Phylogenetic relationships of the genus Sibynophis

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1 Volume 52(2):4, 202 Phylogenetic relationships of the genus Sibynophis (Serpentes: Colubroidea) Hussam Zaher,6 Felipe G. Grazziotin,2 Roberta Graboski,2 Ricardo G. Fuentes Paola Sánchez-Martinez Giovanna G. Montingelli Ya-Ping Zhang 3,4 Robert W. Murphy 3,5 Abstract We present the results of the first molecular analysis of the phylogenetic affinities of the Asian colubroid genus Sibynophis. We recovered a sister-group relationship between Sibynophis and the New World Scaphiodontophis. Although Liophidium sometimes is associated with these genera, the relationship is distant. Morphological characters that Liophidium shares with Sibynophis and Scaphiodontophis are resolved as homoplasies that probably reflect the similarities of their specialized feeding habits. The traditional subfamily Sibynophiinae is elevated to the family-level, and Scaphiodontophiinae is placed in its synonymy. Key-Words: Sibynophiidae; Sibynophis; Scaphiodontophis; Phylogeny. Introduction The genera Liophidium, Sibynophis, and Scaphiodontophis occur on three distinct landmasses Madagascar, Asia, and Central America, respectively. Despite their isolation, these snakes long have been thought to be closely related to each other. In each genus, the dentary bears a peculiar posterior dentigerous process that is completely detached from the compound bone and teeth are numerous and closely set (Duméril et al., 854; Boulenger, 0, 8). Duméril et al. (854) were the first authors to place the four species that share these morphological characteristics in the subgenus Enicognathus of their genus Ablabes. Later, Boulenger (0) substituted Enicognathus, preoccupied, with Polyodontophis. Museu de Zoologia, Universidade de São Paulo. Caixa Postal 42.4, , São Paulo, SP, Brasil. 2. Programa de Pós-Graduação em Zoologia, Universidade Estadual Paulista, Rio Claro, SP, Brasil. 3. Laboratory for Conservation and Utilization of Bio-resources, Yunnan University, Kunming 6500, P.R. China. 4. State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, the Chinese Academy of Sciences, Kunming , P.R. China. 5. Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, Queen s Park, Toronto, Canada M5S 2C6. 6. E mail correspondence to: hussam.zaher@gmail.com

2 42 Zaher, H. et al.: Phylogenetic affinities of Sibynophis that he recognized as a genus. He added four additional species from India and southeastern Asia to Polyodontophis, as well as P. comorensis (= Liophidium mayottensis) and P. sumichrasti from the Comoros Islands and Central America, respectively. Boulenger (8) further expanded Polyodontophis with three additional species, and described the genus Liophidium, to accommodate L. trilineatum from Madagascar. Polyodontophis was posteriorly synonymized with Fitzinger s (3) Sibynophis when Stejneger (0) noted the latter genus has priority over the former, an arrangement followed by all subsequent authors (e.g., Dunn, 928; Smith, ; Taylor & Smith, ). Dunn (928) partially formalized Duméril s and Boulenger s perspectives by erecting the subfamily Sibynophiinae to accommodate the Asian and Central American species of the genus Sibynophis, but he did not include the Malagasy taxa. Although Dunn did not explicitly refer to the Malagasy species of Sibynophis, he implicitly considered them to be unrelated to the Asian and Central American species. According to Dunn (928), both Asian and Central American Sibynophis retain a single-lobed hemipenis with a single sulcus spermaticus a condition strikingly distinct from the bilobed hemipenis with a forked sulcus that occurs in the Malagasy species of Sibynophis. Taken together, both dentitional and hemipenial similarities between the Asiatic and New World species were so compelling that Dunn decided to consider them congeneric (Dunn, 928: 20). It was only after Taylor & Smith () that the New World species of Sibynophis were allocated in a distinct genus Scaphiodontophis. After Dunn (928), Bogert (0) briefly discussed the hemipenial differences between the Malagasy Sibynophis torquatus and its Asiatic and Central American congeners, but he did not remove the taxon from Sibynophiinae. Following Bogert (0), Leviton & Munsterman (6) erected the genus Parasibynophis to accommodate the Malagasy and Comoran species Sib. torquatus, Sib. rhodogaster, and Sib. mayottensis, distinguishing them from the Asian and New World taxa on the basis of osteological and hemipenial characters. Like Bogert (0), these authors considered the mandibular characters to be of greater phylogenetic importance than the hemipenial features, reinforcing the allocation of all three genera Scaphiodontophis, Sibynophis, and Parasibynophis into the subfamily Sibynophiinae. Later, Guibé (8) synonymized Parasibynophis with Liophidium, an arrangement followed by subsequent authors (e.g., Underwood, 7; Williams & Wallach, 9; Cadle, 9). Several more recent authors recognized either the subfamily Sibynophiinae or tribe Sibynophiini (e.g., Kuhn, 7; Underwood, 7; Smith et al., 7; Ferrarezzi, 4; Zaher, 9). Further, Taylor & Smith (), Leviton & Munsterman (6), Underwood (7), Ferrarezzi (4), and Zaher (9) explicitly recognized either the Sibynophiinae or Sibynophiini as a natural assemblage of hingedteeth snakes that