REV ISS WEB ZSC 12007 42-3 262.cs2014.cameronsiler.com/wp-content/uploads/2014/09/2013...Zaher et al. 2009; Pyron et al. 2011). To date, the superm-atrix of Pyron et al. (2011) has

Multilocus phylogeny reveals unexpected diversificationpatterns in Asian wolf snakes (genus Lycodon)CAMERON D. SILER, CARL H. OLIVEROS, ANSSI SANTANEN & RAFE M. BROWN

Submitted: 6 September 2012Accepted: 8 December 2012doi:10.1111/zsc.12007

Siler, C. D., Oliveros, C. H., Santanen, A., Brown, R. M. (2013). Multilocus phylogenyreveals unexpected diversification patterns in Asian wolf snakes (genus Lycodon). —ZoologicaScripta, 42, 262–277.The diverse group of Asian wolf snakes of the genus Lycodon represents one of many poorlyunderstood radiations of advanced snakes in the superfamily Colubroidea. Outside of threespecies having previously been represented in higher-level phylogenetic analyses, nothing isknown of the relationships among species in this unique, moderately diverse, group. Thegenus occurs widely from central to Southeast Asia, and contains both widespread species toforms that are endemic to small islands. One-third of the diversity is found in the Philippinearchipelago. Both morphological similarity and highly variable diagnostic characters havecontributed to confusion over species-level diversity. Additionally, the placement of thegenus among genera in the subfamily Colubrinae remains uncertain, although previousstudies have supported a close relationship with the genus Dinodon. In this study, we providethe first estimate of phylogenetic relationships within the genus Lycodon using a new multi-locus data set. We provide statistical tests of monophyly based on biogeographic, morpho-logical and taxonomic hypotheses. With few exceptions, we are able to reject many of thesehypotheses, indicating a need for taxonomic revisions and a reconsideration of the group'sbiogeography. Mapping of color patterns on our preferred phylogenetic tree suggests thatbanded and blotched types have evolved on multiple occasions in the history of the genus,whereas the solid-color (and possibly speckled) morphotype color patterns evolved onlyonce. Our results reveal that the colubrid genus Dinodon is nested within Lycodon—a clearfinding that necessitates the placing of the former genus in synonymy with the latter.Corresponding author: Cameron D. Siler, Department of Biology, University of South Dakota,Vermillion, SD 57069, USA. E-mail: [email protected] H. Oliveros, Department of Ecology and Evolutionary Biology, University of Kansas,Lawrence, KS, 66045-7561, USA. E-mail: [email protected] Santanen, Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence,KS, 66045-7561, USA. E-mail: [email protected] M. Brown, Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence,KS, 66045-7561, USA. E-mail: [email protected]

IntroductionThe superfamily Colubroidea (sensu Pyron et al. 2011), oradvanced snakes, represents one of the most strikinglydiverse terrestrial radiations of living vertebrates (Lawsonet al. 2005; Burbrink & Pyron 2008; Pyron et al. 2011).Currently, more than 2500 species are recognized (Lawsonet al. 2005; Pyron et al. 2011) in this relatively young cladedating only to the Cenozoic (Burbrink & Pyron 2008;Vidal et al. 2009). Seven families of snakes are consideredmembers of the clade (Colubridae, Elapidae, Homalopsi-dae, Lamprophiidae, Pareatidae, Viperidae and Xenoder-matidae; Pyron et al. 2011), which has been determined to

be monophyletic based on both morphological (Rieppel1988; Zaher 1999; Lee & Scanlon 2002) and moleculardata (Cadle 1988; Heise et al. 1995; Kraus & Brown 1998;Gravlund 2001; Slowinski & Lawson 2002, 2005; Wilcoxet al. 2002; Lawson et al. 2005; Pyron et al. 2011). Overthe last decade, numerous studies have investigated rela-tionships among advanced snakes within the superfamilyColubroidea with widely varying degrees of taxonomicinclusion (Lawson et al. 2005; Burbrink & Pyron 2008;Wiens et al. 2008; Kelly et al. 2009; Vidal et al. 2009;Zaher et al. 2009; Pyron et al. 2011). To date, the superm-atrix of Pyron et al. (2011) has been by far the most

262 ª 2013 The Norwegian Academy of Science and Letters, 42, 3, May 2012, pp 262–277

Zoologica Scripta

extensive of these studies, including 761 species represent-ing 299 genera (29% of the global species diversity ofadvanced snakes).The computational obstacles involved in estimating phy-

logenies for this and other highly diverse radiations aresubstantial; moreover, when combined with the absence ofavailable genetic samples for a large number of recognizedspecies, we likely are far from a complete understandingof species-level relationships within advanced snakes.Despite the relative success of new approaches and meth-odologies to employing supermatrices in phylogeneticstudies of species-rich groups (Sanderson et al. 2003;Driskell et al. 2004; Philippe et al. 2004; Wiens et al.2005; de Queiroz & Gatesy 2007; Thomson & Shaffer2010; Pyron et al. 2011), there is still a clear need for con-tinued research aimed at filling many of the gaps in taxo-nomic coverage and resolving more fine-scale relationshipsamong the many poorly understood genera within Colu-broidea. A characteristic example of one of these enig-matic genera is the Wolf Snakes of the genus Lycodonwithin the subfamily Colubrinae (family Colubridae).Although this radiation of non-venomous Asian snakes isonly moderately diverse (36 currently recognized species;The Reptile Database 2012), only three species have everbeen included in phylogenetic studies (L. capucinus, L. lao-ensis, and L. zawi; Heise et al. 1995; Kraus & Brown 1998;Lawson et al. 2005; Kelly et al. 2003, 2009; Pyron et al.2011). Among the studies that have included representa-tives of Lycodon, the genus has been consistently recoveredas the sister species to the genus Dinodon with moderatesupport (Kelly et al. 2003, 2009; Lawson et al. 2005; Pyronet al. 2011). When wider taxonomic sampling has beenincluded in phylogenetic analyses, the genera Lycodon andDinodon are recovered as members of a larger clade thatincludes the colubrid genera Boiga, Dasypeltis, Dipsadoboa,Crotaphopeltis, and Telescopus, albeit with weak support(Kelly et al. 2003, 2009; Lawson et al. 2005; Pyron et al.2011).In just the last two decades, 14 new species of snakes of

