phylogeny-based species delimitation in philippine slender skinks

PHYLOGENY-BASED SPECIES DELIMITATION IN PHILIPPINESLENDER SKINKS (REPTILIA: SQUAMATA: SCINCIDAE:

BRACHYMELES): TAXONOMIC REVISION OF PENTADACTYLSPECIES GROUPS AND DESCRIPTION OF THREE NEW SPECIES

CAMERON D. SILER1

AND RAFE M. BROWN

Biodiversity Institute and Department of Ecology and Evolutionary Biology, University of Kansas, 1345 JayhawkBoulevard, Lawrence, KS 66045-7593, USA

ABSTRACT: We use data from external morphology and mitochondrial gene sequences to provide the basisfor a taxonomic revision of two polytypic, pentadactyl Philippine species of scincid lizards of the genusBrachymeles. Although previous studies have noted significant morphological variation among islandpopulations, the similarities in body size and scale pigmentation and pattern have led to the continuedrecognition of these two ‘‘widespread species.’’ A third, widespread, pentadactyl species, Brachymeles talinis,is known from Jolo Island and the central and northern Philippine islands. We evaluate both morphologicaland genetic data to define species limits in B. boulengeri, B. schadenbergi, and B. talinis. Our molecular andmorphological data indicate each of the four subspecies of B. boulengeri, and both subspecies of B.schadenbergi, are genetically distinct, with ranges biogeographically circumscribed, differ from theircongeners by numerous external morphological features, and therefore should be recognized as full species.Our morphological and genetic data necessitate the recognition of northern populations of B. talinis (fromLuzon Island) as a new species and also reveal an unanticipated new species from Masbate Island. Finally,morphological data require the recognition of the B. talinis population from Jolo Island as a unique, newspecies. These 10 taxa elevate the total known number of species of Brachymeles from 18 to 25.

BUOD: Gamit ang datos mula sa panglabas na kaanyuan at mga gene sequence mula sa mitochondrianagsagawa kami ng rebisyon ng tatlong species ng reptilyanong nabibilang sa genus na Brachymeles. Sa kabilang malaking pagkakaiba sa kaanyuan ng mga populasyon sa iba’t ibang isla, dalawang kalat na species—Brachymeles boulengeri at Brachymeles schadenbergi—ang patuloy na kinikilala dahil sa magkakatulad nalaki ng katawan at pagkakahawig ng kulay at disenyo ng kanilang kaliskis. Ang ikatlong species, angBrachymeles talinis, ay matatagpuan sa isla ng Jolo at sa gitna at hilagang bahagi ng Pilipinas. Ipinapakita ngaming datos na ang bawat isa sa apat na subspecies ng B. boulengeri at ang dalawang subspecies ng B.schadenbergi ay may natatanging kalamnang genetiko, may bukod-bukod na distribusyon, at naiiba sa ibangmiyembro ng genus sa maraming aspekto ng panlabas na kaanyuan at samakatwid ay nararapat na kilalaninbilang ganap na species. Binibigyang katwiran ng aming datos genetiko ang pagkilala sa populasyon ng B.talinis mula sa Luzon bilang bagong species at ang pagtuklas ng isang di-anaasahang bagong species mula saisla ng Masbate. Sa huli, kinakailangan ding makilala ang populasyon ng B. talinis mula sa isla ng Jolo bilangisang bago at ganap na species dahil sa kanilang namumukod na kaanyuan. Sa pamamagitan ng rebisyong ito,tumataas ang bilang ng species na nabibilang sa genus na Brachymeles mula sa Pilipinas sa 25.

Key words: Biodiversity; Brachymeles boulengeri; Brachymeles schadenbergi; Brachymeles talinis;Endemism; Faunal region; Limb reduction; Philippines; Taxonomy

THERE ARE ONLY four genera of scincidlizards possessing both fully limbed andlimbless species (Brachymeles, Chalcides,Lerista, and Scelotes; Brandley et al., 2008;Lande, 1978; Wiens and Slingluff, 2001).Within the genus Brachymeles, all but one ofthe 18 recognized species are endemic to thePhilippines. The exception is B. apus fromnorthern Borneo (Brown and Alcala, 1980;Hikida, 1982; Siler et al., 2009, 2010a,b). Sixspecies are pentadactyl (B. bicolor, B. bou-

lengeri, B. gracilis, B. makusog, B. schaden-bergi, and B. talinis); eight are nonpentadac-tyl, with reduced limbs and numbers ofdigits (B. bonitae, B. cebuensis, B. elerae, B.muntingkamay, B. pathfinderi, B. samarensis,B. tridactylus, and B. wrighti); and four areentirely limbless (B. apus, B. minimus, B.lukbani, and B. vermis). Within the nonpen-tadactyl species (Brown, 1956; Brown andRabor, 1967; Dumeril and Bibron, 1839;Taylor, 1917, 1918, 1925) exist a wide rangeof limb- and digit-reduced states, from minutelimbs that lack full digits (B. bonitae, B.1 CORRESPONDENCE: email, [email protected]

Herpetological Monographs, 24, 2010, 1–54

E 2010 by The Herpetologists’ League, Inc.

1

cebuensis, B. muntingkamay, B. samarensis,and B. tridactylus) to moderately developedlimbs with four to five digits on the hands andfeet (B. elerae, B. pathfinderi, and B. wrighti:Brown and Alcala, 1980; Hikida, 1982; Siler etal., 2009, 2010b). All species are semifossorialand typically found in dry, rotting materialinside decaying logs or in loose soil and leaflitter.

Shared body plans and similar externalmorphological features among populations ofBrachymeles has proven problematic fordiagnosing species (Brown and Alcala, 1980;Siler et al., 2009, 2010a,b). In addition, severalrare, mid- to high-elevation species have longbeen represented by only a few specimens, insome cases without knowledge of their exacttype locality (e.g., B. bicolor, B. elerae, B.wrighti, and B. pathfinderi). Three species arepolytypic: B. boulengeri contains four subspe-cies and B. gracilis and B. schadenbergi eachcontain two subspecies (Brown, 1956; Brownand Rabor, 1967; Brown and Alcala, 1980).Several other species are recognized as havingwidespread distributions that span historicalfaunal demarcations in the Philippines (Brownand Diesmos, 2002; Brown and Guttman,2002; Heaney, 1985, 1986), including B.talinis, B. samarensis, and B. bonitae (Brown,1956; Brown and Alcala, 1980; Brown andRabor, 1967).

TAXONOMIC HISTORY

The genus Brachymeles was first describedby Dumeril and Bibron (1839) for the small,limb-reduced species B. bonitae. Three addi-tional species (Senira bicolor, Gray, 1845;Eumeces (Riopa) gracilis, Fischer, 1885; andE. (R.) schadenbergi, Fischer, 1885) weretransferred to the genus by Boettger (1886)and Boulenger (1887). These four speciesrepresented the known diversity in the genusfor 30 yr, until Taylor published a series ofherpetofaunal descriptions in the early 1900s.In Taylor’s (1917) review of the genus, herevised B. gracilis to not only include popu-lations in the Mindanao Faunal Region butalso populations on Negros and Mindoroislands (Fig. 1). A few years later, Taylor(1922b) described B. boulengeri, based onmaterial from Polillo Island, and includedpopulations from Luzon, Mindoro, and Ne-

gros islands as representatives of the species(Fig. 1). Thirty years later, Brown (1956)described B. gracilis taylori and includedB. boulengeri as one of three subspecies ofthe polytypic species B. gracilis. Brown andRabor’s (1967) description of B. gracilisboholensis and B. g. mindorensis brought thenumber of subspecies within B. gracilis tofive. It was not until 1980 that Brown andAlcala (1980) resurrected the polytypic spe-cies B. boulengeri and included four subspe-cies (B. b. boulengeri, B. b. boholensis, B. b.mindorensis, and B. b. taylori), all believed tobe distinct from B. gracilis. This viewcharacterized the taxonomy of B. boulengerifor the next 30 yr. Numerous studies have

FIG. 1.—Map of the Philippine islands, with islandlabels provided for islands with representative samplesused for this study. The five recognized major Pleistoceneaggregate island complexes (PAICs), major island groups,and additional deep-water islands are labeled for refer-ence. Islands of the Romblon Island group are designatedby the first letter of the island name (T, Tablas Island; R,Romblon Island; and S, Sibuyan Island). Current islandsin the Philippines are shown in medium gray; light grayareas enclosed in black 120-m bathymetric contoursindicate the hypothesized maximum extent of land duringthe mid- to late Pleistocene.

2 HERPETOLOGICAL MONOGRAPHS [No. 24

mentioned the morphological variation amongisland populations of B. boulengeri and otherspecies (Brown, 1956; Brown and Alcala,1980; Brown and Rabor, 1967; Taylor,1922b), but all refrained from elevating thesesubspecies to full species. Brachymeles bou-lengeri boulengeri and B. b. taylori have largergeographic distributions across multiple is-lands within a single faunal region (Brownand Alcala, 1980; Brown and Rabor, 1967;Taylor, 1922b), whereas B. b. boholensis andB. b. mindorensis are single-island endemics(Brown, 1956; Brown and Alcala, 1980; Brownand Rabor, 1967).

Although B. talinis was described originallyas B. schadenbergi talinis (Brown, 1956), andconsidered part of the widespread B. scha-denbergi complex from the Sulu Islands andthe Mindanao, Visayan, and Luzon Pleisto-cene aggregate island complexes (PAICs;Brown and Diesmos, 2002; Brown and Gutt-man, 2002), the subspecies was described onthe basis of material from Negros Island(Brown, 1956). Brown (1956) referred toseries of specimens from Jolo and Luzonislands as likely exemplars of B. s. talinis andhypothesized three explanations for the un-usual distribution of B. schadenbergi in thePhilippines: (1) chance colonization acrossocean barriers into distinct faunal regions bytwo subspecies; (2) morphological conver-gence within a polytypic species; and (3)prolonged maintenance of two morphologi-cally similar, disjunct distributions of siblingspecies within the Philippines. At the time,Brown (1956) supported the first hypothesiswith reservations; however, in Brown andRabor’s (1967) review of Brachymeles, newlyavailable material supported the third hypoth-esis and led to the recognition of B. talinisfrom Jolo, Negros, and Luzon islands as adistinct ‘‘sibling species’’ of B. schadenbergipopulations from Basilan, Mindanao, Bohol,and Leyte islands. When Brown and Alcala(1980) revised the genus, they restricted thegeographic distribution of B. talinis to thecentral and northern Philippine islands andpostponed the assignment of the two speci-mens from Jolo until the morphologicalvariability of that island population was betterunderstood. Although samples from through-out the central and northern range of B.

talinis have been available, the recognition ofthis widespread species has continued for.40 yr (Brown and Alcala, 1980; Siler et al.,2009, 2010a,b).

Following the separation of B. talinis fromthe B. schadenbergi complex, Brown andRabor (1967) recognized two subspecies ofB. schadenbergi: one subspecies from westernand south central Mindanao Island andBasilan Island (B. s. schadenbergi), and theother subspecies from eastern Mindanao,Camiguin Sur, Bohol, and Leyte islands (B.s. orientalis). Fischer (1885) had previouslydesignated the type locality for B. schaden-bergi as southern Mindanao Island, andspecimens from south central MindanaoIsland have thus been identified as B. s.schadenbergi (Brown and Alcala, 1980; Brownand Rabor, 1967). In addition to its alreadybroad geographic distribution, Brown andAlcala (1980) predicted that B. s. orientalisalso would be observed on Samar Island.

Both B. boulengeri and B. talinis aredistributed across several distinct PAICs,including the Luzon, Mindanao, Mindoro,and Visayan island complexes (Fig. 1). Becausemany recent studies have revealed that fewendemic Philippine reptiles actually possessbroad distributions spanning these regionalfaunistic boundaries (Brown and Diesmos,2002, 2009; Siler et al., 2010a; Welton et al.,2009, 2010), we have begun to reevaluate theknown polytypic and widespread species in thegenus Brachymeles. Our goal is to revise thetaxonomy such that individual units (species)represent independently evolving, cohesivelineage segments (sensu Simpson, 1961; deQueiroz, 1998, 1999; Frost and Hillis, 1990;Wiley, 1978). Careful examination of numerousrecently collected specimens from throughoutthe known ranges of B. boulengeri, B. scha-denbergi, and B. talinis, as well as all relevantname-bearing type material, results in thereorganization of B. boulengeri, B. schaden-bergi, and B. talinis into 10 distinct species. Inthis paper, we provide a phylogenetic analysisof most of these taxa, fully describe eachevolutionary lineage, clarify species boundar-ies, and provide the first illustrations of mostthese taxa. We also provide information oneach species’ natural history, ecology, andgeographic distribution.

2010] HERPETOLOGICAL MONOGRAPHS 3

MATERIALS AND METHODS

Fieldwork, Sample Collection, andSpecimen Preservation

Fieldwork was conducted on Bohol, Cala-yan, Camiguin Norte, Camiguin Sur, Catan-duanes, Dinagat, Leyte, Luzon, Masbate,Mindanao, Mindoro, Negros, Panay, Polillo,Romblon, Samar, Sibuyan, and Tablas islands,all in the Philippines (Fig. 1) between 1992and 2009. Specimens were collected between0900 and 1600 h, euthanized with aqueouschloretone, dissected for genetic samples(liver preserved in 95% ethanol or flash frozenin liquid nitrogen), fixed in 10% buffered-formalin, and eventually (,2 mo) transferredto 70% ethanol. Newly sequenced voucherspecimens are deposited in US and Philippinemuseum collections (see Acknowledgmentsand Specimens Examined); if available,voucher information corresponding to datafrom GenBank sequences is included inTable 1.

Taxon Sampling and Outgroup Selection forPhylogenetic Analyses

Because our primary goal was to estimatephylogenetic relationships among the subspe-cies and island populations of B. boulengeri,B. schadenbergi, and B. talinis, we sequencedonly one to two exemplars per species;however, in the case of a species occurringon multiple islands within a single PAIC, oracross a large island such as Mindanao, wesampled multiple populations to providegreater geographic resolution. We chose twoscincid taxa (Plestiodon egregius and Lygo-soma bowringii) based on relationships pre-sented in a recent phylogenetic analysis ofscincid lizards (Brandley et al., 2005) asoutgroups. In total, 39 ingroup samples wereused to construct phylogenetic inferences.There are no tissue samples of the populationof B. talinis from Jolo Island, and thispopulation is therefore not included in thephylogenetic analyses.

DNA Extraction, Purification,and Amplification

We extracted total genomic DNA fromtissues (Table 1) by using the modifiedguanidine thiocyanate extraction method ofEsselstyn et al. (2008). The mitochondrial

ATPase 8 (ATP8) and ATPase 6 (ATP6)protein-coding genes were amplified usingstandard polymerase chain reaction (PCR)methods with the primers ATPf (59-CTCA-GARATCTGCGGGYCAAATCACA-39) andATPr (59-GTGCYTTCTCGRRTAATRTCYC-GTCAT-39; M. Brandley, unpublished data).PCR products were visualized on 1.0%agarose gels and then purified with ExoSAP-IT (US78201, GE Healthcare Biosciences,Piscataway, New Jersey, USA). Purified tem-plates were sequenced with the same primersand the ABI Prism BigDye Terminatorchemistry, version 3.1 (Applied Biosystems,Foster City, California, USA). Cycle-sequenc-ing products were purified with SephadexMedium (NC9406038, GE Healthcare Biosci-ences) in Centri-Sep 96 spin plates (CS-961,Princeton Separations, Princeton, New Jersey,USA). Sequencing products were run on anABI Prism 3130xl Genetic Analyzer (AppliedBiosystems). Gene sequences were assembledwith Sequencher 4.8 (Gene Codes Corp., AnnArbor, Michigan, USA).

Alignment and Phylogenetic Analysis

An initial alignment was produced inMuscle, version 3.7 (Edgar, 2004), and visualinspections were made in MacClade 4.08(Maddison and Maddison, 2005). No instanc-es of insertions or deletions, or ambiguouslyaligned regions, were observed in the data,and all data were used for analyses. The finalalignment consisted of 838 aligned nucleo-tides.

Phylogenetic analyses were conducted us-ing parsimony, likelihood, and Bayesian opti-mality criteria. Parsimony (maximum parsi-mony [MP]) analyses were conducted inPAUP* 4.0 (Swofford, 2002), with all charac-ters weighted equally. Most-parsimonioustrees were estimated using heuristic searcheswith 1000 random addition-sequence repli-cates and tree bisection and reconnection(TBR) branch swapping. To assess heuristicsupport, nonparametric bootstrapping wasconducted using 1000 replicates, each with100 random addition-sequence replicates andTBR branch swapping.

Partitioned maximum likelihood (ML) anal-yses were conducted in RAxML-HPC, version7.04 (Stamatakis, 2006). The alignment was


TABLE 1.—Summary of specimens corresponding to genetic samples included in the study, general locality, andGenBank accession number. PNM/CMNH 5 deposited in the Cincinnati Museum of Natural History; LSUHC 5 LaSierra University Herpetological Collections; * 5 currently uncataloged specimen, deposited in the National Museum of

the Philippines.

Species Voucher Locality

GenbankAccessionNumbers

ATPase 8, 6

Lygosoma bowringi LSUHC 6970 West Malaysia HQ239366Plestiodon egregius MVZ 11013 Not available NC000888Brachymeles taylori ACD 862* Philippines, Cebu Island HQ239368B. taylori KU 307738 Philippines, Negros Island, Municipality of Valencia,

Mt. TalinisHQ239369

B. taylori RMB 3226* Philippines, Negros Island, Municipality of Valencia,Mt. Talinis

HQ239370

B. mindorensis KU 307739 Philippines, Mindoro Island, Municipality of Bongabong HQ239371B. mindorensis KU 307740 Philippines, Mindoro Island, Municipality of Bongabong HQ239372B. boholensis RMB 2866* Philippines, Bohol Island, Municipality of Bilar HQ239373B. boholensis RMB 2875* Philippines, Bohol Island, Municipality of Bilar HQ239374B. boulengeri KU 307753 Philippines, Polillo Island, Municipality of Polillo HQ239375B. boulengeri KU 307754 Philippines, Polillo Island, Municipality of Polillo HQ239376B. boulengeri KU 313831 Philippines, Luzon Island, Municipality of Labo, Mt. Labo HQ239377B. boulengeri KU 313829 Philippines, Luzon Island, Municipality of Labo, Mt. Labo HQ239378B. boulengeri KU 320058 Philippines, Luzon Island, Municipality of Laguna,

Mt. MakilingHQ239379

B. boulengeri KU 320059 Philippines, Luzon Island, Municipality of Laguna,Mt. Makiling

HQ239380

B. kadwa ACD 3290* Philippines, Luzon Island HQ239381B. kadwa RMB 4040* Philippines, Luzon Island, Municipality of Irosin, Mt.

BulusanHQ239382

B. kadwa KU 307965 Philippines, Camiguin Norte Island, Municipality ofCalayan

HQ239383

B. kadwa KU 307985 Philippines, Camiguin Norte Island, Municipality ofCalayan

HQ239384

B. kadwa KU 304906 Philippines, Calayan Island, Municipality of Calayan HQ239385B. kadwa KU 304875 Philippines, Calayan Island, Municipality of Calayan HQ239386B. talinis RMB 3305* Philippines, Negros Island, Municipality of Valencia,

Mt. TalinisHQ239387

B. talinis KU 303990 Philippines, Sibuyan Island, Municipality of Magdiwang HQ239388B. talinis KU 306786 Philippines, Panay Island, Municipality of Sibalom HQ239389B. talinis KU 306781 Philippines, Panay Island, Municipality of Sibalom HQ239390B. talinis KU 315358 Philippines, Tablas Island, Municipality of Calatrava HQ239391B. talinis KU 315359 Philippines, Tablas Island, Municipality of Calatrava HQ239392B. tungaoi KU 323935 Philippines, Masbate Island, Municipality of Mobo HQ239393B. tungaoi KU 323936 Philippines, Masbate Island, Municipality of Mobo HQ239394B. tungaoi KU 323933 Philippines, Masbate Island, Municipality of Mobo HQ239395B. tungaoi KU 323934 Philippines, Masbate Island, Municipality of Mobo HQ239396B. schadenbergi PNM/CMNH

H1457Philippines, Mindanao Island, Municipality of Kiamba,

Mt. BusaHQ239397

B. schadenbergi KU 314980 Philippines, Mindanao Island, Municipality ofZamboanga City

HQ239398

B. schadenbergi KU 314981 Philippines, Mindanao Island, Municipality ofZamboanga City

HQ239399

B. orientalis KU 314092 Philippines, Mindanao Island, Municipality of SanFrancisco

HQ239400

B. orientalis KU 311239 Philippines, Leyte Island, Municipality of Baybay HQ239401B. orientalis KU 311230 Philippines, Leyte Island, Municipality of Baybay HQ239402B. orientalis KU 310945 Philippines, Samar Island, Municipality of Taft HQ239403B. orientalis KU 310357 Philippines, Camiguin Sur Island, Municipality of

MambajaoHQ239404

B. orientalis KU 305469 Philippines, Dinagat Island, Municipality of Loreto HQ239405B. orientalis KU 324028 Philippines, Bohol Island, Municipality of Bilar HQ239406


partitioned into six regions consisting of thecodon positions of ATP8 and ATP6. Analysesthat partition protein-coding genes by codonposition have been shown to improve resultinginferences (Brandley et al., 2005). The parti-tions were run under the same model (GTR +I + C), with 100 replicate best-tree inferences.Each inference was performed with a randomstarting tree and relied on the rapid hill-climbing algorithm (Stamatakis, 2006). Cladesupport was assessed with 1000 bootstrappseudoreplicates. We considered branchesreceiving $70% bootstrap support to be wellsupported (Hillis and Bull, 1993; also seeWilcox et al., 2002). The Akaike InformationCriterion (AIC), as implemented in MrMo-delTest 2.2 (Nylander, 2004), was used to findappropriate models of sequence evolution.The best-fit model for each of the sixpartitions (Table 2) was used for Bayesiananalyses performed in MrBayes 3.1 (Ronquistand Huelsenbeck, 2003). The same partition-ing strategy used for ML analyses was used forBayesian inferences. Searches over tree spacewere conducted with four runs, each with fourchains, and were run for 2 3 107 generations.Trees were sampled every 1000 generations,with 4000 samples discarded as burn-in; thisleft 16,001 post burn-in trees from each runincluded in the summary. Visual inspectionfor chain stationarity and high effectivesample size values was conducted within theprogram Tracer, version 1.4 (Rambaut andDrummond, 2007). In addition, correlationsof split frequencies and cumulative splitfrequencies were examined using the programAWTY (Nylander et al., 2008). We consideredbranches with posterior probabilities $0.95 tobe well supported (Wilcox et al., 2002).

Morphological Data

We examined fluid-preserved specimens(see Appendix) for variation in qualitativeand mensural characters. Sex was determinedby gonadal inspection, and measurementswere taken to the nearest 0.1 mm with digitalcalipers by CDS. Museum abbreviations forspecimens examined follow Leviton et al.(1985).

Meristic and mensural characters werechosen based on Siler et al. (2009, 2010a,b):characters evaluated were snout–vent length(SVL), axilla–groin distance (AGD), totallength (TotL), midbody width (MBW), mid-body height (MBH), tail length (TL), tailwidth (TW), tail height (TH), head length(HL), head width (HW), head height (HH),snout–forearm length (SnFa), eye diameter(ED), eye–narial distance (END), snoutlength (SNL), internarial distance (IND),forelimb length (FLL), hind limb length(HLL), midbody scale-row count (MBSR),paravertebral scale-row count (PVSR), axilla–groin scale-row count (AGSR), Finger-IIIlamellae count (FinIIIlam), Toe-IV lamellaecount (ToeIVlam), supralabial count (SL),infralabial count (IFL), supraciliary count(SC), and supraocular count (SO). In thedescription, ranges are followed by mean 6SD in parentheses.

Species Concept

We follow the general lineage concept ofspecies (de Queiroz, 1998, 1999) as a logicalextension of the evolutionary species concept(Frost and Hillis, 1990; Simpson, 1961; Wiley,1978). We consider as distinct lineages thosepopulations that are morphologically, andgenetically, distinct. Lineage-based speciesconcepts have been successfully used in therecognition of Philippine biodiversity (Brownand Diesmos, 2002; Brown et al., 2000a, 2008,2009; Brown and Guttman, 2002; Gaulke etal., 2007; Welton et al., 2009, 2010) due to thehighly partitioned nature of the archipelago(Brown and Diesmos, 2009), and because thegeological history of the islands has been sowell documented (Hall, 2002; Yumul et al.,2009). In this study, we use an estimate ofphylogenetic relationships as a guide fordelimiting species but restrict our diagnosesof new species to those populations diagnosed

TABLE 2.—Models of evolution selected by AkaikeInformation Criterion (AIC) and applied for partitioned,

Bayesian phylogenetic analyses.a

Partition AIC modelNo. of

characters

ATP8, first codon position HKY + G 53ATP8, second codon position HKY + I + G 53ATP8, third codon position GTR + G 53ATP6, first codon position GTR + I 227ATP6, second codon position GTR + I + G 227ATP6, third codon position GTR + G 227

a The model GTR + I + G was used for partitioned RAxMLHPC analyses.


by nonoverlapping morphological characterstates.

RESULTS

Phylogeny

Of 838 aligned mitochondrial nucleotidepositions, 392 and 306 were variable andparsimony-informative, respectively. Just con-sidering the alignment for Brachymeles se-quence data, 310 and 279 were variable andparsimony-informative, respectively. The MLanalysis resulted in a single optimal tree(2lnL 5 5249.903). The resulting topologyfrom the Bayesian analysis is very similar tothe ML tree. Trees estimated from ML, MP,and Bayesian analyses are consistent withrespect to support for nine unique species ofBrachymeles.

No inferences support the monophyly of B.boulengeri. All analyses recover B. b. boho-lensis as part of a clade with B. s. orientalisand B. s. schadenbergi with high support(Fig. 2). Results of both analyses (ML andBayesian) show a polytomy with four lineages,including B. b. mindorensis, B. b. taylori, B. b.boulengeri, and a clade consisting of theremaining species (Fig. 2). All analyses sup-port a clade of B. b. boholensis, B. s.schadenbergi, and B. s. orientalis as sister toa clade of three distinct lineages of B. talinissamples, whereas MP analyses consistentlysupported the clade of B. talinis as sister to aclade of B. b. mindorensis, B. b. taylori, and B.b. boulengeri. With the inclusion of two shortmitochondrial genes for only three of the 18recognized species in the genus, it is likelythat differences in topologies among ML andBayesian analyses reflect limited taxon andcharacter sampling. Nevertheless, all analysesresult in the strong support of nine geneticallydistinct lineages of Brachymeles (Fig. 2). Inaddition, samples of B. talinis cluster intothree major clades, each of which is stronglysupported in all analyses (Fig. 2).

Uncorrected pairwise sequence divergenc-es are low within named taxa and relativelyhigh between these lineages (Table 3). Lev-els of sequence divergence show that thenine mitochondrial DNA lineages discoveredby our phylogenetic analyses (B. b. boholen-sis, B. b. boulengeri, B. sp. nov. [Masbate

Island], B. sp. nov. [Calayan, CamiguinNorte, and Luzon islands], B. b. mindorensis,B. talinis, B. b. taylori, B. s. orientalis, and B.s. schadenbergi) are distinguished fromcongeners by levels of genetic divergenceequal to, or greater than, those betweenpreviously defined species, viz., B. boulen-geri, B. talinis, and B. schadenbergi (Table 3;Fig. 2). The two most genetically similarlineages (B. sp. nov. [Masbate Island] andB. sp. nov. [northern Philippines]) areseparated by 3.8–5.2% sequence divergence,and sequence divergences among all subspe-cies of B. boulengeri, and among bothsubspecies of B. schadenbergi, are .7.3%(Table 3; Fig. 2). The three lineages with thegreatest range of sequence divergence acrosspopulations are the presently defined sub-specific taxa that occur across multipleislands within a single PAIC (B. b. boulen-geri, B. b. taylori, and B. talinis; Table 3;Figs. 2, 3, 5). Sequence divergence amongpopulations in the lineage of northernpopulations of B. talinis, and among popula-tions of B. schadenbergi orientalis, are muchlower than those of other lineages known tobe distributed across multiple islands (Ta-ble 3; Figs. 3–5).

Morphology

Variation in morphological characters (Ta-bles 4–6) mirrors the results observed inphylogenetic analyses and supports the rec-ognition of nine Brachymeles lineages. Inaddition, comparison of meristic and mensuralmorphological characters identified a 10thunique lineage from Jolo Island, previouslyrecognized as a population of B. talinis.Characters differing among these 10 lineagesinclude body size, degree of limb develop-ment, finger and toe lamellae counts, headand body scale counts and patterns, andpigmentation patterns (Tables 4–6; see spe-cies accounts below). We observed no men-sural or meristic differences between thesexes of any of the 10 species.

Superficially, subspecies of both B. boulen-geri and B. schadenbergi, and island popula-tions of B. talinis, seem morphologicallysimilar, especially in overall body size; how-ever, numerous nonoverlapping differenceswere detected in meristic, mensural, and color


pattern characters for each complex member,readily defining 10 distinct lineages betweenthe three complexes (Tables 4–6).

In summary, each lineage possesses uniqueand nonoverlapping suites of diagnostic char-acter states of morphology, perfectly corre-sponding to nine of the clades defined inphylogenetic analyses of DNA sequence data(tissues unavailable for B. cf. talinis from Jolo

Island). Combined with biogeographic evi-dence, and clearly separate geographicalranges, our data suggest the presence of 10evolutionary lineages, worthy of taxonomicrecognition.