includes Scaphiodontophis and Sibynophis but excludes Liophidium. Underwood (7: ) stated the issue with clarity: this subfamily [Sibynophinae] as proposed by Dunn included the Madagascar species now put into Liophidium; the relationships of these latter snakes lie elsewhere but the remaining two genera Sibynophis and Scaphiodontophis are certainly related and systematically rather isolated. Ferrarezzi (4: 90) provided a diagnosis for the tribe Sibynophiini that, according to the author, includes two putative synapomorphies: posterior process of the dentary free and elongate; maxillary teeth small, robust and flattened laterally. Although shared by Scaphiodontophis and Sibynophis, these two characters also are present in Liophidium, in which they have evolved homoplastically. Zaher (9: 28) further documented that Sibynophis and Scaphiodontophis share an additional putative hemipenial synapomorphy viz., a sharply curved, U shaped convolution of the sulcus spermaticus in the proximal region of the hemipenial body. More recently, Pyron et al. (20) erected a new subfamily of colubroid snakes, the Scaphiodontophiinae, to accommodate the New World genus Scaphiodontophis. Their analysis also successfully tested the phylogenetic position of Liophidium, which appears as a member of their subfamily Pseudoxyrhophiinae, and not as the sister-group of Scaphiodontophis. However, monophyly of the traditional subfamily Sibynophiinae, including the genera Sibynophis and Scaphiodontophis (sensu Zaher, 9), requires testing because Sibynophis has not been sampled in any of the recent molecular phylogenetic analyses. Here, we test this hypothesis by adding Sibynophis collaris to Pyron et al. s (20) data matrix, and discuss the derived osteological and hemipenial characters shared by these genera. Material and Methods DNA from Sibynophis collaris was extracted from liver following the protocol of Hillis et al. (6). Sequences for two nuclear protein-coding genes were amplified via polymerase chain reaction (PCR):

3 Papéis Avulsos de Zoologia, 52(2), oocyte maturation factor (c mos) and recombinationactivating gene (rag). To amplify c mos we used the primers described in Zaher et al. (2009); and for rag we used the primers Mart.FL (5 AGC TGC AGY CAR TAY CAY AAR ATG TA 3 ) and Amp. R (5 AAC TCA GCT GCA TTK CCA ATR TCA 3 ) as described by Chiari et al. (2004). PCRs were performed using standard protocols with the addition of 0.4% of Triton X (Sigma-Aldrich, Saint Louis, MO, USA) and annealing temperature of 54 C. PCRs were purified with shrimp alkaline phosphatase and exonuclease I (GE Healthcare, Piscataway, NJ). Sequences were processed using a BigDye Terminator cycle sequencing kit in an ABI 3700 sequencer (Applied Biosystems, Foster City, CA, USA). Both strands were sequenced and checked. The consensus of the two strands was constructed using Geneious version (Drummond et al., 20). The voucher of Sibynophis collaris used in this study was deposited at the Royal Ontario Museum (ROM 258). Sequences of rga and c mos were submitted to GenBank, receiving accession numbers JQ42 and JQ42, respectively. One sequence of Sibynophis chinensis for cytochrome b (cytb) was downloaded from GenBank (accession JN580). We assumed monophyly of the genus Sibynophis and this allowed the creation of a combined terminal (c mos and rag of S. collaris and cytb of S. chinensis). We included the sequences of Sibynophis in the matrix of Pyron et al. (20), while preserving the existing alignment, by using the command add available in MAFFT version 6.8 (Katoh et al., 2002). We conducted a maximum likelihood analysis using RAxML (Stamatakis, 2006). As performed by Pyron et al. (20), the concatenated data were partitioned by codon position and the GTRGAMMA model was used for all partitions. The rapid bootstrap algorithm was employed, and 500 non-parametric bootstrap replicates were performed using the cluster hosted at the Laboratório de Alto Desempenho Pontifícia Universidade Católica do Rio Grande do Sul (LAD PUCRS). This approach allowed complete searches because it used the tree generated at every fifth bootstrap replicate as the starting tree for an independent LSR (Lazy Subtree Rearrangement) swapping. Nodal support was assessed through bootstrapping (BS). We compared the hemipenial, palato-maxillary, and dentary complexes of Sibynophis chinensis with those of Scaphiodontophis annulatus, Liophidium rhodogaster, and L. torquatus. Taylor & Smith (), Leviton & Munsterman (6) and Zaher (9) previously described these complexes. Cleared-and-stained skulls were prepared following the technique of Dingerkus & Uhler (7) in which cartilage is stained with Alcian blue, bone with Alizarin red, and tissue is macerated with Trypsin. Skeletal nomenclature follows that of Cundall & Irish (2008). Methods for hemipenial preparation and terminology followed those of Zaher (9) and Zaher & Prudente (2003). Figure : Summary tree showing the major clades of Caenophidia as recovered in the maximum likelihood analysis of Pyron et al. s (20) data set with Sibynophis sequences added (full topology in Appendix S). Taxon names in bold indicate high-level taxa (higher than family). Numbers near nodes indicate bootstrap values for unnamed clades. Numbers in parenthesis after a taxon name indicate bootstrap values for that clade.

