Herpetological Monographs, 24, 2010, 1–54 E 2010 by The Herpetologists’ League, Inc.

PHYLOGENY-BASED SPECIES DELIMITATION IN PHILIPPINE SLENDER SKINKS (REPTILIA: SQUAMATA: SCINCIDAE: BRACHYMELES): TAXONOMIC REVISION OF PENTADACTYL SPECIES GROUPS AND DESCRIPTION OF THREE NEW SPECIES

1 CAMERON D. SILER AND RAFE M. BROWN Biodiversity Institute and Department of Ecology and Evolutionary Biology, University of Kansas, 1345 Jayhawk Boulevard, Lawrence, KS 66045-7593, USA

ABSTRACT: We use data from external morphology and mitochondrial gene sequences to provide the basis for a taxonomic revision of two polytypic, pentadactyl Philippine species of scincid lizards of the genus Brachymeles. Although previous studies have noted significant morphological variation among island populations, the similarities in body size and scale pigmentation and pattern have led to the continued recognition 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 morphological and genetic data to define species limits in B. boulengeri, B. schadenbergi, and B. talinis. Our molecular and morphological 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 their congeners 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 (from Luzon 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, new species. 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 mitochondria nagsagawa kami ng rebisyon ng tatlong species ng reptilyanong nabibilang sa genus na Brachymeles. Sa kabila ng 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 na laki ng katawan at pagkakahawig ng kulay at disenyo ng kanilang kaliskis. Ang ikatlong species, ang Brachymeles talinis, ay matatagpuan sa isla ng Jolo at sa gitna at hilagang bahagi ng Pilipinas. Ipinapakita ng aming 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 ibang miyembro ng genus sa maraming aspekto ng panlabas na kaanyuan at samakatwid ay nararapat na kilalanin bilang 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 sa isla ng Masbate. Sa huli, kinakailangan ding makilala ang populasyon ng B. talinis mula sa isla ng Jolo bilang isang 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 scincid lengeri, B. gracilis, B. makusog, B. schaden- lizards possessing both fully limbed and bergi, and B. talinis); eight are nonpentadac- limbless species (Brachymeles, Chalcides, tyl, with reduced limbs and numbers of Lerista, and Scelotes; Brandley et al., 2008; digits (B. bonitae, B. cebuensis, B. elerae, B. Lande, 1978; Wiens and Slingluff, 2001). muntingkamay, B. pathfinderi, B. samarensis, Within the genus Brachymeles, all but one of B. tridactylus, and B. wrighti); and four are the 18 recognized species are endemic to the entirely limbless (B. apus, B. minimus, B. Philippines. The exception is B. apus from lukbani, and B. vermis). Within the nonpen- northern Borneo (Brown and Alcala, 1980; tadactyl species (Brown, 1956; Brown and Hikida, 1982; Siler et al., 2009, 2010a,b). Six Rabor, 1967; Dume´ril and Bibron, 1839; species are pentadactyl (B. bicolor, B. bou- Taylor, 1917, 1918, 1925) exist a wide range of limb- and digit-reduced states, from minute 1 CORRESPONDENCE: email, [email protected] limbs that lack full digits (B. bonitae, B.

