 Thermal Cycler with the following reaction conditions: initial denaturation at 94°C for 2 min, second denaturation at 94°C for 35 s, annealing at 50°C for 35 s followed by an extension cycle at 72°C for 95 s +4 s per cycle for 32 cycles.