4 44 Zaher, H. et al.: Phylogenetic affinities of Sibynophis Figure 2: Palatomaxillary arch. A: Dorsal view of Sibynophis chinensis AMNH 344; B: Ventral view of Sibynophis chinensis AMNH 344; C: Dorsal view of Scaphiodontophis annulatus KU 0; D: Ventral view of Scaphiodontophis annulatus KU 0; E: Dorsal view of Liophidium rhodogaster UMMZ 2027; F: Ventral view of Liophidium rhodogaster UMMZ Abbreviations: ept = ectopterygoid; mx = maxilla; pal = palatine; pt = pterygoid. Scale bar = 2 mm.

5 Papéis Avulsos de Zoologia, 52(2), Institutional codes are, as follow: AMNH, American Museum of Natural History; KU, Museum of Natural History, University of Kansas; and UMMZ, University of Michigan, Museum of Zoology. The following specimens were examined: Sibynophis chinensis (skull: AMNH 344; hemipenis: AMNH 3402); Scaphiodontophis annulatus (skull and hemipenis: KU 0); Liophidium rhodogaster (skull: UMMZ 2027; hemipenis: UMMZ 20); and L. torquatus (skull: UMMZ 2050). Results Our tree-search procedure in RAxML produced a topology with a score of lnl = A summary of the tree topology is given in Figure, and the full tree is provided as supplementary material (Appendix S). Although we retrieved the same well-supported clades (BS > 70%) as did Pyron et al. (20), the relationships at poorly supported nodes frequently differed (e.g., affinities between elapoid families sensu Zaher et al., 2009). Our analysis clustered Sibynophis with Scaphiodontophis in a robustly supported clade within Colubroidea (sensu Zaher et al., 2009), with a bootstrap value of (Fig. ; Appendix S). In contrast, Liophidium maintained its elapoid affinities (Zaher, 9; Nagy et al., 2003; Pyron et al., 20), clustering with the other Malagasy snakes as the sister-group of a clade formed by Heteroliodon, Pseudoxyrhopus, and Liopholidophis (Nagy et al., 2003). This result corroborated the monophyly of the sibynophiines, as previously recognized by Underwood (7), Ferrarezzi (4), and Zaher (9). Sibynophiines, calamariids, and colubrids (including Grayiinae and Colubrinae of Pyron et al., 20) appeared rooted in an unresolved polytomy, forming a weakly supported clade (BS = 30; Fig. ) that also was retrieved by Pyron et al. (20) with a slightly higher bootstrap value (BS = 52). Discussion According to the phylogenetic affinities recovered in our molecular analysis (Fig. ), Scaphiodontophis and Sibynophis share four putative morphological synapomorphies that are absent in Liophidium and all remaining pseudoxyrhophiines. These are, as follow: () distally broadened, plate-like maxillary process of the palatine with an anterior edge projecting from the level of the anterior tip of the palatine and covering completely the palatine process of the maxilla (Fig. 2); Figure 3: Hemipenis. A: asulcate view of Sibynophis chinensis AMNH 3402; B: sulcate view of Sibynophis chinensis AMNH 3402; C: asulcate view of Scaphiodontophis annulatus KU 0; D: sulcate view of Scaphiodontophis annulatus KU 0; E: asulcate view of Liophidium rhodogaster UMMZ 20; F: sulcate view of Liophidium rhodogaster UMMZ 20. Scale bar = 2 mm.

6 46 Zaher, H. et al.: Phylogenetic affinities of Sibynophis (2) maxilla projects posteriorly to the maxillaryectopterygoid contact, forming a significantly long dentigerous process posterior to the ectopterygoid process of the maxilla (Fig. 2); (3) choanal process of the palatine absent (Fig. 2); and (4) sharply curved U shaped convolution of the sulcus spermaticus in the proximal region of the hemipenial body (Fig. 3). Alternately, five morphological characters found in Scaphiodontophis and Sibynophis appear to have evolved homoplastically in Liophidium. These characters include: () numerous, closely set teeth (Figs. 2, 4); (2) compound bone anterior to the mandibular fossa with a long, tubular and dorsally curved shape (Fig. 4); (3) splenial and angular bones reduced (Fig. 4); (4) mandibular fossa reduced and restricted to the posterior one-fifth of the compound bone (Fig. 4); and (5) posterior dentigerous process of the dentary elongated and detached from the compound bone (Fig. 4). These homoplastic characters likely reflect the specialized durophagous diet of these snakes (Savitzky,, ). Cadle (9), following Savitzky s () lead, found hinged and peculiarly shaped teeth in Pseudoxyrhopus, Exallodontophis, Heteroliodon, and Pararhadinaea; all of these Malagasy genera are closely related to Liophidium and known to feed on durophagous prey. Cadle (9) also pointed out that hinged teeth occur in some African and Asian snakes that feed on hard prey (e.g., Lycophidion, Mehelya, Chamaelycus, Tropidonophis). The apparently widespread geographical distribution of hinged teeth represents additional evidence that this peculiar tooth morphology has arisen independently several times. Higher level taxonomic ranking of Colubroides (sensu Zaher et al., 2009) Pyron et al. (20) differ from Zaher et al. (2009) in preferring the traditional meaning of the superfamily Colubroidea, including the Xenodermatidae, Pareatidae, Viperidae, Homalopsidae, Elapidae, Figure 4: Mandible. A: Medial view of Sibynophis chinensis AMNH 344; B: Medial view of Scaphiodontophis annulatus KU 0; C: Medial view of Liophidium rhodogaster UMMZ Abbreviations: an = angular; cp = compound bone; d = dentary; sp = splenial. Scale bar = 2 mm.