Taxonomic Conclusions

Our inferred phylogeny (Fig. 2), biogeogra-phically separate ranges of island endemic

FIG. 2.—Maximum clade credibility tree from a phylogenetic analysis of mitochondrial data (mitochondrial ATPase6[ATP6] and ATP8; 2lnL 5249.903214). Nodes shown with numerical values corresponding to maximum-likelihoodnonparametric bootstrap support, and Bayesian posterior probabilities support values, respectively. Terminals arelabeled with taxonomic names and sampling localities.


species; diagnostic, nonoverlapping morpho-logical character states; and genetic diver-gences between the taxa (Table 3) indicate thedistinctiveness of a new species from Luzon,Calayan, and Camiguin Norte islands, a newspecies from Masbate Island, and a newspecies from Jolo Island. In addition, themolecular and morphological data stronglysupport the elevation of all subspecies of B.boulengeri and B. schadenbergi to full species(Table 3; Fig. 2). Each of the 10 species ismorphologically distinct from each other andall other known species in the genus, and eachof the nine species included in phylogeneticanalyses are also genetically distinct. With theexception of B. talinis, each lineage is endemicto one of four isolated PAICs, therebyproviding additional support for the distinc-tiveness of each lineage’s evolutionary historyand integrity. Accordingly, we recognize allfour subspecies of the former polytypicspecies B. boulengeri, and both subspecies ofthe former polytypic species B. schadenbergi,as full species.

TAXONOMIC ACCOUNTS

Brachymeles boulengeri Taylor 1922b: 246Figs. 3, 6, 11A

Brachymeles boulengeri (part), Taylor, 1922b.Type locality: Polillo Island, Philippines(holotype presumed lost).

Brachymeles gracilis Boulenger (part), Brown,1956; Brown and Rabor, 1967.

Brachymeles gracilis (part), Brown and Alcala,1970.

Brachymeles boulengeri boulengeri (part),Brown and Alcala, 1980.

Designation of a neotype for B. boulen-geri.—Taylor’s holotype for B. boulengeri(Philippine Bureau of Science PublicationNo. 17: 246, collected 15 July 1920) wasdestroyed in the destruction of the PhilippineBureau of Science in World War II, with no

TABLE 3.—Uncorrected pairwise sequence divergence (percent) for mitochondrial data for Brachymeles boholensis, B.boulengeri, B. mindorensis, B. taylori, B. talinis, B. kadwa, B. tungaoi, B. orientalis, and B. schadenbergi (see Fig. 3).

Percentages on the diagonal represent intraspecific genetic diversity (bolded for emphasis).

boholensis boulengeri mindorensis taylori talinis kadwa tungaoi orientalis schadenbergi

boholensis 0.6boulengeri 15.3–15.9 0.0–4.7mindorensis 16.1–16.7 10.6–11.5 0.6taylori 15.6–16.0 10.2–11.6 11.0–11.2 0.3–4.8talinis 13.0–14.6 10.4–11.8 11.8–12.4 10.3–12.4 0.1–4.5kadwa 12.7–13.6 10.4–11.7 11.5–12.1 9.3–10.3 6.9–8.6 0.1–2.2tungaoi 13.2–13.9 10.5–12.1 10.7–11.6 9.7–10.1 7.2–8.3 3.8–5.2 0.0–1.5orientalis 13.4–14.7 14.7–16.5 15.6–16.7 15.5–16.9 14.3–15.8 15.2–16.2 15.3–16.1 0.6–2.1schadenbergi 12.6–13.8 14.5–16.0 16.0–17.3 15.2–16.3 13.8–15.4 15.2–16.5 15.2–16.2 7.3–8.5 0.1–2.6

FIG. 3.—Hypothesized distributions of Brachymelesboholensis, B. boulengeri, B. mindorensis, and B. taylori inthe Philippines. The sampling localities are indicated byblack shapes, and the hypothesized geographic range ofeach species indicated by shaded islands, with shapes andshades of islands corresponding to the map’s key.


mention of a repository for the holotype. Inthe absence of an existing holotype and inaccordance with article No. 75 of the Inter-national Code of Zoological Nomenclature(ICZN, 1999), we designate a neotype for thisspecies. Accordingly, we choose an adult malespecimen from the type locality of PolilloIsland. Preserved adult specimens from Tay-lor’s (1922b) type locality (Polillo Island) havebeen examined in the collections at CaliforniaAcademy of Sciences (CAS); unfortunately,these original specimens are either poorlypreserved, incomplete, or not sexually mature.However, recent collections from PolilloIsland (Fig. 1) have resulted in well-preservedadult individuals that clearly exhibit thediagnostic characters for the species. Fromthese collections, we have chosen a maleneotype collected as part of a series that

contains adults of both sexes and agrees withTaylor’s (1922a) holotype description.

Neotype.—PNM 9720 (RMB Field No.5647, formerly KU 307756), adult male,collected from a fallen, rotting log in second-ary growth forest (1000 to 1230 h) in BarangayPinaglubayan, Municipality of Polillo, QuezonProvince, Polillo Island, Philippines (14u 459090 N, 121u 589 060 E; WGS-84), by RMB, J.Fernandez, Y. Vicente, and M. Vicente.

Diagnosis.—Brachymeles boulengeri can bedistinguished from congeners by the followingcombination of characters: (1) body sizemoderate (SVL 60.5–93.1 mm); (2) pentadac-tyl; (3) Finger-III lamellae five or six; (4) Toe-IV lamellae nine or 10; (5) moderate limblength; (6) supralabials six or seven; (7)infralabials seven; (8) pineal eye spot present;

FIG. 4.—Hypothesized distributions of Brachymelesorientalis, B. schadenbergi, and B. vindumi in thePhilippines. The sampling localities are indicated by blackshapes, and the hypothesized geographic range of eachspecies indicated by shaded islands, with shapes andshades of islands corresponding to the map’s key.Unknown Mindanao Island range boundaries indicatedby dashed lines.

FIG. 5.—Hypothesized distributions of Brachymelestungaoi, B. kadwa, and B. talinis in the Philippines. Thesampling localities are indicated by black shapes, and thehypothesized geographic range of each species indicatedby shaded islands, with shapes and shades of islandscorresponding to the map’s key. Islands of the RomblonIsland group are designated by the first letter of the islandname (T, Tablas Island; R, Romblon Island; and S,Sibuyan Island).


(9) supranasals not contacting on midline; (10)prefrontals not contacting on midline; (11)midline contact of first pair of chin shields;(12) enlarged chin shields in two pairs; (13)nuchal scales undifferentiated; (14) fourth andfifth supralabial below eye; (15) auricularopening present; (16) continuous, light dorso-lateral stripes present; and (17) middorsalstripes absent (Tables 4 and 5).

Comparisons.—Characters distinguishingB. boulengeri from all pentadactyl species ofBrachymeles are summarized in Tables 4 and5. Brachymeles boulengeri most closely re-sembles B. boholensis, B. mindorensis, and B.taylori, but it differs from these three taxa byhaving six or seven supralabials and five or sixsupraciliaries, and by the absence of contin-uous, dark middorsal stripes (Tables 4 and 5).Brachymeles boulengeri further differs fromB. boholensis by having a relatively longer tail,five or six Finger-III lamellae, the fourth andfifth supralabial below the eye, midlinecontact between the first pair of enlargedchin shields, and by lacking a third pair ofenlarged chin shields (Tables 4 and 5); fromB. mindorensis it is differentiated by itssmaller body size, shorter hind limbs, andpossession of nine or 10 Toe-IV lamellae,seven infralabials, and by having the fourthand fifth supralabial below the eye (Tables 4and 5); and from B. taylori by the midlinecontact between the first pair of enlarged chinshields, and the presence of continuous, lightdorsolateral stripes (Tables 4 and 5).

From all nonpentadactyl species of Brachy-meles (B. apus, B. bonitae, B. cebuensis, B.elerae, B. lukbani, B. minimus, B. muntingka-may, B. pathfinderi, B. samarensis, B. tridac-tylus, B. vermis, and B. wrighti), B. boulengeridiffers by having a pentadactyl body form (vs.nonpentadactyl), greater forelimb lengths(.8.2 mm vs. ,6.9 mm), greater hind limblengths (.14.3 mm vs. ,12.9 mm), Toe-IVlamellae nine or 10 (vs. eight or fewer), amidbody scale row count of 26 or 27 (vs. ,24in all nonpentadactyl species except for 28 inB. wrighti; Taylor, 1925), and by the presenceof a postnasal scale (vs. absence). With theexception of B. pathfinderi, B. boulengeridiffers further from all nonpentadactyl speciesby the absence of a third pair of enlarged chinshields (vs. presence) and the presence of

auricular openings (vs. absence). From allnonpentadactyl species except for B. pathfin-deri, B. boulengeri differs by having aparavertebral scale count of 63–66 (vs. .84).From B. apus, B. lukbani, B. minimus, and B.vermis, B. boulengeri is distinguished by thepresence (vs. absence) of limbs.

Description of neotype.—Adult male, hemi-penes everted (Fig. 6); SVL 93.1 mm; bodymoderate relative to other Brachymeles; headweakly differentiated from neck, nearly aswide as body, HW 10.3% SVL, 109.0% HL;HL 36.5% SnFa; SnFa 9.4% SVL; snoutmoderately long, rounded in dorsal andlateral profile, SNL 52.2% HL; auricularopening present, moderate; eyes moderate,ED 2.2% SVL, 23.5% HL, 72.0% END, pupilsubcircular; body slightly depressed, MBW141.8% MBH; body scales smooth, glossy,imbricate; longitudinal scale rows at midbody26; paravertebral scale rows 64; axilla–groinscale rows 43; limbs well developed, penta-dactyl, digits small; FinIIIlam 5; ToeIVlam 9;FLL 19.2% AGD, 12.5% SVL; HLL 28.7%AGD, 18.7% SVL; order of digits fromshortest to longest for hand: V 5 I , IV 5II , III, for foot: I , V , II , III , IV; tailoriginal, not as wide as body, sharply taperedtoward the end, TW 70.1% MBW, TL 84.6%SVL.

Rostral projecting dorsoposteriorly to pointin line with anterior edge of nasal scale,broader than high, forming a narrow suturewith frontonasal; frontonasal wider than long;nostril ovoid, centered in a single rectangularnasal; nasals well separated; supranasals pres-ent, large, moderately separated by frontona-sal; postnasals present; prefrontals narrowlyseparated by frontal; frontal suboctagonal,anterior margin in moderate contact withfrontonasal and first two anterior supraocu-lars, 53 wider than anteriormost supraocular;supraoculars five; frontoparietals moderate, inbroad medial contact, contact 2–4 supraocu-lars; interparietal diamond shaped, slightlywider than long, nearly one half frontal length;parietal eyespot present in posterior one thirdof scale; parietals separated by interparietal;nuchals nonenlarged, undifferentiated fromdorsal scales; loreals two, decreasing in sizefrom anterior to posterior; anterior lorealapproximately as long as and slightly higher


than posterior loreal, in contact with prefron-tal, postnasal, supranasal, second supralabial;posterior loreal and frontonasal; preocularsingle, nearly two thirds the height ofposterior loreal; presubocular single; supraci-liaries six, most anterior contacting prefrontaland separating posterior loreal from firstsupraocular, most posterior extending tomidline of last supraocular; subocular rowcomplete; lower eyelid with one row of scales,lacking an enlarged oval window, largelytransparent; supralabials seven, fourth andfifth below the eye, infralabials seven.

Mental wider than long, in contact with firstinfralabials; postmental single, wider thanmental, followed by two pairs of enlargedchin shields; first pair in broad medial contact,

second pair slightly wider than first, separatedby a single medial scale.

Scales on limbs smaller than body scales;scales on dorsal surfaces of digits large,wrapping around lateral edges of digits; lamel-lae undivided; palmar surfaces of hands andplantar surfaces of feet covered by small,irregular scales, each with raised anterior edges;scales on dorsal surface of hands and feetsmaller than limb scales, lacking raised edges.

Coloration of neotype in preservative.—Ground color of body medium brown; lateraland ventral surfaces of body lacking darkpigment; dorsum, from posterior edge ofsupranasals to tail tip, uniformly dark brown,with color spanning six full and two half rows ofscales at midbody and narrowing to cover four

TABLE 4.—Summary of meristic and mensural characters in all known pentadactyl species of Brachymeles. Sample size,body length, and total length among males and females, and general geographical distribution (PAIC 5 Pleistoceneaggregate island complexes, sensu Brown and Diesmos, 2002) are included for reference (snout–vent length [SVL], totallength [TotL], midbody width [MBW], forelimb length [FLL], and hind limb length [HLL] given as range over mean 6SD; all body proportions given as percentage over mean 6 SD). In cases of scale count variation within species,numbers of individuals showing specific counts are given in parentheses. Other character abbreviations are as follows:

TL 5 tail length; TW 5 tail width; FinIIIlam 5 Finger-III lamellae count; ToeIVlam 5 Toe-IV lamellae count.

Range

boholensis(5 male,

14 female)

boulengeri(7 male,

8 female)

mindorensis(6 male,

12 female)

taylori(8 male,

13 female)gracilis gracilis

(7 male,11 female)

gracilis hilong(8 male,

12 female)

Bohol Island Luzon PAIC Mindoro IslandNegros and

Cebu islands Mindanao PAIC Mindanao PAIC

SVL (female) 83.8–94.0 60.5–95.5 90.0–106.8 65.8–93.2 62.8–75.8 59.9–81.5(88.4 6 3.1) (84.0 6 11.2) (98.8 6 5.3) (83.9 6 7.4) (67.6 6 3.7) (71.8 6 7.0)

SVL (male) 84.1–93.6 72.3–93.1 93.9–104.2 83.1–99.2 60.1–82.3 61.5–78.5(89.1 6 4.1) (82.5 6 6.7) (100.2 6 4.1) (87.0 6 5.2) (67.5 6 7.6) (70.5 6 5.1)

TotL (female) 129.6–174.8 129.7–167.4 162.5–206.7 130.3–168.5 116.4–134.4 94.3–159.2(154.1 6 14.7) (159.3 6 13.1) (180.2 6 14.2) (149.9 6 13.0) (126.3 6 5.5) (126.4 6 17.1)

TotL (male) 154.5–166.2 124.3–173.1 165.3–197.0 149.6–176.7 113.9–133.7 116.7–139.4(160.7 6 5.9) (151.4 6 19.4) (184.9 6 11.5) (164.3 6 11.3) (123.2 6 8.8) (127.4 6 7.8)

MBW 11.9–15.0 9.9–14.7 12.8–20.8 11.0–16.8 8.1–11.6 7.9–12.1(13.4 6 1.0) (12.4 6 1.7) (16.0 6 1.8) (13.8 6 1.7) (9.5 6 0.9) (10.1 6 1.2)

TL/SVL 53–90 67–114 60–99 69–103 69–100 57–98(76 6 13) (89 6 16) (85 6 11) (83 6 10) (88 6 10) (78 6 14)

TW/MBW 62–78 60–81 54–80 54–80 57–79 57–81(70 6 5) (70 6 6) (69 6 8) (69 6 7) (67 6 6) (69 6 7)

FLL 9.0–11.2 8.2–11.7 10.0–13.0 9.0–10.4 5.9–7.9 7.1–9.3(10.1 6 0.7) (10.5 6 0.8) (11.4 6 0.8) (9.8 6 0.4) (6.9 6 0.6) (8.3 6 0.6)

FLL/SVL 10–13 12–14 10–13 10–14 9–12 10–14(11 6 1) (13 6 1) (11 6 1) (12 6 1) (10 6 1) (12 6 1)

HLL 15.4–18.7 14.3–18.7 18.8–23.1 15.6–18.7 10.3–13.6 12.2–16.0(17.2 6 1.0) (17.2 6 1.1) (20.6 6 1.2) (17.0 6 1.0) (12.0 6 0.7) (14.3 6 1.0)

HLL/SVL 18–22 18–24 18–24 18–25 15–20 17–23(19 6 1) (21 6 2) (21 6 2) (20 6 2) (18 6 1) (20 6 1)

FinIIIlam 6 (19) 5 (14) 5 (16) 5 (12) 4 (8) 5 (20)6 (1) 6 (2) 6 (9) 5 (10)

ToeIVlam 9 (7) 9 (13) 8 (14) 9 (10) 7 (4) 8 (13)10 (12) 10 (2) 9 (4) 10 (11) 8 (14) 9 (7)


rows of scales posterior to parietals; darkerpigmentation covers entire surface of dorsalscales, with exception of pigmentation on halfrows of scales; head scales uniform brown;lateral half of supraoculars lacking dark pig-mentation; rostral, nasal, postnasal, supranasal,and first supralabial dark gray with light brownblotches; pineal eyespot charcoal, surroundedby cream border. Faint, indistinct light dorso-lateral stripes, formed by the absence of darkpigmentation, extending from level of anterioredge of eye to base of tail, spanning one full andtwo half rows of scales; small blotch of darkbrown pigment dorsal to auricular opening.Limbs mottled light and medium browndorsally, yellowish brown ventrally; dorsal andventral surface of digits dark brown.

Coloration in life.—Dorsal ground colorhomogeneous medium brown to yellowishbrown (Fig. 11A); gradual lateral dorsolateraldemarcation between dorsal (dark) and ven-tral (light) coloration; lateral and ventralsurfaces of body homogeneous mediumbrown to yellowish brown; dark brown spotsand longitudinal lines of spots absent fromlateral surfaces.

Measurements of neotype (in millimeters).—SVL 93.1; AGD 60.7; TotL 171.9; MBW14.7; MBH 10.4; TL 78.8; TW 10.3; TH8.3; HL 8.8; HW 9.6; HH 7.2; SnFa 24.0; ED2.1; END 2.9; SNL 4.6; IND 3.1; FLL 11.7;HLL 17.5; MBSR 26; PVSR 64; AGSR 43;FinIIIlam 5; ToeIVlam 9; SL 7; IFL 7; SC 6;SO 5.

bicolor(7 male,

6 female)

talinis(11 male,

10 female)

kadwa(12 male,

15 female) tungaoi(2 male,

6 female)

makusog(7 male,

4 female)

orientalis(21 male,

19 female)

schadenbergi(14 male,

10 female)

vindumi(1 male,

1 female)

Luzon Island Visayan PAIC

Luzon, Babuyan,Claro, and Calayan

islands Masbate Island Luzon PAIC Mindanao PAIC Mindanao PAIC Jolo Island

125.9–153.3 103.8–126.7 90.6–135.7 78.2–106.2 98.6–118.0 97.6–115.2 93.1–113.5 104.9(139.4 6 10.1) (116.5 6 6.8) (109.7 6 11.7) (95.6 6 10.7) (108.8 6 8.0) (104.6 6 5.9) (104.8 6 6.7)

120.4–140.3 103.1–123.1 97.0–128.2 89.9, 104.8 82.5–123.5 84.7–112.3 94.4–115.8 113.6(134.0 6 7.6) (113.6 6 7.1) (109.3 6 9.1) (110.9 6 13.7) (99.0 6 8.4) (104.5 6 6.1)

225.3–290.7 187.5–236.2 170.4–221.0 168.9–206.5 183.8–217.3 159.7–213.0 180.8–217.4 —(260.7 6 30.7) (209.4 6 18.0) (196.3 6 18.7) (187.2 6 15.9) (201.5 6 17.7) (194.3 6 17.1) (202.4 6 17.6)

233.4–301.4 191.7–238.4 181.9–255.2 178.3, 203.1 162.3–241.3 154.6–221.1 179.2–219.7 —(280.1 6 24.9) (209.0 6 12.4) (208.3 6 20.3) (201.5 6 25.1) (184.8 6 22.3) (203.0 6 14.4)

13.6–15.3 15.9–20.9 14.3–21.0 11.9–17.0 13.0–19.9 11.0–18.9 13.5–19.1 14.2,14.8(14.4 6 0.6) (18.5 6 1.5) (16.9 6 1.8) (14.4 6 1.6) (17.5 6 2.1) (15.1 6 1.8) (15.2 6 1.7)

62–116 61–107 68–106 82–116 61–117 62–106 65–103 —(97 6 22) (84 6 12) (87 6 12) (97 6 11) (84 6 16) (85 6 12) (92 6 11)

69–84 63–94 66–84 62–80 69–83 56–87 57–77 83, 79(76 6 6) (75 6 6) (75 6 5) (70 6 6) (75 6 4) (70 6 6) (68 6 6)

9.0–12.0 11.3–17.7 10.7–15.0 11.0–13.8 12.8–17.0 10.4–15.6 11.1–13.5 13.2,13.3(10.5 6 1.1) (14.2 6 1.4) (13.0 6 1.2) (12.2 6 0.9) (14.5 6 1.1) (13.4 6 1.5) (12.5 6 0.8)

6.6–8.8 10–15 10–16 11–14 12–17 11–15 10–14 12, 13(7.7 6 0.7) (12 6 1) (12 6 1) (13 6 1) (13 6 1) (13 6 1) (12 6 1)16.1–20.5 20.5–27.9 17.9–24.1 17.0–22.4 19.4–25.3 18.6–25.3 18.5–21.9 22.7,

22.7(17.6 6 1.3) (23.7 6 2.2) (21.8 6 1.6) (20.0 6 1.8) (22.1 6 1.9) (22.1 6 2.2) (20.6 6 1.2)11.3–14.9 18–25 17–26 20–23 18–25 18–24 17–22 20, 22

(13.0 6 1.1) (21 6 2) (20 6 2) (21 6 1) (20 6 2) (22 6 2) (20 6 1)4 (8) 5 (19) 5 (25) 5 (5) 5 (6) 6 (36) 5 (13) 6 (2)5 (5) 6 (2) 6 (2) 6 (3) 6 (5) 7 (4) 6 (11)6 (2) 8 (2) 7 (1) 9 (5) 9 (6) 8 (4) 8 (14) 9 (1)7 (11) 9 (12) 8 (14) 10 (3) 10 (5) 9 (29) 9 (10) 10 (1)

10 (7) 9 (11) 10 (7)10 (1)

TABLE 4.—Extended.


TA

BL

E5.

—S

um

mar

yof

qu

alit

ativ

ed

iagn

osti

cch

arac

ters

(pre

sen

t,ab

sen

t)in

all

know

np

enta

dac

tyl

spec

ies

ofB

rach

ymel

es.

Th

ep

airs

ofen

larg

edsc

ales

pos

teri

orto

the

pos

tmen

tal

scal

ear

eab

bre

viat

edas

chin

shie

ldp

airs

wit

hre

fere

nce

toth

efi

rst,

seco

nd

,an

dth

ird

pai

rs(w

hen

pre

sen

t).

Ch

arac

ter

abb

revi

atio

ns

are

asfo

llow

s:M

BS

R5

mid

bod

ysc

ale-

row

cou

nt;

AG

SR

5ax

illa–

groi

nsc

ale-

row

cou

nt;

SL

5su

pra

lab

ial

cou

nt;

IFL

5in

fral

abia

lco

un

t;S

C5

sup

raci

liary

cou

nt;

SO

5su

pra

ocu

lar

cou

nt.

boho

lens

is(5

mal

e,14

fem

ale)

boul

enge

ri(7

mal

e,8

fem

ale)

min

dore

nsis

(6m

ale,

12fe

mal

e)

tayl

ori

(8m

ale,

13fe

mal

e)

grac

ilis

grac

ilis

(7m

ale,

11fe

mal

e)

grac

ilis

hilo

ng(8

mal

e,12

fem

ale)

bico

lor

(7m

ale,

6fe

mal

e)

talin

is(1

1m

ale,

10fe

mal

e)

kadw

a(1

2m

ale,

15fe

mal

e)

tung

aoi

(2m

ale,

6fe

mal

e)

mak

usog

(7m

ale,

4fe

mal

e)

orie

ntal

is(2

1m

ale,

19fe

mal

e)

scha

denb

ergi

(14

mal

e,10

fem

ale)

vind

umi

(1m

ale,

1fe

mal

e)

MB

SR

26–2

826

–27

26–2

826

–28

24–2

627

–30

27–2

926

–30

26–2

826

–28

25–2

826

–28

26–2

830

,31

AG

SR

42–4

642

–46

42–4

542

–47

46–4

944

–50

64–7

243

–48

47–4

946

–49

42–4

746

–49

45–5

049

,49

PV

SR

63–6

663

–66

63–6

562

–69

67–7

066

–73

88–9

267

–72

68–7

066

–68

60–6

969

–72

67–7

274

,74

SL

7(1

9)6

(12)

7(1

8)6

(21)

6(1

8)6

(20)

6(1

3)7

(20)

7(2

7)7

(4)

6(7

)6

(31)

6(2

)7

(2)

7(3

)7

(4)

7(9

)7

(22)

IFL

7(1

9)7

(15)

6(1

8)7

(21)

7(1

8)6

(20)

6(1

3)7

(21)

6(2

7)6

(4)

6(1

0)6

(7)

6(1

0)6

(2)

SC

6(1

9)5

(1)

6(1

8)6

(21)

6(1

8)6

(20)

6(1

3)7

(1)

7(3

3)7

(14)

6(1

4)6

(21)

6(2

7)6

(4)

6(1

1)6

(40)

6(2

4)6

(2)

SO

5(19

)5

(15)

5(1

8)5

(21)

5(1

8)5

(20)

5(1

3)5

(21)

5(2

7)5

(4)

5(1

1)5

(40)

5(2

4)5

(2)

Pin

eal

eyes

pot

++

++

2+

++

++

++

++

Su

pra

nas

alco

nta

ct2

22

or+

22

2+

++

+2

2+

+P

refr

onta

lco

nta

ct2

22

or+

2+

22

22

22

22

2F

ron

top

arie

tal

con

tact

++

++

+2

or+

++

2or

++

++

++

Par

ieta

lsco

nta

ct2

or+

2or

+2

or+

2or

++

++

2or

+2

or+

2or

+2

2or

+2

or+

+F

irst

chin

shie

ldp

air

con

tact

2or

++

+2

or+

22

or+

+2

or+

2or

++

2or

+2

or+

2or

+2

En

larg

edch

insh

ield

pai

rs3

22

23

32

22

22

22

2

Ch

insh

ield

pai

rsi

ze3

,1

,2

1,

21

,2

1,

23

,1

,2

3,

1,

22

,1

2,

11

,2

15

22

,1

2,

12

,1

2,

1C

hin

shie

ldp

air

sep

arat

ion

a1(

0/1)

;2(

1);

3(3)

1(0/

1);

2(1)

1(0)

; 2(1)

1(0)

; 2(1)

1(1)

; 2(1)

;3(

3)

1(0/

1);

2(1)

;3(

3/5)

1(0)

; 2(3)

1(0/

1);

2(3)

1(0/

1);

2(1)

1(0)

; 2(1)

1(1)

;2(

3)1(

0/1)

;2(

3)1(

0); 2(3)

1(1)

; 2(1)

Su

boc

ula

rsu

pra

lab

ial

Fif

than

dsi

xth

Fou

rth

and

fift

h

Fif

than

dsi

xth

Fou

rth

and

fift

h

Fou

rth

and

fift

h

Fou

rth

and

fift

h

Fou

rth

and

fift

h

Fif

th and

sixt

h

Fif

th and

sixt

h

Fif

th and

sixt

h

Fou

rth

and

fift

hor

fift

han

dsi

xth

Fou

rth

and

fift

h

Fif

th and

sixt

h

Fif

th and

sixt

h

Dif

fere

nti

ated

nu

chal

s2

22

22

2+

22

22

22

2U

nif

orm

bod

yco

lor

22

22

++

22

22

22

22

Con

tin

uou

s,lig

ht

dor

sola

tera

lst

rip

es+

++

2+

+2

2or

++,

poo

rly

def

ined

+,p

oorl

yd

efin

ed2

22

+

Con

tin

uou

sd

ark

mid

dor

sal

stri

pes

b+

2+

++

2or

+2

2or

++

+2

22

or+

2

Dar

kla

tera

lst

rip

esb

++

++

++

2+

++

22

2or

++

Dar

kve

ntr

alp

igm

enta

tion

+2

22

++

22

+2

22

2+

aP

aren

thes

essh

owth

en

um

ber

ofsm

all

ven

tral

scal

ero

ws

sep

arat

ing

each

enla

rged

pai

rof

chin

shie

lds.

bC

har

acte

rre

fers

tolo

ngi

tud

inal

row

sof

dar

ksc

ale

pig

men

tati

on,

ofte

nin

the

form

ofsp

ots,

alig

ned

into

row

s.


Variation.—Variation in mensural charac-ters is summarized in Table 6. Among the 19specimens examined for the degree of contactbetween parietal scales, nine specimens pos-sessed parietals separated by the interparietal(KU 307750, 307758, 320060, 322315–20),and 10 specimens possessed parietals inmoderate-to-broad medial contact (KU307438–9, 307751–4, 307757, 320058–9,322314) behind the interparietal.

Scale counts were observed to vary amongthe measured series. With the exception of asingle specimen with five supraciliaries (KU307752), all specimens examined had sixsupraciliaries. The number of supralabialsvaried between six (CAS 61297, 62272–3,62276–7, KU 307438–9, 307752–4, 307757–8)and seven (CAS 61096, KU 307751). Speci-mens were observed to have midbody scalerow counts of 26 (CAS 61096, 31297, 62273,62276–7, KU 307439, 307751–4, 307758) and27 (CAS 62272, KU 307750, 307757); axilla–groin scale row counts of 42 (KU 307439,307758), 43 (KU 307750–4, 307757), 44 (CAS61096, 62272–3, 62276–7), and 46 (CAS61297); and paravertebral scale row countsof 63 (KU 307439, 307750, 307752, 307754,307757–8), 64 (CAS 61096, 62272–3, 62277,KU 307751, 307753), 65 (CAS 62276), and 66(CAS 61297).