1 2 HERPETOLOGICAL MONOGRAPHS [No. 24 cebuensis, B. muntingkamay, B. samarensis, and B. tridactylus) to moderately developed limbs with four to five digits on the hands and feet (B. elerae, B. pathfinderi,andB. wrighti: Brown and Alcala, 1980; Hikida, 1982; Siler et al., 2009, 2010b). All species are semifossorial and typically found in dry, rotting material inside decaying logs or in loose soil and leaf litter. Shared body plans and similar external morphological features among populations of Brachymeles has proven problematic for diagnosing species (Brown and Alcala, 1980; Siler et al., 2009, 2010a,b). In addition, several rare, mid- to high-elevation species have long been represented by only a few specimens, in some cases without knowledge of their exact type locality (e.g., B. bicolor, B. elerae, B. wrighti, and B. pathfinderi). Three species are polytypic: B. boulengeri contains four subspe- cies and B. gracilis and B. schadenbergi each contain two subspecies (Brown, 1956; Brown and Rabor, 1967; Brown and Alcala, 1980). Several other species are recognized as having widespread distributions that span historical faunal demarcations in the Philippines (Brown FIG. 1.—Map of the Philippine islands, with island and Diesmos, 2002; Brown and Guttman, labels provided for islands with representative samples 2002; Heaney, 1985, 1986), including B. used for this study. The five recognized major Pleistocene aggregate island complexes (PAICs), major island groups, talinis, B. samarensis,andB. bonitae (Brown, and additional deep-water islands are labeled for refer- 1956; Brown and Alcala, 1980; Brown and ence. Islands of the Romblon Island group are designated Rabor, 1967). by the first letter of the island name (T, Tablas Island; R, Romblon Island; and S, Sibuyan Island). Current islands in the Philippines are shown in medium gray; light gray TAXONOMIC HISTORY areas enclosed in black 120-m bathymetric contours The genus Brachymeles was first described indicate the hypothesized maximum extent of land during the mid- to late Pleistocene. by Dume´ril and Bibron (1839) for the small, limb-reduced species B. bonitae. Three addi- tional species (Senira bicolor, Gray, 1845; gros islands as representatives of the species Eumeces (Riopa) gracilis, Fischer, 1885; and (Fig. 1). Thirty years later, Brown (1956) E.(R.) schadenbergi, Fischer, 1885) were described B. gracilis taylori and included transferred to the genus by Boettger (1886) B. boulengeri as one of three subspecies of and Boulenger (1887). These four species the polytypic species B. gracilis. Brown and represented the known diversity in the genus Rabor’s (1967) description of B. gracilis for 30 yr, until Taylor published a series of boholensis and B. g. mindorensis brought the herpetofaunal descriptions in the early 1900s. number of subspecies within B. gracilis to In Taylor’s (1917) review of the genus, he five. It was not until 1980 that Brown and revised B. gracilis to not only include popu- Alcala (1980) resurrected the polytypic spe- lations in the Mindanao Faunal Region but cies B. boulengeri and included four subspe- also populations on Negros and Mindoro cies (B. b. boulengeri, B. b. boholensis, B. b. islands (Fig. 1). A few years later, Taylor mindorensis, and B. b. taylori), all believed to (1922b) described B. boulengeri, based on be distinct from B. gracilis. This view material from Polillo Island, and included characterized the taxonomy of B. boulengeri populations from Luzon, Mindoro, and Ne- for the next 30 yr. Numerous studies have 2010] HERPETOLOGICAL MONOGRAPHS 3 mentioned the morphological variation among talinis have been available, the recognition of island populations of B. boulengeri and other this widespread species has continued for species (Brown, 1956; Brown and Alcala, .40 yr (Brown and Alcala, 1980; Siler et al., 1980; Brown and Rabor, 1967; Taylor, 2009, 2010a,b). 1922b), but all refrained from elevating these Following the separation of B. talinis from subspecies to full species. Brachymeles bou- the B. schadenbergi complex, Brown and lengeri boulengeri and B. b. taylori have larger Rabor (1967) recognized two subspecies of geographic distributions across multiple is- B. schadenbergi: one subspecies from western lands within a single faunal region (Brown and south central Mindanao Island and and Alcala, 1980; Brown and Rabor, 1967; Basilan Island (B. s. schadenbergi), and the Taylor, 1922b), whereas B. b. boholensis and other subspecies from eastern Mindanao, B. b. mindorensis are single-island endemics Camiguin Sur, Bohol, and Leyte islands (B. (Brown, 1956; Brown and Alcala, 1980; Brown s. orientalis). Fischer (1885) had previously and Rabor, 1967). designated the type locality for B. schaden- Although B. talinis was described originally bergi as southern Mindanao Island, and as B. schadenbergi talinis (Brown, 1956), and specimens from south central Mindanao considered part of the widespread B. scha- Island have thus been identified as B. s. denbergi complex from the Sulu Islands and schadenbergi (Brown and Alcala, 1980; Brown the Mindanao, Visayan, and Luzon Pleisto- and Rabor, 1967). In addition to its already cene aggregate island complexes (PAICs; broad geographic distribution, Brown and Brown and Diesmos, 2002; Brown and Gutt- Alcala (1980) predicted that B. s. orientalis man, 2002), the subspecies was described on also would be observed on Samar Island. the basis of material from Negros Island Both B. boulengeri and B. talinis are (Brown, 1956). Brown (1956) referred to distributed across several distinct PAICs, series of specimens from Jolo and Luzon including the Luzon, Mindanao, Mindoro, islands as likely exemplars of B. s. talinis and and Visayan island complexes (Fig. 1). Because hypothesized three explanations for the un- many recent studies have revealed that few usual distribution of B. schadenbergi in the endemic Philippine reptiles actually possess Philippines: (1) chance colonization across broad distributions spanning these regional ocean barriers into distinct faunal regions by faunistic boundaries (Brown and Diesmos, two subspecies; (2) morphological conver- 2002, 2009; Siler et al., 2010a; Welton et al., gence within a polytypic species; and (3) 2009, 2010), we have begun to reevaluate the prolonged maintenance of two morphologi- known polytypic and widespread species in the cally similar, disjunct distributions of sibling genus Brachymeles. Our goal is to revise the species within the Philippines. At the time, taxonomy such that individual units (species) Brown (1956) supported the first hypothesis represent independently evolving, cohesive with reservations; however, in Brown and lineage segments (sensu Simpson, 1961; de Rabor’s (1967) review of Brachymeles, newly Queiroz, 1998, 1999; Frost and Hillis, 1990; available material supported the third hypoth- Wiley, 1978). Careful examination of numerous esis and led to the recognition of B. talinis recently collected specimens from throughout from Jolo, Negros, and Luzon islands as a the known ranges of B. boulengeri, B. scha- distinct ‘‘sibling species’’ of B. schadenbergi denbergi, and B. talinis, as well as all relevant populations from Basilan, Mindanao, Bohol, name-bearing type material, results in the and Leyte islands. When Brown and Alcala reorganization of B. boulengeri, B. schaden- (1980) revised the genus, they restricted the bergi, and B. talinis into 10 distinct species. In geographic distribution of B. talinis to the this paper, we provide a phylogenetic analysis central and northern Philippine islands and of most of these taxa, fully describe each postponed the assignment of the two speci- evolutionary lineage, clarify species boundar- mens from Jolo until the morphological ies, and provide the first illustrations of most variability of that island population was better these taxa. We also provide information on understood. Although samples from through- each species’ natural history, ecology, and out the central and northern range of B. geographic distribution. 4 HERPETOLOGICAL MONOGRAPHS [No. 24