7 Papéis Avulsos de Zoologia, 52(2), 202 an expanded Lamprophiidae, and a highly speciose Colubridae. We acknowledge the fact that this scheme promotes taxonomic stability for the long-standing name Colubroidea. However, such traditional taxonomic hierarchy no longer accommodates new knowledge of phylogenetic affinities within the group appropriately (Vidal et al., 2007; Kelly et al., 2009; Zaher et al., 2009). As knowledge of the phylogenetic affinities of colubroideans improves, additional taxonomic changes that create a certain amount of instability will be required to accommodate the newly resolved relationships (Lawson et al., 2005; Vidal et al., 2007; Zaher et al., 2009). Such taxonomic adjustments, provided that they are stated explicitly (Zaher et al., 2009), are necessary to incorporate the diversity and newly discovered monophyletic groupings of the colubroidean radiation. As pointed out by Frost et al. (2006: 43) We expect that regulated nomenclature will increasingly be pushed toward the terminal taxa and that unregulated taxa will increasingly be rankless. The reason for this is that there really is a practical limit to the number of ranks that workers are willing to use So, our observation is that sociological pressures will push workers towards ever smaller families Regardless, we think that this process will correspond with enormous progress in phylogenetic understanding. Zaher et al. s (2009) new taxonomic scheme represents a step in that direction for the largest, most speciose group of snakes. In the present study, Scaphiodontophis and Sibynophis form a well-supported clade rooted in a polytomy along with families Colubridae and Calamariidae (Fig. ). Therefore, we recognize Sibynophiidae Dunn, 928 (ex Sibynophiinae) as a distinct family that includes the genera Scaphiodontophis Taylor & Smith, and Sibynophis Fitzinger, 3 (type genus) within Zaher et al. s (2009) taxonomic scheme. Because Sibynophiidae Dunn, 928, has priority over Scaphiodontophiinae Pyron et al., 20, we synonymize the latter into the former. Resumo Os resultados apresentados correspondem à primeira análise molecular feita acerca das afinidades filogenéticas do gênero Sibynophis, um colubróideo de origem asiática. Sibynophis aparece como sendo o grupo-irmão do gênero Neotropical Scaphiodontophis. Embora Liophidium esteja por vezes associado a estes dois gêneros, a sua relação é apenas distante. Os caracteres morfológicos compartilhados entre Liophidium, Sibynophis e Scaphiodontophis correspondem a homoplasias que refletem provavelmente os hábitos alimentares especializados semelhantes, presentes nos três gêneros em questão. A tradicional subfamília Sibynophiinae é elevada ao nível de família, enquanto que Scaphiodontophiinae passa para sua sinonímia. Palavras-Chave: Sibynophiidae; Sibynophis; Scaphiodontophis; Filogenia. Acknowledgments The authors wish to thank Charles Myers, Darrel Frost and David Kizirian (AMNH), Ronald Nussbaum and Greg Schneider (UMMZ), William Duellman and Linda Trueb (KU) for their support and for allowing us to prepare and analyze specimens from the collections under their care. We are grateful to the staff of the Laboratório de Alto Desempenho, Pontifícia Universidade Católica do Rio Grande do Sul (LAD PUCRS), who generously allowed access to the cluster under their care. We are also indebted to John Cadle, Linda Trueb, and Gustavo Scrocchi for their insightful review of a previous version of the work. Funding for this study was provided by the Fundação de Amparo à Pesquisa do Estado de São Paulo (BIOTA/FAPESP; grants number 02/602 4 and / ) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq; grants number /200 and /200 0) to HZ. This study also benefitted from a Visiting Professorship for Senior International Scientists from the Chinese Academy of Sciences and a Natural Sciences and Engineering Research Council of Canada Discovery Grant (348) to RWM, a Postdoctoral fellowship from CNPq (number 52069/200 0) to GGM, and a PhD scholarship from FAPESP to FGG (number 07/527 5). References Bogert, C.M. 0. Herpetological results of the Vernay Angola Expedition. Part. Snakes, including an arrangement of African Colubridae. Bulletin of the American Museum of Natural History, 77: 07. Boulenger, G.A. 0. The fauna of British India, including Ceylon and Burma. Reptilia and Batrachia. Taylor & Francis, London. xviii, 54p. Boulenger, G.A. 8. Catalogue of the snakes in the British Museum, Vol. 3. Taylor & Francis, London. xiv + 727p. Cadle, J.E. 9. The dentition, systematics, and phylogeny of Pseudoxyrhopus and related genera from Madagascar, with descriptions of a new species and a new genus. Bulletin of the Museum of Comparative Zoology, 55(8):3 443.