We also observed lamellae counts to varyamong the measured series. With the excep-tion of a single specimen with six Finger-IIIlamellae (KU 307750), all specimens examinedhad five. Two specimens examined had 10 Toe-IV lamellae (KU 307750, 307757), whereas theremaining specimens examined had nine.

Distribution.—Brachymeles boulengeri oc-curs in central and southern Luzon and onMarinduque and Polillo islands (Fig. 3). Thespecies has been collected in the CamarinesNorte Province of the Bicol Peninsula andmay eventually be found to occur furthersouth on the Bicol Peninsula and even onCatanduanes Island.

Ecology and natural history.—Brachymelesboulengeri occurs in disturbed and secondarygrowth forest. Little or no original, low-elevation forest remains throughout the rangeof B. boulengeri, but we assume the speciesonce also occurred in first-growth forest whenthis forest type extended into low-elevation

areas. Individuals have been observed underpiles of rotting coconut husks, in the humusmaterial within rotting logs, and in loose soiland leaf litter surrounding the root networksof trees. This species is quite common at thetype locality, which has been virtually com-pletely converted to coconut plantations.When disturbed, individuals immediatelymove in a rapid serpentine manner, attempt-ing to burrow into loose soil or humus.

Sympatric lizard species observed on Lu-zon, Polillo, and Marinduque islands includethe following: (Agamidae) Bronchocela crista-tella, Draco spilopterus, Gonocephalus so-phiae, Hydrosaurus pustulatus; (Gekkonidae)Cyrtodactylus philippinicus, Gehyra mutilata,Gekko gecko, Gekko mindorensis, Hemidacty-lus frenatus, H. garnoti, H. luzonensis, H.platyurus, Pseudogekko compressicorpus, P.smaragdina; (Scincidae) B. bonitae, B. bicolor,B. elerae, B. lukbani, B. makusog, B. mun-tingkamay, B. samarensis, B. kadwa, B.wrighti, Emoia atrocostata, Eutropis bonto-censis, E. multicarinata, E. multifasciata,Lamprolepis smaragdina, Lipinia pulchella,Sphenomorphus cumingi, S. decipiens, S.jagori, S. leucospilos, S. luzonensis, S. steerei,S. stejnegeri, Tropidophorus grayi; and (Var-anidae) Varanus marmoratus and V. olivaceus.

Brachymeles boholensis Brown and Rabor1967

Figs. 3, 6, 11C

Brachymeles gracilis boholensis, Brown andRabor, 1967. Type locality: 6 km southeastof Sierra Bullones, Teacher’s Park, BoholIsland, Philippines, 400–433-m elevation,9u 479 32.530 N, 124u 189 14.40 E (holotype:CAS-SU24528).


Brachymeles boulengeri boholensis (part),Brown and Alcala, 1980.Diagnosis.—Brachymeles boholensis can be

distinguished from congeners by the followingcombination of characters: (1) body sizemoderate (SVL 83.8–93.6 mm); (2) limbspentadactyl; (3) Finger-III lamellae six; (4)Toe-IV lamellae nine or 10; (5) limb lengthmoderate; (6) supralabials seven; (7) infra-labials seven; (8) pineal eye spot present; (9)supranasals not contacting on midline; (10)


TA

BL

E6.

—S

um

mar

yof

un

ivar

iate

mor

ph

olog

ical

vari

atio

nam

ong

men

sura

lch

arac

ters

inse

ries

ofB

rach

ymel

esbo

hole

nsis

,B.b

oule

nger

i,B

.min

dore

nsis

,B.t

aylo

ri,B

.tal

inis

,B

.kad

wa,

B.t

unga

oi,B

.ori

enta

lis,B

.sch

aden

berg

i,an

dB

.vin

dum

i.C

har

acte

rab

bre

viat

ion

sar

eas

follo

ws:

SV

L5

snou

t–ve

nt

len

gth

;AG

D5

axill

a–gr

oin

dis

tan

ce;

Tot

L5

tota

lle

ngt

h;

MB

W5

mid

bod

yw

idth

;M

BH

5m

idb

ody

hei

ght;

TL

5ta

ille

ngt

h;

TW

5ta

ilw

idth

;T

H5

tail

hei

ght;

HL

5h

ead

len

gth

;H

W5

hea

dw

idth

;H

H5

hea

dh

eigh

t;S

nF

a5

snou

t–fo

rear

mle

ngt

h;

ED

5ey

ed

iam

eter

;E

ND

5ey

e–n

aria

ld

ista

nce

;S

NL

5sn

out

len

gth

;IN

D5

inte

rnar

ial

dis

tan

ce;

FL

L5

fore

limb

len

gth

;H

LL

5h

ind

limb

len

gth

.

boho

lens

is(5

mal

e;14

fem

ale)

boul

enge

ri(7

mal

e;8

fem

ale)

min

dore

nsis

(6m

ale;

12fe

mal

e)

tayl

ori

(8m

ale;

13fe

mal

e)

talin

is(1

1m

ale;

10fe

mal

e)

kadw

a(1

2m

ale;

15fe

mal

e)

tung

aoi

(2m

ale;

6fe

mal

e)

orie

ntal

is(2

1m

ale;

19fe

mal

e)

scha

denb

ergi

(14

mal

e;10

fem

ale)

vind

umi

(1m

ale;

1fe

mal

e)

SV

L(m

ale)

84.1

–93.

672

.3–9

3.1

93.9

–104

.283

.1–9

9.2

103.

1–12

3.1

97.0

–128

.289

.2,

104.

884

.7–1

12.3

94.4

–115

.811

3.6

(89.

16

4.1)

(82.

56

6.7)

(100

.26

4.1)

(87.

06

5.2)

(113

.66

7.1)

(109

.36

9.1)

(99.

06

8.4)

(104

.56

6.1)

SV

L(f

emal

e)83

.8–9

4.0

60.5

–95.

590

.0–1

06.8

65.8

–93.

210

3.8–

126.

790

.6–1

35.7

78.2

–106

.297

.6–1

15.2

93.1

–113

.510

4.9

(88.

46

3.1)

(84.

06

11.2

)(9

8.8

65.

3)(8

3.9

67.

4)(1

16.5

66.

8)(1

09.7

611

.7)

(95.

66

10.7

)(1

04.6

65.

9)(1

04.8

66.

7)A

GD

(mal

e)54

.7–6

1.9

46.3

–60.

759

.5–6

7.6

52.8

–66.

064

.2–9

0.3

54.1

–84.

556

.8,

69.3

51.0

–75.

360

.3–7

6.8

72.7

(58.

16

2.8)

(53.

06

5.0)

(63.

36

3.3)

(55.

96

4.2)

(74.

66

7.5)

(70.

46

7.3)

(62.

76

7.0)

(67.

76

4.9)

AG

D (fem

ale)

53.0

–61.

236

.2–6

1.3

56.3

–74.

946

.6–6

0.6

66.5

–81.

448

.0–9

3.7

49.8

–69.

657

.9–7

5.9

64.8

–92.

367

.4(5

7.0

62.

7)(5

4.3

68.

3)(6

4.8

65.

5)(5

4.7

64.

6)(7

4.7

64.

9)(7

1.8

610

.7)

(61.

96

7.7)

(66.

66

4.8)

(72.

06

7.7)

Tot

L(m

ale)

154.

5–16

6.2

124.

3–17

3.1

165.

3–19

7.0

149.

6–17

6.7

191.

7–23

8.4

181.

9–25

5.2

178.

3,20

3.1

154.

6–22

1.1

179.

2–21

9.7

—(1

60.7

65.

9)(1

51.4

619

.4)

(184

.96

11.5

)(1

64.3

611

.3)

(209

.06

12.4

)(2

08.3

620

.2)

(184

.86

22.3

)(2

03.0

614

.4)

Tot

L (fem

ale)

129.

6–17

4.8

129.

7–16

7.4

162.

5–20

6.7

130.

3–16

8.5

187.

5–23

6.2

170.

4–22

1.0

168.

9–20

6.5

159.

7–21

3.0

180.

8–21

7.4

—(1

54.1

614

.7)

(159

.36

13.1

)(1

80.2

614

.2)

(149

.96

13.0

)(2

09.4

618

.0)

(196

.36

18.7

)(1

87.2

615

.9)

(194

.36

17.1

)(2

02.4

617

.6)

MB

W(m

ale)

12.2

–13.

910

.1–1

4.7

12.8

–16.

011

.6–1

6.8

15.9

–20.

114

.3–1

9.6

13.9

,15

.211

.0–1

7.7

13.5

–17.

214

.2(1

3.0

60.

7)(1

2.1

61.

9)(1

4.7

61.

1)(1

3.3

61.

7)(1

7.7

61.

3)(1

6.5

61.

6)(1

4.4

61.

6)(1

4.8

61.

3)M

BW (fem

ale)

11.9

–15.

09.

9–14

.614

.5–2

0.8

11.0

–16.

617

.3–2

0.9

14.3

–21.

011

.9–1

7.0

12.3

–18.

913

.9–1

9.1

14.8

(13.

66

1.1)

(12.

76

1.7)

(16.

76

1.8)

(14.

06

1.8)

(19.

36

1.1)

(17.

26

2.0

(14.

46

1.9)

(15.

86

1.8)

(15.

96

2.4)

MB

H(m

ale)

11.7

–12.

57.

9–10

.410

.2–1

2.9

9.4–

14.6

10.4

–15.

99.

2–14

.98.

6,14

.69.

6–15

.212

.1–1

4.8

11.6

(12.

16

0.4)

(9.0

61.

0)(1

2.0

61.

0)(1

1.8

61.

7)(1

3.5

61.

7)(1

1.5

61.

6)(1

2.6

61.

3)(1

3.3

60.

9)M

BH (fem

ale)

10.4

–14.

26.

2–10

.211

.4–1

6.6

10.4

–15.

212

.7–1

8.6

8.8–

14.7

7.7–

13.1

10.8

–16.

710

.2–1

6.2

11.5

(11.

86

1.0)

(8.9

61.

3)(1

4.2

61.

8)(1

2.2

61.

5)(1

6.2

62.

1)(1

1.1

61.

7)(1

0.9

62.

4)(1

3.9

61.

6)(1

3.0

62.

8)T

L(m

ale)

70.4

–74.

852

.0–8

8.6

62.2

–94.

263

.0–8

5.7

86.8

–115

.376

.1–1

26.9

89.1

,98

.363

.8–1

08.8

70.3

–112

.5—

(72.

96

2.3)

(69.

26

14.7

)(8

4.6

612

.0)

(76.

46

9.2)

(96.

16

8.5)

(99.

06

14.4

)(8

5.4

615

.1)

(99.

56

11.3

)T

L(f

emal

e)45

.3–8

2.3

69.0

–86.

365

.5–9

9.9

55.4

–76.

170

.7–1

15.2

70.5

–109

.279

.5–1

05.4

61.2

–106

.087

.7–1

07.9

—(6

5.6

613

.0)

(76.

06

6.7)

(82.

66

11.6

)(6

7.0

67.

3)(9

4.2

617

.6)

(88.

96

13.1

)(9

1.6

69.

2)(8

9.2

613

.7)

(97.

96

9.0)

TW

(mal

e)8.

5–10

.07.

2–10

.310

.2–1

1.9

7.4–

12.2

11.8

–15.

911

.0–1

5.1

8.8,

12.1

8.6–

11.1

9.6–

11.1

11.8

(9.4

60.

5)(8

.76

1.0)

(10.

96

0.6)

(9.5

61.

4)(1

3.5

61.

2)(1

3.0

61.

3)(1

0.0

60.

8)(1

0.6

60.

6)T

W(f

emal

e)8.

4–10

.47.

0–9.

910

.0–1

2.5

7.4–

12.1

12.9

–15.

610

.3–1

5.7

7.3–

12.5

9.2–

12.5

9.0–

11.0

11.6

(9.4

60.

5)(8

.66

1.2)

(11.

06

0.7)

(9.4

61.

3)(1

4.2

60.

9)(1

2.4

61.

5)(1

0.0

61.

7)(1

0.9

60.

9)(9

.96

0.9)

TH

(mal

e)7.

6–8.

15.

4–8.

38.

0–9.

56.

3–10

.09.

7–11

.48.

7–12

.18.

0,10

.57.

6–9.

97.

7–9.

58.

4(7

.86

0.2)

(6.8

60.

9)(8

.56

0.6)

(8.0

61.

0)(1

0.7

60.

6)(1

0.2

61.

0)(8

.56

0.7)

(8.6

60.

7)T

H(f

emal

e)6.

9–8.

55.

1–8.

07.

3–10

.06.

9–10

.310

.6–1

3.2

8.2–

12.7

6.0–

10.1

7.3–

10.5

7.8–

9.7

9.3

(7.6

60.

5)(6

.56

0.9)

(8.5

60.

7)(8

.06

1.0)

(11.

76

0.9)

(9.9

61.

3)(8

.36

1.4)

(9.1

60.

8)(8

.76

0.8)

HL

(mal

e)8.

3–9.

07.

3–8.

98.

6–10

.58.

4–9.

09.

2–12

.28.

6–12

.18.

7,10

.19.

3–11

.59.

8–11

.59.

6(8

.76

0.4)

(8.3

60.

7)(9

.66

0.7)

(8.7

60.

2)(1

0.7

60.

9)(1

0.3

61.

0)(1

1.2

60.

6)(1

0.7

60.

7)


boho

lens

is(5

mal

e;14

fem

ale)

boul

enge

ri(7

mal

e;8

fem

ale)

min

dore

nsis

(6m

ale;

12fe

mal

e)

tayl

ori

(8m

ale;

13fe

mal

e)

talin

is(1

1m

ale;

10fe

mal

e)

kadw

a(1

2m

ale;

15fe

mal

e)

tung

aoi

(2m

ale;

6fe

mal

e)

orie

ntal

is(2

1m

ale;

19fe

mal

e)

scha

denb

ergi

(14

mal

e;10

fem

ale)

vind

umi

(1m

ale;

1fe

mal

e)

HL

(fem

ale)

8.6–

9.8

7.9–

9.2

8.9–

10.9

7.6–

9.9

9.9–

12.6

8.9–

11.4

7.7–

9.7

9.0–

13.8

9.4–

11.9

9.8

(9.0

60.

4)(8

.36

0.5)

(9.9

60.

7)(8

.76

0.8)

(11.

36

0.9)

(9.8

60.

8)(8

.86

0.7)

(10.

96

1.0)

(10.

66

1.0)

HW

(mal

e)9.

3–10

.99.

1–10

.310

.3–1

2.1

9.6–

11.6

11.9

–14.

511

.2–1

4.3

10.1

,11

.89.

9–12

.010

.7–1

3.2

10.8

(10.

26

0.6)

(9.5

60.

5)(1

1.2

60.

6)(1

0.4

60.

8)(1

3.2

60.

8)(1

2.4

61.

0)(1

1.1

60.

6)(1

1.8

60.

9)H

W(f

emal

e)9.

3–10

.57.

5–9.

910

.7–1

2.4

9.3–

12.0

12.8

–15.

810

.4–1

3.3

9.1–

12.6

10.6

–13.

79.

5–12

.310

.6(1

0.0

60.

4)(9

.16

0.8)

(11.

46

0.5)

(10.

56

0.9)

(14.

06

0.9)

(11.

66

1.0)

(10.

86

1.4)

(11.

76

0.7)

(11.

46

1.3)

HH

(mal

e)7.

2–7.

96.

2–7.

58.

2–10

.17.

3–8.

88.

8–12

.77.

9–10

.97.

2,10

.46.

9–9.

67.

6–10

.17.

9(7

.76

0.3)

(6.9

60.

4)(9

.06

0.6)

(7.9

60.

5)(9

.96

1.1)

(9.3

60.

9)(8

.46

0.6)

(8.8

60.

8)H

H(f

emal

e)7.

2–8.

05.

7–7.

28.

1–9.

97.

3–9.

79.

3–12

.57.

3–10

.06.

2–10

.47.

9–10

.87.

9–9.

27.

8(7

.56

0.3)

(6.6

60.

5)(8

.96

0.5)

(8.1

60.

7)(1

0.7

61.

0)(8

.46

0.9)

(8.3

61.

6)(9

.26

0.9)

(8.7

60.

6)S

nF

a(m

ale)

22.9

–25.

119

.1–2

4.0

25.9

–28.

120

.7–2

3.3

26.2

–34.

126

.5–3

2.7

22.8

,26

.223

.9–3

1.1

25.8

–29.

928

.5(2

3.7

61.

0)(2

2.4

61.

7)(2

7.0

60.

7)(2

2.1

61.

0)(3

0.6

62.

3)(2

8.9

61.

7)(2

7.4

62.

1)(2

7.9

61.

7)S

nF

a (fem

ale)

21.3

–24.

217

.9–2

4.4

24.3

–27.

520

.8–2

5.1

29.1

–34.

725

.4–3

1.9

20.9

–28.

625

.6–3

2.3

24.8

–30.

326

.8(2

2.7

60.

8)(2

2.7

62.

1)(2

5.9

61.

0)(2

2.2

61.

1)(3

1.4

61.

9)(2

8.2

61.

9)(2

5.0

62.

8)(2

8.8

62.

0)(2

7.9

62.

3)E

D(m

ale)

1.7–

2.1

1.8–

2.3

2.0–

2.5

1.4–

2.0

2.0–

2.3

1.6–

2.3

1.7,

1.8

1.6–

2.0

1.8–

2.3

2.3

(1.8

60.

2)(2

.06

0.2)

(2.2

60.

2)(1

.86

0.2)

(2.2

60.

1)(1

.96

0.2)

(1.8

61.

0)(2

.16

0.2)

ED

(fem

ale)

1.6–

2.0

1.5–

2.3

2.0–

2.6

1.5–

2.1

2.0–

2.5

1.6–

2.2

1.3–

1.9

1.6–

2.5

2.0–

2.4

2.0

(1.8

60.

1)(1

.96

0.3)

(2.2

60.

2)(1

.86

0.2)

(2.2

60.

1)(1

.96

0.2)

(1.7

60.

2)(1

.96

0.2)

(2.2

60.

2)E

ND

(mal

e)2.

4–2.

62.

7–3.

02.

5–2.

92.

8–3.

43.

7–5.

13.

7–4.

33.

3,3.

83.

5–4.

54.

0–4.

23.

9(3

.56

0.1)

(2.8

60.

1)(2

.76

0.2)

(3.1

60.

2)(4

.36

0.4)

(3.9

60.

2)(4

.06

0.3)

(4.1

60.

1)E

ND (fem

ale)

3.0–

3.7

2.1–

3.0

3.4–

4.2

2.7–

3.5

3.7–

5.3

3.5–

4.3

2.5–

3.7

3.7–

5.0

3.4–

4.3

3.7

(3.4

60.

2)(2

.86

0.3)

(3.6

60.

2)(3

.06

0.2)

(4.5

60.

4)(3

.86

0.3)

(3.2

60.

4)(4

.26

0.3)

(4.0

60.

4)S

NL

(mal

e)5.

0–5.

34.

2–4.

65.

4–6.

04.

1–4.

85.

7–7.

55.

3–6.

85.

2,5.

45.

1–6.

55.

3–6.

15.

7(5

.26

0.1)

(4.4

60.

1)(5

.76

0.2)

(4.4

60.

3)(6

.46

0.6)

(6.0

60.

4)(5

.96

0.4)

(5.8

60.

3)S

NL

(fem

ale)

4.8–

5.6

3.4–

4.8

5.0–

6.4

4.1–

5.0

5.5–

7.5

3.1–

6.5

4.9–

5.7

5.4–

7.1

5.2–

6.2

5.6

(5.3

60.

2)(4

.26

0.4)

(5.5

60.

4)(4

.66

0.3)

(6.6

60.

5)(5

.66

0.8)

(5.3

60.

4)(6

.26

0.4)

(5.9

60.

5)IN

D(m

ale)

3.0–

3.2

2.8–

3.1

3.2–

3.6

2.7–

3.2

3.3–

4.3

3.2–

4.2

3.0,

3.4

3.0–

3.8

3.0–

3.6

3.4

(3.1

60.

1)(2

.96

0.1)

(3.4

60.

1)(2

.96

0.2)

(3.9

60.

3)(3

.76

0.3)

(3.3

60.

2)(3

.46

0.2)

IND

(fem

ale)

3.0–

3.3

2.3–

3.2

3.1–

3.8

2.7–

3.2

3.6–

4.4

3.3–

4.1

2.7–

3.8

3.2–

3.9

2.8–

3.6

3.6

(3.1

60.

1)(2

.86

0.3)

(3.4

60.

2)(2

.96

0.2)

(3.9

60.

2)(3

.66

0.2)

(3.2

60.

4)(3

.56

0.3)

(3.3

60.

4)F

LL

(mal

e)9.

6–11

.110

.0–1

1.7

10.7

–12.

89.

3–10

.411

.3–1

7.7

10.9

–14.

812

.5,

12.8

10.4

–15.

611

.8–1

3.4

13.2

(10.

56

0.7)

(10.

56

0.6)

(11.

46

0.7)

(9.9

60.

4)(1

3.9

61.

8)(1

3.1

61.

2)(1

2.8

61.

6)(1

2.6

60.

6)F

LL

(fem

ale)

9.0–

11.2

8.2–

11.3

10.0

–13.

09.

0–10

.313

.5–1

8.9

10.7

–15.

011

.0–1

3.8

11.6

–15.

611

.1–1

3.5

13.3

(9.9

60.

7)(1

0.4

61.

0)(1

1.4

60.

9)(9

.76

0.4)

(14.

66

0.9)

(12.

96

1.3)

(12.

06

1.0)

(13.

96

1.2)

(12.

26

1.1)

HL

L(m

ale)

16.2

–17.

516

.6–1

8.5

20.1

–22.

616

.1–1

7.9

20.5

–27.

920

.5–2

3.7

20.1

,21

.018

.6–2

4.3

19.4

–21.

922

.7(1

6.8

60.

5)(1

7.4

60.

6)(2

0.8

60.

9)(1

6.9

60.

7)(2

3.0

62.

4)(2

2.2

61.

1)(2

1.2

62.

0)(2

0.9

60.

9)H

LL (fem

ale)

15.4

–18.

714

.3–1

8.7

18.8

–23.

115

.6–1

8.7

20.5

–26.

117

.9–2

4.1

17.0

–22.

418

.7–2

5.3

18.5

–21.

922

.7(1

7.3–

1.2)

(17.

06

1.4)

(20.

66

1.3)

(17.

16

1.1)

(24.

46

1.7)

(21.

56

1.8)

(19.

96

2.1)

(22.

86

2.1)

(20.

06

1.6)

TA

BL

E6.

—C

onti

nu

ed.


prefrontals not contacting on midline; (11)enlarged chin shields in three pairs; (12) nuchalscales undifferentiated; (13) fifth and sixthsupralabial below eye; (14) auricular openingpresent; (15) continuous, light, dorsolateralstripes present; and (16) continuous, darkmiddorsal stripes present (Tables 4 and 5).

Comparisons.—Characters distinguishingB. boholensis from all pentadactyl species ofBrachymeles are summarized in Tables 4 and5. Brachymeles boholensis most closely re-sembles B. boulengeri, B. mindorensis, and B.taylori, but it differs from them by having sixlamellae on Finger-III and by the presence of

FIG. 6.—Illustration of head of adult female Brachymeles boholensis (KU 323972) and adult male neotype ofBrachymeles boulengeri (PNM 9720; formerly KU 307756) in dorsal, lateral, and ventral views. Taxonomically diagnostichead scales are labeled as follows: C, chin shield; F, frontal; FN, frontonasal; FP, frontoparietal; IL, infralabial; IP,interparietal; L, loreal; M, mental; N, nasal; P, parietal; PF, prefrontal; PM, postmental; PN, postnasal; PO, preocular;PSO, presubocular; R, rostral; SC, supraciliary; SL, supralabial; SN, supranasal; and SO, supraocular. Roman numeralsindicate scales in the supraocular series, with Arabic numbers indicating scales in the supraciliary series. Illustrationsby CDS.


three pairs of enlarged chin shields (Tables 4and 5). Brachymeles boholensis differs furtherfrom B. boulengeri by having seven suprala-bials, six supraciliaries, and the fifth and sixthsupralabial below the eye, and by the presenceof continuous, dark middorsal stripes (Ta-bles 4 and 5); from B. mindorensis by having asmaller body size, shorter hind limbs, nine or10 Toe-IV lamellae, seven infralabials, and bythe absence of contact between supranasalsand the absence of contact between prefron-tals (Tables 4 and 5); and from B. taylori byhaving seven supralabials, the fifth and sixthsupralabial below the eye, and the presence ofcontinuous, light dorsolateral stripes (Tables 4and 5).

From all nonpentadactyl species of Brachy-meles (B. apus, B. bonitae, B. cebuensis, B.elerae, B. lukbani, B. minimus, B. muntingka-may, B. pathfinderi, B. samarensis, B. tridac-tylus, B. vermis, and B. wrighti), B. boholensisdiffers by having pentadactyl (vs. nonpenta-dactyl) limbs, greater forelimb lengths(.9.0 mm vs. ,6.9 mm), greater hind limblengths (.15.4 mm vs. ,12.9 mm), Toe-IVlamellae nine or 10 (vs. eight or fewer), fifthand sixth supralabial below the eye (vs. fourthand fifth), and by presence of a postnasal scale(vs. absence). In addition, B. boholensis differsfrom all nonpentadactyl species except B.wrighti by having 26–28 midbody scale rows(vs. ,24); from all nonpentadactyl speciesexcept B. pathfinderi by having 63–66 para-vertebrals (vs. .84) and by the presence ofauricular openings (vs. absence); and from B.apus, B. lukbani, B. minimus, and B. vermisby the presence of limbs (vs. absence).

Description (based on holotype and 38referred specimens, including 12 paratypesat CAS).—Details of the head scalation of anadult female are shown in Fig. 6. Measure-ments of the holotype are provided below inbrackets. Body moderate relative to otherBrachymeles; maximum SVL 93.6 mm formales, 94.0 mm for females [89.5, female](Tables 4 and 5); head weakly differentiatedfrom neck, nearly as wide as body, HW 10.0–12.4% (11.3 6 0.7) SVL [11.8], 104.2–131.9%(112.6 6 6.7) HL [110.7]; HL 35.2–43.6%(38.9 6 2.3) SnFa [39.3]; SnFa 24.1–28.0%(26.0 6 1.0) SVL [27.0]; snout moderatelylong, broadly rounded in dorsal profile,

rounded in lateral profile, SNL 54.6–64.6%(59.3 6 2.8) HL [57.4]; auricular openingpresent, moderate; eyes small, ED 1.8–2.2%(2.0 6 0.1) SVL [2.1], 17.6–23.9% (20.3 61.6) HL [19.5], 45.1–61.8% (52.5 6 3.7) END[52.9], pupil subcircular; body slightly de-pressed, MBW 98.1–136.2% (113.0 6 10.3)MBH [131.0]; scales smooth, glossy, imbri-cate; longitudinal scale rows at midbody 26–28[28]; paravertebral scale rows 63–66 [64];axilla–groin scale rows 42–46 [44]; limbs welldeveloped, pentadactyl, digits small; FinIII-lam 6 [6]; ToeIVlam 7–10 [7]; FLL 15.6–20.5% (17.6 6 1.4) AGD [20.5], 10.2–12.9%(11.4 6 0.8) SVL [12.4]; HLL 27.6–34.6%(30.0 6 2.3) AGD [32.9], 17.8–21.5% (19.4 61.2) SVL [20.0]; order of digits from shortestto longest for hand: I 5 V , II 5 III 5 IV, forfoot: V , I , II , III 5 IV; tail not as wide asbody, sharply tapered toward the end, TW61.6–78.4% (70.3 6 5.0) MBW [63.5], TL52.7–90.0% (75.7 6 13.0) SVL [86.8].

Rostral projecting dorsoposteriorly to pointin line with center of nasal; broader than high,in contact with frontonasal; frontonasal widerthan long; nostril ovoid, centered in a singlerectangular nasal; nasals well separated; supra-nasals present, large, moderately separated byfrontonasal; postnasals present; prefrontalsbroadly separated by frontal; frontal nearlydiamond shaped, in moderate contact withfrontonasal, first two anterior supraoculars, 43wider than anterior supraocular; supraocularsfive; frontoparietals moderate in size, in contact,each frontoparietal in contact with posteriorthree or four supraoculars; interparietal mod-erate in size, quadrilaterally shaped, longer thanwide, its length slightly greater than midlinelength of frontoparietals; parietal eyespot pres-ent in posterior half of scale; parietals separatedby interparietal; nuchals undifferentiated; lor-eals two, anterior loreal largest, in contact withprefrontal, postnasal, supranasal, second supra-labial, posterior loreal and frontonasal; preocu-lar single, nearly two thirds as high as posteriorloreal; supraciliaries six, anterior supraciliarycontacting prefrontal and separating posteriorloreal from first supraocular; subocular rowcomplete; lower eyelid with one row of scales,lacking an enlarged oval window, largelytransparent; supralabials seven, fifth and sixthbeneath center of eye; infralabials seven.


Mental wider than long, in contact withfirst infralabials; postmental single, enlarged,slightly wider than mental, followed by threepairs of enlarged chin shields, first pair inbroad medial contact, second pair equal inwidth to first pair, broadly separated by singlemedial scale, third pair separated by threemedial scales.

Scales on limbs smaller than body scales;scales on dorsal surfaces of digits large,wrapping around lateral edges of digits;lamellae undivided; palmar surfaces of handsand plantar surfaces of feet covered by small,variably shaped scales, each bearing variablyraised anterior edges; scales on dorsal surfaceof hands and feet smaller than limb scales,lacking raised edges.