the genus Lycodon have been described, increasing theknown diversity of this Indian and Southeast Asian colubridgenus by nearly 40% (Ota & Ross 1994; Lanza 1999;Slowinski et al. 2001; Daltry & Wüster 2002; Gaulke 2002;Mukherjee & Bhupathy 2007; Vogel et al. 2009; Vogel &David 2010; Vogel & Luo 2011; Zhang et al. 2011; Vogelet al. 2012). The 36 recognized species of Lycodon occurthroughout central to Southeast Asia, from regions east ofthe Caspian Sea, eastern Iran and India to southern China,the Indo-Australian Archipelago, the Ryukyu Islands ofJapan and the Philippines (Lanza 1999; Fig. 1). Althoughmany of these newly described species have been namedfrom just a few specimens, and solely on the basis of

morphological data, sufficient genetic sampling has recentlybeen amassed to allow for an initial molecular appraisal ofthe genus’ phylogenetic relationships and affinities to otherSE Asian genera. The geographic ranges of species in thegenus Lycodon fall along a divergent spectrum, fromwidespread species whose range spans nearly the entire dis-tribution of the genus (i.e., L. aulicus) to a number ofrange-restricted, microendemic lineages (i.e., L. alcalai,L. bibonius, L. chrysoprateros, L. fausti, L. ferroni, L. solivagus,L. tesselatus). Coloration varies greatly across this uniquegroup of snakes; however, most species can be groupedinto one of four distinct colour pattern categories: banded,blotched, solid and speckled (Fig. 2). Variation in colourpatterns among species representing each of thesemorphotypes has consistently led to confusion over speciesboundaries (Lanza 1999).Nearly one-third of the diversity within the genus occurs

in the Philippines, with nine of the 11 species now consid-ered endemic to this island archipelago (Lanza 1999; Gau-lke 2002). Faunal demarcations in the archipelago havetraditionally been explained by the geography of Pleisto-cene aggregate island complexes (PAICs: Brown & Dies-mos 2002, 2009; Heaney 1985; Heaney et al. 1998, 2005;Siler et al. 2010, 2012b). During glacial periods, adjacentislands separated by shallow waters experienced greaterconnectivity as decreased sea levels (100–140 m below cur-rent levels) resulted in increased land-positive regions. Thecyclical nature of this process has provided an explanatorytool for explaining the distribution of biotic diversity in thePhilippines. Although recent studies have resulted in mixedsupport for the PAIC model of diversification (review: Sileret al. 2010, 2012b), this model remains a heuristic tool forexploring and understanding many of the evolutionary pro-cesses underlying the accumulation of biodiversity in thePhilippines.In this study, we investigate the patterns of diversifica-

tion among species of Lycodon from a phylogenetic perspec-tive, providing the first estimate of genealogicalrelationships for this unique radiation of Asian snakes.First, we attempt to estimate the phylogenetic position ofLycodon among recognized, closely related colubrid snakesto provide insight into patterns of morphological evolutionand regional diversification. Second, we provide a firstassessment of the evolution of major colour patterns in Lyc-odon. Finally, we test the following suite of biogeographic,morphological and taxonomic hypotheses aimed at betterunderstanding how diversity is partitioned among WolfSnakes. Here, we address three general questions: (i) Arecurrently recognized species boundaries supported by phy-logeny? (ii) Do previously defined species groups withcommon color pattern types form monophyletic clades?And (iii) do biogeographic patterns observed in other Asian

ª 2013 The Norwegian Academy of Science and Letters, 42, 3, May 2012, pp 262–277 263

C. D. Siler et al. � Multilocus phylogeny of Asian wolf snakes

squamates also apply to Wolf Snakes and do these explainspecies boundaries?Our data reveal patterns of lineage diversification at odds

with currently recognized taxonomy; we conclude thatalthough there is some support for a few instances thatmay result in the eventual recognition of additional crypticspecies, diversity within some portions of the genus actuallymay be overestimated as a result of taxonomic decisionsbased on colour pattern and untested biogeographic expec-tations. This article represents a first step towards anunderstanding of the major evolutionary trends in colourpatterns, biogeographic relationships and phylogeny-informed taxonomy of Asian wolf snakes.

Materials and methodsTaxon sampling and data collection

Ingroup sampling included 44 individuals representing atleast 16 of the 37 currently recognized species in Lycodon(Figs 1 and 2; Appendix 1). To the best of our knowl-

edge, tissues are unavailable for more than half of therecognized species diversity in the genus. To assessmonophyly of the genus, a broad sampling (17 taxarepresenting 11 genera) from the family Colubridae wasincluded (Appendix 1) based on recent higher-levelphylogenetic studies of Colubroidea (Kraus & Brown1998; Lawson et al. 2005; Kelly et al. 2009; Pyron et al.2011).Genomic DNA was extracted from liver tissues stored in

95% ethanol. We sequenced the mitochondrial gene Cyto-chrome b (cyt-b) in 50 vouchered specimens using pub-lished primers and protocols (Burbrink et al. 2000; Lawsonet al. 2005; Pyron et al. 2011). For 39 and 41 of these sam-ples (Appendix 1), we also sequenced a 569 base pairregion of the nuclear oocyte maturation factor Mos (c-mos) gene and a 1018 base pair region of the fifth intronof the nuclear Vimentin (vim) gene, respectively, usingpublished primers and protocols (Pyron and Burbrink2009; Pyron et al. 2011). Amplified products were

Fig. 1 Distribution of Lycodon samplesused for this study. Species-specificlocality markers correspond to theinternal figure key. Philippine samples areshown on the enlarged inset map. Therecognized distribution of Dinodon speciesincluded in this study is shaded forreference.


Multilocus phylogeny of Asian wolf snakes � C. D. Siler et al.

visualized on 1.5% agarose gels, and PCR products werepurified with 1 lL of a 20% dilution of ExoSAP-IT(US78201, Amersham Biosciences, Piscataway, NJ, USA).Cycle sequencing reactions were run using ABI Prism Big-Dye Terminator chemistry (Ver. 3.1; Applied Biosystems,Foster City, CA, USA) and purified with Sephadex

(NC9406038, Amersham Biosciences) in Centri-Sep 96spin plates (CS-961, Princeton Separations, Princeton, NJ,USA). Purified products were analysed with an ABI Prism3130xl Genetic Analyzer (Applied Biosystems). Continuousgene sequences were assembled and edited using SEQUEN-

CHER 4.8 (Gene Codes Corp., Ann Arbor, MI, USA). To

Fig. 2 Hypothesized relationships among species of Lycodon for the three genes sampled in this study (mitochondrial cyt-b; nuclear c-mosand vim), illustrated by ML bootstrap topologies. Nodes supported by � 95% Bayesian PP and � 70% MLBP are indicated by dots onnodes. Terminals are labeled with taxonomic names, followed by sampling localities when required for clarity. The phylogenetic position ofthe genus Dinodon is highlighted for reference.



this data set, we added 11 sequences of cyt-b and onesequence of c-mos of outgroup taxa available on genbank(Appendix 1). All novel sequences were deposited in Gen-Bank (Appendix 1).