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9 Appendix S Tropidophis haetianus Anilius scytale Boa constrictor Cylindrophis ruffus Xenopeltis unicolor Acrochordus granulatus Achalinus rufescens Achalinus meiguensis Stoliczkaia borneensis Xenodermus javanicus Asthenodipsas vertebralis Pareas nuchalis Aplopeltura boa Pareas macularius 3 Pareas hamptoni Pareas margaritophorus Proatheris superciliaris Eristicophis macmahoni Pseudocerastes persicus Pseudocerastes fieldi Macrovipera schweizeri Macrovipera lebetina Vipera xanthina Vipera wagneri 78 Vipera bornmuelleri 35 Vipera albizona 76 Vipera raddei Daboia russelii 7 Vipera palaestinae 72 Macrovipera mauritanica Macrovipera deserti Vipera ammodytes Vipera aspis Vipera latastei Vipera kaznakovi 55 Vipera ursinii 76 Vipera dinniki Vipera seoanei 88 Vipera nikolskii 92 Vipera berus Vipera barani Echis carinatus Echis ocellatus 67 Echis pyramidum Echis coloratus Cerastes vipera Cerastes cerastes 37 Causus resimus Causus defilippii 85 Causus rhombeatus 9 Atheris chlorechis Atheris barbouri 87 Atheris ceratophora 40 Atheris hispida Atheris squamigera 20 Atheris desaixi 3 Atheris nitschei Bitis arietans Bitis worthingtoni Bitis gabonica 50 Bitis nasicornis Bitis peringueyi Bitis caudalis Bitis xeropaga Bitis atropos Bitis cornuta Bitis rubida Azemiops feae Garthius chaseni Tropidolaemus wagleri Deinagkistrodon acutus 8 Hypnale hypnale Calloselasma rhodostoma Trimeresurus puniceus Trimeresurus borneensis Trimeresurus malabaricus Trimeresurus trigonocephalus 90 Trimeresurus gramineus Cryptelytrops insularis Cryptelytrops septentrionalis Cryptelytrops albolabris Cryptelytrops cantori Cryptelytrops purpureomaculatus Cryptelytrops erythrurus Himalayophis tibetanus Viridovipera medoensis Cryptelytrops macrops 29 Cryptelytrops venustus Viridovipera gumprechti Viridovipera stejnegeri 4 83 Viridovipera vogeli Popeia popeiorum 2 Parias hageni 88 Parias malcolmi Parias flavomaculatus Parias sumatranus 63 Parias schultzei Trimeresurus gracilis Ovophis okinavensis 42 Gloydius halys Gloydius shedaoensis 77 Gloydius strauchi Gloydius blomhoffii Gloydius ussuriensis Ovophis monticola 2 Triceratolepidophis sieversorum Zhaoermia mangshanensis Protobothrops cornutus Protobothrops jerdonii Protobothrops tokarensis Protobothrops flavoviridis Protobothrops mucrosquamatus Protobothrops elegans Ophryacus undulatus Ophryacus melanurus Lachesis muta Lachesis stenophrys 4 Bothriechis schlegelii Bothriechis nigroviridis Bothriechis lateralis Bothriechis thalassinus Bothriechis marchi 46 4 Bothriechis bicolor Bothriechis rowleyi Bothriechis aurifer Agkistrodon contortrix Agkistrodon piscivorus Agkistrodon bilineatus 25 Agkistrodon taylori Crotalus ravus Crotalus pusillus Sistrurus miliarius 25 Sistrurus catenatus Crotalus cerastes Crotalus enyo 6 Crotalus horridus Crotalus transversus Crotalus tortugensis Crotalus atrox 55 8 Crotalus durissus Crotalus basiliscus Crotalus totonacus 29 Crotalus molossus Crotalus willardi Crotalus mitchellii 2 46 Crotalus adamanteus 38 Crotalus tigris Crotalus scutulatus Crotalus viridis Crotalus oreganus Atropoides occiduus Atropoides nummifer Atropoides olmec Atropoides picadoi 33 Cerrophidion godmani Cerrophidion tzotzilorum 55 Cerrophidion petlalcalensis Porthidium dunni Porthidium ophryomegas Porthidium yucatanicum 46 Porthidium nasutum Porthidium porrasi Porthidium lansbergii Bothrocophias campbelli 59 Bothrocophias microphthalmus Bothrocophias hyoprora Bothrops pictus Bothrops ammodytoides Bothrops itapetiningae Bothrops alternatus Bothrops cotiara Bothrops fonsecai Bothrops jararaca 90 Bothrops insularis Bothrops alcatraz Bothrops erythromelas Bothrops neuwiedi Bothrops diporus Bothriopsis bilineata Bothriopsis pulchra Bothriopsis chloromelas Bothriopsis taeniata Bothrops jararacussu Bothrops brazili Bothrops punctata Bothrops caribbaeus 63 Bothrops lanceolatus Bothrops asper Bothrops colombiensis Bothrops marajoensis Bothrops moojeni Bothrops atrox Bothrops leucurus Enhydris matannensis Enhydris plumbea Enhydris enhydris 5 Enhydris chinensis Enhydris punctata 4 54 Myron richardsonii Enhydris polylepis Enhydris bocourti Homalopsis buccata 54 Cerberus australis Cerberus microlepis 25 Cerberus rynchops Erpeton tentaculatum Bitia hydroides 27 Cantoria violacea 76 Fordonia leucobalia 59 Gerarda