Coloration in preservative.—Ground colorof body medium brown, lateral surfacelacking dark pigment, dorsal surface bearingcontinuous, longitudinal rows of olive greento brown pigment, spanning six full and twohalf rows of scales at midbody, narrowing tofour full and two half rows of scales posteriorto parietals, extending from posterior edge ofparietals to tail tip; dark pigmentation cover-ing one half to three fourths of dorsal scales;lateral surface of body with six continuousto discontinuous longitudinal rows of olivegreen to brown stripes, extending from post-erior edge of eye to base of tail; dorsolateralstripes present, lacking dark pigmentation,spanning two half rows of scales, extendingfrom posterior edge of supraoculars to base oftail. Ventral scales with or without dark spots.Head scales uniform medium brown, darkerbrown than ventral scales; rostral, nasal,postnasal, supranasal, first supralabial, men-tal, and first infralabial dark gray; pinealeyespot poorly defined, small, and light todark brown. Limbs mottled light and mediumbrown dorsally, yellowish brown ventrally;dorsal and ventral surfaces of digits darkbrown.

Coloration in life.—Ground color of bodymedium brown to tan (Fig. 11C); longitudinalrows of darker brown pigmentation; dorsolat-eral stripes light brown to tan; limbs bearingdark brown mottling dorsally.

Variation.—Morphometric variation issummarized in Table 6. We observed varia-tion among the 20 specimens examined for the

degree of contact between head scales. Sixspecimens were observed to have parietalsseparated by the interparietal (KU 323944,323949, 323953, 323962, 323975–6), and 14specimens possessed parietals in moderate tobroad medial contact (KU 323948, 323952,323954–6, 323960, 323963, 323966, 323970,323972, 323981, 323982, 323990, 324001)behind the interparietal. In addition, sevenspecimens do not have the first pair ofenlarged chin shields in medial contact(KU 323954–5, 323970, 323975, 323981–2,324001), and 13 specimens with the first pairof enlarged chin shields in moderate to broadmedial contact (KU 323944, 323948–9, 323952–3, 323956, 323960, 323962–3, 323966, 323972,323976, 323990).

Scale counts were observed to vary amongthe measured series. Specimens were ob-served to have midbody scale row counts of26 (CAS-SU 18709, 18717, 24502, 24523–5,24541, 24543, 25443–4), 27 (CAS-SU 24503–4, 24522, 24867), and 28 (CAS-SU 24518,24520–21, 24528, 25447); axilla–groin scalerow counts of 42 (CAS-SU 18717, 24502,24520, 24524), 43 (CAS-SU 24523, 24541,24543), 44 (CAS-SU 24504, 24518, 24525,24528, 24867, 25447), 45 (CAS-SU 24503,24521, 25443), and 46 (CAS-SU 18709, 24522,25444); and paravertebral scale row counts of63 (CAS-SU 18717, 24502, 24520, 24524,24541, 24543), 64 (CAS-SU 24504, 24523,24525, 24528, 24867, 25447), 65 (CAS-SU24518), and 66 (CAS-SU 18709, 24503,24521–2, 25443–4).

We also observed Toe-IV lamellae counts tovary among the measured series. With theexception of a single specimen with sevenTwo-IV lamellae (CAS-SU 24528, holotype),all specimens examined had either nine (CAS-SU 18709, 24504, 24523–4, 24867, 25444) or10 (CAS-SU 18717, 24502–3, 24518, 24520–2,24525, 24541, 24543, 25443, 25447). Weobserved the holotype to have malformeddigits that we believe resulted in fewer Toe-IVlamellae than are observed for all otherindividuals of this species.

Distribution.—Brachymeles boholensis isknown only from Bohol Island (Fig. 3).

Ecology and natural history.—Brachymelesboholensis occurs in agricultural habitats, aswell as in disturbed and secondary growth


forests. No original, low-elevation forest re-mains on Bohol Island, but we assume thespecies once also occurred in primary forest atlow elevations. Individuals have been ob-served under piles of rotting coconut husks,in the humus material within rotting logs, andin loose soil and leaf litter surrounding theroot networks of trees. Interestingly, thisspecies seems to be a ubiquitous habitatgeneralist on Bohol, whereas its congener B.orientalis seems to be restricted to fallen androtting logs in secondary growth forest on thesame island (CDS, personal observation). As istypical for species in the genus, individualsimmediately attempt to evade capture bymoving to quickly burrow into loose soil orhumus.

Although only two species of Brachymeleshave been confirmed to occur on Bohol Island(B. boholensis and B. orientalis), populationsof B. samarensis are known to occur onLapinig Grande and Lapinig Chico islandsjust off the northeast coast of Bohol Island(Brown and Alcala, 1980). No individuals ofthis species lacking fully formed digits occuron Bohol; however, given proximity of thesesmall islands to mainland Bohol, it seemslikely that this neighboring, limb-reducedspecies of Brachymeles eventually may bediscovered on Bohol.

Sympatric lizard species observed on BoholIsland include the following: (Agamidae)Bronchocela cristatella, Draco bimaculatus,D. ornatus, D. reticulates, Gonocephalussemperi, Hydrosaurus pustulatus; (Gekkoni-dae) Cyrtodactylus annulatus, Gehyra muti-lata, Gekko gecko, Hemidactylus frenatus, H.platyurus, Hemiphyllodactylus typus, Lepi-dodactylus aureolineatus, L. planicaudus,Pseudogekko compressicorpus, P. brevipes;(Scincidae) Brachymeles schadenbergi orien-talis, Emoia atrocostata, Eutropis multicar-inata, E. multifasciata, Lamprolepis smarag-dina, Lipinia pulchella, L. quadrivittata,Sphenomorphus acutus, S. cumingi, S. fas-ciatus, S. jagori, S. minanensis, S. steerei, S.variegatus; and (Varanidae) Varanus cumingi.Also, Brachymeles samarensis (Scincidae) isknown to occur on Lipinig Grande andLipinig Chico islands just off the northeastcoast of Bohol Island (Brown and Alcala,1980).

Brachymeles mindorensis Brown and Rabor1967

Figs. 3, 7

Brachymeles gracilis mindorensis, Brown andRabor, 1967. Type locality: Bank of TaroginRiver, 30 km southeast of Calapan, Mind-oro Oriental Province, Mindoro Island, Phi-lippines, 0–33-m elevation, 13u 119 25.440 N,121u 89 52.80 E (holotype: CAS-SU 24487).


Brachymeles boulengeri mindorensis (part),Brown and Alcala, 1980.Diagnosis.—Brachymeles mindorensis can

be distinguished from congeners by thefollowing combination of characters: (1) bodysize moderate (SVL 90.0–104.2 mm); (2)pentadactyl; (3) Finger-III lamellae five orsix; (4) Toe-IV lamellae eight or nine; (5)moderate limb length; (6) supralabials seven;(7) infralabials six; (8) pineal eye spot present;(9) enlarged chin shields in two pairs; (10)nuchal scales undifferentiated; (11) fifth andsixth supralabial below eye; (12) auricularopening present; (13) continuous, light, dorso-lateral stripes present; and (14) continuous,dark middorsal stripes present (Tables 4 and 5).

Comparisons.—Characters distinguishingB. mindorensis from all pentadactyl speciesof Brachymeles are summarized in Tables 4and 5. Brachymeles mindorensis most closelyresembles B. boholensis, B. boulengeri, and B.taylori, but it differs from these three taxa byhaving a larger body size, longer hind limbs,eight or nine Toe-IV lamellae, and six infra-labials (Tables 4 and 5). Brachymeles mind-orensis further differs from B. boholensis byhaving five or six Finger-III lamellae and bythe absence of a third pair of enlarged chinshields (Tables 4 and 5); from B. boulengeriby having seven supralabials, six supraciliaries,and the fifth and sixth supralabial below theeye, and by the presence of continuous, darkmiddorsal stripes (Tables 4 and 5); and fromB. taylori by having seven supralabials and thefifth and sixth supralabial below the eye, andby the presence of continuous, light dorsolat-eral stripes (Tables 4 and 5).

From all nonpentadactyl species of Brachy-meles (B. apus, B. bonitae, B. cebuensis, B.elerae, B. lukbani, B. minimus, B. muntingka-may, B. pathfinderi, B. samarensis, B. tridac-


tylus, B. vermis, and B. wrighti), B. mind-orensis differs by having a pentadactyl bodyform (vs. nonpentadactyl), longer forelimblengths (.10.0 mm vs. ,6.9 mm), greaterhind limb lengths (.18.8 mm vs. ,12.9 mm),and the fifth supralabial below the eye (vs.fourth), and by the presence of a postnasalscale (vs. absence). In addition, Brachymelesmindorensis differs from all nonpentadactylspecies except B. wrighti by having a midbody

scale row count 26–28 (vs. ,24); from allnonpentadactyl species except B. pathfinderiby having a paravertebral count 63–65 (vs..84) and by the presence of auricularopenings (vs. absence); and from B. apus, B.lukbani, B. minimus, and B. vermis by thepresence of limbs (vs. absence).

Description (based on holotype and 33referred paratypes at CAS).—Details of thehead scalation of an adult male are shown in

FIG. 7.—Illustration of head of adult male Brachymeles mindorensis (KU 304343) and adult female Brachymelestaylori (KU 324049) in dorsal, lateral, and ventral views. Labels for taxonomically diagnostic head scales follow thoseshown in Figure 6. Illustrations by CDS.


Fig. 7. Measurements of the holotype areincluded below in brackets. Body moderaterelative to other Brachymeles, elongate withrespect to other lizards; maximum SVL104.2 mm for males, 106.8 mm for females[106.8, female] (Tables 4 and 5); head weaklydifferentiated from neck, nearly as wide asbody, HW 10.5–12.9% (11.4 6 0.7) SVL[10.9], 104.2–130.1% (116.1 6 6.0) HL[114.6]; HL 31.6–42.9% (37.3 6 3.1) SnFa[39.2]; SnFa 24.1–29.9% (26.5 6 1.3) SVL[24.1]; snout moderately long, rounded indorsal and lateral profile, SNL 50.6–68.2%(57.3 6 4.9) HL [52.2]; auricular openingpresent, small; eyes moderate, ED 2.0–2.6%(2.2 6 0.2) SVL [2.0], 19.4–27.4% (22.7 62.0) HL [21.1], 51.8–72.2% (60.7 6 5.1) END[58.5], pupil nearly round; body slightlydepressed, MBW 105.4–156.0% (120.2 614.0) MBH [125.0]; scales smooth, glossy,imbricate; longitudinal scale rows at midbody26–28 [26]; paravertebral scale rows 63–65[65]; axilla–groin scale rows 42–45 [45]; limbswell developed, pentadactyl, digits small;FinIIIlam 5–6 [5]; ToeIVlam 8–9 [9]; FLL14.5–21.3% (17.8 6 1.9) AGD [15.5], 10.1–13.1% (11.5 6 0.9) SVL [10.9]; HLL 16.7–37.9% (31.5 6 5.0) AGD [25.1], 10.5–24.0%(20.3 6 2.9) SVL [17.6]; order of digits fromshortest to longest for hand: I 5 V , II 5 IV, III, for foot: I , V , II , III 5 IV; tail notas wide as body, sharply tapered toward theend, TW 54.4–80.1% (69.1 6 8.1) MBW[54.4], TL 60.4–99.3% (84.6 6 11.5) SVL[93.5].

Rostral projecting dorsoposteriorly to pointin line with anterior edge of nasal, broaderthan high, in narrow contact with frontonasal;frontonasal wider than long; nostril ovoid,centered in a single rectangular nasal; nasalswell separated; supranasals present, large,narrowly separated by frontonasal; postnasalspresent; prefrontals narrowly separated byfrontal; frontal octagonal, narrowly contactingfrontonasal and first two supraoculars anteri-orly, 43 wider than anteriormost supraocular;supraoculars five; frontoparietals moderate, inbroad medial contact, each frontoparietal incontact with supraoculars 2–4; interparietallarge, quadrilaterally shaped, slightly longerthan wide, its length slightly greater thanmidline length of frontoparietal; parietal

eyespot present in posterior one half of scale;parietals moderately separated behind inter-parietal; nuchals undifferentiated from adja-cent dorsal scales; loreals two, anterior lorealin contact with prefrontal, postnasal, suprana-sal, second supralabial, posterior loreal andfrontonasal; preocular, single, nearly one thirdas high as posterior loreal; presubocularsingle; supraciliaries six, the anteriormostcontacting prefrontal and separating posteriorloreal from first supraocular; subocular rowcomplete; lower eyelid with one row of scales,lacking an enlarged oval window, largelytransparent; supralabials seven, fifth and sixthbelow eye; infralabials six.

Mental wider than long, in contact with firstinfralabials; single enlarged postmental, equalin width to mental followed by two pairs ofenlarged chin shields; first pair in moderatecontact, second pair wider than first, moder-ately separated by single medial scale.

Scales on limbs smaller than body scales;scales on dorsal surfaces of digits large,wrapping around lateral edges of digits;lamellae undivided; palmar surfaces of handsand plantar surfaces of feet covered by small,variably shaped scales, each with variablyraised anterior edges; scales on dorsal surfaceof hands and feet smaller than limb scales,lacking raised edges.

Coloration in preservative.—Ground colorof body medium brown; dorsal surface of bodywith eight longitudinal rows of dark brownstripes, six continuous medial rows, twodiscontinuous lateral rows, spanning eight fullrows of scales at midbody, narrowing to six fullrows of scales posterior to parietals, extendingfrom posterior edge of parietals to tail tip;pigmentation covering middle one third ofdorsal scales; dorsolateral stripes present,lacking dark pigmentation, spanning onewhole and two half rows of scales, extendingfrom posterior edge of supraoculars to base oftail. Lateral surface of body light brownground color with three or four rows of nearlycontinuous spots of dark brown pigmentation,extending from auricular opening to tail tip.Tail striped with longitudinal rows of darkpigmentation. Ventral scales lacking darkspots. Head scales uniform medium brown,darker brown than ventral scales; posterior-most supraocular lacking pigmentation; dark


pigmentation surrounding auricular opening,connected to dark pigmentation on headscales; rostral, nasal, postnasal, supranasal,and first supralabial dark gray; pineal eyespotpoorly defined, small and light cream. Limbsmottled dark brown dorsally, yellowish brownventrally; dorsal and ventral surface of digitsdark brown.

Coloration in life.—Ground color of bodydark to medium brown; continuous, darkmiddorsal stripes dark brown to black; dorso-lateral stripes light brown to tan, graduallybecome predominately tan on tail; limbs darkbrown dorsally.

Variation.—Morphometric variation of theseries is summarized in Table 6. We observedvariation among the 16 specimens examinedfor the degree of contact between head scales.Five specimens have parietals separated bythe interparietal (KU 304352, 304354, 304488,308447–8), and 11 specimens have parietalsin moderate to broad medial contact (KU304351, 304353, 304355, 304412–3, 307739–42, 308404, 308534) behind the interparietal;four specimens have supranasals narrowlyseparated by the frontonasal (KU 307740–1,308404, 308448), six specimens have supra-nasals in medial point contact (KU 304351–4,307742, 308447), and six specimens havesupranasals in moderate medial contact (KU304355, 304412–3, 304488, 307739, 308534);13 specimens have prefrontals moderatelyseparated by the frontal (KU 304351–4,304412–3, 304488, 307739–41, 308447–8,308534), one specimen has prefrontals nar-rowly separated by the frontal ( KU 308404),and two specimens have prefrontals in medialpoint contact (KU 304355, 307742).

Scale counts were observed to vary amongthe measured series. Specimens were ob-served to have midbody scale row counts of26 (CAS-SU 24487, 24551, 24561, 24566,24574, 24577, 24579), 27 (CAS-SU 24549,24573), and 28 (CAS-SU 24550, 24552–4,24562, 24564, 24568, 24570, 24578); axilla–groin scale row counts of 42 (CAS-SU 24564),43 (CAS-SU 24573), 44 (CAS-SU 24553–4,24561–2, 24568, 24570, 24578), and 45(CAS-SU 24487, 24549–52, 24566, 24574,24577, 24579); and paravertebral scale rowcounts of 63 (CAS-SU 24564, 24573), 64(CAS-SU 24561–2, 24568), and 65 (CAS-SU

24487, 24549–54, 24566, 24570, 24574,24577–9).

We also observed lamellae counts to varyamong the series. With the exception of twospecimens with six Finger-III lamellae (CAS-SU 24561, 24574), all specimens examinedhad five. Specimens were observed to haveToe-IV lamellae counts of eight (CAS-SU24549–54, 24561–2, 24564, 24566, 24570,24573, 24577–8) or nine (CAS-SU 24487,24568, 24574, 24579).

Distribution.—Brachymeles mindorensis isknown only from Mindoro Island (Fig. 3).

Ecology and natural history.—Brachymelesmindorensis occurs in disturbed and second-ary growth forest. Individuals have beenobserved under piles of rotting coconut husks,in the humus material within rotting logs, andin loose soil and leaf litter surrounding theroot networks of trees. The species has beenobserved to be quite common in certainhabitats on Mindoro Island (CDS and RMB,personal observations). Brachymeles mindor-ensis occurs sympatrically with B. bonitae(Brown, 1956; Brown and Alcala, 1980; Brownand Rabor, 1967), and it is the largest speciesof the B. boulengeri complex. When individ-uals were disturbed, they attempted to quicklyburrow back into loose soil or humus.

Sympatric lizard species observed on Mind-oro Island include the following: (Agamidae)Bronchocela marmoratus, Draco quadrasi,Gonocephalus interruptus, Hydrosaurus pustu-latus; (Gekkonidae) Cyrtodactylus philippini-cus, Gehyra mutilata, Gekko gecko, G. mind-orensis, Hemidactylus frenatus, H. garnoti, H.platyurus, Hemiphyllodactylus typus, Lepido-dactylus naujanensis; (Scincidae) Brachymelesbonitae, Dasia olivaceum, Emoia atrocostata,Eutropis multicarinata, E. multifasciata, Lam-prolepis smaragdina, Lipinia auriculatum,Sphenomorphus cumingi, S. jagori, S. steerei;and (Varanidae) Varanus marmoratus.

Brachymeles taylori Brown 1956Figs. 3, 7, 11B

Brachymeles gracilis taylori, Brown, 1956.Type locality: Sitio Lunga, 13 km WestDumaguete, 3 km West Valencia, on lowridge on the north side of the Maite River,Negros Oriental Province, Negros Island,Philippines, 600-m elevation, 9u 179 32.960


N, 123u 149 2.40 E (holotype: CAS-SU24487).


Brachymeles boulengeri taylori (part), Brownand Alcala, 1980.Diagnosis.—Brachymeles taylori can be

distinguished from congeners by the followingcombination of characters: (1) body sizemoderate (SVL 65.8–99.2 mm); (2) pentadac-tyl; (3) Finger-III lamellae five or six; (4) Toe-IV lamellae nine or 10; (5) moderate limblength; (6) supralabials six; (7) infralabialsseven; (8) pineal eye spot present; (9)supranasals not contacting on midline; (10)prefrontals not contacting on midline; (11)enlarged chin shields in two pairs; (12) nuchalscales undifferentiated; (13) fourth and fifthsupralabial below eye; (14) auricular openingpresent; (15) dorsolateral stripes absent; and(16) continuous, dark middorsal stripes pres-ent (Tables 4 and 5).

Comparisons.—Characters distinguishingBrachymeles taylori from all pentadactylspecies of Brachymeles are summarized inTables 4 and 5. Brachymeles taylori mostclosely resembles B. boholensis, B. boulengeri,and B. mindorensis, but it differs from thesethree taxa by the absence of continuous, lightdorsolateral stripes (Tables 4 and 5). Brachy-meles taylori can further be distinguishedfrom B. boholensis by having five or sixFinger-III lamellae, six supralabials, and thefourth and fifth supralabial below the eye, andby the absence of a third pair of enlarged chinshields (Tables 4 and 5); from B. boulengeriby having six supralabials and six supracili-aries, and the presence of continuous, darkmiddorsal stripes (Tables 4 and 5); and fromB. mindorensis by having a smaller body size,shorter hind limbs, nine or 10 Toe-IVlamellae, six supralabials, seven infralabials,the fourth and fifth supralabial below the eye,and no contact between supranasals andprefrontals (Tables 4 and 5).

From all nonpentadactyl species of Brachy-meles (B. apus, B. bonitae, B. cebuensis, B.elerae, B. lukbani, B. minimus, B. muntingka-may, B. pathfinderi, B. samarensis, B. tridac-tylus, B. vermis, and B. wrighti), B. tayloridiffers by having pentadactyl limbs (vs. non-pentadactyl), longer forelimbs (.9.0 mm vs.

,6.9 mm), longer hind limbs (.15.6 mm vs.,12.9 mm), and Toe-IV lamellae nine or 10(vs. eight or fewer), and by the presence of apostnasal scale (vs. absence). In addition,Brachymeles taylori differs from all nonpen-tadactyl species except B. wrighti by having amidbody scale row count 26–28 (vs. ,24);from all nonpentadactyl species except B.pathfinderi by having a paravertebral count62–69 (vs. .84) and by the presence ofauricular openings (vs. absence); and from B.apus, B. lukbani, B. minimus, and B. vermisby the presence of limbs (vs. absence).

Description (based on holotype and 33referred specimens, including five paratypesat CAS).—Details of the head scalation of anadult female are shown in Fig. 7. Measure-ments of the holotype are provided below inbrackets. Body moderate relative to otherBrachymeles, elongate with respect to otherlizards; maximum SVL 99.2 mm for males,93.2 mm for females [65.8, female] (Tables 4and 5); head weakly differentiated from neck,nearly as wide as body, HW 11.3–15.3% (12.36 1.0) SVL [15.3], 104.4–139.5% (120.5 610.2) HL [132.8]; HL 34.8–44.6% (39.2 6 2.5)SnFa [36.4]; SnFa 23.4–31.7% (26.2 6 1.7)SVL [31.7]; snout moderately long, broadlyrounded in dorsal profile, depressed in lateralprofile, SNL 44.1–62.5% (52.2 6 4.6) HL[62.5]; auricular opening present, moderate;eyes moderate, ED 1.6–2.8% (2.1 6 0.2) SVL[2.8], 15.9–24.4% (20.8 6 2.4) HL [24.4], 48.8–69.5% (59.6 6 6.1) END [62.7], pupil nearlyround; body slightly depressed, MBW 89.1–148.4% (115.1 6 14.7) MBH [112.9]; scalessmooth, glossy, imbricate; longitudinal scalerows at midbody 26–28 [27]; paravertebralscale rows 62–69 [64]; axilla–groin scale rows42–47 [43]; limbs well developed, pentadactyl,digits small; FinIIIlam 5–6 [6]; ToeIVlam 8–10[9]; FLL 15.7–20.4% (17.8 6 1.2) AGD [19.9],10.4–14.1% (11.5 6 0.9) SVL [14.1]; HLL26.7–36.8% (31.1 6 2.5) AGD [35.6], 17.8–25.2% (20.1 6 1.7) SVL [25.2]; order of digitsfrom shortest to longest for hand: I 5 V , IV ,II 5 III, for foot: I , V , II , III 5 IV; tail notas wide as body, sharply tapered posteriorly,TW 54.0–80.3% (68.9 6 6.7) MBW [75.0], TL69.2–103.1% (83.3 6 10.4) SVL [98.2].

Rostral projecting dorsoposteriorly to pointin line with anterior edge of nasal, broader


than high, in moderate contact with frontona-sal; frontonasal wider than long; nostril ovoid,in center of single rectangular nasal; nasalswell separated; supranasals present, large,narrowly separated by frontonasal; postnasalspresent; prefrontals moderately separated byfrontal; frontal nearly octagonal, its anteriormargin in moderate contact with frontonasal,in contact with first two anterior supraoculars,4.53 wider than anteriormost supraocular;supraoculars five; frontoparietals moderate, inbroad medial contact, each frontoparietal incontact with supraoculars 2–4; interparietalmoderate, quadrilaterally shaped, longer thanwide, its length nearly equal to midline lengthof frontoparietal; parietal eyespot presentin posterior one half of scale; parietals inmoderate contact behind interparietal ornarrowly separated; nuchals undifferentiated;loreals two, decreasing in size from anterior toposterior, subequal, anterior loreal in contactwith prefrontal, postnasal, supranasal, secondsupralabial, posterior loreal and frontonasal;preocular single, nearly two thirds as high asposterior loreal; single presubocular; supraci-liaries six, the anteriormost contacting pre-frontal and separating posterior loreal fromfirst supraocular; subocular row complete;lower eyelid with one row of scales, lackingan enlarged oval window, largely transparent;supralabials six, fourth and fifth below the eye;infralabials seven.

Mental wider than long, in contact with firstinfralabial; postmental single, enlarged, slight-ly wider than mental, followed by two pairs ofenlarged chin shields, scales of first pairseparated or in moderate contact, second pairslightly wider than first, broadly separated bya single medial scale.

Scales on limbs smaller than body scales;scales on dorsal surfaces of digits large,wrapping around lateral edges of digits;lamellae undivided; palmar surfaces of handsand plantar surfaces of feet covered by small,irregular scales, each with irregular raisedanterior edges; scales on dorsal surface ofhands and feet smaller than limb scales,lacking raised edges.

Coloration in preservative.—Ground colorof body medium olive brown; dorsal pigmen-tation nearly all dark brown, gradually fadinginto olive brown lateral surface and yellowish

brown ventral surface of body, ventral surfacewithout dark pigmentation; dark dorsal pig-mentation in nearly continuous block acrossdorsal surface or at times forming 10 contin-uous longitudinal rows of dark brown pig-ment, spanning six full rows of scales acrossdorsal surface, extending from posterior edgeof parietals to tail tip, additional three to fourscale rows on each lateral surface with darkcoloration, covering one half to two thirds ofdorsal and lateral scales; dorsolateral stripesabsent. Tail coloration matches body colora-tion. Head scales uniform dark brown, darkerbrown than ventral scales; rostral, nasal,postnasal, supranasal, and first supralabialdark gray; pineal eyespot poorly defined, smalland light cream. Limbs mottled mediumbrown dorsally, yellowish brown ventrally;dorsal and ventral surface of digits darkbrown.

Coloration in life.—Ground color of bodylight to medium brown (Fig. 11B); medium todark brown dorsal pigmentation graduallyfading into medium brown lateral pigmenta-tion; limbs mottled medium to dark browndorsally.

Variation.—Morphometric variation of theseries is summarized in Table 6. We observedvariation among the 14 specimens examinedfor the degree of contact between head scales.Six specimens have parietals moderatelyseparated by the interparietal (KU 324048,324050, 324052–5), one specimen has parie-tals narrowly separated by the interparietal(KU 324051), and seven specimens haveparietals in moderate medial contact (KU306651, 324044–7, 324049, 324056) behindthe interparietal; eight specimens do not havethe first pair of enlarged chin shields in medialcontact (KU 324045, 324048–50, 324053–6),one specimen has the first pair of enlargedchin shields in point medial contact (KU324046), and five specimens have the first pairof enlarged chin shields in moderate medialcontact (KU 306651, 324044, 324047,324051–2).

Scale counts were observed to vary amongthe measured series. Specimens were ob-served to have midbody scale row counts of26 (CAS-SU 18641, 18656–7, 18748, 21873,21877, 21880, 21884, 22355, CAS 154971,154673, 154680–2, 154686), 27 (CAS-SU


18615, 22356, CAS 154679), and 28 (CAS-SU18649, 21883, CAS 154678); axilla–groin scalerow counts of 42 (CAS 154680, 154686), 43(CAS-SU 18615, 18656, CAS 154682), 44(CAS-SU 18641, 18748, 21873, 21877, 21880,22356, CAS 154678, 154679, 154681), 45(CAS-SU 21884, 22355, CAS 154671), 46(CAS-SU 18649, 21883, CAS 154673), and 47(CAS-SU 18657); and paravertebral scale rowcounts of 62 (CAS 154680, 154686), 64 (CAS-SU 18615, 18656, CAS 154682), 66 (CAS-SU18641, 18748, 21873, 21877, 21880, 21884,22356, CAS 154678–9, 154681), 67 (CAS-SU18649, CAS 154671, 154673), 68 (CAS-SU21883, 22355), and 69 (CAS-SU 18657).

We also observed lamellae counts to varyamong the measured series. Specimens wereobserved to have Finger-III lamellae counts offive (CAS-SU 18641, 18649, 18656, 18748,21883–4, 22356, CAS 154671, 154673,154678–80) or six (CAS-SU 18615, 18657,21873, 21877, 21880, 22355, CAS 154681–2,154686); Toe-IV lamellae counts of eight(CAS 154678), nine (CAS-SU 18615, 18649,18656, 21880, 22356, CAS 154671, 154673,154682, 154686), and 10 (CAS-SU 18641,18657, 18748, 21873, 21877, 21883–4, 22355,CAS 15479–81).

There is some color variation in theexamined series, with the degree and defini-tion of continuous, dark middorsal stripes. Allspecimens have continuous, dark lines run-ning down the middorsal surface of the body;however, the continuous lines in some spec-imens are present without a dark, middorsalbackground coloration (KU 324045–6, 324049–51, 324053–4). The dark lines in other specimensoverlay a dark, middorsal region covered by along streak of dark background pigmentation(KU 324044, 324047–8, 324052, 324055–6).

Distribution.—Brachymeles taylori isknown from Negros, Cebu, Inampulugan,Pan de Azucar, Danjugan, Ponson, and Poroislands (Fig. 3).