Sequence alignment and phylogenetic analyses

Initial alignments were produced in Muscle (Edgar 2004)with minimal subsequent manual adjustments. To assessphylogenetic congruence between the mitochondrial andnuclear data, we inferred the phylogeny for each geneindependently using likelihood and Bayesian analyses. Fol-lowing the observation of no moderate to highly sup-ported incongruence between data sets (Fig. 2), we feltjustified in using the combined, concatenated, data forsubsequent analyses. Exploratory analyses of the combineddata set of 61 individuals (including all taxa, some ofwhich were missing data for c-mos and vim) and areduced data set of individuals with no missing data exhib-ited identical relationships; we therefore chose to includeall available data (61 individuals) for subsequent analysesof the concatenated data set. Alignments and resultingtopologies are deposited in Dryad (doi:10.5061/dryad.cp6gg).Concatenated, partitioned Bayesian analyses were con-

ducted in MRBAYES v3.2.1 (Ronquist & Huelsenbeck 2003).The mitochondrial data set was partitioned by codon posi-tion for the protein-coding region of cyt-b, c-mos and vimwere analysed as single subsets. The Akaike InformationCriterion (AIC), as implemented in JMODELTEST v0.1.1(Posada 2008), was used to select the best model of nucleo-tide substitution for each partition (Table 1). A rate multi-plier model was used to allow substitution rates to varyamong subsets, and default priors were used for all modelparameters. We ran four independent MCMC analyses,each with four Metropolis-coupled chains, an incrementalheating temperature of 0.02, and an exponential distribu-tion with a rate parameter of 25 as the prior on branchlengths (Marshall 2010). All analyses were run for 15 mil-lion generations, with parameters and topologies sampled

every 5000 generations. We assessed stationarity and con-vergence of parameters with TRACER v1.4 (Rambaut &Drummond 2007) and confirmed convergence of tree splitswith AWTY (Wilgenbusch et al. 2004). Stationarity wasachieved after three million generations (i.e., the first20%), and we conservatively discarded the first 25% ofsamples as burn-in.Partitioned maximum likelihood (ML) analyses were

conducted in RAXMLHPC v7.0 (Stamatakis 2006) on the con-catenated data set using the same partitioning strategy asfor Bayesian analysis. The more complex model (GTR +Γ) was applied for all subsets (Table 1), and 100 replicateML inferences were performed for each analysis. Eachinference was initiated with a random starting tree, andnodal support was assessed with 100 bootstrap pseudorepli-cates (Stamatakis et al. 2008).To compare the results of concatenated analyses to a

multilocus, coalescent analysis, we conducted partitioned,coalescent-based analyses in the program BEAST v1.7.4(Drummond & Rambaut 2007). Four independent BEAST

runs of 80 million generations were completed under thesame partitioning strategy as for Bayesian analyses, impos-ing an uncorrelated lognormal relaxed clock prior on sub-stitution rate (Drummond et al. 2006) and Yule speciationprior. Parameters were sampled every 20 000 generations,and the initial 50% of each run was discarded as burn-in.Convergence was evaluated under the same strategy as forBayesian analyses.

Hypotheses testing, biogeography, colour patterns and

taxonomy

We tested a series of hypotheses based on taxonomic,morphological and biogeographic expectations to addressthe following three major questions concerning the pat-terns of Lycodon diversification (Table 2). (i) Are currentlyrecognized species boundaries among widespread and mic-roendemic species supported within a phylogenetic con-text? (ii) Are the predominant color patterns exhibited byspecies of the genus Lycodon monophyletic groups? And(iii) do monophyletic species assemblages conform toregional biogeographic expectations? More specifically, wewere interested in addressing several questions concerningbiogeographic patterns of Lycodon diversification: (A) Arethe radiations of species of the genus Lycodon in the Phil-ippines and in Mainland Asia monophyletic? And (B)within the Philippines, are recognized island groups(Babuyan and Batanes Island Groups) regional centers ofdiversity for monophyletic radiations of microendemic spe-cies? To statistically evaluate the probability of eachexperimentally constrained topology, we estimated theprobability of each hypothesis within a Bayesian frame-work using the proportion of 3004 post-burn-in trees

Table 1 Models of evolution selected by AIC (as implemented inJMODELTEST) and those applied in partitioned, model-based, analy-ses of mitochondrial (cyt-b) and nuclear (c-mos, vim) data

Partition AIC modelNumber ofcharacters

Cytb, 1st codon position GTR + Γ 370Cytb, 2nd codon position GTR + Γ 370Cytb, 3rd codon position GTR 370Cmos HKY + Γ 569Vimentin HKY + Γ 1018



consistent with each topology as an estimate of the poster-ior probability of that hypothesis. This was accomplishedby filtering the pool of post-burn-in trees for each con-straint topology using the program PAUP* v4.0b10 (Swof-ford 1999).

ResultsTaxon sampling, data collection and sequence alignment

The complete, aligned matrix contains 44 samples of Lyc-odon, representing 43% of the currently recognized species.Seventeen additional samples are included from the familyColubridae, including representative taxa of the followinggenera: Ahaetulla, Boiga, Dasypeltis, Dinodon, Dipsadoboa,Ptyas, Gonyosoma, Lampropeltis, Oligodon, Pantherophis andPituophis. Following initial unrooted analyses, and in thelight of recent estimates of colubrid phylogeny (Pyron et al.2011), we rooted the tree using Ahaetulla prasina. A 364base pair insertion was observed in the middle of the vimgene for Lycodon effraensis. Exploratory analyses excludingthis region from the data set had no observable impact onthe results. The number of variable and parsimony-infor-mative characters are as follows : 584 and 518 of 1110 (cyt-b); 67 and 45 of 569 (c-mos); and 137 and 99 of 1018 (vim).

Phylogenetic analyses

With few exceptions, analyses of the combined data resultin topologies with high ML bootstrap support and poster-ior probabilities among our ingroup taxa. Although analy-ses resulted in poor to moderate support for severalrelationships among outgroup taxa, general topological pat-terns are congruent across these analyses (Figs 2 and 3;Fig. S1). As has been observed in recent higher-level stud-ies of the family Colubridae, poor basal support in theclade containing Wolf Snakes results in uncertainty regard-ing the close relatives of the genus Lycodon (Pyron et al.2011). Our analyses do recover a clade of colubrid snakes(Boiga + Dasypeltis + Dipsadoboa + Crotaphopeltis + Telescopus)as sister to a clade of Lycodon + Dinodon as was similarlyreported in Pyron et al. (2011), but with weak support(Fig. 3, Clade 1). Previous studies that had limited sam-pling of Lycodon have found high support for a sister rela-tionship between the genera Lycodon and Dinodon (Kellyet al. 2003, 2009; Lawson et al. 2005; Pyron et al. 2011).However, all concatenated and coalescent-based analysesbased on denser sampling of the genus recover Dinodontaxa nested within Lycodon [Figs 2 and 3 (Clades 2, 7); Fig.S1].