MATERIALS AND METHODS ATPase 8 (ATP8) and ATPase 6 (ATP6) Fieldwork, Sample Collection, and protein-coding genes were amplified using Specimen Preservation standard polymerase chain reaction (PCR) methods with the primers ATPf (59-CTCA- Fieldwork was conducted on Bohol, Cala- GARATCTGCGGGYCAAATCACA-39) and yan, Camiguin Norte, Camiguin Sur, Catan- ATPr (59-GTGCYTTCTCGRRTAATRTCYC- duanes, Dinagat, Leyte, Luzon, Masbate, GTCAT-39; M. Brandley, unpublished data). Mindanao, Mindoro, Negros, Panay, Polillo, PCR products were visualized on 1.0% Romblon, Samar, Sibuyan, and Tablas islands, agarose gels and then purified with ExoSAP- all in the Philippines (Fig. 1) between 1992 IT (US78201, GE Healthcare Biosciences, and 2009. Specimens were collected between Piscataway, New Jersey, USA). Purified tem- 0900 and 1600 h, euthanized with aqueous plates were sequenced with the same primers chloretone, dissected for genetic samples and the ABI Prism BigDye Terminator (liver preserved in 95% ethanol or flash frozen chemistry, version 3.1 (Applied Biosystems, in liquid nitrogen), fixed in 10% buffered- Foster City, California, USA). Cycle-sequenc- formalin, and eventually (,2 mo) transferred to 70% ethanol. Newly sequenced voucher ing products were purified with Sephadex specimens are deposited in US and Philippine Medium (NC9406038, GE Healthcare Biosci- museum collections (see Acknowledgments ences) in Centri-Sep 96 spin plates (CS-961, and Specimens Examined); if available, Princeton Separations, Princeton, New Jersey, voucher information corresponding to data USA). Sequencing products were run on an from GenBank sequences is included in ABI Prism 3130xl Genetic Analyzer (Applied Table 1. Biosystems). Gene sequences were assembled with Sequencher 4.8 (Gene Codes Corp., Ann Taxon Sampling and Outgroup Selection for Arbor, Michigan, USA). Phylogenetic Analyses Alignment and Phylogenetic Analysis Because our primary goal was to estimate phylogenetic relationships among the subspe- An initial alignment was produced in cies and island populations of B. boulengeri, Muscle, version 3.7 (Edgar, 2004), and visual B. schadenbergi, and B. talinis, we sequenced inspections were made in MacClade 4.08 only one to two exemplars per species; (Maddison and Maddison, 2005). No instanc- however, in the case of a species occurring es of insertions or deletions, or ambiguously on multiple islands within a single PAIC, or aligned regions, were observed in the data, across a large island such as Mindanao, we and all data were used for analyses. The final sampled multiple populations to provide alignment consisted of 838 aligned nucleo- greater geographic resolution. We chose two tides. scincid taxa (Plestiodon egregius and Lygo- Phylogenetic analyses were conducted us- soma bowringii) based on relationships pre- ing parsimony, likelihood, and Bayesian opti- sented in a recent phylogenetic analysis of mality criteria. Parsimony (maximum parsi- scincid lizards (Brandley et al., 2005) as mony [MP]) analyses were conducted in outgroups. In total, 39 ingroup samples were PAUP* 4.0 (Swofford, 2002), with all charac- used to construct phylogenetic inferences. ters weighted equally. Most-parsimonious There are no tissue samples of the population trees were estimated using heuristic searches of B. talinis from Jolo Island, and this with 1000 random addition-sequence repli- population is therefore not included in the cates and tree bisection and reconnection phylogenetic analyses. (TBR) branch swapping. To assess heuristic support, nonparametric bootstrapping was DNA Extraction, Purification, conducted using 1000 replicates, each with and Amplification 100 random addition-sequence replicates and We extracted total genomic DNA from TBR branch swapping. tissues (Table 1) by using the modified Partitioned maximum likelihood (ML) anal- guanidine thiocyanate extraction method of yses were conducted in RAxML-HPC, version Esselstyn et al. (2008). The mitochondrial 7.04 (Stamatakis, 2006). The alignment was 2010] HERPETOLOGICAL MONOGRAPHS 5