Ecology and natural history.—Brachymelestaylori occurs in agricultural areas as well asdisturbed and secondary growth forest. Littleor no original, low-elevation forest remains inthe Visayas, but we assume the species oncealso occurred in primary forest. Individualshave been observed under piles of rottingcoconut husks, in the humus material within

rotting logs, and in loose soil and leaf littersurrounding the root networks of trees. Thespecies is quite common throughout its range(CDS, personal observation) and occurs sym-patrically with three other species of Brachy-meles (B. cebuensis, B. talinis, and B. tridac-tylus; Brown, 1956; Brown and Alcala, 1980;Brown and Rabor, 1967). Similar to B.boulengeri, B. taylori seems to have a widergeographic distribution that spans multiplePhilippine islands. This is in contrast to theisland endemic species B. boholensis and B.mindorensis that are known from just Boholand Mindoro islands, respectively. As with allmembers of the genus, when disturbed,individuals move rapidly attempting to burrowinto the loose soil or humus.

Sympatric lizard species observed withinthe range of B. taylori include the following:(Agamidae) Bronchocela marmoratus, Dracospilopterus, Gonocephalus sophiae, Hydro-saurus pustulatus; (Dibamidae) Dibamus ar-genteus; (Gekkonidae) Cyrtodactylus philip-pinicus, Gehyra mutilata, Gekko gecko, G.mindorensis, G. enrstkelleri, Luperosauruscorfieldi, Pseudogekko brevipes, Hemidactylusfrenatus, H. platyurus, Hemiphyllodactylustypus, Lepidodactylus christiani, L. herrei,L. lugubris; (Scincidae) Brachymeles cebuen-sis, B. talinis, B. cebuensis, Emoia atrocostata,Eutropis multicarinata, E. multifasciata, Lam-prolepis smaragdina, Lipinia auriculata, L.pulchella, L. quadrivittata, L. rabori, Spheno-morphus arborens, S. coxi, S. jagori, S. steerei,Tropidophorus grayi; and (Varanidae) Vara-nus nuchalis.

Brachymeles orientalis Brown and Rabor1967

Figs. 4, 8, 11F,G

Brachymeles schadenbergi orientalis, Brownand Rabor, 1967. Type locality: BarioDusita, 11 km southeast of Sierra Bullones,Bohol Province, Bohol Island, Philippines,533-m elevation, 9u 469 57.50 N, 124u 18910.80 E (holotype: CAS-SU 24436).

Brachymeles schadenbergi (part), Brown andAlcala, 1970.

Brachymeles schadenbergi (part), Brown andAlcala, 1980.Diagnosis.—Brachymeles orientalis can be

distinguished from congeners by the following


combination of characters: (1) body large(SVL 97.6–112.3 mm); (2) pentadactyl; (3)Finger-III lamellae six or seven; (4) Toe-IVlamellae 8–10; (5) limbs relatively long; (6)supralabials six or seven; (7) infralabials six orseven; (8) pineal eye spot present; (9)supranasal contact absent; (10) prefrontalsnot contacting on midline; (11) enlarged chinshields in two pairs; (12) nuchal scales

undifferentiated; (13) fourth and fifth supra-labial below eye; (14) auricular openingpresent; (15) dorsolateral stripes absent; and(16) middorsal stripes absent (Tables 4 and 5).

Comparisons.—Characters distinguishingBrachymeles orientalis from all pentadactylspecies of Brachymeles are summarized inTables 4 and 5. Brachymeles orientalis mostclosely resembles B. makusog and B. scha-

FIG. 8.—Illustration of head of adult male Brachymeles orientalis (KU 311241) and adult male Brachymelesschadenbergi (KU 314992) in dorsal, lateral, and ventral views. Labels for taxonomically diagnostic head scales followthose shown in Fig. 6. Illustrations by CDS.


denbergi, but it differs from both taxa byhaving six or seven Finger-III lamellae, 8–10Toe-IV lamellae, and the fourth and fifthsupralabial below the eye (Tables 4 and 5),and by the presence (vs. absence) of reddishorange to salmon-colored scales on the lateralsurfaces of the body. Brachymeles orientaliscan further be distinguished from B. makusogby having a greater maximum axilla–groinscale row count and a greater maximumparavertebral scale row count (Table 5) andfrom B. schadenbergi by having no contactbetween supranasals, and by the absence ofcontinuous, dark middorsal stripes and darklateral stripes (Table 5).

From all nonpentadactyl species of Brachy-meles (B. apus, B. bonitae, B. cebuensis, B.elerae, B. lukbani, B. minimus, B. muntingka-may, B. pathfinderi, B. samarensis, B. tridac-tylus, B. vermis, and B. wrighti), B. orientalisdiffers by having pentadactyl limbs (vs.nonpentadactyl), longer forelimb lengths(.10.4 mm vs. ,6.9 mm), greater hind limblengths (.18.6 mm vs. ,12.9 mm), and by thepresence (vs. absence) of a postnasal scale. Inaddition, B. orientalis differs from all non-pentadactyl species except B. pathfinderi byhaving Toe-IV lamellae 8–10 (vs. four orfewer), a paravertebral scale row count 69–72 (vs. .84), and by the presence (vs.absence) of auricular openings; from allnonpentadactyl species except B. wrighti byhaving a midbody scale row count 26–28 (vs.,24); and from B. apus, B. lukbani, B.minimus, and B. vermis by the presence oflimbs (vs. absence).

Description (based on holotype and 52referred specimens, including 13 paratypesfrom CAS).—Details of the head scalation ofan adult male are shown in Fig. 8. Measure-ments of the holotype are provided below inbrackets. Body large relative to other Brachy-meles, elongate with respect to other lizards;maximum SVL 112.3 mm for males, 115.2 mmfor females [99.9, female] (Tables 4 and 5);head weakly differentiated from neck, nearlyas wide as body, HW 10.2–12.7% (11.2 6 0.7)SVL [11.9] 87.5–120.0% (108.4 6 7.3) HL[120.0]; HL 33.7–45.0% (37.6 6 2.6) SnFa[38.0]; SnFa 25.3–30.2% (27.6 6 1.1) SVL[26.1]; snout long, rounded in dorsal andlateral profile, SNL 46.6–65.2% (57.5 6 4.2)

HL [62.5]; auricular opening present, moder-ate; eyes moderate, ED 1.5–2.3% (1.8 6 0.2)SVL [1.8], 14.8–19.6% (17.3 6 1.3) HL [18.2],38.2–56.7% (44.8 6 3.7) END [45.9], pupilsubcircular; body slightly depressed, MBW93.1–138.2% (114.5 6 11.4) MBH [124.9];scales smooth, glossy, imbricate; longitudinalscale rows at midbody 26–30 [28]; paraverte-bral scale rows 69–73 [72]; axilla–groin scalerows 46–49 [48]; limbs well developed,pentadactyl, digits small; FinIIIlam 6–7 [6];ToeIVlam 8–10 [8]; FLL 17.0–24.8% (20.7 62.0) AGD [18.8], 10.7–15.4% (13.1 6 1.1)SVL [12.3]; HLL 27.6–42.7% (34.2 6 3.4)AGD [32.2], 18.4–24.2% (21.6 6 1.7) SVL[21.2]; order of digits from shortest to longestfor hand: I , V , II 5 IV , III, for foot: I ,V , II , III 5 IV; tail nearly as wide as body,sharply tapered toward the end, TW 55.6–86.9% (69.7 6 6.4) MBW [70.9], TL 62.2–106.0% (85.2 6 11.6) SVL [106.0].

Rostral projecting dorsoposteriorly to pointin line with anterior edge of nasal, broaderthan high, in narrow contact with frontonasal;frontonasal wider than long; nostril ovoid,centered in a single rectangular nasal; supra-nasals present, large, narrowly separated byfrontonasal; postnasals present; prefrontalsmoderately separated by frontal; frontal nearlydiamond shaped, its anterior margin inmoderate contact with frontonasal, in contactwith first two anterior supraoculars, 43 widerthan anteriormost supraocular; supraocularsfive; frontoparietals moderate, in broad medi-al contact, each frontoparietal in contact withsupraoculars 2–4; interparietal moderate,quadrilaterally shaped, longer than wide, itslength slightly greater than midline length offrontoparietal; parietal eyespot present inposterior half of scale; parietals in narrowcontact or separated behind interparietal;nuchals nonenlarged, undifferentiated; lorealstwo, decreasing in size from anterior toposterior, anterior loreal about as long asand 1.83 higher than posterior loreal, incontact with prefrontal, postnasal, supranasal,second supralabial, posterior loreal and fron-tonasal, and occasionally with first supralabial;preocular single, nearly two thirds as high asposterior loreal; presubocular single; supraci-liaries six, the anteriormost contacting pre-frontal and separating posterior loreal from


first supraocular; subocular row complete;lower eyelid with one row of scales, lackingan enlarged oval window, largely transparent;supralabials six or seven (six), first 23 size ofother supralabials, fourth and fifth below theeye; infralabials six or seven (seven).

Mental wider than long, in contact with firstinfralabial; single enlarged postmental, widerthan mental; followed by two pairs of enlargedchin shields, first pair narrowly separated bysingle row of undifferentiated scales or inmoderate medial contact, scales of second pairnarrower than first, broadly separated bythree medial scales.


Coloration in preservative.—Ground colorof body cream; lateral and ventral surfaces ofbody lacking dark pigment; dorsum of body,from posterior edge of supranasals to tail tip,uniformly dark brown with dark pigmentationspanning six and two half rows of scales atmidbody and narrowing to cover four and twohalf rows of scales posterior to parietals; bodydark brown dorsally abruptly changing tocream laterally and ventrally; head scalesuniform dark brown; rostral, nasal, postnasal,supranasal, and first supralabial light gray;pineal eyespot charcoal; small dark brownblotch dorsal to auricular openings. Limbsmottled light and medium brown dorsally,cream colored ventrally; dorsal and ventralsurface of digits light brown.

Coloration in life.—Dorsal ground colorhomogeneous medium brown (Fig. 11F,G);sharp lateral demarcation between dorsal andlateral and ventral coloration; lateral and ventralsurfaces of body bright burnt orange, orange-brown, or salmon colored; dark brown spotsand longitudinal lines of spots absent fromlateral surfaces. Limbs medium brown dorsally,burnt orange to orange-brown ventrally. Dorsalhead scales uniform medium brown.

Variation.—Morphometric variation of theseries is summarized in Table 6. We observed

variation among the 19 specimens examinedfor the degree of contact between head scales.Twelve specimens were observed to haveparietals moderately separated by the inter-parietal (KU 305470, 310734–6, 310942,310944, 310949, 310951, 311232–5), onespecimen has parietals in point medial contact(KU 311231), and six specimens have parietalsin moderate medial contact (KU 310739,310943, 310945–6, 310955, 311241) behindthe interparietal; nine specimens do not havethe first pair of enlarged chin shields in medialcontact (KU 210736, 310942, 310945–6,310949, 310951, 311231, 311234–5), and 10specimens have the first pair of enlargedchin shields in moderate medial contact(KU 305470, 310734–5, 310739, 310943–4,310955, 311232–3, 311241).

Scale counts were observed to vary amongthe measured series. The number of suprala-bials varied between six (CAS-SU 18702,24428, 24434, 24436–7, 24442, 24446–9,24451, 24458, 25452, 25460, 28332, CAS102404, 110978–81, 133301, 133616, 133749,133752, 133754, KU 310734, 310736, 310942,311231–2, 311234) and seven (CAS-SU24450, 28320–1, 28338, 28370, CAS 110976–7, 110982–3); infralabials varied between six(CAS-SU 24446, KU 310734, 310736, 310942,311231–2, 311234) and seven (CAS-SU18702, 24428, 24434, 24436–7, 24442,24447–51, 24458, 25452, 25460, 28320–1,28332, 28338, 28370, CAS 102404, 110976–83, 133301, 133616, 133749, 133752, 133754).Specimens were observed to have midbodyscale row counts of 26 (CAS-SU 24428,24446), 27 (CAS-SU 24458, 25460), 28(CAS-SU 18702, 24434, 24436–7, 24442,24447–52, CAS 102404, 133301, 133752,133754), 29 (CAS-SU 28320, 28338, CAS110976, 110981, 133616, 133749), and 30(CAS-SU 28231–2, 28370, CAS 110977–80,110982–3); axilla–groin scale row counts of 46(CAS-SU 25452, CAS 110983, 133616), 47(CAS-SU 24450, 24458, 25460, CAS 110979–81, 133301), 48 (CAS-SU 18702, 24428,24436, 24442, 24446, 24449, 28332, 28338,CAS 110976, 110978, 110982, 133749), and 49(CAS-SU 24434, 24437, 24447–8, 24451,28320–1, 28370, CAS 102404, 110977,133752, 133754); and paravertebral scale rowcounts of 69 (CAS-SU 25452, CAS 133616),


70 (CAS-SU 24450, 24458, 25460, CAS133301), 71 (CAS-SU 18702, 24428, 24442,24446, 24449, 28332, 28338, CAS 110980,110983, 133749), 72 (CAS-SU 24434, 24436–7, 24447–8, 24451, 28320–1, CAS 102404,110976–9, 110981–2, 133752, 133754), and 73(CAS-SU 28370).

We also observed lamellae counts to varyamong the measured series. Specimens wereobserved to have Finger-III lamellae counts ofsix (CAS-SU 18702, 24428, 24434, 24436–7,24442, 24446–7, 24449–51, 24458, 25460,28320–1, 28332, 28338, 28370, CAS 102404,110976–83, 133301, 133616, 133749, 133754,KU 310734, 311231–2, 311234) or seven(CAS-SU 24448, CAS 133752, KU 310736,310942); Toe-IV lamellae counts of eight(CAS-SU 24436, 25452, 28320, CAS 102404),nine (CAS-SU 18702, 24428, 24434, 24437,24442, 24446–51, 24458, 25460, 28321, 28332,28370, CAS 110976–7, 110979, 110981–3,133301, 133616, 133754, KU 310734, 310942,311232, 311234), or 10 (CAS-SU 28338, CAS110978, 110980, 133749, 133752, KU 310736,311231).

Distribution.—Brachymeles orientalis isknown from Bohol, Samar, Leyte, Dinagat,Camiguin Sur islands and the eastern andcentral portions of Mindanao Island (Fig. 4).

Ecology and natural history.—Brachymelesorientalis occurs in agricultural areas as wellas disturbed and secondary growth forest. OnSamar, Leyte, Mindanao, and Camiguin Surislands, we have collected this species inprimary forest, and on Bohol Island it ispresent in mature secondary growth. Individ-uals have been observed under piles of rottingcoconut husks, in the humus material withinrotting logs, and in loose soil and leaf littersurrounding the root networks of trees. Thespecies is quite common throughout its rangewith the exception of Bohol Island (CDS,personal observation), and it occurs sympatri-cally with four other species of Brachymeles indifferent parts of its range (Brown, 1956;Brown and Alcala, 1980; Brown and Rabor,1967). Brachymeles orientalis occurs sympat-rically with B. boholensis on Bohol Island, B.gracilis hilong and B. samarensis on Samarand Leyte islands, B. gracilis hilong onMindanao Island, and B. cf. gracilis hilongon Camiguin Sur Island (CDS, personal

observation). Similar to B. boulengeri, B.orientalis seems to have a wider geographicdistribution that spans multiple Philippineislands. This is in contrast to the pentadactyl,island endemic species B. boholensis, B.tungaoi, and B. mindorensis that are knownfrom just Bohol, Masbate, and Mindoroislands, respectively. As do all members ofthe genus, disturbed individuals move in arapid serpentine manner and always attemptto burrow back into loose soil or humus.

Sympatric lizard species observed withinthe range of Brachymeles orientalis includethe following: (Agamidae) Bronchocela crista-tella, Draco bimactulatus, D. cyanopterus, D.mindanensis, D. ornatus, Gonocephalus inter-ruptus, G. semperi, Hydrosaurus pustulatus;(Gekkonidae) Cyrtodactylus agusanensis, C.annulatus, C. jambangan, Gehyra mutilata,Gekko gecko, Gekko mindorensis, Hemidacty-lus frenatus, H. platyurus, Pseudogekko com-pressicorpus; (Scincidae) Brachymeles grac-ilis hilong, B. cf. g. gracilis, B. samarensis,Eutropis indeprensa, E. multifasciata, Lam-prolepis smaragdina, L. pulchella, L. quad-rivittata, Sphenomorphus abdictus abdictus,S. acutus, S. cumingi, S. cf. mindanensis, S.coxi, S. fasciatus, S. jagori, S. llanosi, S.steerei, S. variegatus, Tropidophorus misami-nus; and (Varanidae) Varanus cumingi.

Brachymeles schadenbergi (Fischer 1885)Figs. 4, 8, 11E

Senira bicolor (part), Gray, 1845.Eumeces (Riopa) schadenbergi, Fischer, 1885.

Type locality: ‘‘Southern Mindanao Island,Philippines’’ (reported by Fischer [1885] asNo. 845 housed in the Dresden Museum).

Brachymeles schadenbergi (part), Boettger,1886; Boulenger, 1887; Boettger, 1893;Taylor, 1917, 1922a,b; Brown and Alcala,1970.

Brachymeles schadenbergi schadenbergi (part),Brown, 1956; Brown and Rabor, 1967.Diagnosis.—Brachymeles schadenbergi can

be distinguished from congeners by thefollowing combination of characters: (1) bodylarge (SVL 93.1–115.8 mm); (2) pentadactyl;(3) Finger-III lamellae five or six; (4) Toe-IVlamellae eight or nine; (5) limbs relativelylong; (6) supralabials six or seven; (7) infra-labials six or seven; (8) pineal eye spot present;


(9) supranasals in contact; (10) prefrontals notcontacting on midline; (11) enlarged chinshields in two pairs; (12) nuchal scalesundifferentiated; (13) fifth and sixth suprala-bial below eye; (14) auricular opening present;and (15) continuous, light dorsolateral stripesabsent (Tables 4 and 5).

Comparisons.—Characters distinguishingBrachymeles schadenbergi from all pentadac-tyl species of Brachymeles are summarized inTables 4 and 5. Brachymeles schadenbergimost closely resembles B. makusog and B.orientalis, but it differs from both taxa byhaving eight or nine Toe-IV lamellae, and thefifth and sixth supralabial below the eye, andby contact between supranasals (Tables 4 and5). Brachymeles schadenbergi can further bedistinguished from B. makusog by having agreater maximum axilla–groin scale row countand a greater maximum paravertebral scalerow count (Table 5) and from B. orientalis bythe absence (vs. presence) of reddish orangeto salmon-colored scales on the lateral surfac-es of the body.

From all nonpentadactyl species of Brachy-meles (B. apus, B. bonitae, B. cebuensis, B.elerae, B. lukbani, B. minimus, B. muntingka-may, B. pathfinderi, B. samarensis, B. tridac-tylus, B. vermis, and B. wrighti), B. schaden-bergi differs by having a pentadactyl bodyform (vs. nonpentadactyl), longer forelimblengths (.11.1 mm vs. ,6.9 mm), greaterhind limb lengths (.18.5 mm vs. ,12.9 mm),and by the presence of a postnasal scale (vs.absence). In addition, B. schadenbergi differsfrom all nonpentadactyl species except B.pathfinderi by having Toe-IV lamellae eight ornine (vs. four or fewer), 67–72 paravertebrals(vs. .84), and by the presence (vs. absence) ofauricular openings; from all nonpentadactylspecies except B. wrighti by having a midbodyscale row count 26–28 (vs. ,24); and from B.apus, B. lukbani, B. minimus, and B. vermisby the presence (vs. absence) of limbs.

Description (based on holotype descriptionand 34 referred specimens).—Details of thehead scalation of an adult male are shown inFig. 8. The holotype was not examined by us;however, measurements of the holotype takenfrom the original description are providedbelow in brackets. Body large relative to otherBrachymeles, elongate with respect to other

lizards; maximum SVL 115.8 mm for males,113.5 mm for females [85] (Tables 4 and 5);head weakly differentiated from neck, nearlyas wide as body, HW 10.2–11.7% (11.2 6 0.5)SVL, 102.0–116.8% (108.7 6 4.2) HL; HL37.0–40.2% (38.4 6 0.9) SnFa; SnFa 25.7–27.4% (26.8 6 0.5) SVL; snout moderatelylong, rounded in dorsal and lateral profile,SNL 48.8–58.8% (54.3 6 2.9) HL; auricularopening present, moderate; eyes moderate,ED 1.8–2.2% (2.1 6 0.1) SVL, 18.4–21.8%(20.0 6 1.0) HL, 44.7–57.5% (52.3 6 4.3)END, pupil nearly round; body slightlydepressed, MBW 94.8–135.4% (115.8 613.0) MBH; scales smooth, glossy, imbricate;longitudinal scale rows at midbody 26–28 [28fide Fischer, 1885]; paravertebral scale rows67–72; axilla–groin scale rows 45–50 [46 fideFischer, 1885]; limbs well developed, penta-dactyl, digits small; FinIIIlam 5–6; ToeIVlam8–9; FLL 12.5–21.6% (18.0 6 2.4) AGD,10.3–13.7% (12.0 6 0.9) SVL [12.9 fideFischer, 1885]; HLL 20.1–24.4% (29.7 63.9) AGD, 17.4–22.0% (19.8 6 1.4) SVL[22.4 fide Fischer, 1885]; order of digits fromshortest to longest for hand: V 5 I , II 5 IV, III, for foot: I , V , II , III , IV; tail notas wide as body, gradually tapered toward theend, TW 57.4–76.8% (68.5 6 5.7) MBW, TL64.6–102.6% (91.6 6 10.8) SVL.

Rostral projecting dorsoposteriorly to pointin line with anterior edge of nasal, broaderthan high, separated from frontonasal; fronto-nasal wider than long; nostril ovoid, centeredin a single rectangular nasal; supranasalspresent, large, in broad medial contact;postnasals present; prefrontals moderatelyseparated by frontal; frontal nearly diamondshaped, its anterior margin in moderatecontact with frontonasal, in contact with firsttwo anterior supraoculars, 53 wider thananteriormost supraocular; supraoculars five;frontoparietals moderate, broad contact me-dially, each frontoparietal in contact withsupraoculars 2–4; interparietal moderate,quadrilaterally shaped, longer than wide, itslength greater than midline length of fronto-parietal; parietal eyespot present in posteriorhalf of scale; parietals in moderate to broadcontact behind interparietal or moderatelyseparated; nuchals undifferentiated; lorealstwo, decreasing in size from anterior to


posterior, subequal, in contact with prefrontal,postnasal, supranasal, second supralabial, pos-terior loreal and frontonasal; preocular single,nearly two thirds as high as posterior loreal;presubocular single; supraciliaries six, theanteriormost contacting prefrontal and sepa-rating posterior loreal from first supraocular;subocular row complete; lower eyelid withone row of scales, lacking an enlarged ovalwindow, largely transparent; supralabials sixor seven, fifth and sixth beneath center of eye;infralabials six or seven.

Mental wider than long, in contact with firstinfralabials; single enlarged postmental, widerthan mental, followed by two pairs of enlargedchin shields; first pair in slight contact ornarrowly separated by single undifferentiatedscale, second pair narrower than first, broadlyseparated by undifferentiated scales.


Coloration in preservative.—Ground colorof body medium brown; dorsal surfaces nearlyall dark brown, gradually fading into mediumbrown lateral and ventral surfaces of body;dark dorsal pigmentation in nearly continuousblock across dorsal surface, spanning six fulland two half rows of scales at midbody andnarrowing to cover four full and two half rowsof scales posterior to parietals; lateral surfaceswith one to two irregular dark brown lines onposterior half of axilla–groin region; headscales uniform dark brown; rostral, nasal,postnasal, supranasal, first supralabial, mental,and first infralabial dark gray; pineal eyespotpoorly defined, surrounded by light creamborder. Tail coloration matches body colora-tion. Limbs mottled dark brown dorsally,medium brown ventrally; dorsal and ventralsurface of digits dark brown.

Coloration in life.—Dorsal ground colorhomogeneous dark brown (Fig. 11E);blotched, irregular, lateral demarcation be-tween dorsal and lighter lateral and ventralcoloration; lateral and ventral surfaces of body

medium brown; lateral surfaces with irregu-larly shaped rows of dark brown spots. Limbsdark brown dorsally, medium brown ventrally.Dorsal head scales blotched dark and mediumbrown.

Variation.—Morphometric variation of theseries is summarized in Table 6. We observedvariation among the 36 specimens examinedfor the degree of contact between head scales.Twenty-one specimens were observed to haveparietals moderately separated by the inter-parietal (KU 314969, 314976, 314984–5,314988–9, 314992, 314997; MCZ 26552–3,26556–8, 26561, 26563, 26566, 26568, 26571–2, 26574), one specimen has parietals narrow-ly separated by the interparietal (KU 314996),and 14 specimens have parietals in moderatemedial contact (KU 314967, 314970–5,314977–8, 314980, 314990–1, 314994; MCZ26555) behind the interparietal; seven speci-mens do not have the first pair of enlargedchin shields in medial contact (KU 314971,314973, 314976, 314990, 314992, 314994;MCZ 26552, 26554, 26563), one specimenhas the first pair of enlarged chin shields inpoint medial contact (KU 314997), and 26specimens have the first pair of enlarged chinshields in moderate medial contact (KU314967, 314969, 314970, 314972, 314974–5,314977–8, 314980, 314984–5, 314988–9,314991, 314996; MCZ 26553, 26555–8,26561, 26566, 26568, 26571–2, 26574).

Scale counts were observed to vary amongthe measured series. The number of suprala-bials varied between six (CAS 23495, KU314991) and seven (CAS 23468–9, 23471,23479–81, 23484–5, 23494, 23496, 60493, KU314967, 314969, 314974–5, 314977–8,314980, 314984–5, 314994, 314996); infrala-bials varied between six (KU 314967, 314969,314974–5, 314977, 314980, 314984–5,314991, 314996) and seven (CAS 23468–9,23471, 23479–81, 23484–5, 23494–6, 60493).Specimens were observed to have midbodyscale row counts of 26 (CAS 23468, 23479–81,23494–6, 60493), 27 (CAS 23469, 23484), and28 (CAS 23471, 23485); axilla–groin scale rowcounts of 45 (CAS 23495), 46 (CAS 23469,23494, 23496), 47 (CAS 23468, 23484), 48(CAS 23485, 60493), 49 (CAS 23471, 23480),and 50 (CAS 23479, 23481); and paravertebralscale row counts of 67 (CAS 23495), 68 (CAS


23494, 23496), 70 (CAS 23484), 71 (CAS23468–9, 23471, 23480, 23485, 60493), and 72(CAS 23479, 23481).

We also observed lamellae counts to varyamong the measured series. Specimens wereobserved to have Finger-III lamellae counts offive (CAS 23469, 23495, 60493, KU 314967,314969, 314974, 314977–8, 314984–5,314991, 314994, 314996) or six (CAS 23468,23471, 23479–81, 23484–5, 23494, 23496, KU314975, 314980); Toe-IV lamellae counts ofeight (CAS 23468, 23494–5, 60493, KU314967, 314969, 314974–5, 314977, 314984–5, 314991, 314994, 314996) or nine (CAS23469, 23471, 23479–81, 23484–5, 23496, KU314978, 314980).

There is a small degree of color variation inthe examined series, with the degree anddefinition of continuous, dark middorsalpigmentation. Most of the examined speci-mens show patterns consistent with a contin-uous, dark streak of pigmentation covering themiddorsal region of the body (KU 314969,314974–5, 314977–8, 314980, 314984–5,314991, 314994, 314996). In several speci-mens, continuous, dark middorsal stripes areevident overlaying the dark ground coloration(KU 314967).

Distribution.—Brachymeles schadenbergiis known from Basilan and western Mindanaoislands (Fig. 4).

Ecology and natural history.—Brachymelesschadenbergi occurs in a variety of habitatsfrom disturbed and secondary growth toprimary forest and intact climax forest.Individuals have been observed in the humusmaterial within rotting logs and in loose soiland leaf litter surrounding the root networksof trees. Individuals are moderately commonin populations sampled (CDS and RMB,personal observations) and occur sympatrical-ly with B. gracilis gracilis in western Mind-anao Island (Brown, 1956; Brown and Alcala,1980; Brown and Rabor, 1967). We collectednumerous specimens in pitfall traps, indicat-ing some level of surface activity. Although B.schadenbergi occurs on multiple islands in thesouthern Philippines, the species seems tohave a more restricted geographic distributioncompared with more widespread pentadactylspecies, such as B. boulengeri, B. talinis, B.orientalis, and B. kadwa. As in other members

of the genus, disturbed individuals move in arapid serpentine manner and always attemptto burrow back into loose soil or humus.

Sympatric lizard species observed within therange of Brachymeles schadenbergi include thefollowing: (Agamidae) Bronchocela cristatella,Draco bimaculatus, D. cyanopterus, D. mind-anensis, Gonocephalus interruptus, Hydro-saurus amboinensis; (Gekkonidae) Cyrtodac-tylus jambangan, Gehyra mutilata, Gekkogecko, Hemidactylus frenatus, H. platyurus,Hemiphyllodactylus typus, Lepidodactylus sp.,L. quadrivittata, Luperosaurus joloensis, Pseu-dogekko compressicorpus; (Scincidae) Brachy-meles gracilis gracilis, Eutropis indeprensa, E.multicarinata, E. multifasciata, E. englei, Lam-prolepis smaragdina, Sphenomorphus atrigu-laris, S. fasciatus, S. jagori, S. steerei, S.variegatus, Tropidophorus misaminus, T. par-telloi; and (Varanidae) Varanus cumingi.

Brachymeles talinis Brown 1956Figs. 5, 9, 11H

Brachymeles schadenbergi talinis, Brown,1956. Type locality: ‘‘On the low ridgenorth side of the Maite River, 5 to 6 kmwest of Valencia,’’ Negros Oriental Prov-ince, Negros Island, Philippines, 933-melevation, 9u 179 19.250 N, 123u 119 56.40E (holotype: CAS-SU 18358).

Brachymeles talinis, Brown and Rabor, 1967.Brachymeles talinis, Brown and Alcala, 1980.