Table 2 Description of topology tests conducted using Bayesian methods

Hypothesis Description of constraint Posterior probability

Biogeographic testsH1 Monophyly of Philippine Lycodon All samples of Philippine Lycodon 0.0H2 Monophyly of Mainland Asia Lycodon All samples of Lycodon from Mainland

Asia (including Peninsular Malaysia)0.0

H3 Monophyly of Babuyan Island Group Lycodon All samples of Lycodon from the BabuyanIsland Group, Philippines

0.0

H4 Monophyly of Batanes Island Group Lycodon All samples of Lycodon from the BatanesIsland Group, Philippines

0.17

Tests of hypotheses based on morphologyH5 Monophyly of solid morphotype All samples of L. alcalai, L. chrysoprateros,

L. sp. (Babuyan Claro Island), and L. sp. (Calayan Island)11.0

H6 Monophyly of banded morphotype All samples of Lycodon bibonius, L. dumerilii, L. effraensis,L. cf. effraensis, L. fasciatus, L. laoensis, L. ruhstrati,L. stormi, and L. subcinctus

0.0

H7 Monophyly of blotched morphotype All samples of Lycodon aulicus, L. butleri, L. muelleri, andL. zawi

0.0

Tests of hypotheses based on current taxonomyInterspecificH8 Monophyly of Lycodon All samples of Lycodon 0.0H9 Reciprocal Monophyly of Lycodon and Dinodon All samples of Lycodon constrained to be reciprocally monophyletic

to a clade containing all samples of Dinodon0.0

IntraspecificH10 Monophyly of Lycodon alcalai All samples of Lycodon alcalai 0.18H11 Monophyly of Philippine L. aulicus All samples of Lycodon aulicus from the Philippines 0.0H12 Monophyly of Mainland Asia L. aulicus All samples of Lycodon aulicus from Mainland Asia 0.0

1The population of Lycodon on Babuyan Claro Island possesses a solid to lightly banded color pattern. For the purposes of this study we consider the population generally topossess a solid color pattern.



Fig. 3 Hypothesized relationships among species of Lycodon included in this study, illustrated by the maximum clade credibility treeresulting from Bayesian analyses of the concatenated molecular data set (mtDNA + nuDNA). Nodes supported by � 95% Bayesian PP and� 70% MLBP are indicated by dots on nodes. Terminals are labeled with taxonomic names and sampling localities, with representativephotographs showing the diversity of color patterns across the phylogeny. The phylogenetic position of the genus Dinodon is highlighted inred for reference. Numerical labels correspond to clades referred to in the Results and Discussion.



Four major clades were recovered among the focal taxawith strong support in all analyses (Fig. 3, Clades 3, 6, 10,and Lycodon stormi). All analyses recovered five Philippineendemic species in a single monophyletic clade (Fig. 3,Clade 3). The microendemic species from the Babuyan andBatanes Island Groups in the northern extreme of the Phil-ippines (L. alcalai, L. bibonius, L. chrysoprateros and L. sp.Babuyan Claro) were recovered as part of a single clade(Fig. 2, Clade 4); however, suprisingly members of thisclade showed very low interspecific genetic diversity. Thisclade of microendemics was recovered sister to L. muelleri,which occurs in northern Philippines. Sister to L. muelleriand the microendemics of the Babuyans and Batanes isL. dumerilii, a widespread Philippine endemic. Two majorclades recovered in all analyses consisted primarily ofMainland Asia and Peninsular Malaysia taxa, although pop-ulations of L. subcinctus and L. capucinus from the Philip-pines were nested within Clade 6 and 10, respectively(Fig. 3). The two species of Dinodon included in this study(D. semicarinatum and D. rufozonatum) are supported to bemonophyletic and are recovered as part of a polytomy withL. butleri, L. fasciatus and the sampled populations ofL. subcinctus (Fig. 3, Clade 6). All analyses recover the twosampled populations of L. effraensis from Thailand andPeninsular Malaysia as deeply divergent (Fig. 2, Clade 11).Although both ML and Bayesian analyses resulted in con-sistent topological relationships, one exception wasobserved. In ML analyses, L. stormi is recovered as the sis-ter species to the clade composed of L. effraensis, L. laoensis,L. jara, L. zawi and the widespread L. aulicus Complex(results not shown), while Bayesian analyses recoveredL. stormi as the sister species to Clade 6 + 10 (Fig. 3, Clade5). Interestingly, several widespread species show littlegenetic divergence among sampled populations (Lycodon au-licus Complex, L. dumerilii, L. laoensis), while other speciesshow considerable intraspecific genetic divergence (L. muel-leri, L. subcinctus).

Evaluating biogeographic, morphological and taxonomic

hypotheses

Among the biogeography-based hypotheses, the Bayesianmethod provided no support [posterior probability (pp)approaching zero] for the monophyly of species of Lycodonfrom the Philippines, Mainland Asia, or the Babuyan andBatanes Island Groups, respectively (Table 2). However,the tests recovered weak support for the monophyly ofisland populations of Lycodon from the Batanes IslandGroup (pp = 0.17; Table 2). Among the morphology-basedhypotheses, the results did not support the monophyly ofbanded or blotched morphotypes, but strongly supportedthe monophyly of the solid or near-solid morphotype(pp = 1.0; Table 2). It should be noted that the solid or

near-solid morphotype is present only in taxa in the islandsnorth of Luzon, among which there is little genetic diver-gence. The majority of taxonomy-based hypothesesreceived no support; however, the results weakly supportedthe monophyly of island populations of L. alcalai(pp = 0.18; Table 2).

DiscussionAffinities of Lycodon

This study represents the first phylogenetic analysis of thegenus Lycodon, and although genetic samples were availableonly for roughly half of the recognized species, our analy-ses resolved relationships among many of the included spe-cies with strong support (Figs 2 and 3). Although previousstudies have implied that Lycodon and Dinodon are recipro-cally monophyletic (Kelly et al. 2003, 2009; Lawson et al.2005; Pyron et al. 2011), this finding is an artifact of sparsesampling from Lycodon. Our results do not support themonophyly of the genus Lycodon (Figs 2 and 3; Table 2);rather, we find that Lycodon is paraphyletic with respect toDinodon. This result is not surprising considering thatmembers of these two genera are morphologically verysimilar, being distinguished only by a single, somewhatambiguously defined character, the degree of curvature ofthe maxillary bone (Smith 1943). Genetic samples areneeded for 21 species of Lycodon and six species of Dinodonunsampled in this study to estimate relationships among allspecies in these two genera of colubrid snakes.Previous studies that have included samples of Lycodon

have not been able to confidently resolve many of the sisterrelationships to the genus within the subfamily Colubrinae(Heise et al. 1995; Kraus & Brown 1998; Kelly et al. 2003,2009; Lawson et al. 2005; Pyron et al. 2011). Although ourattempts to sample widely from outgroup taxa results insome well-supported relationships, the most closely relatedgenera to the clade Lycodon + Dinodon remains weaklysupported.