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

Genbank Accession Numbers

Species Voucher Locality ATPase 8, 6 Lygosoma bowringi LSUHC 6970 West Malaysia HQ239366 Plestiodon egregius MVZ 11013 Not available NC000888 Brachymeles taylori ACD 862* Philippines, Cebu Island HQ239368 B. taylori KU 307738 Philippines, Negros Island, Municipality of Valencia, HQ239369 Mt. Talinis B. taylori RMB 3226* Philippines, Negros Island, Municipality of Valencia, HQ239370 Mt. Talinis B. mindorensis KU 307739 Philippines, Mindoro Island, Municipality of Bongabong HQ239371 B. mindorensis KU 307740 Philippines, Mindoro Island, Municipality of Bongabong HQ239372 B. boholensis RMB 2866* Philippines, Bohol Island, Municipality of Bilar HQ239373 B. boholensis RMB 2875* Philippines, Bohol Island, Municipality of Bilar HQ239374 B. boulengeri KU 307753 Philippines, Polillo Island, Municipality of Polillo HQ239375 B. boulengeri KU 307754 Philippines, Polillo Island, Municipality of Polillo HQ239376 B. boulengeri KU 313831 Philippines, Luzon Island, Municipality of Labo, Mt. Labo HQ239377 B. boulengeri KU 313829 Philippines, Luzon Island, Municipality of Labo, Mt. Labo HQ239378 B. boulengeri KU 320058 Philippines, Luzon Island, Municipality of Laguna, HQ239379 Mt. Makiling B. boulengeri KU 320059 Philippines, Luzon Island, Municipality of Laguna, HQ239380 Mt. Makiling B. kadwa ACD 3290* Philippines, Luzon Island HQ239381 B. kadwa RMB 4040* Philippines, Luzon Island, Municipality of Irosin, Mt. HQ239382 Bulusan B. kadwa KU 307965 Philippines, Camiguin Norte Island, Municipality of HQ239383 Calayan B. kadwa KU 307985 Philippines, Camiguin Norte Island, Municipality of HQ239384 Calayan B. kadwa KU 304906 Philippines, Calayan Island, Municipality of Calayan HQ239385 B. kadwa KU 304875 Philippines, Calayan Island, Municipality of Calayan HQ239386 B. talinis RMB 3305* Philippines, Negros Island, Municipality of Valencia, HQ239387 Mt. Talinis B. talinis KU 303990 Philippines, Sibuyan Island, Municipality of Magdiwang HQ239388 B. talinis KU 306786 Philippines, Panay Island, Municipality of Sibalom HQ239389 B. talinis KU 306781 Philippines, Panay Island, Municipality of Sibalom HQ239390 B. talinis KU 315358 Philippines, Tablas Island, Municipality of Calatrava HQ239391 B. talinis KU 315359 Philippines, Tablas Island, Municipality of Calatrava HQ239392 B. tungaoi KU 323935 Philippines, Masbate Island, Municipality of Mobo HQ239393 B. tungaoi KU 323936 Philippines, Masbate Island, Municipality of Mobo HQ239394 B. tungaoi KU 323933 Philippines, Masbate Island, Municipality of Mobo HQ239395 B. tungaoi KU 323934 Philippines, Masbate Island, Municipality of Mobo HQ239396 B. schadenbergi PNM/CMNH Philippines, Mindanao Island, Municipality of Kiamba, HQ239397 H1457 Mt. Busa B. schadenbergi KU 314980 Philippines, Mindanao Island, Municipality of HQ239398 Zamboanga City B. schadenbergi KU 314981 Philippines, Mindanao Island, Municipality of HQ239399 Zamboanga City B. orientalis KU 314092 Philippines, Mindanao Island, Municipality of San HQ239400 Francisco B. orientalis KU 311239 Philippines, Leyte Island, Municipality of Baybay HQ239401 B. orientalis KU 311230 Philippines, Leyte Island, Municipality of Baybay HQ239402 B. orientalis KU 310945 Philippines, Samar Island, Municipality of Taft HQ239403 B. orientalis KU 310357 Philippines, Camiguin Sur Island, Municipality of HQ239404 Mambajao B. orientalis KU 305469 Philippines, Dinagat Island, Municipality of Loreto HQ239405 B. orientalis KU 324028 Philippines, Bohol Island, Municipality of Bilar HQ239406 6 HERPETOLOGICAL MONOGRAPHS [No. 24