Diagnosis.—Brachymeles talinis can bedistinguished from congeners by the followingcombination of characters: (1) body size large(SVL 103.8–123.1 mm); (2) pentadactyl; (3)Finger-III lamellae five or six; (4) Toe-IVlamellae 8–10; (5) limbs relatively long; (6)paravertebral scale rows 67–72; (7) suprala-bials seven; (8) infralabials seven; (9) pinealeye spot present; (10) supranasals in contact;(11) prefrontals not contacting on midline;(12) enlarged chin shields in two pairs; (13)nuchal scales undifferentiated; (14) fifth andsixth supralabial below eye; (15) auricularopening present; (16) dark lateral stripespresent; and (17) venter devoid of darkpigmentation (Tables 4 and 5).

Comparisons.—Characters distinguishingBrachymeles talinis from all pentadactylspecies of Brachymeles are summarized inTables 4 and 5. Brachymeles talinis most


closely resembles B. kadwa, B. makusog, B.tungaoi, and B. vindumi, but it differs fromthese four taxa by having the range ofparavertebral scale rows reaching .70 but,74, and seven infralabials (Table 5). Bra-chymeles talinis can further be distinguishedfrom B. kadwa by having 8–10 Toe-IVlamellae, the first enlarged chin shield wider

than the second, frontoparietals in contact,and by the absence of dark ventral pigmenta-tion (Tables 4 and 5); from B. makusog byhaving seven supralabials the fifth and sixthsupralabial below the eye, supranasals incontact, and by the presence of dark lateralstripes (Tables 4 and 5); from B. tungaoi byhaving a larger body size, shorter relative tail

FIG. 9.—Illustration of head of adult male Brachymeles talinis (KU 306769) and adult male holotype of Brachymeleskadwa (PNM 9721; formerly KU 323091) in dorsal, lateral, and ventral views. Labels for taxonomically diagnostic headscales follow those shown in Fig. 6. Illustrations by CDS.


length, 8–10 Toe-IV lamellae, and the firstenlarged chin shield wider than the second(Tables 4 and 5); and from B. vindumi byhaving fewer axilla–groin scale rows, fewerparavertebral scale rows, and by the absenceof dark ventral pigmentation (Table 5).

From all nonpentadactyl species of Brachy-meles (B. apus, B. bonitae, B. cebuensis, B.elerae, B. lukbani, B. minimus, B. muntingka-may, B. pathfinderi, B. samarensis, B. tridac-tylus, B. vermis, and B. wrighti), B. talinisdiffers by having a pentadactyl body form (vs.nonpentadactyl), longer forelimb lengths(.11.3 mm vs. ,6.9 mm), and greater hindlimb lengths (.20.5 mm vs. ,12.9 mm), andby the presence of a postnasal scale (vs.absence). In addition, B. talinis differs from allnonpentadactyl species except B. wrighti byhaving a midbody scale row count 26–30 (vs.,24); from all nonpentadactyl species exceptB. pathfinderi by having a paravertebral scalerow count 68–70 (vs. .84), and by thepresence of auricular openings (vs. absence);from all nonpentadactyl species except B.apus and B. wrighti by having a larger bodysize (SVL .103.1 mm vs. ,81.3 mm); andfrom B. apus, B. lukbani, B. minimus, and B.vermis by the presence (vs. absence) of limbs.

Description (based on holotype and 30referred specimens, including two paratypesfrom CAS).—Details of the head scalation ofan adult male are shown in Fig. 9. Measure-ments of the holotype are provided below inbrackets. Body large relative to other Brachy-meles, elongate with respect to other lizards;maximum SVL 123.1 mm for males, 116.5 mmfor females [118.7, male] (Tables 4 and 5);head weakly differentiated from neck, nearlyas wide as body, HW 10.6–13.2% (11.8 6 0.5)SVL [12.1], 111.7–136.2% (124.1 6 6.8) HL[130.3]; HL 30.6–40.7% (35.4 6 2.7) SnFa[33.5]; SnFa 25.1–29.8% (27.0 6 1.3) SVL[27.6]; snout moderately long, broadly round-ed in dorsal and lateral profile, SNL 51.5–65.8% (58.9 6 4.0) HL [65.8]; auricularopening present, moderate; eyes moderate,ED 1.7–2.2% (1.9 6 0.1) SVL [1.7], 17.4–24.8% (20.0 6 1.9) HL [18.2], 40.9–62.1%(50.3 6 5.7) END [41.5], pupil nearly round;body slightly depressed, MBW 109.3–153.8%(126.7 6 14.4) MBH [109.6]; scales smooth,glossy, imbricate; longitudinal scale rows at

midbody 26–30 [29]; paravertebral scale rows67–72 [72]; axilla–groin scale rows 43–48 [48];limbs well developed, pentadactyl, digitsmoderate; FinIIIlam 5–6 [6]; ToeIVlam 8–10[10]; FLL 15.1–23.9% (19.1 6 1.8) AGD[19.6], 10.1–15.3% (12.4 6 1.2) SVL [14.9];HLL 26.8–38.9% (31.8 6 3.0) AGD [30.9],18.0–24.9% (20.6 6 1.8) SVL [23.5]; order ofdigits from shortest to longest for hand: V , I, IV , II , III, for foot: I 5 V , II , III 5IV; tail nearly as wide as body at base, sharplytapered toward the end, TW 63.4–94.0% (75.16 6.2) MBW [94.0], TL 60.6–107.2% (83.9 612.0) SVL [73.2].

Rostral projecting dorsoposteriorly to pointin line with anterior edge of nasal, broaderthan high, completely separated from fronto-nasal by broad supranasal contact; frontonasalwider than long; nostril ovoid, in center ofsingle trapezoidal nasal; supranasals present,large, in broad medial contact; postnasalspresent; prefrontals moderately separated byfrontal; frontal nearly octagonal shaped, itsanterior margin in moderate contact withfrontonasal, in contact with first two anteriorsupraoculars, 43 wider than anteriormostsupraocular; supraoculars five; frontoparietalsmoderate, in moderate medial contact, eachfrontoparietal in contact with supraoculars2–4; interparietal moderate, quadrilaterallyshaped, width nearly equal to length, itslength nearly equal to midline length offrontoparietal; parietal eyespot present inposterior one third of scale; parietals in pointto moderate contact behind interparietal ornarrowly separated; nuchals undifferentiated;loreals two, decreasing in size from anterior toposterior, subequal, in contact with prefrontal,postnasal, supranasal, second supralabial, pos-terior loreal and frontonasal; preocular single,nearly three fourths as high as posterior loreal;single presubocular; supraciliaries six, theanteriormost contacting prefrontal and sepa-rating posterior loreal from first supraocular;subocular row complete; lower eyelid withone row of scales, lacking an enlarged ovalwindow, largely transparent; supralabials sev-en, fifth and sixth below the eye; infralabialsseven.

Mental wider than long, in contact with firstinfralabials; single enlarged postmental, widerthan mental; followed by two pairs of enlarged


chin shields; first pair in moderate contact ormoderately separated by a single medial scale,wider than second pair; second pair separatedby three undifferentiated scales.


Coloration in preservative.—Ground colorof body medium brown; longitudinal stripeson dorsal surface of body present or absent;when present a total of eight longitudinal darkbrown spot rows, extending from posterioredge of parietals to base of tail: six continuousmedial rows and two discontinuous postero-lateral rows, together spanning eight full rowsof scales at midbody, narrowing to six full rowsof scales posterior to parietals; when dark spotrows are absent, pigmentation forms nearlycontinuous dark dorsal surface, covering onehalf to entire surface of dorsal scales; dorso-lateral stripes present or absent, when pres-ent, well defined, continuous, lacking darkpigmentation, spanning two whole and twohalf row of scales from auricular opening tobase of tail. Lateral and ventral surface ofbody medium brown. Lateral surface withthree to six discontinuous longitudinal rows ofdark brown spots, rows often extending toedge of ventral surface. Ventral surfacewithout dark pigmentation. Tail colorationequal to body coloration, dorsal surfacecovered with dark brown blotches, ventralsurface covered with scattered dark brownspots, fewer than dorsal surface. Head scaleshomogeneous dark brown; rostral, nasal,postnasal, supranasal, first supralabial, mentaland first infralabial dark gray; pigment sur-rounding pineal eyespot reduced to indistinct,small and medium brown. Limbs mottledmedium brown dorsally, yellowish brownventrally; dorsal and ventral surface of digitsdark brown.

Coloration in life.—Dorsal ground colormedium brown (Fig. 11H); when present,longitudinal rows of spots dark brown toblack; dorsolateral stripes light to medium

brown, bordered middorsally by rows of darkspots; lateral surface ground color light brownto tan; ventral surfaces of body light brown totan. Dorsal surfaces of limbs dark to mediumbrown, ventral surfaces light brown. Dorsalhead scales blotched dark and medium brown.

Variation.—Morphometric variation of theseries is summarized in Table 6. We observedvariation among the 19 specimens examinedfor the degree of contact between head scales.Four specimens were observed to haveparietals moderately separated by the inter-parietal (KU 306757, 306763, 306767,306786), one specimen has parietals narrowlyseparated by the interparietal (KU 306758),and 14 specimens have parietals in moderatemedial contact (KU 306756, 306759–60,306762, 306764–6, 306769–71, 306773–6)behind the interparietal; 13 specimens donot have the first pair of enlarged chin shieldsin medial contact (KU 304756–7, 306759–60,306762, 306764–6, 306767, 306769–70,306774–5) and six specimens have the firstpair of enlarged chin shields in moderatemedial contact (KU 306758, 306763, 306771,306773, 306776, 306786).

Scale counts were observed to vary amongthe measured series. Specimens were ob-served to have midbody scale row counts of26 (KU 306766), 28 (CAS-SU 22311, 22317,37996, KU 306765), 29 (CAS-SU 12225,18358, 22323, 27972, 89813, CAS 133871,KU 306758, 306774), and 30 (CAS-SU 22312,27997, KU 306756, 306760, 306769, 306772–3,306786); axilla–groin scale row counts of 43(KU 306786), 44 (CAS-SU 12225, 22311,27996–7), 45 (CAS-SU 22323, KU 306756,306758, 306760, 306765–6, 306772), 46 (CAS-SU 22312, 22317, 27972, 89813, CAS 133871,KU 306773), 47 (KU 306774), and 48 (CAS-SU18358, KU 306769); and paravertebral scalerow counts of 67 (KU 306786), 68 (CAS-SU12225, 22311, 27996–7, KU 306756, 306758),69 (CAS-SU 22312, 22323, KU 306760,306765–6, 306772–3), 70 (CAS-SU 22317,37972, 89813, CAS 133871, KU 306769), 71(KU 306774), and 72 (CAS-SU 18358).

We also observed lamellae counts to varyamong the measured series. With the excep-tion of two specimens observed to have sixFinger-III lamellae (CAS-SU 18358, CAS133871), all other examined specimens were


observed to have five. We also observed Toe-IV lamellae counts of eight (KU 306769,306786), nine (CAS-SU 22311, 22317,89813, KU 306756, 306758, 306760, 306765–6, 306772–4), and 10 (CAS-SU 12225, 18358,22312, 22323, 27972, 27996–7, CAS 133871).

Color variation exists in the degree anddefinition of continuous, dark middorsalstripes. Many specimens show patterns con-sistent with continuous, middorsal dark lines(KU 306651, 306756–7, 306759, 30676,2306765–7, 306769–72, 306776, 306786,306763). The dark lines are obscured in someand irregular in others, where the middorsalregion is covered by a long streak of darkpigmentation, with little to moderate linedefinition (KU 306759–60, 306764, 306774).

Distribution.—Brachymeles talinis isknown from Negros, Panay, Romblon, Sibu-yan, and Tablas islands (Fig. 5). It is also likelyto occur on Guimaras Island.

Ecology and natural history.—Brachymelestalinis occurs in a variety of habitats fromagricultural areas, to disturbed and second-ary growth forest. Little or no original,lowland forest remains in the Visayas, butwe assume the species originally occurred inprimary forest. Individuals have been ob-served under piles of rotting coconut husks,in the humus material within rotting logs,and in loose soil and leaf litter surroundingthe root networks of trees. The species ismoderately common throughout its range(CDS, personal observation) and occurssympatrically with three other species (B.bonitae, B. talinis, and B. tridactylous;Brown, 1956; Brown and Alcala, 1980;Brown and Rabor, 1967). Individuals wereoften encountered in pitfall traps, indicatingsome level of activity outside of fossorialmicrohabitats. Similar to B. boulengeri, B.talinis seems to have a wider geographicdistribution that spans multiple Philippineislands. This is in contrast to the islandendemic species B. boholensis, B. tungaoi,and B. mindorensis that are known from justBohol, Masbate, and Mindoro islands, re-spectively. As do all members of the genus,when disturbed, individuals attempt toescape by moving in a rapid serpentinemanner and attempting to burrow back intoloose soil or humus.

Sympatric lizard species observed withinthe range of Brachymeles talinis include thefollowing: (Agamidae) Bronchocela marmor-atus, Draco spilopterus, Hydrosaurus pustu-latus; (Dibamidae) Dibamus argenteus; (Gek-konidae) Cyrtodactylus philippinicus, Gehyramutilata, Gekko gecko, Gekko mindorensis,Gonocephalus sophiae, Hemidactylus frena-tus, H. platyurus, Hemiphyllodactylus typus,Lepidodactylus christiani, L. herrei, L. lugu-bris, Luperosaurus corfieldi, Pseudogekkobrevipes; (Scincidae) Brachymeles tridactylus,B. taylori, Emoia atrocostata, Eutropis multi-carinata, E. multifasciata, Lamprolepis smar-agdina, Lipinia auriculata, L. pulchella, L.quadrivittata, L. rabori, Sphenomorphus ar-borens, S. coxi, S. jagori, S. steerei, Tropido-phorus grayi; and (Varanidae) Varanus nu-chalis.

Brachymeles kadwa sp. nov.Figs. 5, 9, 11D

Holotype.—PNM 9721 (RMB FieldNo. 12466, formerly KU 323091), adult male,collected under rotting logs in secondary-growth forest (1000 to 1230 h) on 4 June 2009,on the campus of Aurora State College ofTechnology, Barangay Zabali, Municipality ofBaler, Aurora Province, Luzon Island, Philip-pines (15u 449 310 N, 121u 349 340 E; WGS-84), by CDS, RMB, J. Fernandez, L. Welton,J. Brown, J. Siler, Y. Vicente, and M. Vicente.

Paratopotypes.—Three adult males (KU323092, 323095, 323096) and four adult fe-males (KU 323106, 323094, 323104, 323100),collected between 4 and 7 June 2009.

Paratypes.—Four adult males (KU 304875,304900, 304915, 304941) and six adult females(KU 304897, 304902–3, 304905–6, 304929)collected between 15 and 22 March 2006 (19u179 380 N, 121u 249 320 E; WGS-84; 245 mabove sea level) Barangay Magsidel, Munici-pality of Calayan, Cagayan Province, CalayanIsland, Philippines, by RMB, C. Oliveros, andJ. Fernandez; four adult males (KU 304575,307984, 307996, 307998) and five adultfemales (KU 304559, 304593, 304708,304754, 308011), four adult males and fiveadult females collected between 3 and 11March 2006 from 300-m elevation (18u 559 450N, 121u 539 560 E; WGS-84) BarangayBalatubat, Municipality of Calayan, Cagayan


Province, Camiguin Norte Island, Philippines,by RMB, C. Oliveros, and J. Fernandez.

Referred specimens.—CALAYAN IS-LAND: CAGAYAN PROVINCE: Municipality ofCalayan: Barangay Magsidel: KU 304908,304899, 304907, 304909, 304921, 304941;CAMIGUIN NORTE ISLAND: CAGAYAN

PROVINCE: Municipality of Calayan: BarangayBalatubat: KU 304558, 304562–65, 304569,304571–74, 304627–30, 304643, 304647,304696–99, 304704–07, 304709–12, 304714,304753, 304755–59, 307965–66, 307985–86,307997, 307999–8003, 308006–10, 308012–15, 308017–18; LUZON ISLAND: AURORA

PROVINCE: Municipality of Baler: BarangayZabali, ASCOT: KU 323090–91, 323093,323097–99, 323101–03, 323105, 323107; Mu-nicipality of Casiguran, IDC property: KU323108–48; Municipality of San Luis: Baran-gay Real, Sitio Minoli: KU 322320.

Diagnosis.—Brachymeles kadwa can bedistinguished from congeners by the followingcombination of characters: (1) body size large(SVL 90.6–128.2 mm); (2) pentadactyl; (3)Finger-III lamellae five or six; (4) Toe-IVlamellae 7–10; (5) limbs relatively long; (6)paravertebrals 68–70; (7) supralabials seven;(8) infralabials six; (9) pineal eye spot present,small; (10) supranasals in contact; (11) pre-frontals not contacting on midline; (12)enlarged chin shields in two pairs; (13) nuchalscales undifferentiated; (14) fifth and sixthsupralabial below eye; (15) auricular openingpresent; (16) continuous, light dorsolateralstripes present, indistinct; (17) continuous,dark middorsal stripes present; (18) darklateral stripes present; and (19) dark ventralpigmentation present (Tables 4 and 5).

Comparisons.—Characters distinguishingthe new species from all pentadactyl speciesof Brachymeles are summarized in Tables 4and 5. Brachymeles kadwa most closelyresembles B. makusog, B. tungaoi, B. talinis,and B. vindumi, but it differs from these fourtaxa by having 7–10 Toe-IV lamellae and thesecond enlarged chin shield wider than thefirst (Tables 4 and 5). Brachymeles kadwa canfurther be distinguished from B. makusog byhaving seven supralabials, the fifth and sixthsupralabial below the eye, six infralabials, thepresence of supranasal contact, the presenceof continuous, light dorsolateral stripes, con-

tinuous, dark middorsal stripes, dark lateralstripes, and dark ventral pigmentation (Ta-ble 5); from B. tungaoi by having a greatermidbody width, shorter relative tail length,paravertebrals 68–70, and the presence ofdark ventral pigmentation (Tables 4 and 5);from B. talinis by having 28 or fewer midbodyscale rows, 70 or fewer paravertebrals, infra-labials six, and by the presence dark ventralpigmentation (Table 5); and from B. vindumiby having five or six Finger-III lamellae, 26–28 midbody scale rows, paravertebrals 68–70,and by the presence of continuous, darkmiddorsal stripes (Tables 4 and 5).

From all nonpentadactyl species of Brachy-meles (B. apus, B. bonitae, B. cebuensis, B.elerae, B. lukbani, B. minimus, B. muntingka-may, B. pathfinderi, B. samarensis, B. tridac-tylus, B. vermis, and B. wrighti), B. kadwadiffers by having a pentadactyl body form (vs.nonpentadactyl), longer forelimb lengths(.10.7 mm vs. ,6.9 mm), and greater hindlimb lengths (.17.9 mm vs. ,12.9 mm), andby the presence of a postnasal scale (vs.absence). In addition, B. kadwa differs fromall nonpentadactyl species except B. wrightiby having 26–28 midbody scales (vs. ,24);from all nonpentadactyl species except B.pathfinderi by having 68–70 paravertebrals(vs. .84), and by the presence of auricularopenings (vs. absence); from all nonpentadac-tyl species except B. apus and B. wrighti byhaving a larger body size (SVL .90.6 mm vs.,81.3 mm); and from B. apus, B. lukbani, B.minimus, and B. vermis by the presence oflimbs (vs. absence).

Description of holotype.—Mature male,hemipenes everted (Fig. 10); SVL 106.2 mm;body moderately large relative to otherBrachymeles, elongate with respect to otherlizards; head weakly differentiated from neck,nearly as wide as body, HW 11.0% SVL,111.1% HL; HL 38.1% SnFa; SnFa 26.0%SVL; snout moderately long, rounded indorsal and lateral profile, SNL 56.1% HL;auricular opening present, small; eyes moder-ate, ED 1.9% SVL, 19.6% HL, 54.8% END,pupil nearly round; body slightly depressed,MBW 157.3% MBH; body scales smooth,glossy, imbricate; longitudinal scale rows atmidbody 28; paravertebral scale rows 68;axilla–groin scale rows 47; limbs well devel-


oped, pentadactyl, digits moderate; FinIIIlam5; ToeIVlam 10; FLL 20.3% AGD, 13.0%SVL; HLL 32.0% AGD, 20.6% SVL; order ofdigits from shortest to longest for hand: I 5 V, II 5 IV , III, for foot: V , I , II , III 5IV; tail nearly as wide as body at base,gradually tapered toward the end, TW 73.5%MBW, TL 101.6% SVL.

Rostral projecting dorsoposteriorly to pointin line with anterior edge of nostril, broaderthan high, separated from frontonasal bymoderate contact of supranasals; frontonasalwider than long; nostril ovoid, centered in asingle rectangular nasal; supranasals large, inmoderate medial contact; postnasals present;prefrontals moderately separated by frontal;

FIG. 10.—Illustration of head of adult male holotype of Brachymeles tungaoi (PNM 9722; formerly KU 323933) andadult male holotype of Brachymeles vindumi (CAS 60724) in dorsal, lateral, and ventral views. Labels for taxonomicallydiagnostic head scales follow those shown in Fig. 6. Illustrations by CDS.


FIG. 11.—Photographs in life of (A) Brachymeles boulengeri (KU 307756), snout–vent length (SVL) 5 98.0 mm; (B)B. taylori (RMB 3283, deposited at Philippine National [PNM]), SVL 5 81.0 mm; (C) B. boholensis (RMB 2877,deposited at PNM), female, SVL 5 89.0 mm; (D) B. kadwa (KU 304593), SVL 5 101.0 mm; (E) B. schadenbergi (KU314973), female, SVL 5 107.0 mm; (F) B. orientalis (KU 311240), juvenile, SVL 5 51.0 mm; (G) B. orientalis (KU324029), female, SVL 5 91.0 mm; and (H) Brachymeles talinis (RMB 3305; deposited at PNM), SVL 5 139.0 mm.Photographs by CDS and RMB. To view figure in color, please refer to digital online version.


frontal nearly diamond shaped, its anteriormargin in moderate contact with frontonasal,in contact with first two anterior supraoculars,43 wider than anteriormost supraocular;supraoculars five; frontoparietals moderate,point contact medially or moderately separat-ed, each frontoparietal in contact with suprao-culars 2–4; interparietal moderate, quadrilat-erally shaped, its length slightly greater thanmidline length of frontoparietal; parietaleyespot present in posterior one third of scale,indistinct; parietals in moderate contact be-hind interparietal; nuchals undifferentiated;loreals two, decreasing in size from anterior toposterior, subequal, in contact with prefrontal,postnasal, supranasal, second supralabial, pos-terior loreal and frontonasal; preocular single,nearly two thirds as high as posterior loreal;single presubocular; supraciliaries six, theanteriormost contacting prefrontal and sepa-rating posterior loreal from first supraocular,posteriormost extending to midline of lastsupraocular; subocular row complete; lowereyelid with one row of scales, lacking anenlarged oval window, largely transparent;supralabials seven, fifth and sixth below theeye; infralabials six.

Mental wider than long, in contact with firstinfralabials; single enlarged postmental, slight-ly wider than mental; followed by two pairs ofenlarged chin shields, first pair in moderatemedial contact, second pair slightly wider thanfirst, separated by a single undifferentiatedscale.

Scales on limbs smaller than body scales;scales on dorsal surfaces of digits large,wrapping around lateral edges of digits;lamellae undivided; palmar surfaces of handsand plantar surfaces of feet covered by small,irregular scales, each with raised anterioredges; scales on dorsal surface of hands andfeet smaller than limb scales, lacking raisededges.

Coloration in preservative.—Ground colorof body dark brown; dorsal surface of bodywith eight longitudinal rows of dark brownspots spanning eight full rows of scales atmidbody and extending from posterior edge ofparietals to base of tail: six rows in middorsalregion, flanked by discontinuous dorsolateralrows; spot rows narrowing to six full rows ofscales posterior to parietals; dark coloration

covering middle three fourths of dorsal scales;dorsolateral stripes somewhat indistinct, dis-continuous, spanning two half rows of scalesfrom auricular opening point just posterior toforelimb insertion; dark dorsal colorationblends gradually into medium brown lateraland ventral surface of body. Lateral surfacewith six discontinuous, dark brown spot rows,extending to edge of ventral surface. Ventralsurface with scattered dark brown spots. Tailcoloration similar to body coloration, dorsalsurface covered with dark brown blotches,ventral surface covered with few dark brownspots. Head scales homogeneous dark brown;rostral, nasal, postnasal, supranasal, firstsupralabial, mental, and first infralabial darkgray; pineal eyespot indistinct, small and lightbrown. Limbs mottled dark brown dorsally,yellowish brown ventrally; dorsal and ventralsurface of digits dark brown.

Coloration in life.—Ground color of bodylight to medium brown (Fig. 11D); dorsalsurfaces of limbs medium brown.

Measurements of holotype (in millime-ters).—SVL 106.2; AGD 68.2; TotL 214.0;MBW 15.9; MBH 10.1; TL 107.9; TW 11.7;TH 9.2; HL 10.5; HW 11.7; HH 8.6; SnFa27.6; ED 2.1; END 3.8; SNL 5.9; IND 3.7;FLL 13.8; HLL 21.8; MBSR 28; PVSR 68;AGSR 47; FinIIIlam 5; ToeIVlam 10; SL 7;IFL 6; SC 6; SO 5.

Variation.—Morphometric variation of theseries is summarized in Table 6. We observedvariation among the 25 specimens examinedfor the degree of contact between head scales.Fourteen specimens were observed to haveparietals moderately separated by the inter-parietal (KU 304559, 304574–5, 304593,304630, 304708, 304754–5, 304759, 304906,307984–5, 307996, 308007), one specimen hasparietals in point medial contact (KU 308011),and 10 specimens have parietals in moderatemedial contact (KU 304875, 304897, 304900,304902–3, 304905, 304915, 304929, 304941,307998) behind the interparietal; two speci-mens have frontoparietals moderately sepa-rated by the frontal (KU 304559, 307984), onespecimen has frontoparietals narrowly sepa-rated by the frontal (KU 304574), and 22specimens have frontoparietals in moderatemedial contact (KU 304575, 304593, 304630,304708, 304754–5, 304759, 304875, 304897,


304900, 304902–3, 304905–6, 304915,304929, 304941, 307985, 307996, 307998,308007, 308011). We observed the first pairof enlarged chin shields narrowly separated ina single specimen (KU 307996) and inmoderate contact for all other examinedspecimens.

Scale counts were observed to vary amongthe measured series. Specimens were ob-served to have midbody scale row counts of 26(KU 304559, 307996, 323104), 27 (KU304593, 307984, 307998), and 28 (KU304575, 304708, 304754, 304875, 304897,304900, 304902–3, 304905–6, 304915,304929, 304941, 308011, 323091, 323091–2,323094–6, 323100, 323106); axilla–groin scalerow counts of 47 (KU 304559, 304575,304593, 304875, 304902, 304929, 307996,307998, 308011, 323091, 323096), 48 (KU304708, 304754, 304897, 304900, 304915,304941, 307984, 323092, 323094–5, 323100,323104, 323106), 49 (KU 304903, 304905–6);and paravertebral scale row counts of 68 (KU304559, 304593, 304900, 307996, 323091,323096), 69 (KU 304575, 304708, 304754,304875, 304929, 304941, 307984, 307998,308011, 323092, 323094–5, 323100, 323104,323106), and 70 (KU 304897, 304902–3,304905–6, 304915).

We also observed lamellae counts to varyamong the measured series. With the excep-tion of two specimens observed to have sixFinger-III lamellae (KU 304903, 304906), allother examined specimens were observed tohave five. We also observed Toe-IV lamellaecounts of seven (KU 304593), eight (KU304559, 304575, 304708, 304754, 304875,304897, 304900, 304915, 304929, 304941,307984, 307996, 307998, 308011), nine (KU304902–3, 304905–6, 323092, 323094–6,323100, 323104, 323106), or 10 (KU 323091).

Distribution.—Brachymeles kadwa isknown from numerous localities on LuzonIsland as well as from Calayan and CamiguinNorte Islands of the Babuyan Island group(Fig. 5).

Ecology and natural history.—Brachymeleskadwa occurs in agricultural areas, disturbedsecondary growth forest, and first growthforests of Luzon, Camiguin Norte, andCalayan. Individuals have been observedunder piles of rotting coconut husks, in the

humus material within rotting logs, and inloose soil and leaf litter surrounding the rootnetworks of trees. This species is quitecommon in all sampling localities, and wehave taken large series in pitfall traps,indicating some level of surface activity. Whendisturbed, individuals immediately moved in arapid serpentine manner and attempted toburrow back into loose soil or humus.

Sympatric lizard species observed on Lu-zon, Camiguin Norte, and Calayan islandsinclude the following: (Agamidae) Broncho-cela cristatella, Draco spilopterus, Gonoce-phalus sophiae, Hydrosaurus pustulatus;(Gekkonidae) Cyrtodactylus philippinicus,Gehyra mutilata, Gekko gecko, Gekko mind-orensis, Hemidactylus frenatus, H. garnoti, H.luzonensis, H. platyurus, Luperosaurus cf.cumingi, L. kubli, Pseudogekko compressicor-pus, P. smaragdina; (Scincidae) Brachymelesbonitae, B. bicolor, B. elerae, B. lukbani, B.makusog, B. muntingkamay, B. samarensis, B.cf. talinis, B. wrighti, Emoia atrocostata,Eutropis bontocensis, E. multicarinata, E.multifasciata, Lamprolepis smaragdina, Lipi-nia pulchella, Sphenomorphus cumingi, S.decipiens, S. jagori, S. leucospilos, S. luzonen-sis, S. steerei, S. stejnegeri, Tropidophorusgrayi; and (Varanidae) Varanus marmoratus.