Phylogeny and evaluation of species diversity

The discovery of species new to science cryptically masquer-ading among widespread species complexes has been exten-sively documented in Asia and beyond (Stuart et al. 2006;Bickford et al. 2007; Pfenninger & Schwenk 2007; Brown &Stuart 2012). In the archipelagos of Southeast Asia, the phe-nomenon has been shown to characterize many vertebrategroups (for reviews, see Stuart et al. 2006; Siler et al. 2012a)and is consistent with several findings supported in thisstudy. Our results support cases of deeply divergent lineageswithin some taxa (L. effraensis, L. subcinctus; Fig. 3) thatlikely represent unrecognized and/or undescribed species.Some of the lineage diversity we uncovered appears to corre-spond to taxonomic entities previously identified (currently



recognized subspecies or synonyms) and some do not. Forexample, three subspecies currently are recognized withinthe widespread Lycodon subcinctus (L. s. subcinctus, L. s. sealei,and L. s. maculatus; Lanza 1999; The Reptile Database2012). Our analyses revealed deep genetic divergencesamong at three populations of this species sampled fromThailand, Peninsular Malaysia, and the Philippines (Fig. 3,Clade 8). Additionally, the three divergent lineages recov-ered in all analyses not only appear to be morphologicallydistinct (Fig. 3; Lanza 1999; L. Grismer, B. Stuart, personalcommunication), but previously have been described as threedistinct taxonomic entities in the literature.Reinhardt described the nominal species in the early

1800s (in F. Boie 1827), and the subspecies L. s. subcinctusis currently recognized to occur in Vietnam, Laos, Cambo-dia, Thailand, Peninsular Malaysia, Indonesia (Bali, borneo,Java, Lombok, Mentaway Archipelago, Nias, Simeulue,Sumatra, Sumbawa) and possibly Singapore (Lanza 1999).Nutphand (1986) described L. suratensis without designat-ing type specimens in the description, and the species wassubsequently synonymized with L. subcinctus subcinctus(Pauwels et al. 2006). Leviton (1955) described a uniquesubspecies of L. subcinctus (L. s. sealei), which occurs inIndonesia (Borneo), Malaysia (Borneo), Brunei and thePhilippines (Palawan Island) (Lanza 1999; Gaulke 2002).Whether the three lineages we reveal correspond to eachof these previously recognized names (requiring theelevation of the three currently recognized subspecies) is asubject for future study, involving careful examination offreshly collected material, use of genetic samples from thetype localities, and examination of the name-bearing types.On the other hand, a few patterns observed in this study

contradict the commonly observed phenomenon of wide-spread cryptic diversity in Southeast Asia and, in fact, indi-cate that species diversity within several clades in the genusLycodon may be overestimated, rather than underestimated.For example, the entire clade of microendemic speciesfrom the Babuyan and Batanes Island Groups are separatedby an average of 1.3% (uncorrected p-distance for cyt-b)sequence divergence among island populations (Fig. 3,Clade 4), and if we exclude the one apparently divergentlineage (L. bibonius), the average genetic divergence amongthese putative species is reduced to a mere 0.3%. Thesegenetically similar island populations of the Babuyan andBatanes Island Groups also share similar colour patterns,and with the exception of the banded colour pattern ofL. bibonius, the remaining island populations possess quitesimilar non-banded to weakly banded color patterns (Ota& Ross 1994; CHO and RMB, personal observation). Withthe exception of L. chrysoprateros that appears to have fewerventral scales (186–194 vs. >198; Ota & Ross 1994) thanthe remaining microendemic species of the northern

Philippines, few diagnostic, morphological characters havebeen presented in the literature that (i) are not recognizedto be highly variable characters in the genus (e.g., colourpattern, relative sizes of scales), (ii) unambiguously distin-guish these recognized species from each other and (iii) arenon-overlapping in their variable states with the rangesobserved in the other island populations (Ota & Ross1994). The absence of genetic divergence observed in thisstudy among the extreme northern Philippine lineages,coupled with few diagnostic morphological characters (butsee Ota & Ross 1994), may indicate a need for future taxo-nomic revisions of this clade of Philippine endemic WolfSnakes (Fig. 3, Clade 4), with a possible concomitantreduction in recognized species diversity. This anticipatedaction would have important conservation implications;currently, L. chrysoprateros, a Dalupiri Island endemic, isconsidered ‘Critically Endangered’ (IUCN 2012).Between these two extremes lie species with moderate

genetic structure observed among populations (L. muelleri,L. aulicus Complex). Populations of L. muelleri from theBicol Faunal Region of Luzon Island and nearby Catandu-anes Island are 4.4–4.9% divergent from populations ineast-central Luzon (Fig. 3). It remains to be determinedwhether this genetic split also corresponds to the Mid-Sierra Filter Zone, a recently recognized biogeographicbarrier on Luzon Island (Welton et al. 2010; Balete et al.2011). Many more population-level samples are needed toproperly evaluate whether this species has a continuous dis-tribution across its range. It is interesting to note the largenumber of squamate species endemic to the Luzon faunalregion that are restricted either to the Bicol Faunal Region,including Polillo and/or Catanduanes Island and nearbyQuezon and Bulacan provinces of Luzon (Brachymeles bicol-andia, B. brevidactylus, B. cobos, B. lukbani, B. makusog,B. minimus, Cerberus microlepis, Luperosaurus cumingii, Parvo-scincus laterimaculatus, Pseudogekko smaragdinus, Varanus oli-vaceus) or to Luzon proper to the exclusion of the BicolPeninsula (Brachymeles bicolor, B. elerae, B. muntingkamay,B. wright, Eutropis bontocensis, E. multicarinata borealis, Gekkocarusadensis, Lipinia pulchella levitoni, Luperosaurus angliit,L. kubli, Lycodon solivagus, L. tessellatus, Parvoscincus beyeri,P. boyingi, P. hadros, P. igoratorum, P. lawtoni, P. leucospilos,P. luzonensis, P. tagapayo, Varanus bitatawa), with few spe-cies still recognized to have distributions spanning the tran-sition between faunal regions (Boiga dendrophila divergens,B. philippina, Brachymeles boulengeri, B. kadwa, Calamaria bi-torques, Otosaurus cumingi, Pinoyscincus abdictus aquilonius,V. marmoratus; PhilBREO 2012 LifeDesk,).The morphologically variable colour patterns within and

between populations of L. aulicus have led to long-standingconfusion over species boundaries. Many authors have longrecognized two subspecies, L. a. aulicus and L. a. capucinus