prevostiana Oxyrhabdium leporinum Calliophis bivirgata Sinomicrurus japonicus Sinomicrurus kelloggi Sinomicrurus macclellandi Micruroides euryxanthus Micrurus fulvius 28 Micrurus corallinus Micrurus mipartitus 5 Micrurus surinamensis 54 Micrurus hemprichii 4 Micrurus lemniscatus Micrurus narduccii 20 Micrurus decoratus Micrurus pyrrhocryptus Micrurus baliocoryphus 92 Micrurus altirostris 9 Micrurus ibiboboca 44 Micrurus spixii 58 Micrurus frontalis 72 Micrurus brasiliensis Ophiophagus hannah 45 Hemibungarus calligaster 8 Aspidelaps scutatus Walterinnesia aegyptia 88 Hemachatus haemachatus Naja annulata Naja multifasciata 54 Naja nivea Naja annulifera Naja nigricollis Naja mossambica Naja pallida 67 Naja nubiae 33 Naja kaouthia 45 Naja naja 5 Naja atra 7 Naja mandalayensis Naja siamensis Dendroaspis angusticeps 90 Dendroaspis polylepis Bungarus flaviceps 9 Bungarus bungaroides 9 Bungarus fasciatus Bungarus caeruleus Bungarus sindanus Bungarus niger 88 Bungarus multicinctus Bungarus candidus Elapsoidea nigra Elapsoidea sundevallii Elapsoidea semiannulata Laticauda colubrina Toxicocalamus preussi 22 Micropechis ikaheka 2 Demansia vestigiata Demansia papuensis Simoselaps semifasciatus Simoselaps anomalus Simoselaps bertholdi 4 Oxyuranus microlepidotus Oxyuranus scutellatus 78 Pseudonaja modesta Pseudonaja textilis 2 Simoselaps calonotus Vermicella intermedia Aspidomorphus muelleri 3 Acanthophis praelongus Acanthophis antarcticus 8 Pseudechis porphyriacus Pseudechis butleri 38 Pseudechis australis Pseudechis papuanus 6 Pseudechis guttatus 77 Pseudechis colletti Cacophis squamulosus 50 Furina diadema Furina ornata 8 Suta fasciata Suta monachus Suta suta 52 Denisonia devisi 29 Elapognathus coronata 58 Rhinoplocephalus nigrescens 79 Rhinoplocephalus bicolor Hemiaspis signata Hemiaspis damelii Echiopsis curta 44 Drysdalia mastersii 69 Drysdalia coronoides Austrelaps superbus Austrelaps labialis 75 Echiopsis atriceps 48 Hoplocephalus bitorquatus Notechis scutatus Tropidechis carinatus Emydocephalus annulatus Aipysurus eydouxii Aipysurus laevis Aipysurus fuscus Aipysurus apraefrontalis Aipysurus duboisii Parahydrophis mertoni Hydrelaps darwiniensis Hydrophis elegans 69 Astrotia stokesii Hydrophis czeblukovi 33 Disteira major 3 Acalyptophis peronii 59 Hydrophis ornatus 23 Disteira kingii 8 Hydrophis brooki Hydrophis macdowelli Lapemis curtus Pelamis platura Hydrophis lapemoides Hydrophis spiralis Hydrophis cyanocinctus Hydrophis pacificus Hydrophis semperi Buhoma depressiceps 69 Buhoma procterae Lycophidion nigromaculatum Lycophidion laterale Lycophidion capense Lycophidion ornatum 70 Lamprophis swazicus Hormonotus modestus 76 Gonionotophis brussauxi Mehelya nyassae Mehelya stenophthalmus 63 Mehelya capensis Mehelya poensis Pseudoboodon lemniscatus Bothrolycus ater Bothrophthalmus brunneus Bothrophthalmus lineatus Lamprophis virgatus Lamprophis fuliginosus Lamprophis lineatus 69 Lamprophis olivaceus Lamprophis guttatus 29 Lamprophis fiskii Lamprophis inornatus Lycodonomorphus rufulus 46 Lycodonomorphus whytii Lycodonomorphus laevissimus Prosymna ruspolii Prosymna janii Prosymna visseri Homoroselaps lacteus 88 Atractaspis microlepidota 23 Atractaspis boulengeri Atractaspis bibronii Atractaspis corpulenta Atractaspis micropholis Macrelaps microlepidotus 92 Amblyodipsas dimidiata Xenocalamus transvaalensis Polemon notatus Polemon collaris Polemon acanthias Aparallactus werneri Aparallactus guentheri Aparallactus capensis 7 Psammodynastes pulverulentus Pseudaspis cana Pythonodipsas carinata Rhamphiophis rubropunctatus Malpolon monspessulanus Malpolon moilensis Dipsina multimaculata Psammophylax acutus Psammophylax rhombeatus Psammophylax tritaeniatus Psammophylax variabilis Mimophis mahfalensis Hemirhagerrhis viperina 5 58 Hemirhagerrhis hildebrandtii Hemirhagerrhis kelleri Psammophis crucifer Psammophis condanarus Psammophis lineolatus Psammophis trigrammus Psammophis jallae 70 Psammophis leightoni 75 Psammophis notostictus Psammophis angolensis Psammophis schokari 7 Psammophis punctulatus Psammophis praeornatus Psammophis biseriatus Psammophis tanganicus Psammophis lineatus Psammophis subtaeniatus 59 Psammophis sudanensis Psammophis orientalis 92 Psammophis rukwae Psammophis sibilans Psammophis leopardinus Psammophis mossambicus Psammophis phillipsi Ditypophis