Etymology.—CDS is pleased to name thisnew species for his loving wife Jessi M. Silerfor her endless support that has made all ofthis research possible. The name of the newspecies is derived from one of the localdialects spoken in the Philippines. The word‘‘kadwa’’ is the Ilonggo term for friend andcompanion. Suggested common name: Jessi’sslender skink.

Brachymeles tungaoi sp. nov.Figs. 5, 10

Holotype.—PNM 9722 (CDS FieldNo. 5125, formerly KU 323933), adult male,collected in rotting stump in disturbed,residential habitat (1000 to 1230 h) 4 Sep-tember 2009, at 61-m elevation in Municipal-ity of Masbate City, Masbate Province,Masbate Island, Philippines (12u 219 010 N,123u 379 420 E; WGS-84), by CDS and J.Fernandez.

Paratopotypes.—KU 323934–36, threeadult females, collected between 3 and 7


September 2009, from 61–99-m elevation byCDS and J. Fernandez.

Paratypes.—One adult male (CAS 144313),three adult females (CAS 144229–30,144341), and four juvenile specimens ofunknown sex (CAS 144290, 144306–7,144342), collected 2 June 1976 ‘‘in humusunder rotting log,’’ in Barangay Tugbo,Municipality of Mobo, Masbate Province,Masbate Island, Philippines (12u 209 11.040N, 123u 379 58.80 E; WGS-84; 400-m eleva-tion) by A. Alcala.

Diagnosis.—Brachymeles tungaoi can bedistinguished from congeners by the followingcombination of characters: (1) body sizemoderate (SVL 78.2–106.2 mm); (2) relativetail length long; (3) pentadactyl; (3) Finger-IIIlamellae five or six; (4) Toe-IV lamellae nineor 10; (5) limb length moderate; (6) paraver-tebral scale rows 66–68; (7) supralabials seven;(8) infralabials six; (9) pineal eye spot present,large; (10) supranasals in contact; (11) pre-frontals not contacting on midline; (12)contact between first pair of chin shields;(13) enlarged chin shields in two pairs; (14)nuchal scales undifferentiated; (15) fifth andsixth supralabial below eye; (16) auricularopening present; (17) continuous, light dorso-lateral stripes present, indistinct; (18) contin-uous, dark middorsal stripes present; (19) darklateral stripes present; and (20) dark ventralpigmentation absent (Tables 4 and 5).

Comparisons.—Characters distinguishingthe new species from all pentadactyl speciesof Brachymeles are summarized in Tables 4and 5. Brachymeles tungaoi most closelyresembles B. kadwa, B. makusog, B. talinis,and B. vindumi, but it differs from these fourtaxa by having a smaller body size, smallermidbody width, greater relative tail length, thefirst and second pairs of enlarged chin shieldsequal in width, and contact between the firstpair of enlarged chin shields (Tables 4 and 5).Brachymeles tungaoi can be further distin-guished from B. kadwa by having nine or 10Toe-IV lamellae, paravertebral scale rows 66–68, frontoparietal in contact, and by theabsence of dark ventral pigmentation (Ta-bles 4 and 5); from B. makusog by havingseven supralabials, six infralabials, the fifthand sixth supralabial below the eye, suprana-sals in moderate contact, the presence of

continuous, light dorsolateral stripes, contin-uous, dark middorsal stripes, and dark lateralstripes (Table 5); from B. talinis by havingnine or 10 Toe-IV lamellae, 66–68 paraverte-brals, infralabials six (Tables 4 and 5); andfrom B. vindumi by having five or six Finger-III lamellae, 26–28 midbody scale rows, and66–68 paravertebrals (Tables 4 and 5).

From all nonpentadactyl species of Brachy-meles (B. apus, B. bonitae, B. cebuensis, B.elerae, B. lukbani, B. minimus, B. muntingka-may, B. pathfinderi, B. samarensis, B. tridac-tylus, B. vermis, and B. wrighti), B. tungaoidiffers by having a pentadactyl body form (vs.nonpentadactyl), longer forelimb lengths(.11.0 mm vs. ,6.9 mm), and greater hindlimb lengths (.17.0 mm vs. ,12.9 mm), andby the presence of a postnasal scale(vs. absence). In addition, Brachymeles tun-gaoi differs from all nonpentadactyl speciesexcept B. wrighti by having a midbody scalerow count 26–28 (vs. ,24); from all non-pentadactyl species except B. pathfinderi byhaving a paravertebral scale row count 66 (vs..84), and by the presence of auricularopenings (vs. absence); and from B. apus, B.lukbani, B. minimus, and B. vermis by thepresence of limbs (vs. absence).

Description of holotype.—Mature male,hemipenes everted (Fig. 10); SVL 89.2 mm;body moderate relative to other Brachymeles,elongate with respect to other lizards; headweakly differentiated from neck, nearly aswide as body, HW 11.3% SVL, 115.9% HL;HL 38.0% SnFa; SnFa 25.6% SVL; snoutmoderately long, bluntly rounded in dorsalprofile, sharply rounded in lateral profile,SNL 60.1% HL; auricular opening present,moderate; eyes small, ED 1.9% SVL, 19.2%HL, 50.0% END, pupil nearly round; bodyslightly depressed, MBW 162.0% MBH; bodyscales smooth, glossy, imbricate; longitudinalscale rows at midbody 28; paravertebral scalerows 66; axilla–groin scale rows 46; limbs welldeveloped, pentadactyl, digits moderate; Fin-IIIlam 6; ToeIVlam 10; FLL 22.5% AGD,14.4% SVL; HLL 35.4% AGD, 22.6% SVL;order of digits from shortest to longest forhand: I 5 V , II 5 IV , III, for foot: I , V ,II , IV , III; tail not as wide as body,gradually tapered toward the end, TW 63.7%MBW, TL 99.9% SVL.


Rostral projecting dorsoposteriorly to pointin line with anterior edge of nasal, broaderthan high, moderately separated from fronto-nasal; frontonasal wider than long; nostrilovoid, centered in a single rectangular nasal;supranasals present, large, in narrow medialcontact; postnasals present; prefrontals broad-ly separated by frontal; frontal nearly octago-nal, its anterior margin in broad contact withfrontonasal, in contact with first two anteriorsupraoculars, 53 wider than anteriormostsupraocular; supraoculars five; frontoparietalsmoderate, in broad contact medially, eachfrontoparietal in contact with supraoculars2–4; interparietal moderate, quadrilaterallyshaped, its length nearly equal to midlinelength of frontoparietal; distinct parietaleyespot present, large, in posterior half ofscale; parietals broadly separated by interpa-rietal; nuchals undifferentiated; loreals two,decreasing in size from anterior to posterior,subequal, in contact with prefrontal, postna-sal, supranasal, second supralabial, posteriorloreal and frontonasal; preocular single, nearlyone half as high as posterior loreal; singlepresubocular; supraciliaries six, the anterior-most contacting prefrontal and separatingposterior loreal from first supraocular, poste-riormost extending to midline of last suprao-cular; subocular row complete; lower eyelidwith one row of scales, lacking an enlargedoval window, largely transparent; supralabialsseven, fifth and sixth below the eye; infra-labials six.

Mental wider than long, in contact with firstinfralabials; single enlarged postmental, widerthan mental; followed by two pairs of enlargedchin shields; first pair in broad medial contact,second pair separated by single undifferenti-ated scale.


Coloration of holotype in preservative.—Ground color of body medium brown; dorsalsurface of body with eight continuous, longi-

tudinal rows of dark brown spots, extendingfrom posterior edge of parietals to base of tail;spot rows span six full and two half rows ofscales at midbody, narrowing to four full andtwo half rows of scales posterior to parietals;pigmentation covering middle one third ofdorsal scales; dorsolateral stripes indistinct,discontinuous, spanning one whole and twohalf row of scales from auricular opening tomidbody. Lateral and ventral surface of bodylight brown. Lateral surface with threediscontinuous rows of dark brown spots,spanning posterior two thirds of axilla–groindistance. Ventral surface without dark pig-mentation. Tail with dark dorsal blotches andspots; dark pigment reduced ventrally. Headscales homogeneous dark brown; rostral,nasal, postnasal, supranasal, first supralabial,mental, and first infralabial light gray; pinealeyespot large distinct, light cream. Limbsmottled medium to dark brown dorsally,yellowish brown ventrally; dorsal and ventralsurface of digits dark brown.

Coloration of holotype in life.—Colorationin life is unrecorded; however, becauseBrachymeles specimens do not change signif-icantly during preservation (CDS and RMB,personal observations), we suspect that thepreserved coloration and patterns are muchlike those in life.

Measurements of holotype (in millime-ters).—SVL 89.2; AGD 56.8; TotL 178.3;MBW 13.9; MBH 8.6; TL 89.1; TW 8.8; TH8.0; HL 8.7; HW 10.1; HH 7.2; SnFa 22.8; ED1.7; END 3.3; SNL 5.2; IND 3.0; FLL 12.8;HLL 20.1; MBSR 28; PVSR 66; AGSR 46;FinIIIlam 6; ToeIVlam 10; SL 7; IFL 6; SC 6;SO 5.

Variation.—Morphometric variation of theseries is summarized in Table 6. Specimenswere observed to have parietals moderatelyseparated by the interparietal (CAS 144229–30, 144341, KU 323933, 323935–6) or in pointmedial contact (CAS 144313, KU 323934).

Scale counts were observed to vary amongthe measured series. Specimens were ob-served to have midbody scale row counts of 26(KU 323935), 27 (CAS 144313), and 28 (CAS144229–30, 144341, KU 323933–4, 323936);axilla–groin scale row counts of 46 (KU323933, 323935–6), 47 (CAS 144229–30,144341, KU 323934), and 49 (CAS 144313);


and paravertebral scale row counts of 66 (KU323933–6), 67 (CAS 144229), and 68 (CAS144230, 144313, 144341).

We also observed lamellae counts to varyamong the measured series. Specimens wereobserved to have Finger-III lamellae counts offive (CAS 144229–30, 144341, KU 323934–5)or six (CAS 144313, KU 323933, 323936);Toe-IV lamellae counts of nine (CAS 144229,144313, 144341, KU 323935–6) or 10 (CAS144230, KU 323933–4).

Distribution.—Brachymeles tungaoi isknown only from Masbate Island (Fig. 5).

Ecology and natural history.—Brachymelestungaoi occurs in agricultural areas as well asdisturbed and secondary growth forest habi-tat. Little or no original, low-elevation forestremains on Masbate Island, but we assumethe species once also occurred in primaryforest. Individuals were collected in thehumus material within the rotting stumps oftrees. When disturbed, individuals immedi-ately moved in a rapid serpentine manner andattempted to burrow back into loose soil orhumus.

Sympatric lizard species observed on Mas-bate Island include the following: (Agamidae)Bronchocela cristatella, Draco spilopterus,Gonocephalus sophiae; (Gekkonidae) Cyrto-dactylus philippinicus, Gehyra mutilata,Gekko gecko, Hemidactylus frenatus, H. pla-tyurus; (Scincidae) Brachymeles bonitae,Emoia atrocostata, E. multicarinata, E. multi-fasciata, Lamprolepis smaragdina, Lipiniapulchella, Sphenomorphus decipiens, S. ja-gori; and (Varanidae) Varanus marmoratus.

Etymology.—We take pleasure in namingthe new species after our friend and dedicatedfield collaborator Jason B. ‘‘Tungao’’ Fernan-dez, with thanks for years of hard work towardthe research of semifossorial lizards. Suggest-ed common name: Tungao’s slender skink.

Brachymeles vindumi sp. nov.Figs. 4, 10

Holotype.—CAS 60724 (EHT Field No.1718), adult male, collected between 25 Octo-ber and 17 November 1920, in Sulu Province,Jolo Island, Philippines, by Edward H. Taylor.

Paratypes.—One adult female (CAS60725), one juvenile female (MCZ 26577),and one juvenile of unknown sex (CAS

60723), collected over the same dates and inthe same locality as holotype.

Diagnosis.—Brachymeles vindumi can bedistinguished from congeners by the followingcombination of characters: (1) body sizemoderate (SVL 104.9–113.6 mm); (2) penta-dactyl; (3) Finger-III lamellae six; (4) Toe-IVlamellae nine or 10; (5) moderate limb length;(6) midbody scale rows 30 or 31; (7) axilla–groin scale rows 49; (8) paravertebral scalerows 74; (9) supralabials seven; (10) infra-labials six; (11) pineal eye spot present,indistinct; (12) supranasals in contact; (13)prefrontals separate; (14) parietal in contact;(15) enlarged chin shields in two pairs; (16)first pair of chin shields separated; (17)nuchals undifferentiated; (18) fifth and sixthsupralabials below the eye; (19) auricularopening present; (20) continuous, light dorso-lateral stripes present, distinct; (21) darklateral stripes present; and (22) dark ventralpigmentation present (Tables 4 and 5).

Comparisons.—Characters distinguishingthe new species from all pentadactyl speciesof Brachymeles are summarized in Tables 4and 5. Brachymeles vindumi most closelyresembles B. kadwa, B. talinis, and B. tungaoi,but it differs from these three taxa by havingsix Finger-III lamellae, six supralabials, mid-body scale rows 30 or 31, axilla–groin scalerows 49, paravertebral scale rows 74, the firstpair of enlarged chin shields separated, andthe presence of continuous, light dorsolateralstripes (Tables 4 and 5). Brachymeles vindumican further be distinguished from B. kadwaand B. talinis by having nine or 10 Toe-IVlamellae (Tables 4 and 5); from B. talinis byhaving six infralabials (Table 5); from B.kadwa by contact between frontoparietals(Table 5); and from B. kadwa and B. tungaoiby contact between parietals (Table 5).

From all nonpentadactyl species of Brachy-meles (B. apus, B. bonitae, B. cebuensis, B.elerae, B. lukbani, B. minimus, B. muntingka-may, B. pathfinderi, B. samarensis, B. tridac-tylus, B. vermis, and B. wrighti), B. vindumidiffers by having a pentadactyl body form (vs.nonpentadactyl), longer forelimb lengths(.13.2 mm vs. ,6.9 mm), greater hind limblengths (.22.7 mm vs. ,12.9 mm), andgreater number of midbody scale rows (30or 31 vs. ,28), and by the presence of a


postnasal scale (vs. absence). In addition, B.vindumi differs from all nonpentadactyl spe-cies except B. pathfinderi by having aparavertebral scale row count 74 (vs. .84)and by the presence of auricular openings (vs.absence); and from B. apus, B. lukbani, B.minimus, and B. vermis by the presence oflimbs (vs. absence).

Description of holotype.—Mature male,hemipenes not everted (Fig. 9); SVL113.6 mm; body moderate relative to otherBrachymeles, elongate with respect to otherlizards; head weakly differentiated from neck,nearly as wide as body, HW 9.5% SVL,112.5% HL; HL 33.7% SnFa; SnFa 25.1%SVL; snout moderately long, rounded indorsal and lateral profile, SNL 59.7% HL;auricular opening present, moderate; eyessmall, ED 2.0% SVL, 23.5% HL, 58.0%END, pupil nearly round; body slightlydepressed, MBW 122.9% MBH; body scalessmooth, glossy, imbricate; longitudinal scalerows at midbody 31; paravertebral scale rows74; axilla–groin scale rows 49; limbs welldeveloped, pentadactyl, digits moderate; Fi-nIIIlam 6; ToeIVlam 9; FLL 18.1% AGD,11.6% SVL; HLL 31.2% AGD, 20.0% SVL;order of digits from shortest to longest forhand: I , V , II , IV , III, for foot: I 5 V ,II 5 III , IV; tail regenerated, not as wide asbody, sharply tapered toward the end, TW83.1% MBW.

Rostral projecting dorsoposteriorly to pointin line with anterior edge of nasal, broaderthan high, moderately separated from fronto-nasal by supranasal contact; frontonasal widerthan long; nostril ovoid, centered in a singlerectangular nasal; supranasals present, large,in moderate medial contact; postnasals pres-ent; prefrontals narrowly separated by frontal;frontal nearly octagonal, its anterior margin innarrow contact with frontonasal, in contactwith first two anterior supraoculars, 43 widerthan anteriormost supraocular; supraocularsfive; frontoparietals moderate, in broad medi-al contact, each frontoparietal in contact withsupraoculars 2–4; interparietal small, diamondshaped, its length equal in size to midlinelength of frontoparietal; parietal eyespotabsent; parietals in broad contact behindinterparietal; nuchals undifferentiated; lorealstwo, decreasing in size from anterior to

posterior, subequal, in contact with prefrontal,postnasal, supranasal, second supralabial, pos-terior loreal and frontonasal; preocular single,nearly two thirds as high as posterior loreal;single presubocular; supraciliaries six, theanteriormost contacting prefrontal and sepa-rating posterior loreal from first supraocular,posteriormost extending to midline of lastsupraocular; single subocular row complete;lower eyelid with one row of scales, lacking anenlarged oval window, largely transparent;supralabials seven, fifth and sixth below theeye; infralabials six.

Mental wider than long, in contact with firstinfralabials; single enlarged postmental, slight-ly wider than mental; followed by two pairs ofenlarged chin shields, scales of first pairseparated by a single undifferentiated scale,second pair separated by three undifferenti-ated scales.


Coloration of holotype in preservative.—Ground color of body medium to dark brown;middorsal surface of body covered with darkpigmentation, extending from posterior edgeof supranasals to base of tail, made of eightirregular, longitudinal middorsal rows of darkbrown spots, spanning six full and two halfrows of scales at midbody, narrowing to six fullrows of scales posterior to parietals, pigmen-tation covering middle one third of dorsalscales; dorsolateral stripes present, clearlydefined, continuous, lacking dark pigmenta-tion, spanning one whole and one half row ofscales from anterior-most supraocular to baseof tail. Lateral and ventral surface of bodymedium to dark brown. Lateral surface withsix to eight irregular dark spot rows, graduallybecoming fainter on ventral surface. Ventralsurface with irregular dark spots and blotches.Tail with continuous dark blotches and spotsdorsally, dark pigment reduced ventrally.Head scales homogeneous mottled mediumand dark brown dorsally; rostral, nasal,


postnasal, supranasal, first supralabial, mental,and first infralabial light brown to tan; pinealeyespot absent; dark brown blotch of pigmen-tation on lateral surfaces of head, spanningfrom posterior edge of eye to posterior edge ofauricular openings. Limbs mottled medium todark brown; dorsal surface of digits darkbrown, ventral surface of digits mediumbrown.

Coloration of holotype in life.—Colorationin life is unrecorded; however, becauseBrachymeles specimens do not change signif-icantly during preservation (CDS and RMB,personal observations), we suspect that thepreserved coloration and patterns are muchlike those in life.

Measurements of holotype (in millime-ters).—SVL 113.6; AGD 72.7; TotL N/A;MBW 14.2; MBH 11.6; TL N/A; TW 11.8;TH 8.4; HL 9.6; HW 10.8; HH 7.9; SnFa 28.5;ED 2.3; END 3.9; SNL 5.7; IND 3.4; FLL13.2; HLL 22.7; MBSR 31; PVSR 74; AGSR49; FinIIIlam 6; ToeIVlam 9; SL 7; IFL 6; SC6; SO 5.

Variation.—Morphometric variation of theseries is summarized in Table 6. Specimenswere observed to have midbody scale rowcounts of 30 (CAS 60725) or 31 (CAS 60724),and Toe-IV lamellae counts of nine (CAS60724) or 10 (CAS 60725).

Distribution.—Brachymeles vindumi isknown only from Jolo Island (Fig. 4).

Ecology and natural history.—Brachymelesvindumi presumably occurs in disturbedhabitat as well as secondary growth forest onJolo Island. Due to security concerns, norecent surveys have been conducted on JoloIsland; therefore, no information is availableon the ecology of this species.

Sympatric lizard species observed on JoloIsland include the following: (Agamidae)Draco guentheri; (Gekkonidae) Cyrtodactylusannulatus, Gehyra mutilata, Gekko gecko, G.mindorensis, Hemidactylus frenatus, H. pla-tyurus, Luperosaurus joloensis; (Scincidae)Brachymeles vermis, Eutropis multifasciata,E. rudis, Lamprolepis smaragdina, Lipiniaquadrivittata, Lygosoma bowringi, Spheno-morphus biparietalis, S. variegatus; and (Var-anidae) Varanus cumingi.

Etymology.—We take pleasure in namingthe new species for our close friend and

colleague Jens Vindum. The specific epithet isa patronym in the genitive singular, chosen inthanks for the many years of support andassistance he has provided during our re-search on Philippine amphibians and reptiles.Suggested common name: Jens’ slender skink.

DISCUSSION

Phylogenetic analyses of the mitochondrialATP8 and ATP6 genes resulted in strongsupport for nine lineages of Brachymeles(Fig. 2). The phylogeny, combined with mor-phological data, supports the elevation of allsubspecies of the polytypic species B. boulen-geri and B. schadenbergi to full species.However, the inferred relationships betweenseveral of the species sampled are weaklysupported. This may be indicative of rapiddiversification of Brachymeles or simply indi-cate a lack of character support at someinternal nodes. Given the use of only mito-chondrial data for our phylogenetic analyses,caution must be taken when interpretinginterspecies relationships, because a singlelocus can be subject to random variation, deepcoalescence, lineage sorting, and naturalselection (Brown et al., 2010; Edwards andBeerli, 2002; Galtier et al., 2009). Regardlessof the potential weaknesses of our single-locusapproach, our results are strongly supportedby an independent, comprehensive dataset ofmorphological characters.

No analyses supported the monophyly ofspecies formerly part of Brachymeles boulen-geri (B. boholensis, B. boulengeri, B. mind-orensis, and B. taylori; Fig. 2). Another cladeincluding B. talinis, B. kadwa, and B. tungaoiwas estimated with strong support (Fig. 2). Aspreviously recognized, B. talinis spanned twodistinct, recognized faunal regions (Luzon andVisayas). Given this formerly wide geograph-ical distribution, it is not surprising that thenorthern populations (Luzon and the Babuyanislands) constitute a genetically distinct line-age that we describe here as B. kadwa. Wewere surprised, however, to discover anadditional genetically distinct lineage onMasbate Island (Fig. 2). The fauna of MasbateIsland is recognized as part of the Visayan orcentral Philippine islands and has beenhypothesized to have shared land bridgeconnections with the central islands during


periods of glacial maxima (Dickerson, 1928;Heaney, 1985; Inger, 1954; Voris, 2000).Although we expected Masbate populationsto be more closely related to Visayan (Negros+ Panay) populations, all analyses stronglysupported the sister relationship between B.tunagoi (Masbate) and B. kadwa (Luzon),providing additional biogeographic supportfor the distinctiveness of B. tungaoi. We areunaware of phylogeographic or phylogeneticstudies including other vertebrate taxa fromMasbate. Comparison of the systematic affin-ities of other Masbate species may provideinteresting exceptions to the prevailing PAIC-oriented perspective of Masbate as a faunisticextension of the central Visayas (Heaney,1985).

The species recognized in this paperincrease the total number of known speciesof Brachymeles to 25, all but one of which areendemic to the Philippines. During the past2 yr, our knowledge of the species diversity inthe genus has expanded rapidly as the result oflarge-scale sampling efforts across the Philip-pines and the detailed analyses of morpho-logical variation among species and popula-tions (Siler et al., 2009, 2010a,b). Before thiseffort, estimates of Brachymeles species di-versity remained nearly constant for .30 yr(but see Brown and Alcala, 1995), which is atestament to the extent of morphologicalsimilarity among species within the genusand a lack of systematic studies of the group.It comes as little surprise that allopatricpopulations of ‘‘B. boulengeri’’ from theLuzon, Mindanao, Mindoro, and VisayanPAICs have proven to be morphologicallydiagnosable with increased sampling. To date,few studies have provided evidence of truly‘‘widespread’’ reptile species that have geo-graphic distributions spanning recognizedzoogeographic boundaries in the Philippines(but see Siler et al., 2010c) and as is quiteoften the case, these species frequently turnout to constitute multiple evolutionary lineag-es (Brown and Diesmos, 2002; Brown et al.,2009; Gaulke et al., 2007; McGuire andAlcala, 2000; Welton et al., 2009, 2010).

All species of Brachymeles have a semi-fossorial life style, specializing in dry rottingmaterial within rotten logs. Many are habitatspecialists found exclusively in rotting logs,

loose soil, or leaf litter, whereas others arecommon beneath piles of rotting coconuthusks in disturbed, agricultural habitat. Thespecies now found in residential and agricul-tural areas were once native to forestedhabitats. Before recent, focused survey efforts,the relatively low numbers of specimens ofBrachymeles in museum collections handi-capped our efforts at delimiting species. Therarity of Brachymeles in collections was due totheir secretive, semifossorial lifestyle.

This is the first, species-level phylogeneticstudy of Brachymeles. To date, taxonomicreviews of Brachymeles have focused solely onmorphological variation (Brown, 1956; Brownand Alcala, 1980; Brown and Alcala, 1995;Brown and Rabor, 1967; Hikida, 1982). It isapparent that species diversity in the genushas been considerably underestimated; ac-cordingly, discovery of additional undocu-mented (possibly cryptic) diversity is antici-pated in other species groups (e.g., Siler et al.,2009, 2010a,b). Several studies have shownthat the evolution of a burrowing lifestyle iscorrelated with decreasing dispersal abilities(Nevo, 1979; Patton and Feder, 1978; Pattonand Yang, 1977; Selander et al., 1974; Wienset al., 2006). Many Brachymeles lineages haveexperienced reduction or loss of limbs, whichmay further reduce vagility (Daniels et al.,2005; Mulvaney et al., 2005; Wiens et al.,2006). Through time, reduced dispersal abil-ities may lead to increasingly patchy distribu-tions, reduced gene flow between populations,and the accumulation of interpopulationdifferences (Nevo, 1979). However, the rolethat geological history and complex geographyplay on the dispersal abilities and diversifica-tion patterns of Brachymeles species remainsunknown. Regardless of what processes pro-duce species diversity, we expect that addi-tional species await discovery. With severalspecies represented by only a few voucheredspecimens, and frequent morphological con-vergence, it is clear that a comprehensivephylogenetic analysis of the genus will berequired to assess, with accuracy, the speciesdiversity within Brachymeles.

Following the recognition of Brachymelesboholensis, B. boulengeri, B. tungaoi, B.kadwa, B. mindorensis, B. orientalis, B.schadenbergi, B. taylori, and B. vindumi,


there are now 13 pentadactyl species ofBrachymeles. Of these species, eight arelarge-bodied (B. bicolor, B. tungaoi, B. kadwa,B. makusog, B. orientalis, B. schadenbergi, B.talinis, and B. vindumi) and five (B. boholen-sis, B. boulengeri, B. gracilis, B. mindorensis,and B. taylori) have moderately sized bodies.The distribution of pentadactyl species in thePhilippines is relatively even across the majorrecognized faunal regions, with four speciesknown to occur in the Luzon Faunal Region,five in the Mindanao Faunal Region, three inthe Visayan Faunal Region, one in the Mind-oro Faunal Region, and one in the Suluarchipelago (Brown and Alcala, 1980, Brownand Alcala, 1995; Brown and Diesmos, 2002;Siler et al., 2010a). In contrast, the distribu-tion of total species diversity in the genus isless uniform, with 11 species known from theLuzon Faunal Region versus six in the Mind-anao Faunal Region, six in the Visayan FaunalRegion, and only one and two in the Suluarchipelago and Mindoro Faunal Region,respectively (Brown and Alcala, 1980; Brownand Alcala, 1995; Brown and Diesmos, 2002;Siler et al., 2009, 2010a,b). New speciesdiscoveries on Luzon Island have occurredwith consistency during the last two decades;given the island’s complex mountain ranges(Sierra Madres, Cordillera, Zambales, BicolPeninsula volcanoes) and geographic com-plexity (Defant et al., 1989; Yumul et al.,2009), the increase in the region’s faunaldiversity is likely to continue (Brown et. al.1995a,b, 1999, 2000a,b, 2007; Ross andGonzales, 1992; Siler et al., 2009, 2010a,b).It is worth noting that efforts to surveyMindanao have been less extensive thanefforts on Luzon; this may account for someof the differences in diversity between theregions, which may be artifacts of samplingbiases.

At present, there remains one polytypicspecies (B. gracilis) and two ‘‘widespread’’species (B. bonitae and B. samarensis), all withdistributions spanning boundaries betweenrecognized faunal regions (Brown and Alcala,1980). Closer investigation of island popula-tions within each of these species may resultin the discovery of new diversity in the genus.As our understanding of the total diversitywithin Brachymeles increases, it is important

that continued efforts be made to conductsurveys focused on rotting log and leaf littermicrohabitats throughout the ranges of allspecies. Accurate data on the distributionsof these species will allow for a completeassessment of the geographic ranges of thespecies and appropriate decision of conserva-tion status and actions can be made. Atpresent, all 10 species are known or believedto be common throughout their ranges.Although these species currently inhabithighly disturbed, agricultural and residentialareas, no studies on the long-term effect ofdeforestation on populations of Brachymelesexist. Therefore, according to the Internation-al Union for Conservation of Nature catego-ries and classification structure, we considerthe conservation status of these species as‘‘least concern,’’ pending the collection ofadditional information that might suggestotherwise.