TABLE 2.—Models of evolution selected by Akaike Morphological Data Information Criterion (AIC) and applied for partitioned, Bayesian phylogenetic analyses.a We examined fluid-preserved specimens (see Appendix) for variation in qualitative No. of Partition AIC model characters and mensural characters. Sex was determined by gonadal inspection, and measurements ATP8, first codon position HKY + G53were taken to the nearest 0.1 mm with digital ATP8, second codon position HKY + I + G53 ATP8, third codon position GTR + G53calipers by CDS. Museum abbreviations for ATP6, first codon position GTR + I 227 specimens examined follow Leviton et al. ATP6, second codon position GTR + I + G 227 (1985). ATP6, third codon position GTR + G 227 Meristic and mensural characters were a The model GTR + I + G was used for partitioned RAxMLHPC analyses. chosen based on Siler et al. (2009, 2010a,b): characters evaluated were snout–vent length (SVL), axilla–groin distance (AGD), total partitioned into six regions consisting of the length (TotL), midbody width (MBW), mid- codon positions of ATP8 and ATP6. Analyses body height (MBH), tail length (TL), tail that partition protein-coding genes by codon width (TW), tail height (TH), head length position have been shown to improve resulting (HL), head width (HW), head height (HH), inferences (Brandley et al., 2005). The parti- snout–forearm length (SnFa), eye diameter tions were run under the same model (GTR + (ED), eye–narial distance (END), snout I + C), with 100 replicate best-tree inferences. length (SNL), internarial distance (IND), Each inference was performed with a random forelimb length (FLL), hind limb length starting tree and relied on the rapid hill- (HLL), midbody scale-row count (MBSR), climbing algorithm (Stamatakis, 2006). Clade paravertebral scale-row count (PVSR), axilla– support was assessed with 1000 bootstrap groin scale-row count (AGSR), Finger-III pseudoreplicates. We considered branches lamellae count (FinIIIlam), Toe-IV lamellae receiving $70% bootstrap support to be well count (ToeIVlam), supralabial count (SL), supported (Hillis and Bull, 1993; also see infralabial count (IFL), supraciliary count Wilcox et al., 2002). The Akaike Information (SC), and supraocular count (SO). In the Criterion (AIC), as implemented in MrMo- description, ranges are followed by mean 6 delTest 2.2 (Nylander, 2004), was used to find SD in parentheses. appropriate models of sequence evolution. The best-fit model for each of the six Species Concept partitions (Table 2) was used for Bayesian We follow the general lineage concept of analyses performed in MrBayes 3.1 (Ronquist species (de Queiroz, 1998, 1999) as a logical and Huelsenbeck, 2003). The same partition- extension of the evolutionary species concept ing strategy used for ML analyses was used for (Frost and Hillis, 1990; Simpson, 1961; Wiley, Bayesian inferences. Searches over tree space 1978). We consider as distinct lineages those were conducted with four runs, each with four populations that are morphologically, and chains, and were run for 2 3 107 generations. genetically, distinct. Lineage-based species Trees were sampled every 1000 generations, concepts have been successfully used in the with 4000 samples discarded as burn-in; this recognition of Philippine biodiversity (Brown left 16,001 post burn-in trees from each run and Diesmos, 2002; Brown et al., 2000a, 2008, included in the summary. Visual inspection 2009; Brown and Guttman, 2002; Gaulke et for chain stationarity and high effective al., 2007; Welton et al., 2009, 2010) due to the sample size values was conducted within the highly partitioned nature of the archipelago program Tracer, version 1.4 (Rambaut and (Brown and Diesmos, 2009), and because the Drummond, 2007). In addition, correlations geological history of the islands has been so of split frequencies and cumulative split well documented (Hall, 2002; Yumul et al., frequencies were examined using the program 2009). In this study, we use an estimate of AWTY (Nylander et al., 2008). We considered phylogenetic relationships as a guide for branches with posterior probabilities $0.95 to delimiting species but restrict our diagnoses be well supported (Wilcox et al., 2002). of new species to those populations diagnosed 2010] HERPETOLOGICAL MONOGRAPHS 7 by nonoverlapping morphological character Island], B. sp. nov. [Calayan, Camiguin states. Norte, and Luzon islands], B. b. mindorensis, B. talinis, B. b. taylori, B. s. orientalis,andB. RESULTS s. schadenbergi) are distinguished from congeners by levels of genetic divergence Phylogeny equal to, or greater than, those between Of 838 aligned mitochondrial nucleotide previously defined species, viz., B. boulen- positions, 392 and 306 were variable and geri, B. talinis,andB. schadenbergi (Table 3; parsimony-informative, respectively. Just con- Fig. 2). The two most genetically similar sidering the alignment for Brachymeles se- lineages (B. sp. nov. [Masbate Island] and quence data, 310 and 279 were variable and B. sp. nov. [northern Philippines]) are parsimony-informative, respectively. The ML separated by 3.8–5.2% sequence divergence, analysis resulted in a single optimal tree and sequence divergences among all subspe- (2lnL 5 5249.903). The resulting topology cies of B. boulengeri,andamongboth from the Bayesian analysis is very similar to subspecies of B. schadenbergi,are.7.3% the ML tree. Trees estimated from ML, MP, (Table 3; Fig. 2). The three lineages with the and Bayesian analyses are consistent with greatest range of sequence divergence across respect to support for nine unique species of populations are the presently defined sub- Brachymeles. specific taxa that occur across multiple No inferences support the monophyly of B. islands within a single PAIC (B. b. boulen- boulengeri. All analyses recover B. b. boho- geri, B. b. taylori,andB. talinis;Table3; lensis as part of a clade with B. s. orientalis Figs. 2, 3, 5). Sequence divergence among and B. s. schadenbergi with high support populations in the lineage of northern (Fig. 2). Results of both analyses (ML and populations of B. talinis, and among popula- Bayesian) show a polytomy with four lineages, tions of B. schadenbergi orientalis,aremuch including B. b. mindorensis, B. b. taylori, B. b. lower than those of other lineages known to boulengeri, and a clade consisting of the be distributed across multiple islands (Ta- remaining species (Fig. 2). All analyses sup- ble 3; Figs. 3–5). port a clade of B. b. boholensis, B. s. schadenbergi, and B. s. orientalis as sister to Morphology a clade of three distinct lineages of B. talinis Variation in morphological characters (Ta- samples, whereas MP analyses consistently bles 4–6) mirrors the results observed in supported the clade of B. talinis as sister to a phylogenetic analyses and supports the rec- clade of B. b. mindorensis, B. b. taylori,andB. ognition of nine Brachymeles lineages. In b. boulengeri. With the inclusion of two short addition, comparison of meristic and mensural mitochondrial genes for only three of the 18 morphological characters identified a 10th recognized species in the genus, it is likely unique lineage from Jolo Island, previously that differences in topologies among ML and recognized as a population of B. talinis. Bayesian analyses reflect limited taxon and Characters differing among these 10 lineages character sampling. Nevertheless, all analyses include body size, degree of limb develop- result in the strong support of nine genetically ment, finger and toe lamellae counts, head distinct lineages of Brachymeles (Fig. 2). In and body scale counts and patterns, and addition, samples of B. talinis cluster into pigmentation patterns (Tables 4–6; see spe- three major clades, each of which is strongly cies accounts below). We observed no men- supported in all analyses (Fig. 2). sural or meristic differences between the Uncorrected pairwise sequence divergenc- sexes of any of the 10 species. es are low within named taxa and relatively Superficially, subspecies of both B. boulen- high between these lineages (Table 3). Lev- geri and B. schadenbergi, and island popula- els of sequence divergence show that the tions of B. talinis, seem morphologically nine mitochondrial DNA lineages discovered similar, especially in overall body size; how- by our phylogenetic analyses (B. b. boholen- ever, numerous nonoverlapping differences sis, B. b. boulengeri, B. sp. nov. [Masbate were detected in meristic, mensural, and color 8 HERPETOLOGICAL MONOGRAPHS [No. 24

FIG. 2.—Maximum clade credibility tree from a phylogenetic analysis of mitochondrial data (mitochondrial ATPase 6[ATP6] and ATP8; 2lnL 5249.903214). Nodes shown with numerical values corresponding to maximum-likelihood nonparametric bootstrap support, and Bayesian posterior probabilities support values, respectively. Terminals are labeled with taxonomic names and sampling localities. pattern characters for each complex member, Island). Combined with biogeographic evi- readily defining 10 distinct lineages between dence, and clearly separate geographical the three complexes (Tables 4–6). ranges, our data suggest the presence of 10 In summary, each lineage possesses unique evolutionary lineages, worthy of taxonomic and nonoverlapping suites of diagnostic char- recognition. acter states of morphology, perfectly corre- sponding to nine of the clades defined in Taxonomic Conclusions phylogenetic analyses of DNA sequence data Our inferred phylogeny (Fig. 2), biogeogra- (tissues unavailable for B. cf. talinis from Jolo phically separate ranges of island endemic 2010] HERPETOLOGICAL MONOGRAPHS 9