vivax Amplorhinus multimaculatus Duberria variegata Duberria lutrix Alluaudina bellyi 27 Compsophis albiventris Compsophis boulengeri 27 Compsophis laphystia 83 Compsophis infralineatus Liophidium rhodogaster Liophidium vaillanti Liophidium chabaudi 5 Liophidium torquatum Heteroliodon occipitalis Pseudoxyrhopus ambreensis Liopholidophis dimorphus Liopholidophis sexlineatus Liopholidophis dolicocercus 34 Dromicodryas bernieri Dromicodryas quadrilineatus Bibilava infrasignatus Bibilava epistibes 78 Bibilava stumpffi 5 Bibilava martae 4 Bibilava lateralis Leioheterodon geayi Leioheterodon madagascariensis Leioheterodon modestus Micropisthodon ochraceus 56 Langaha madagascariensis 85 Ithycyphus miniatus 55 Ithycyphus oursi Madagascarophis colubrinus Madagascarophis meridionalis 29 Stenophis betsileanus Stenophis pseudogranuliceps Stenophis citrinus Lycodryas sanctijohannis Plagiopholis styani Pseudoxenodon karlschmidti Thermophis baileyi Contia tenuis 44 Heterodon simus Heterodon platirhinos Diadophis punctatus 42 Farancia abacura 90 Carphophis amoenus Tantalophis discolor Amastridium veliferum 43 Coniophanes fissidens Rhadinaea flavilata 60 Rhadinaea fulvivittis Imantodes inornatus Imantodes lentiferus 88 Imantodes gemmistratus Imantodes cenchoa 79 Leptodeira nigrofasciata Leptodeira frenata Leptodeira punctata 36 Leptodeira splendida Leptodeira septentrionalis Leptodeira bakeri Leptodeira annulata Leptodeira maculata Leptodeira rubricata Pseudoleptodeira latifasciata Hypsiglena slevini Hypsiglena jani 63 Hypsiglena ochrorhyncha Hypsiglena torquata Hypsiglena chlorophaea Adelphicos quadrivirgatus Hydromorphus concolor Tretanorhinus nigroluteus Cryophis hallbergi Atractus elaps 5 Atractus wagleri Geophis carinosus Dipsas catesbyi Dipsas pratti 63 Ninia atrata 38 Tropidodipsas sartorii 62 Sibon nebulatus Conophis vittatus Hydrodynastes gigas Xenopholis scalaris Apostolepis flavotorquata Phalotris nasutus 2 Tomodon dorsatus 42 Pseudoeryx plicatilis Hydrops triangularis Helicops angulatus Tropidodryas striaticeps Tropidodryas serra 7 7 Philodryas baroni Philodryas viridissima Philodryas olfersii 4 Philodryas nattereri 3 Xenoxybelis boulengeri 27 Philodryas patagoniensis Pseudablabes agassizii Siphlophis cervinus Oxyrhopus guibei Oxyrhopus petola Phimophis iglesiasi 65 Drepanoides anomalus 77 Pseudoboa neuwiedii Boiruna maculata Psomophis joberti Uromacer catesbyi 7 6 Uromacer oxyrhynchus Uromacer frenatus Liophis lineatus 35 Erythrolamprus aesculapii Waglerophis merremi Lystrophis semicinctus 92 Xenodon neuwiedii 5 Arrhyton taeniatum Arrhyton supernum Arrhyton vittatum 52 Arrhyton landoi Arrhyton dolichura Arrhyton procerum 56 Arrhyton tanyplectum Arrhyton exiguum 50 Alsophis antillensis Alsophis rijgersmaei Alsophis rufiventris 72 Alsophis antiguae Alsophis portoricensis Hypsirhynchus ferox Antillophis parvifrons Arrhyton callilaemum Arrhyton polylepis 9 Arrhyton funereum Darlingtonia haetiana 70 Ialtris dorsalis 2 Antillophis andreae Alsophis vudii Alsophis cantherigerus Afronatrix anoscopus Natriciteres olivacea Xenochrophis punctulatus Xenochrophis flavipunctatus Xenochrophis vittatus Rhabdophis tigrinus Rhabdophis nuchalis Amphiesma sauteri Natrix natrix Natrix tessellata 78 Natrix maura 48 4 Sinonatrix annularis Storeria occipitomaculata Storeria dekayi 77 Virginia striatula 38 Clonophis kirtlandii 55 Seminatrix pygaea Regina rigida Regina alleni Regina grahami 23 Nerodia floridana Nerodia cyclopion Tropidoclonion lineatum 3 Regina septemvittata 30 Nerodia rhombifer Nerodia taxispilota Nerodia erythrogaster Nerodia fasciata Nerodia harteri 80 Nerodia sipedon Thamnophis sirtalis Thamnophis sauritus Thamnophis proximus Thamnophis rufipunctatus Adelophis foxi 28 Thamnophis melanogaster Thamnophis valida Thamnophis godmani Thamnophis exsul Thamnophis scaliger Thamnophis mendax Thamnophis sumichrasti Thamnophis chrysocephalus Thamnophis fulvus 80 Thamnophis cyrtopsis Thamnophis eques Thamnophis marcianus Thamnophis brachystoma Thamnophis radix Thamnophis butleri 59 Thamnophis hammondii 36 Thamnophis ordinoides Thamnophis elegans 92 Thamnophis atratus Thamnophis gigas 78 Thamnophis couchii Sibynophis collaris/chinensis Scaphiodontophis annulatus 4 Pseudorabdion oxycephalum Calamaria pavimentata Grayia tholloni Grayia smithii Grayia ornata 30 Chrysopelea