(Ota & Ross 1994; Lanza 1999; Ferner et al. 2000; Slowin-ski et al. 2001; Gaulke 2002; Das 2003; Jackson & Fritts2004), while others have recognized L. capucinus Boie, 1827as a full species, distinct from L. aulicus (Linnaeus 1758;Taylor 1965; David & Vogel 1996; Daltry & Wüster 2002;Pauwels et al. 2005; Mukherjee & Bhupathy 2007; Vogelet al. 2009; Reza 2010; Vogel & David 2010; McLeod et al.2011; Siler et al. 2011, 2012c; Zhang et al. 2011; Brownet al. 2012a,b; The Reptile Database 2012). Previous sys-tematic studies of Lycodon have noted that both taxa differprimarily in coloration alone, with single populations docu-mented to have a full spectrum of colour forms, from indi-viduals with no dorsal markings to individuals to completeand well-defined series of dorsal crossbars (Wall 1921;Lanza 1999; CDS, RMB, unpublished data). The majorityof studies have consistently divided the geographic rangesof L. a. aulicus and L. a. capucinus in Indochina (Smith1943; Leviton 1965; Fritts 1993; Zhao & Adler 1993; Das1994; David & Vogel 1996; Lanza 1999), with populationsof both forms reported to be sympatric in Myanmar (Lanza1999). Lycodon aulicus aulicus is recognized to primarilyoccur in southern Asia (Pakistan, India, Nepal, Myanmarand Sri Lanka; Smith 1943; David & Vogel 1996; Lanza1999), and L. a. capucinus in eastern Asia (Cook Islands[Australia], Myanmar, Cambodia, Thailand, Vietnam, Sin-gapore, Laos, China, Indonesia, Malaysia, Maldives, Masca-renes, and the Philippines; Smith 1943; Leviton 1965;Fritts 1993; Zhao & Adler 1993; Das 1994; David & Vogel1996; Lanza 1999; Whitaker & Captain 2004), althoughL. a. capucinus is also recognized to occur on the Nicobarand Andaman Islands of India (Smith 1943; Lanza 1999;Whitaker & Captain 2004).Several possibilities remain: (i) L. aulicus may represent a

single, widespread and morphologically variable species.Our results do not show a clear split between two recipro-cally monophyletic clades, nor do the few well-supportedsplits conform to recognized biogeographic regions (e.g.,non-monophyly of the Philippines and Indochina; Fig. 3,Clade 12). Furthermore, only moderate genetic diversity isobserved across the sampled populations, with an averagedivergence of only 3.6% (range = 0.0–5.0%). This level ofgenetic structure among populations is consistent with thedegree of intraspecific genetic variation observed withinother widespread Southeast Asian squamate reptiles (Gam-ble et al. 2012; Brown et al. 2012a,b; CDS, RMB, unpub-lished data) and yet is substantially more divergent than therecognized full species of the Babuyan and Batanes Islands(Ota & Ross 1994). (ii) Lycodon a. aulicus and L. a. capucinusmay represent two distinct lineages, and we have simplysampled from only one of these lineages, including thepopulation from Myanmar where they presumably are sym-patric. Our moderately robust sampling of populations

ranging from Myanmar to the Philippines may actuallyrepresent a single subspecies: L. a. capucinus. Until addi-tional samples from southern Asia (i.e., Andaman Islands,India, Pakistan, Nepal, China) become available, we areunable to evaluate this possibility. Finally (iii), it remainspossible that L. aulicus is actually a complex of unique,cryptic species, and future fine-scale studies of genetic andmorphological patterns across the species’ entire range mayresult in the recognition of additional taxa. Given the lowgenetic diversity observed between populations on Main-land Asia and the Philippines, particularly for the nuclearloci sequenced in this study (Fig. 2), it seems improbablethat many undescribed cryptic lineages are masqueradingamong populations of L. aulicus.

Biogeographic patterns among Wolf snakes

With few exceptions, the results observed in this study areconsistent with many of the biogeographic expectations forvertebrates in Asia and the Philippines. Phylogenetic analy-ses recover Philippine endemic species and Philippine pop-ulations of widespread species as part of three distinctclades (Fig. 3, Clades 3, 8, 12), suggesting that wolf snakeshave colonized the archipelago on at least three, and possi-bly four occasions. All endemic and microendemic Philip-pine species included in this study show a southeast tonorth progression via the eastern Philippine island arcwithin a single clade of the phylogeny (Fig. 3, Clade 3),while the endemic Palawan subspecies L. subcinctus sealeiappears to have arrived in the Philippines by means of thewestern Philippine island arc (Fig. 3, Clade 8). These pat-terns are consistent with many terrestrial vertebrates in thePhilippines, with these southern routes (eastern and wes-tern Philippine island arcs) recognized to be the two domi-nant dispersal pathways into the archipelago (e.g.,Diamond & Gilpin 1983; Heaney 1985, 1986; Brown &Guttman 2002; Evans et al. 2003; Jansa et al. 2006; Jones& Kennedy 2008; Brown et al. 2009; Esselstyn et al. 2009,2010; Esselstyn & Oliveros 2010).Without more complete taxon sampling, we are left with

many unanswered questions. (i) Do the Philippine speciesrecovered as members of Clade 3 (Fig. 3) replace eachother in allopatry? (ii) Is there an explanation for theapparent disparity between the extent of sympatry withinthe Philippines (low) vs. Sundaland (high)? Finally, giventhat one-third of the diversity of Lycodon occurs in thePhilippines, what role did the archipelago play in speciesdiversification among wolf snakes?