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 0.6 boulengeri 15.3–15.9 0.0–4.7 mindorensis 16.1–16.7 10.6–11.5 0.6 taylori 15.6–16.0 10.2–11.6 11.0–11.2 0.3–4.8 talinis 13.0–14.6 10.4–11.8 11.8–12.4 10.3–12.4 0.1–4.5 kadwa 12.7–13.6 10.4–11.7 11.5–12.1 9.3–10.3 6.9–8.6 0.1–2.2 tungaoi 13.2–13.9 10.5–12.1 10.7–11.6 9.7–10.1 7.2–8.3 3.8–5.2 0.0–1.5 orientalis 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 0.6–2.1 schadenbergi 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 species; diagnostic, nonoverlapping morpho- Designation of a neotype for B. boulen- logical character states; and genetic diver- geri.—Taylor’s holotype for B. boulengeri gences between the taxa (Table 3) indicate the (Philippine Bureau of Science Publication distinctiveness of a new species from Luzon, No. 17: 246, collected 15 July 1920) was Calayan, and Camiguin Norte islands, a new destroyed in the destruction of the Philippine species from Masbate Island, and a new Bureau of Science in World War II, with no species from Jolo Island. In addition, the molecular and morphological data strongly support the elevation of all subspecies of B. boulengeri and B. schadenbergi to full species (Table 3; Fig. 2). Each of the 10 species is morphologically distinct from each other and all other known species in the genus, and each of the nine species included in phylogenetic analyses are also genetically distinct. With the exception of B. talinis, each lineage is endemic to one of four isolated PAICs, thereby providing additional support for the distinc- tiveness of each lineage’s evolutionary history and integrity. Accordingly, we recognize all four subspecies of the former polytypic species B. boulengeri, and both subspecies of the former polytypic species B. schadenbergi, as full species.

TAXONOMIC ACCOUNTS Brachymeles boulengeri Taylor 1922b: 246 Figs. 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. FIG. 3.—Hypothesized distributions of Brachymeles Brachymeles gracilis (part), Brown and Alcala, boholensis, B. boulengeri, B. mindorensis, and B. taylori in the Philippines. The sampling localities are indicated by 1970. black shapes, and the hypothesized geographic range of Brachymeles boulengeri boulengeri (part), each species indicated by shaded islands, with shapes and Brown and Alcala, 1980. shades of islands corresponding to the map’s key. 10 HERPETOLOGICAL MONOGRAPHS [No. 24

FIG. 4.—Hypothesized distributions of Brachymeles FIG. 5.—Hypothesized distributions of Brachymeles orientalis, B. schadenbergi, and B. vindumi in the tungaoi, B. kadwa, and B. talinis in the Philippines. The Philippines. The sampling localities are indicated by black sampling localities are indicated by black shapes, and the shapes, and the hypothesized geographic range of each hypothesized geographic range of each species indicated species indicated by shaded islands, with shapes and by shaded islands, with shapes and shades of islands shades of islands corresponding to the map’s key. corresponding to the map’s key. Islands of the Romblon Unknown Mindanao Island range boundaries indicated Island group are designated by the first letter of the island by dashed lines. name (T, Tablas Island; R, Romblon Island; and S, Sibuyan Island). mention of a repository for the holotype. In the absence of an existing holotype and in contains adults of both sexes and agrees with accordance with article No. 75 of the Inter- Taylor’s (1922a) holotype description. national Code of Zoological Nomenclature Neotype.—PNM 9720 (RMB Field No. (ICZN, 1999), we designate a neotype for this 5647, formerly KU 307756), adult male, species. Accordingly, we choose an adult male collected from a fallen, rotting log in second- specimen from the type locality of Polillo ary growth forest (1000 to 1230 h) in Barangay Island. Preserved adult specimens from Tay- Pinaglubayan, Municipality of Polillo, Quezon lor’s (1922b) type locality (Polillo Island) have Province, Polillo Island, Philippines (14u 459 been examined in the collections at California 090 N, 121u 589 060 E; WGS-84), by RMB, J. Academy of Sciences (CAS); unfortunately, Fernandez, Y. Vicente, and M. Vicente. these original specimens are either poorly Diagnosis.—Brachymeles boulengeri can be preserved, incomplete, or not sexually mature. distinguished from congeners by the following However, recent collections from Polillo combination of characters: (1) body size Island (Fig. 1) have resulted in well-preserved moderate (SVL 60.5–93.1 mm); (2) pentadac- adult individuals that clearly exhibit the tyl; (3) Finger-III lamellae five or six; (4) Toe- diagnostic characters for the species. From IV lamellae nine or 10; (5) moderate limb these collections, we have chosen a male length; (6) supralabials six or seven; (7) neotype collected as part of a series that infralabials seven; (8) pineal eye spot present; 2010] HERPETOLOGICAL MONOGRAPHS 11