paradisi Ahaetulla fronticincta 2 Dendrelaphis caudolineatus Ptyas mucosa Ptyas korros Opheodrys aestivus 56 Opheodrys vernalis Chironius carinatus 2 Oxybelis aeneus 7 Leptophis ahaetulla 34 Drymobius rhombifer Dendrophidion dendrophis 8 Coluber constrictor Masticophis flagellum 6 Spilotes pullatus 34 Phyllorhynchus decurtatus Trimorphodon biscutatus 4 Drymarchon corais 6 Drymoluber dichrous Mastigodryas melanolomus Mastigodryas boddaerti 7 Salvadora mexicana Chionactis occipitalis Sonora semiannulata 30 Chilomeniscus stramineus Tantilla relicta 8 Stenorrhina freminvillei Conopsis biserialis Conopsis nasus Pseudoficimia frontalis Sympholis lippiens Oligodon cinereus Scaphiophis albopunctatus Gonyosoma oxycephalum Gonyosoma jansenii Rhadinophis prasina Rhynchophis boulengeri Rhadinophis frenatum Lytorhynchus diadema Coluber zebrinus 75 Coluber dorri 79 Macroprotodon cucullatus 69 Hemorrhois hippocrepis Hemorrhois algirus Hemorrhois ravergieri 79 Hemorrhois nummifer Spalerosophis diadema Platyceps karelini Platyceps florulentus Platyceps najadum Platyceps collaris 36 Platyceps rhodorachis 37 Platyceps rogersi Hemerophis socotrae Dolichophis jugularis 43 Dolichophis caspius Dolichophis schmidti Hierophis viridiflavus Hierophis gemonensis Hierophis spinalis 2 Pseudocyclophis persicus 27 8 Eirenis aurolineatus Eirenis modestus Eirenis decemlineatus Eirenis lineomaculatus Eirenis coronelloides 28 Eirenis rothii Eirenis thospitis 56 Eirenis medus 8 Eirenis levantinus Eirenis punctatolineatus 54 Eirenis barani 36 Eirenis collaris Eirenis eiselti Coelognathus radiata Coelognathus helena 7 Coelognathus erythrurus Coelognathus flavolineatus Coelognathus subradiatus Philothamnus heterodermus Hapsidophrys smaragdina 5 Thrasops jacksonii 63 Dispholidus typus Thelotornis capensis 5 Lycodon zawi 75 Dinodon rufozonatum Dinodon semicarinatum Crotaphopeltis tornieri Dipsadoboa unicolor Telescopus fallax 4 26 Boiga pulverulenta 33 Boiga dendrophila 43 Dasypeltis scabra Dasypeltis atra Maculophis bella Oreocryptophis porphyraceus Euprepiophis mandarinus Euprepiophis conspicillata 4 Orthriophis hodgsoni 26 Orthriophis taeniurus Orthriophis cantoris 20 Orthriophis moellendorffi Zamenis hohenackeri 62 Zamenis persica Rhinechis scalaris Zamenis situla Zamenis lineata Zamenis longissimus Elaphe quatuorlineata Elaphe climacophora 58 Elaphe bimaculata 60 Elaphe dione Elaphe carinata 4 Elaphe schrenckii Elaphe quadrivirgata Coronella girondica Elaphe rufodorsata Coronella austriaca Senticolis triaspis Pantherophis spiloides 23 Pantherophis alleghaniensis Pantherophis obsoletus Pantherophis bairdi Pantherophis vulpinus Pantherophis guttatus Pantherophis slowinskii Pantherophis emoryi Pituophis deppei Pituophis lineaticollis Pituophis vertebralis Pituophis melanoleucus 5 Pituophis ruthveni Pituophis catenifer Rhinocheilus lecontei Pseudelaphe flavirufa 62 Arizona elegans 2 Bogertophis rosaliae 80 Bogertophis subocularis Cemophora coccinea 26 Lampropeltis calligaster Lampropeltis pyromelana Lampropeltis zonata Lampropeltis elapsoides 72 Lampropeltis mexicana Lampropeltis ruthveni Lampropeltis triangulum Lampropeltis extenuata 4 Lampropeltis alterna 88 Lampropeltis getula Lampropeltis nigra Lampropeltis holbrooki 7 Lampropeltis splendida Lampropeltis californiae 0.2

10 EDITORIAL COMMITTEE Publisher: Museu de Zoologia da Universidade de São Paulo. Avenida Nazaré, 4, Ipiranga, CEP , São Paulo, SP, Brasil. Editor-in-Chief: Carlos José Einicker Lamas, Serviço de Invertebrados, Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42.4, CEP , São Paulo, SP, Brasil. E mail: editormz@usp.br. Associate Editors: Mário César Cardoso de Pinna (Museu de Zoologia, Universidade de São Paulo, Brasil); Luís Fábio Silveira (Museu de Zoologia, Universidade de São Paulo, Brasil); Marcos Domingos Siqueira Tavares (Museu de Zoologia, Universidade de São Paulo, Brasil); Sérgio Antonio Vanin (Instituto de Biociências, Universidade de São Paulo, Brasil); Hussam El Dine Zaher (Museu de Zoologia, Universidade de São Paulo, Brasil). Editorial Board: Rüdiger Bieler (Field Museum of Natural History, U.S.A.); Walter Antonio Pereira Boeger (Universidade Federal do Paraná, Brasil); Carlos Roberto Ferreira Brandão (Universidade de São Paulo, Brasil); James M. 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