Affinities of unsampled taxa

On the basis of the results of this study, it is tempting tospeculate on anticipated phylogenetic affinities ofunsampled species diversity. Some unsampled species



relationships are almost certain, given colour pattern andbiogeographic relationships. For example, the unsampledPanay endemic species L. fausti and the northern LuzonIsland L. solivagus are morphologically very similar toL. muelleri and L. dumerilii (Ota & Ross 1994; Ota 2000;Gaulke 2002); accordingly, we expect these species to bemembers of the Philippine Clade 3 (Fig. 3) if genetic sam-ples become available in the future. In contrast, some rela-tionships are difficult to predict. The endemic Philippinespecies from Samar Island (L. ferroni) is brightly bandedand shares morphological similarity with L. stormi, L. sub-cinctus and L. dumerilii, three species that come out in dis-tinctively different parts of our preferred tree (Fig. 3).However, again, as many of the Philippine endemic andmicroendemic species are recovered as part of a singleclade, we might expect L. ferroni to be closely related tothis same group of species as well (Fig. 3, Clade 3).The reoccurrence of the banded morphotype throughout

our tree (Fig. 3), however, renders speculation tenuousconcerning the numerous unsampled banded species(L. cardamomensis, L. fasciatus, L. flavomaculatus, L. futsing-ensis, L. gongshan, L. liuchengchaoi, L. multifasciatus, L. ophi-ophagus, L. paucifasciatus, L. striatus and L. synaptor).Similarly, darkly blotched or near-solid colour patternsappear to have arisen numerous times (Fig. 3), suggestingthat the phylogenetic placement of unsampled taxa withsimilar colour patterns (L. flavicollis, L. osmanhilli, L. tiwariiand L. travancoricus) cannot be confidently predicted bycolour pattern alone.

Taxonomic conclusions

We consider Dinodon to be a junior synonym of Lycodonbased on the following: (i) the results of concatenated andcoalescent-based analyses are conclusive in the placementof Dinodon nested with strong support within the genusLycodon. Not only do statistical tests of topology reject thereciprocal monophyly of the two genera (Table 2), but it isclear that the addition of unsampled taxa from either ofthese genera would not result in their reciprocal mono-phyly. (ii) The two genera are separated on the basis ofonly a single, weak character (degree of curvature of themaxillary bone; Smith 1943). (iii) In this study, we haveincluded the type species of Dinodon, D. rufozonatum (Can-tor, 1842) as well as the type species of Lycodon, L. aulicus(Linnaeus, 1758), and in the case of L. aulicus, we havesampled the species somewhat densely (Fig. 3). (iv) Thegeneric name Lycodon H. Boie, 1826 is the oldest availablename and is the proper generic name for Asian wolf snakes(Adler & Zhao 1995), clearly indicating that DinodonDum�eril, 1853 would be most appropriately regarded as itsjunior synonym. Thus, we place Dinodon back in thesynonymy of Lycodon. Furthermore, it is important to note

that unsampled species of Lycodon and Dinodon, and otherunsampled colubrid genera, may be found to be allied withthis clade in future studies (A. Pyron, personal communica-tion).

AcknowledgementsThanks are due to several institutions, government agenciesand individuals who facilitated this study or provided logis-tical or material support crucial to the results presentedhere. The majority of Philippine sampling that contributedto this work was funded by a National Science FoundationBiotic Surveys and Inventories finding (DEB 0743491 toRMB) and a NSF Doctoral Dissertation Improvementgrant (DEB 0804115 to CDS). We thank the ProtectedAreas and Wildlife Bureau (PAWB) of the PhilippineDepartment of Environment and Natural Resources(DENR) for collection and export permits necessary forthis and related studies. The Economic Planning Unit ofMalaysia, the Chinese central government, and the govern-ment of Myanmar provided research and export permits toL. Grismer (LSUHC), R. Murphy, RMB (ROM and KU,respectively), and J. Vindum (CAS). We thank J. McGuire,L. Grismer, B. Stuart, R. Murphy, A. Diesmos, J. Vindum,and D. Blackburn for access to genetic material and A.Diesmos, V. Yngente, and J. Fernandez for help in thefield. Bryan Stuart, Alex Pyron and an anonymous reviewerprovided constructive comments on earlier drafts of themanuscript. Thanks are due to B. Stuart, L. Grismer andJ. McGuire for use of photographs reproduced in Fig. 2.

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Appendix 1 Summary of specimens and GenBank accession numbers corresponding to genetic samples included inthe study

Species Voucher LocalityGenbank Accession Numberscyt-b c-mos vim

Ahaetulla prasina KU 302954 Philippines, Polillo Island, Quezon Province,Barangay Pinaglubayan

KC010338 KC010299 KC010388

Ahaetulla prasina KU 326673 Philippines, Palawan Island, Palawan Province,Barangay Estrella, Estrella Falls Park

KC010339 KC010300 KC010389

Boiga cynodon KU 324614 Philippines, Negros Island, Negros OccidentalProvince, Barangay Patag, city of Silay,Mt. Bungol

KC010340 KC010301 KC010390

Boiga dendrophila KU 302957 Philippines, Panay Island, Antique Province,Barangay Duyong

KC010341 — KC010391

Dasypeltis atra CAS 201641 Uganda, Kabale district AF 471065 — —

Dinodon rufozonatum CIB 098274 Not reported JF827672 JF827695 —

Dinodon semicarinatus Not reported Ryukyu Archipelago, Japan AB008539 — —

Dipsadoboa unicolor CAS 201660 Uganda, Rukungiri District AF 471062 — —

Gonyosoma oxycephala KU 321724 Philippines, Mindanao Island, Zamboanga City“Province”, Pasonanca Natural Park, TumagaRiver

KC010343 KC010302 KC010393

Lampropeltis alterna Not reported Not reported AF 337128 — —

Lampropeltis triangulum Not reported Not reported AF 337134 — —

Lycodon alcalai KU 327847 Philippines, Bataan Peninsula, Batanes Province,Barangay San Antonio, Sitio Chadpidan

KC010344 KC010303 KC010394

Lycodon alcalai KU 327848 Philippines, Sabtang Island, Batanes Province,Barangay Chavayan

KC010345 KC010304 KC010395

Lycodon aulicus CAS 205000 Myanmar, Rakhine State KC010346 KC010305 KC010396Lycodon aulicus CDSGS 4621 Philippines, Bohol Island, Bohol Province KC010347 — KC010397Lycodon aulicus KU 305141 Philippines, Semirara Island, Antique Province,

Municipality of Caluya, Barangay TinobocKC010348 KC010306 KC010398

Lycodon aulicus KU 315170 Philippines, Mindanao Island, Zamboanga City,Pasonanca Natural Park, Tumaga River

KC010359 KC010317 KC010409

Lycodon aulicus KU 315378 Philippines, Tablas Island, Romblon Province,Barangay Balogo, Sitio Piqueno

KC010350 KC010308 KC010400

Lycodon aulicus KU 328524 Thailand, SERS, Nakhon Ratchasima KC010358 KC010316 KC010408Lycodon aulicus LSUHC 5725 West Malaysia, Johor, Pulau Rawa, KC010355 KC010313 KC010405Lycodon aulicus LSUHC 9277 Vietnam, Kien Giang, Nam Du Island KC010356 KC010314 KC010406Lycodon aulicus LSUHC 9695 Cambodia KC010357 KC010315 KC010407Lycodon aulicus PNM 7705 Philippines, Leyte Island, Leyte province, Leyte state