(9) supranasals not contacting on midline; (10) auricular openings (vs. absence). From all prefrontals not contacting on midline; (11) nonpentadactyl species except for B. pathfin- midline contact of first pair of chin shields; deri, B. boulengeri differs by having a (12) enlarged chin shields in two pairs; (13) paravertebral scale count of 63–66 (vs. .84). nuchal scales undifferentiated; (14) fourth and From B. apus, B. lukbani, B. minimus, and B. fifth supralabial below eye; (15) auricular vermis, B. boulengeri is distinguished by the opening present; (16) continuous, light dorso- presence (vs. absence) of limbs. lateral stripes present; and (17) middorsal Description of neotype.—Adult male, hemi- stripes absent (Tables 4 and 5). penes everted (Fig. 6); SVL 93.1 mm; body Comparisons.—Characters distinguishing moderate relative to other Brachymeles;head B. boulengeri from all pentadactyl species of weakly differentiated from neck, nearly as Brachymeles are summarized in Tables 4 and wide as body, HW 10.3% SVL, 109.0% HL; 5. Brachymeles boulengeri most closely re- HL 36.5% SnFa; SnFa 9.4% SVL; snout sembles B. boholensis, B. mindorensis, and B. moderately long, rounded in dorsal and taylori, but it differs from these three taxa by lateral profile, SNL 52.2% HL; auricular having six or seven supralabials and five or six opening present, moderate; eyes moderate, supraciliaries, and by the absence of contin- ED 2.2% SVL, 23.5% HL, 72.0% END, pupil uous, dark middorsal stripes (Tables 4 and 5). subcircular; body slightly depressed, MBW Brachymeles boulengeri further differs from 141.8% MBH; body scales smooth, glossy, B. boholensis by having a relatively longer tail, imbricate; longitudinal scale rows at midbody five or six Finger-III lamellae, the fourth and 26; paravertebral scale rows 64; axilla–groin fifth supralabial below the eye, midline scale rows 43; limbs well developed, penta- contact between the first pair of enlarged dactyl, digits small; FinIIIlam 5; ToeIVlam 9; chin shields, and by lacking a third pair of FLL 19.2% AGD, 12.5% SVL; HLL 28.7% enlarged chin shields (Tables 4 and 5); from AGD, 18.7% SVL; order of digits from B. mindorensis it is differentiated by its shortest to longest for hand: V 5 I , IV 5 smaller body size, shorter hind limbs, and II , III, for foot: I , V , II , III , IV; tail possession of nine or 10 Toe-IV lamellae, original, not as wide as body, sharply tapered seven infralabials, and by having the fourth toward the end, TW 70.1% MBW, TL 84.6% and fifth supralabial below the eye (Tables 4 SVL. and 5); and from B. taylori by the midline Rostral projecting dorsoposteriorly to point contact between the first pair of enlarged chin in line with anterior edge of nasal scale, shields, and the presence of continuous, light broader than high, forming a narrow suture dorsolateral stripes (Tables 4 and 5). with frontonasal; frontonasal wider than long; From all nonpentadactyl species of Brachy- nostril ovoid, centered in a single rectangular meles (B. apus, B. bonitae, B. cebuensis, B. nasal; nasals well separated; supranasals pres- elerae, B. lukbani, B. minimus, B. muntingka- ent, large, moderately separated by frontona- may, B. pathfinderi, B. samarensis, B. tridac- sal; postnasals present; prefrontals narrowly tylus, B. vermis, and B. wrighti), B. boulengeri separated by frontal; frontal suboctagonal, differs by having a pentadactyl body form (vs. anterior margin in moderate contact with nonpentadactyl), greater forelimb lengths frontonasal and first two anterior supraocu- (.8.2 mm vs. ,6.9 mm), greater hind limb lars, 53 wider than anteriormost supraocular; lengths (.14.3 mm vs. ,12.9 mm), Toe-IV supraoculars five; frontoparietals moderate, in lamellae nine or 10 (vs. eight or fewer), a broad medial contact, contact 2–4 supraocu- midbody scale row count of 26 or 27 (vs. ,24 lars; interparietal diamond shaped, slightly in all nonpentadactyl species except for 28 in wider than long, nearly one half frontal length; B. wrighti; Taylor, 1925), and by the presence parietal eyespot present in posterior one third of a postnasal scale (vs. absence). With the of scale; parietals separated by interparietal; exception of B. pathfinderi, B. boulengeri nuchals nonenlarged, undifferentiated from differs further from all nonpentadactyl species dorsal scales; loreals two, decreasing in size by the absence of a third pair of enlarged chin from anterior to posterior; anterior loreal shields (vs. presence) and the presence of approximately as long as and slightly higher 12 HERPETOLOGICAL MONOGRAPHS [No. 24

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 Pleistocene aggregate island complexes, sensu Brown and Diesmos, 2002) are included for reference (snout–vent length [SVL], total length [TotL], midbody width [MBW], forelimb length [FLL], and hind limb length [HLL] given as range over mean 6 SD; 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.

taylori boholensis boulengeri mindorensis (8 male, gracilis gracilis gracilis hilong (5 male, (7 male, (6 male, 13 female) (7 male, (8 male, 14 female) 8 female) 12 female) 11 female) 12 female) Negros and Range Bohol Island Luzon PAIC Mindoro Island 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)

than posterior loreal, in contact with prefron- second pair slightly wider than first, separated tal, postnasal, supranasal, second supralabial; by a single medial scale. posterior loreal and frontonasal; preocular Scales on limbs smaller than body scales; single, nearly two thirds the height of scales on dorsal surfaces of digits large, posterior loreal; presubocular single; supraci- wrapping around lateral edges of digits; lamel- liaries six, most anterior contacting prefrontal lae undivided; palmar surfaces of hands and and separating posterior loreal from first plantar surfaces of feet covered by small, supraocular, most posterior extending to irregular scales, each with raised anterior edges; midline of last supraocular; subocular row scales on dorsal surface of hands and feet complete; lower eyelid with one row of scales, smaller than limb scales, lacking raised edges. lacking an enlarged oval window, largely Coloration of neotype in preservative.— transparent; supralabials seven, fourth and Ground color of body medium brown; lateral fifth below the eye, infralabials seven. and ventral surfaces of body lacking dark Mental wider than long, in contact with first pigment; dorsum, from posterior edge of infralabials; postmental single, wider than supranasals to tail tip, uniformly dark brown, mental, followed by two pairs of enlarged with color spanning six full and two half rows of chin shields; first pair in broad medial contact, scales at midbody and narrowing to cover four 2010] HERPETOLOGICAL MONOGRAPHS 13