Acknowledgments.—We thank the Protected Areas andWildlife Bureau of the Philippine Department ofEnvironment and Natural Resources for facilitatingcollecting and export permits necessary for this andrelated studies; we are particularly grateful to M. Lim, C.Custodio, and A. Tagtag. Financial support for fieldworkfor CDS was provided by a Panorama Fund grant fromThe University of Kansas Biodiversity Institute, a Madisonand Lila Self Fellowship from the University of Kansas, aFulbright-Hayes Fellowship, and a grant from theNational Science Foundaton (NSF; DEB 0804115).Financial support for RMB and ACD was provided byNSF (EF-0334952 and DEB 0743491) funds to RMB.For the loans of specimens, we thank J. Vindum and A.Leviton (CAS), R. Sison and A. C. Diesmos (PhilippineNational Museum), J. Ferner (Cincinnati Museum ofNatural History), A. Resetar and H. Voris (Field Museumof Natural History), R. Crombie (United States NaturalHistory Museum), and T. LaDuc (Texas MemorialNatural History Museum). Critical reviews of themanuscript were provided by L. Trueb, D. McLeod,and J. Esselstyn, and C. Oliveros assisted with the Tagalogabstract. CDS thanks the CAS’s Stearns Fellowship andthe Museum of Comparative Zoology’s Ernst MayrFellowship for funding recent visits to examine compar-ative material. CDS and RMB extend a special thanks toArvin and Mae Diesmos for continued support.

LITERATURE CITED

BOETTGER, O. 1886. Aufzahlung der von Philippinenbekannten Reptilien und Batrachier. Bericht derSenckenbergischen Naturforschenden Gesellschaft fur1886, Frankfurt am Main. Germany.

BOETTGER, O. 1893. Katalog der Reptilien. Sammlung imMuseum de Senckenbergischer Naturforschenden Ge-sellschaft in Frankfurt am Main, Frankfurt, Germany.


BOULENGER, G. A. 1887. Catalogue of the lizards in theBritish Museum (Natural History. Volume III. Lacer-tidae, Gerrhosauridae, Scincidae, Anelytropidae, Diba-midae, Chamaeleontidae). Taylor & Francis, London,UK.

BRANDLEY, M. C., A. SCHMITZ, AND T. W. REEDER. 2005.Partitioned Bayesian analyses, partition choice, and thephylogenetic relationships of scincid lizards. SystematicBiology 54:373–390.

BRANDLEY, M. C., J. P. HUELSENBECK, AND J. J. WIENS.2008. Rates and patterns in the evolution of snake-likebody form in squamate reptiles: Evidence for repeatedre-evolution of lost digits and long-term persistence ofintermediate body forms. Evolution 62:2042–2064.

BROWN, R. M., AND A. C. DIESMOS. 2002. Application oflineage-based species concepts to oceanic island frogpopulations: The effects of differing taxonomic philos-ophies on the estimation of Philippine biodiversity.Silliman Journal 42:133–162.

BROWN, R. M., AND A. C. DIESMOS. 2009. Philippines,Biology. Pp. 723–732. In R. Gillespie and D. Clague(Eds.), Encyclopedia of Islands. University of CaliforniaPress, Berkeley, California, USA.

BROWN, R. M., AND S. I. GUTTMAN. 2002. Phylogeneticsystematics of the Rana signata complex of Philippineand Bornean stream frogs: Reconsideration of Huxley’smodification of Wallace’s Line at the Oriental–Austra-lian faunal zone interface. Biological Journal of theLinnean Society 76:393–461.

BROWN, R. M., J. W. FERNER, AND L. A. RUEDAS. 1995a. Anew species of lygosomine lizard (Reptilia: Lacertilia:Scincidae; Sphenomorphus) from Mt. Isarog, LuzonIsland, Philippines. Proceedings of the BiologicalSociety of Washington 108:18–28.

BROWN, R. M., J. W. FERNER, AND R. V. SISON. 1995b.Rediscovery and redescription of Sphenomorphusbeyeri Taylor (Reptilia: Lacertilia: Scincidae) from theZambales mountains of Luzon, Philippines. Proceed-ings of the Biological Society of Washington 108:6–17.

BROWN, R. M., J. A. MCGUIRE, J. W. FERNER, AND A. C.ALCALA. 1999. A new diminutive species of skink(Squamata; Scincidae; Lygosominae: Sphenomorphus)from Luzon Island, Republic of the Philippines. Copeia1999:362–370.

BROWN, R. M., J. A. MCGUIRE, AND A. C. DIESMOS. 2000a.Status of some Philippine frogs referred to Ranaeveretti (Anura: Ranidae), description of a new species,and resurrection of R. igorota Taylor 1922. Herpetolo-gica 56:81–104.

BROWN, R. M., J. A. MCGUIRE, J. W. FERNER, N.ICARANGAL, JR., AND R. S. KENNEDY. 2000b. Amphibiansand reptiles of Luzon Island, II: Preliminary report onthe herpetofauna of Aurora Memorial National Park,Philippines. Hamadryad 25:175–195.

BROWN, R. M., A. C. DIESMOS, AND M. V. DUYA. 2007. Anew species of Luperosaurus (Squamata: Gekkonidae)from the Sierra Madre mountain range of northernLuzon Island, Philippines. Raffles Bulletin of Zoology55:153–160.

BROWN, R. M., C. OLIVEROS, C. D. SILER, AND A. C.DIESMOS. 2008. A new Gekko from the Babuyan Islands,northern Philippines. Herpetologica 64:305–320.

BROWN, R. M., C. OLIVEROS, C. D. SILER, AND A. C.DIESMOS. 2009. Phylogeny of Gekko from the northern

Philippines, and description of a new species fromCalayan Island. Journal of Herpetology 43:620–635.

BROWN, R. M., C. W. LINKEM, C. D. SILER, J. SUKUMARAN,J. A. ESSELSTYN, A. C. DIESMOS, D. J. ISKANDAR, D.BICKFORD, B. J. EVANS, J. A. MCGUIRE, L. GRISMER, J.SUPRIATNA, AND N. ANDAYANI. 2010. Phylogeography andhistorical demography of Polypedates leucomystax inthe islands of Indonesia and the Philippines: Evidencefor recent human-mediated range expansion? Molecu-lar Phylogenetics and Evolution. doi:10.1016/j.ympev.2010.06.015.

BROWN, W. C. 1956. A revision of the genus Brachymeles(Scincidae), with descriptions of new species andsubspecies. Breviora 54:1–19.

BROWN, W. C., AND A. C. ALCALA. 1970. The zoogeographyof the herpetofauna of the Philippine Islands, a fringingarchipelago. Proceeding of the California Academy ofSciences 38:105–130.

BROWN, W. C., AND A. C. ALCALA. 1980. Pp. 6–53. In D.Law (Ed.), Philippine Lizards of the Family Scincidae.Silliman University Press, Dumaguete City, Philippines.

BROWN, W. C., AND E. L. ALCALA. 1995. A new species ofBrachymeles (Reptilia: Scincidae) from CatanduanesIsland, Philippines. Proceedings of the BiologicalSociety of Washington 108:392–394.

BROWN, W. C., AND D. S. RABOR. 1967. Review of thegenus Brachymeles (Scincidae), with descriptions ofnew species and subspecies. Proceedings of theCalifornia Academy of Sciences 15:525–548.

DANIELS, S. R., N. J. L. HEIDEMAN, M. G. J. HENDRICKS,M. E. MOKONE, AND K. A. CRANDALL. 2005. Unravelingevolutionary lineages in the limbless fossorial skinkgenus Acontias (Sauria: Scincidae): Are subspeciesequivalent systematic units? Molecular Phylogeneticsand Evolution 34:645–654.

DEFANT, M. J., D. JACQUES, R. C. MAURY, J. DE BOER, AND

J.-L. JORON. 1989. Geochemistry and tectonic setting ofthe Luzon arc, Philippines. Geological Society ofAmerica Bulletin 101:663–672.

DE QUEIROZ, K. 1998. The general lineage concept ofspecies. species criteria, and the process of speciation.Pp. 57–75. In D. J. Howard and S. H. Berlocher (Eds.),Endless Forms: Species and Speciation. Oxford Uni-versity Press, New York, USA.

DE QUEIROZ, K. 1999. The general lineage concept ofspecies and the defining properties of the speciescategory. Pp. 49–89. In R. A. Wilson (Ed.), Species:New Interdisciplinary Essays. Massachusetts Instituteof Technology Press, Cambridge, Massachusetts, USA.

DICKERSON, R. E. 1928. Distribution of life in thePhilippines. Philippine Bureau of Science, Manila.

DUMERIL, A. M. C., AND G. BIBRON. 1839. Erpetologiegeneral ou histoire naturelle complete des reptiles,Tome cinqieme. Librairie Encyclopedique de Roret,Paris, viii + 855 pp.

EDGAR, R. C. 2004. MUSCLE: Multiple sequencealignment with high accuracy and high throughput.Nucleic Acids Research 32:1792–1797.

EDWARDS, S. V., AND P. BEERLI. 2002. Perspective: Genedivergence, population divergence, and the variance incoalescence time in phylogeographic studies. Evolution54:1839–1854.

ESSELSTYN, J. A., H. J. D. GARCIA, M. G. SAULOG, AND L. R.HEANEY. 2008. A new species of Desmalopex (Pteropo-


didae) from the Philippines, with a phylogeneticanalysis of the Pteropodini. Journal of Mammalogy89:815–825.

FISCHER, J. G. 1885. Ichthyologische und herpetologischeBemerkungen. V. Herpetologische Bemerkungen. Jahr-buch Wissenschaftliche Anst, Hamburg, Germany.

FROST, D. R., AND D. M. HILLIS. 1990. Species in conceptand practice: Herpetological applications. Herpetolo-gica 46:87–104.

GALTIER, N., B. NABHOLZ, S. GLEMIN, S., AND G. HURST.2009. Mitochondrial DNA as a marker of moleculardiversity: A reappraisal. Molecular Ecology 18:4541–4550.

GAULKE, M., H. ROSLER, AND R. M. BROWN. 2007. A newspecies of Luperosaurus (Squamata: Gekkonidae) fromPanay Island, Philippines, with comments on thetaxonomic status of Luperosaurus cumingii (Gray,1845). Copeia 2007:413–425.

GRAY, J. E. 1845. Catalogue of the specimens of lizards inthe collection of the British Museum. Edward New-man, London, UK.

HALL, R. 2002. Cenozoic geological and plate tectonicevolution of SE Asia and the SW Pacific: Computer-based reconstructions, model and animations. Journalof Asian Earth Sciences 20:353–431.

HEANEY, L. R. 1985. Zoogeographic evidence for middleand late Pleistocene land bridges to the PhilippinesIslands. Modern Quaternary Research in SoutheastAsia 9:127–144.

HEANEY, L. R. 1986. Biogeography of small mammals inSE Asia: Estimates of rates of colonization, extinctionand speciation. Biological Journal of the LinneanSociety 28:127–165.

HIKIDA, T. 1982. A new limbless Brachymeles (Sauria:Scincidae) from Mt. Kinabalu, North Borneo. Copeia4:840–844.

HILLIS, D. M., AND J. J. BULL. 1993. An empirical tests ofbootstrapping as a method for assessing confidence inphylogenetic analysis. Systematic Biology 42:182–192.

INGER, R. F. 1954. Systematics and zoogeography ofPhilippine Amphibia. Fieldiana: Zoology 33:181–531.

(ICZN) INTERNATIONAL COMMISSION OF ZOOLOGICAL NO-

MENCLATURE. 1999, International Code of ZoologicalNomenclature. Fourth Edition. International Trust forZoological Nomenclature, London, UK.

LANDE, R. 1978. Evolutionary mechanisms of limb loss intetrapods. Evolution 32:73–92.

LEVITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E.DAWSON. 1985. Standards in herpetology and ichthyol-ogy: Part I. Standard symbolic codes for institutionalresource collections in herpetology and ichthyology.Copeia 1985:802–821.

MADDISON, D. R., AND W. P. MADDISON. 2005. MacClade:Analysis of phylogeny and character evolution. Version4.08. Sinauer, Sunderland, Massachusetts, USA.

MCGUIRE, J. A., AND A. C. ALCALA. 2000. A taxonomicrevision of the flying lizards of the Philippine Islands(Iguania: Agamidae: Draco), with a description of a newspecies. Herpetological Monographs 14:92–145.

MULVANEY, A., T. A. CASTOE, G. A. KYLE, L. K. KENNETH,AND C. L. PARKINSON. 2005. Evidence of populationgenetic structure within the Florida worm lizard,Rhineura floridana (Amphisbaenia: Rhineuridae). Jour-nal of Herpetology 39:118–124.

NEVO, E. 1979. Adaptive convergence and divergence ofsubterranean mammals. Annual Reviews 10:269–308.

NYLANDER, J. A. A. 2004. MrModeltest v2. Programdistributed by the author. Evolutionary Biology Centre,Uppsala University, Sweden.

NYLANDER, J. A. A., J. C. WILGENBUSCH, D. L. WARREN, AND

D. L. SWOFFORD. 2008. AWTY (are we there yet?): Asystem for graphical exploration of MCMC convergencein Bayesian phylogenetics. Bioinformatics 24:581–583.

PATTON, J. L., AND J. H. FEDER. 1978. Genetic divergencebetween populations of the pocket gopher, Thomomysumbrinus (Richardson). Saugetierkunde 43:12–30.

PATTON, J. L., AND S. Y. YANG. 1977. Genetic variation inThomomys bottae pocket gophers: Macrogeographicpatterns. Evolution 31:697–720.

RAMBAUT, A., AND A. J. DRUMMOND. 2007. Tracer v1.4.Available at: http://beast.bio.ed.ac.uk/Tracer.

RONQUIST, F., AND J. P. HUELSENBECK. 2003. MRBAYES 3:Bayesian phylogenetic inference under mixed models.Bioinformatics 19:1572–1574.

ROSS, C. A., AND P. C. GONZALES. 1992. Amphibians andreptiles of Catanduanes Island, Philippines. PhilippineNational Museum Papers 2:50–76.

SELANDER, R. K., D. W. KAUFMAN, R. J. BAKER, AND S. L.WILLIAMS. 1974. Genic and chromosomal differentia-tion in pocket gophers of the Geomys bursarius group.Evolution 28:557–564.

SILER, C. D., E. L. RICO, M. R. DUYA, AND R. M. BROWN.2009. A new limb-reduced, loam-swimming skink(Reptilia: Squamata: Scincidae: genus Brachymeles)from central Luzon Island, Philippines. Herpetologica65:92–105.

SILER, C. D., A. C. DIESMOS, AND R. M. BROWN. 2010a.A new loam-swimming skink, genus Brachymeles(Reptilia: Squamata: Scincidae) from Luzon andCatanduanes islands, Philippines. Journal of Herpetol-ogy 44:49–60.

SILER, C. D., D. S. BALETE, A. C. DIESMOS, AND R. M.BROWN. 2010b. A new legless loam-swimming lizard(Reptilia: Squamata: Scincidae: genus Brachymeles)from the Bicol Peninsula, Luzon Island, Philippines.Copeia 2010:114–122.

SILER, C. D., J. R. OAKS, J. A. ESSELSTYN, A. C. DIESMOS,AND R. M. BROWN. 2010c. Phylogeny and biogeographyof Philippine bent-toed geckos (Gekkonidae: Cyrtodac-tylus) contradicts a prevailing model of Pleistocenediversification. Molecular Phylogenetics and Evolution55:699–710.

SIMPSON, G. G. 1961. Principles of Animal Taxonomy.Columbia University Press, New York, New York, USA.

STAMATAKIS, A. 2006. RAxML-VI-HPC: Maximum likeli-hood-based phylogenetic analyses with thousands oftaxa and mixed models. Bioinformatics 22:2688–2690.

SWOFFORD, D. L. 2002. PAUP*. Phylogenetic analysisusing parsimony (*and other methods). Version 4.0.Sinauer Associates, Sunderland, Massachusetts, USA.

TAYLOR, E. H. 1917. Brachymeles, a genus of Philippinelizards. Philippine Journal of Science 12:267–279.

TAYLOR, E. H. 1918. Reptiles of Sulu archipelago.Philippine Journal of Science 13:233–267.

TAYLOR, E. H. 1922a. Additions to the herpetologicalfauna of the Philippine Islands, II. Philippine Journal ofScience 21:257–303.


TAYLOR, E. H. 1922b. The lizards of the Philippine Islands.Department of Agriculture and Natual Resources,Bureau of Science, Manila, Publication 17:1–269.

TAYLOR, E. H. 1925. Additions to the herpetological faunaof the Philippine Islands, IV. Philippine Journal ofScience 26:97–111.

VORIS, H. K. 2000. Maps of Pleistocene sea levels inSoutheast Asia: Shorelines, river systems and timedurations. Journal of Biogeography 27:1153–1167.

WELTON, L. J., C. D. SILER, A. C. DIESMOS, AND R. M.BROWN. 2009. A new bent-toed Gekko (Genus Cyrto-dactylus) from southern Palawan Island, Philippinesand clarification of the taxonomic status of C.annulatus. Herpetologica 65:328–343.

WELTON, L. J., C. D. SILER, A. C. DIESMOS, AND R. M.BROWN. 2010. Phylogeny-based species delimitation ofsouthern Philippines bent-toed geckos and a newspecies of Cyrtodactylus (Squamata; Gekkonidae) fromwestern Mindanao and the Sulu Archipelago. Zootaxa2390:49–68.

WIENS, J. J., AND J. L. SLINGLUFF. 2001. How lizards turninto snakes: A phylogenetic analysis of body-formevolution in anguid lizards. Evolution 55:2303–2318.

WIENS, J. J., M. C. BRANDLEY, AND T. W. REEDER. 2006.Why does a trait evolve multiple times within a clade?Repeated evolution of snake-like body form in squa-mate reptiles. Evolution 60:123–141.

WILCOX, T. P., D. J. ZWICKL, T. A. HEATH, AND D. M.HILLIS. 2002. Phylogenetic relationships the dwarf boasand a comparison of Bayesian and bootstrap measuresof phylogenetic support. Molecular Phylogenetics andEvolution 25:361–371.

WILEY, E. O. 1978. The evolutionary species conceptreconsidered. Systematic Zoology 21:17–26.

YUMUL, G. P., JR., C. B. DIMALANTA, K. QUEANO, AND E.MARQUEZ. 2009. Philippines, geology. Pp. 732–738. InR. Gillespie and D. Calgue (Eds.), Encyclopedia ofIslands. University of California Press, Berkeley,California, USA.

APPENDIX

Additional Specimens Examined

Numbers in parentheses indicate the number ofspecimens examined. With the exception of B. apus, allspecimens examined are from the Philippines. Severalsample sizes are greater than those observed in thedescription due to the examination of subadult specimensthat were excluded from morphometric analyses. Bra-chymeles apus.—(1) BORNEO: MALAYSIA: Sabah: Mt.Kinabalu National Park, Sayap Substation: SP 06915.Brachymeles bicolor.—(24) LUZON ISLAND: AuroraProvince: Municipality of Maria Aurora: Barangay VillaAurora, Sitio Dimani, Aurora Memorial National Park:KU 323149–52; CAGAYAN PROVINCE: Municipality ofBaggao: Sitio Hot Springs: CAS 186111, USNM 140847,498829–30, 498833; Isabela Province, Sierra MadresMountain Range: KU 324097–99, PNM 5785, 9568–77;KALINGA PROVINCE: Balbalasang-Balbalan National Park:FMNH 259438. Brachymeles boholensis.—(19) BO-HOL ISLAND: BOHOL PROVINCE: Municipality of SierraBullones, Barangay Danicop: KU 323944, 323948–9,

323952–6, 323960, 323962–3, 323966, 323970, 323972,323975–6, 323981–2, 323990, 324001; BOHOL ISLAND:BOHOL PROVINCE: 6 km S of Municipality of SierraBullones: Teachers Park: CAS-SU (holotype) 24528;13 km SE Municipality of Sierra Bullones: Dusita Barrio:CAS-SU (paratypes) 24502–04, 24518, 24520–25, 24541,24543, CAS-SU 25443–44, 25447; 1 km E Dusita Barrio:Abacjanan: CAS-SU 24867; Municipality of Sierra Bul-lones: Sandayong: CAS-SU 18709, 18717. Brachymelesbonitae.—(13) MASBATE ISLAND: MASBATE PROVINCE:Municipality of Mobo: Tugbo Barrio: CAS 144223;Mapuyo Barrio: Palangkahoy: CAS 144270; MINDOROISLAND: MINDORO ORIENTAL PROVINCE: Mt. Halcon: SEslope Barawanan Peak: CAS-SU 25713, 25793, 25886–88,25904; Sumagui: CAS 62064 (paratype); POLILLOISLAND: QUEZON PROVINCE: Municipality of Polillo:Barangay Pinaglubayan: KU 307747–49, 307755. Brachy-meles boulengeri.—(26) LUZON ISLAND: AURORA

PROVINCE: Municipality of Baler: KU 322314–20; LUZONISLAND: LAGUNA PROVINCE: Municipality of Los Banos,Barangay Batong Malake: KU 32058–60; Municipality ofLos Banos: CAS 61096; Mt. Maquiling: CAS 61297;POLILLO ISLAND: QUEZON PROVINCE: Municipality ofPolillo: CAS (paratypes) 62272–73, 62276–77; BarangayPinaglubayan: KU 307438–9, 307750–54, 307756 (neo-type), 307757–58. Brachymeles cebuensis.—(8) CEBUISLAND: 40 km SW of Cebu City: Tapal Barrio, SitioMantalungon: CAS-SU (holotype) 24400, (paratypes)24396–97, 24399, 24401, 24403; Municipality of Carcar:Tapal Barrio: CAS 102405 (paratype); 3 km NW CebuCity, Buhisan Barrio, Buhisan Reforestation Project:CAS-SU 27537. Brachymeles elerae.—(4) LUZONISLAND: KALINGA PROVINCE: Municipality of Balbalan:CAS 61499–500, PNM 9563–4. Brachymeles gracilisgracilis.—(18) MINDANAO ISLAND: DAVAO DEL SUR

PROVINCE: Municipality of Malalag: Sitio Kibawalan: CAS-SU 24163, 24165, CAS 124811, 139307–09; Davao City:Buhangin, Kabanti-an: CAS 124803–04, 139293–95,139303–05; Digos City: Tres de Mayo Barrio: CAS124806–08, 139300. Brachymeles gracilis hilong.—(20) MINDANAO ISLAND: AGUSAN DEL NORTE PROV-

INCE: Municipality of Cabadbaran: Diuata MountainRange: Mt. Hilonghilong: Balangbalang: CAS-SU (holo-type) 24407, paratype) 102406, 133578, CAS-SU 24411,133577, 133581–82, 133609, 133612, 133692–93, 133703–06, 133743, 133745–47; SURIGAO DEL SUR PROVINCE:Municipality of Lanuza: Diuata Mountain Range: SibuhayBarrio: CAS-SU (paratype) 24315. Brachymeles luk-bani.—(14) LUZON ISLAND: CAMARINES NORTE PROV-

INCE: Municipality of Labo: Barangay Tulay Na Lupa, Mt.Labo: PNM (holotype) 9567, (paratopotypes) 9589–92,KU (paratopotypes) 313597–99, 313601, 313603–04,313606, 313608, FMNH (paratopotype) 270191. Brachy-meles makusog.—(17) CATANDUANES ISLAND: CAT-

ANDUANES PROVINCE: Municipality of Gigmoto: BarangaySan Pedro, Sitio Tungaw: PNM (holotype) 9565, (para-topotypes) 9583–9584, KU (paratopotypes) 308126,308128, 308136, 308208; LUZON ISLAND: CAMARINES

NORTE PROVINCE: Municipality of Labo, Barangay TulayNa Lupa, Mt. Labo: KU (paratypes) 313612–313614,313616, 313617, PNM (paratypes) 9585–9588, FMNH(paratype) 270200. Brachymeles mindorensis.—(34)MINDORO ISLAND: MINDORO OCCIDENTAL PROVINCE:KU 304351–5, 304412–3, 304488, 307739–42, 308404,


308447–8, 308534; MINDORO ISLAND: MINDORO

ORIENTAL PROVINCE: 30 km SE Municipality of Calapan:Bank of Tarogin River: CAS-SU (holotype) 24487; SEslope Mt. Halcon, Tarogin Barrio: CAS-SU (paratypes)24549–54, 24561–62, 24564; 24566, 24568, 24573–74,24577–79; Mt. Halcon, SE slope Barawanan Peak: CAS-SU (paratype) 24570. Brachymeles minimus.—(6) CAT-ANDUANES ISLAND: CATANDUANES PROVINCE: Munic-ipality of Gigmoto: Barangay San Pedro: KU 308129–31,308210–12. Brachymeles muntingkamay.—(17) LU-ZON ISLAND: NUEVA VIZCAYA PROVINCE: Municipalityof Quezon: Barangay Maddiangat, Mt. Palali: PNM(holotype)9566, (paratopotypes) 9578–82, KU (paratopo-types) 308865–66, 308900–06, 308908, 308953. Brachy-meles pathfinderi.—(40) MINDANAO ISLAND: SAR-

ANGANI PROVINCE: Municipality of Glan: Barangay Taluya:CDS 5235–42; Barangay Tanibulad, Sitio Padido: CDS5192–5206, 5210–20, 5222–27. Brachymeles samaren-sis.—(7) SAMAR ISLAND: EASTERN SAMAR PROVINCE:Municipality of Taft: Barangay San Rafael: KU 310849–50, 310852, 311294–6; LEYTE ISLAND: LEYTE PROV-

INCE: Municipality of Baybay: Barangay Pilim: Sitio SanVicente: KU 311225. Brachymeles orientalis.—(53)BOHOL ISLAND: BOHOL PROVINCE: Municipality ofSierra Bullones: Dusita Barrio: CAS-SU (holotype)24436, CAS-SU (paratypes) 24428, 24434, 24437, CAS(paratype) 102404, CAS-SU 25452; Dusita Barrio: Abac-janan: CAS-SU (paratypes) 24446–51, CAS-SU 25460;Cantaub Barrio: CAS-SU (paratypes) 18702, 24442,24458; CAMIGUIN SUR ISLAND: CAMIGUIN PROVINCE:Municipality of Catarman: Mt. Mambajao: Sitio Sangsan-gan: CAS 110976–83; LEYTE ISLAND: Leyte PROVINCE:Municipality of Baybay: KU 311231–5, 311241; MIND-ANAO ISLAND: AGUSAN DEL NORTE PROVINCE: Munici-pality of Cabadbaran: Diuata Mountain Range: Mt.Hilonghilong: Kasinganan: CAS-SU 133301, 133616,133749, 133752, 133754; SAMAR ISLAND: EasternSamar PROVINCE: Municipality of Taft: KU 305470,310734–6, 310739, 310942–6, 310949, 310951, 310955.Brachymeles schadenbergi.—(45) BASILAN ISLAND:BASILAN PROVINCE: Port Holland: Sawmill: CAS 60493;MINDANAO ISLAND: MISAMIS OCCIDENTAL PROVINCE:2 km NW of Masawan: CAS 23468–69; 4 km NW ofMasawan: CAS 23471; 3 km NW Masawan: south bank ofDapitan River: CAS 23479–81, 23484–85; COTABATO

PROVINCE: Municipality of Tatayan: MCZ 26553, 26555–8, 26561, 26566, 26568, 26571–2, 26574; ZAMBOANGA DEL

NORTE PROVINCE: Dapitan River: CAS-SU 23494–96;ZAMBOANGA CITY PROVINCE: Municipality of Pasonanca:Barangay Baluno: Pasonanca Natural Park: KU 314967,314969, 314970–8, 314980, 314984–85, 314988–92,314994, 314996–7. Brachymeles talinis.—(31) NE-GROS ISLAND: NEGROS ORIENTAL PROVINCE: 6 km WMunicipality of Valencia: Cuernos de Negros MountainRange: ridge on north side of Maite River: CAS-SU(holotype) 18358, (paratype) 89813; Cuernos de NegrosMountain Range: Dayungan Ridge: CAS 133871; Duma-guete City: CAS-SU (paratype) 12225; Municipality ofSiaton: 20 km N Bondo Barrio: CAS-SU 22311–12; 22317,22323; INAMPULAGAN ISLAND: GUIMARAS PROVINCE:Municipality of Sibunag: 8 km W Pulupandan Town:CAS-SU 27972, 27996–97; PANAY ISLAND: ANTIQUE

PROVINCE: Municipality of San Remigio: KU 306756–60,306762–7, 306769, 306770–6, 306786. Brachymelestaylori.—(34) NEGROS ISLAND: NEGROS OCCIDENTAL

PROVINCE: Municipality of Silay City, Barangay Patag: KU324044–56; NEGROS ISLAND: NEGROS ORIENTAL PROV-

INCE: 3 km W Municipality of Valencia: Cuernos deNegros Mountain Range: Sitio Lunga: ridge on north sideof Maiti River: CAS-SU (holotype) 18615, CAS-SU 21873;ridge on south side of Maiti River: CAS-SU (paratype)18641, 18656–57, 18748; Cuernos de Negros MountainRange: CAS-SU (paratype) 18649; top of DayunganRidge: CAS-SU 21877, 21880, 21883–84; 24 km NWBondo Barrio: Bantolinao: CAS-SU 22355–56; CEBUISLAND: CEBU PROVINCE: Municipality of Carcar: TapalBarrio: Sitio Mantalongon: CAS 154671, 154673, 154678–82, 154686. Brachymeles tridactylus.—(20) NEGROSISLAND: NEGROS OCCIDENTAL PROVINCE: 16 km EMunicipality of La Castellana: Barrio Cabagna-an:southern slope of Mt. Canlaon: CAS-SU 19424, 19426–27, 19429, 19452, 19458; 20 km E Municipality of LaCastellana: Sitio Kalapnagan: CAS-SU 27082–83; NEGROS

ORIENTAL PROVINCE: Hills North and Northwest ofMayaposi: CAS-SU (holotype) 18354; PANAY ISLAND:ANTIQUE PROVINCE: Municipality of Culasi: BarangayAlojipan: KU 307726–36. Brachymeles vermis.—(5)JOLO ISLAND: SULU PROVINCE: CAS-SU (paratype)62489, CAS-SU 60720–22, 60857.


phylogeny-based species delimitation in philippine slender skinks

Documents