University campus, Barangay Guadalupe, Municipalityof Baybay

KC010349 KC010307 KC010399

Lycodon bibonius KU 304589 Philippines, Camiguin Norte Island, Cagayan Province,Barangay Balatabat, Local name “Limandok”

KC010351 KC010309 KC010401

Lycodon butleri LSUHC 8066 West Malaysia, Pahang, Fraser’s Hill KC010352 KC010310 KC010402Lycodon butleri LSUHC 9136 West Malaysia, Perak, Bukit Larut KC010353 KC010311 KC010403Lycodon butleri LSUHC 9421 West Malaysia, Pahang, Bukit Larut KC010354 KC010312 KC010404Lycodon chrysoprateros KU 307720 Philippines, Dalupiri Island, Cagayan Province,

Manolong CreekKC010360 KC010318 KC010410

Lycodon dumerilii KU 305168 Philippines, Dinagat Island KC010362 — KC010412Lycodon dumerilii KU 319989 Philippines, Mindanao Island, Agusan del Sur Province,

Barangay Bagusan II, Mt. MagdiwataKC010361 KC010319 KC010411

Lycodon dumerilii PNM 7751 Philippines, Leyte Island, Leyte Province, Municipality ofBurauen, Lake Mohagnao

KC010363 KC010320 KC010413

Lycodon cf. effraensis KU 328526 Thailand, Khao Luang NP, Nakhon Si Thammarat KC010364 KC010321 KC010414Lycodon effraensis LSUHC 9670 West Malaysia, Kedah KC010376 KC010328 —

Lycodon fasciatus CAS 234875 Myanmar, Chin State KC010365 — —

Lycodon fasciatus CAS 234957 Myanmar, Chin State KC010366 — —

Lycodon jara CAS 235387 Myanmar, Kachin State KC010367 KC010322 —



Appendix 1 Continued

Species Voucher LocalityGenbank Accession Numberscyt-b c-mos vim

Lycodon laoensis FMNH 258659 Lao PDR, Salavan Province KC010368 KC010323 —

Lycodon laoensis FMNH 262186 Vietnam, Dong Nai KC010369 KC010324 —

Lycodon laoensis KU 328529 Thailand, Khao Luang NP,Nakhon Si Thammarat

KC010371 — KC010416

Lycodon laoensis LSUHC 8481 Cambodia, Pursat Province, O’Lakmeas KC010370 KC010325 KC010415Lycodon muelleri CDSGS 5256 Philippines, Catanduanes Island KC010372 — KC010417Lycodon muelleri DLSUD 031 Philippines, Luzon Island, De La Salle University

reference collection, Cavite Province, Municipalityof Ternate, Sitio Maloayas

KC010373 KC010326 KC010418

Lycodon muelleri KU 313891 Philippines, Luzon Island, Camarines Norte Province,Barangay Tulay Na Lupa

KC010375 KC010327 KC010420

Lycodon muelleri KU 323384 Philippines, Luzon Island, Aurora Province,Barangay Lipimental

KC010374 — KC010419

Lycodon ruhstrati LSUHC 4199 China, Hainan Island, Wuzhi Shan KC010381 KC010332 —

Lycodon sp. KU 304827 Philippines, Baboyau Claro, Cagayan Province,Barangay Babuyan Claro

KC010377 KC010329 KC010421

Lycodon sp. KU 304844 Philippines, Baboyau Claro, Cagayan Province,Barangay Babuyan Claro

KC010378 KC010330 KC010422

Lycodon sp. KU 304870 Philippines, Calayan, Cagayan Province, BarangayMagsidel, Local Name “Macarra”

KC010379 — KC010423

Lycodon stormi JAM 7487 Indonesia, Sulawesi, Propinsi Sulawesi Tenggara,Kabupaten Konawe Selatan, Kecamatan Konda,Desa Moramo, Air Terjun Moramo

KC010380 KC010331 KC010424

Lycodon subcinctus KU 309447 Philippines, Irawan, Palawan Province, BarangayIrawan, Irawan Watershed

KC010385 — KC010428

Lycodon subcinctus KU 327571 Philippines, Palawan Island, Palawan Province,Barangay Estrella, Estrella Falls Park

KC010384 KC010335 KC010427

Lycodon subcinctus KU 328531 Thailand, SERS, Nakhon Ratchasima KC010383 KC010334 KC010426Lycodon subcinctus LSUHC 5016 West Malaysia, Pahang, Sungai Lembing Logging Camp, KC010382 KC010333 KC010425Lycodon zawi CAS 239944 Myanmar, Rakhine State KC010386 KC010336 —

Lycodon zawi CAS 210323 Myanmar, Sagaing Division AF 471040Oligdon maculatus KU 321699 Philippines, Mindanao Island KC010387 KC010337 —

Pantheropis bairdi isolate RDZ183 Not reported GU 073440 — —

Pantheropis obsoletus isolate RDZ264 Not reported GU 073446 — —

Pituophis catenifer JR18 Pet Trade FJ 627819 — —

Pituophis deppei JR12 Pet Trade FJ 627818 — —

Ptyas luzonensis TNHC 63002 Philippines, Luzon Island, Zambales Province,Municipality of Olongapo, SBMA Naval Base,“Nav-mag” area, Ilanin Forest (Triboa Bay)

KC010342 —

CAS = California Academy of Sciences Herpetological Collections; CDSGS = Non-vouchered genetic samples deposited at the University of Kansas Natural History Museum;CIB = Chengdu Institute of Biology; DLSUD = De La Salle University Dasmari~nas Reference Collection; FMNH = Field Museum of Natural History Herpetological Collections;JAM = Jimmy A. McGuire genetic sample, deposited in Museum Zoologicum Bogoriense (National Museum of Indonesia, Cibinong, Java); JR = Javier Rodriguez Robles;KU = University of Kansas Natural History Museum; LSUHC = La Sierra University Herpetological Collections; PNM = Philippine National Museum Herpetological Collections;RDZ = reference identification acronym presented in Vandewege et al. (2012); TNHC = Texas Natural History Collections, University of Texas at Austin.

Supporting InformationAdditional Supporting Information may be found in theonline version of this article:

Fig. S1. Maximum clade credibility chronogram amongspecies of Lycodon calculated in BEAST.



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REV ISS WEB ZSC 12007 42-3 262.cs2014.cameronsiler.com/wp-content/uploads/2014/09/2013...Zaher et al. 2009; Pyron et al. 2011). To date, the superm-atrix of Pyron et al. (2011) has