TABLE 4.—Extended.

kadwa (12 male, bicolor talinis 15 female) tungaoi makusog orientalis schadenbergi vindumi (7 male, (11 male, (2 male, (7 male, (21 male, (14 male, (1 male, 6 female) 10 female) Luzon, Babuyan, 6 female) 4 female) 19 female) 10 female) 1 female) Claro, and Calayan Luzon Island Visayan PAIC 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, (14.4 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) 14.8 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, (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) 13.3 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, (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) 22.7 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) rows of scales posterior to parietals; darker Coloration in life.—Dorsal ground color pigmentation covers entire surface of dorsal homogeneous medium brown to yellowish scales, with exception of pigmentation on half brown (Fig. 11A); gradual lateral dorsolateral rows of scales; head scales uniform brown; demarcation between dorsal (dark) and ven- lateral half of supraoculars lacking dark pig- tral (light) coloration; lateral and ventral mentation; rostral, nasal, postnasal, supranasal, surfaces of body homogeneous medium and first supralabial dark gray with light brown brown to yellowish brown; dark brown spots blotches; pineal eyespot charcoal, surrounded and longitudinal lines of spots absent from by cream border. Faint, indistinct light dorso- lateral surfaces. lateral stripes, formed by the absence of dark Measurements of neotype (in millimeters).— pigmentation, extending from level of anterior SVL 93.1; AGD 60.7; TotL 171.9; MBW edge of eye to base of tail, spanning one full and 14.7; MBH 10.4; TL 78.8; TW 10.3; TH two half rows of scales; small blotch of dark 8.3; HL 8.8; HW 9.6; HH 7.2; SnFa 24.0; ED brown pigment dorsal to auricular opening. 2.1; END 2.9; SNL 4.6; IND 3.1; FLL 11.7; Limbs mottled light and medium brown HLL 17.5; MBSR 26; PVSR 64; AGSR 43; dorsally, yellowish brown ventrally; dorsal and FinIIIlam 5; ToeIVlam 9; SL 7; IFL 7; SC 6; ventral surface of digits dark brown. SO 5. TABLE 5.—Summary of qualitative diagnostic characters (present, absent) in all known pentadactyl species [ of Brachymeles. The pairs of enlarged MONOGRAPHS HERPETOLOGICAL scales posterior to the 14 postmental scale are abbreviated as chin shield pairs with reference to the first, second, and third pairs (when present). Character abbreviations are as follows: MBSR 5 midbody scale-row count; AGSR 5 axilla–groin scale-row count; SL 5 supralabial count; IFL 5 infralabial count; SC 5 supraciliary count; SO 5 supraocular count.

gracilis gracilis boholensis boulengeri mindorensis taylori gracilis hilong bicolor talinis kadwa tungaoi makusog orientalis schadenbergi vindumi (5 male, 14 (7 male, 8 (6 male, 12 (8 male, 13 (7 male, 11 (8 male, 12 (7 male, 6 (11 male, 10 (12 male, 15 (2 male, 6 (7 male, 4 (21 male, (14 male, (1 male, 1 female) female) female) female) female) female) female) female) female) female) female) 19 female) 10 female) female) MBSR 26–28 26–27 26–28 26–28 24–26 27–30 27–29 26–30 26–28 26–28 25–28 26–28 26–28 30, 31 AGSR 42–46 42–46 42–45 42–47 46–49 44–50 64–72 43–48 47–49 46–49 42–47 46–49 45–50 49, 49 PVSR 63–66 63–66 63–65 62–69 67–70 66–73 88–92 67–72 68–70 66–68 60–69 69–72 67–72 74, 74 SL 7 (19) 6 (12) 7 (18) 6 (21) 6 (18) 6 (20) 6 (13) 7 (20) 7 (27) 7 (4) 6 (7) 6 (31) 6 (2) 7 (2) 7 (3) 7 (4) 7 (9) 7 (22) IFL 7 (19) 7 (15) 6 (18) 7 (21) 7 (18) 6 (20) 6 (13) 7 (21) 6 (27) 6 (4) 6 (10) 6 (7) 6 (10) 6 (2) SC 6 (19) 5 (1) 6 (18) 6 (21) 6 (18) 6 (20) 6 (13) 7 (1) 7 (33) 7 (14) 6 (14) 6 (21) 6 (27) 6 (4) 6 (11) 6 (40) 6 (24) 6 (2) SO 5(19) 5 (15) 5 (18) 5 (21) 5 (18) 5 (20) 5 (13) 5 (21) 5 (27) 5 (4) 5 (11) 5 (40) 5 (24) 5 (2) Pineal eyespot ++++2 +++ + + + +++ Supranasal contact 222or + 222 ++ + + 22++ Prefrontal contact 222or + 2 + 222222222 Frontoparietal contact +++++2 or ++ + 2 or ++ + + + + Parietals contact 2 or + 2 or + 2 or + 2 or ++ + +2 or + 2 or + 2 or + 22or + 2 or ++ First chin shield pair 2 or ++ +2 or + 22or ++2 or + 2 or ++ 2 or + 2 or + 2 or + 2 contact Enlarged chin shield 3 2223322 2 2 2 222 pairs Chin shield pair size 3 , 1 , 21, 21, 21, 23, 1 , 23, 1 , 22, 12, 11, 215 22, 12, 12, 12, 1 Chin shield pair 1(0/1); 1(0/1); 1(0); 1(0); 1(1); 1(0/1); 1(0); 1(0/1); 1(0/1); 1(0); 1(1); 1(0/1); 1(0); 1(1); separationa 2(1); 2(1) 2(1) 2(1) 2(1); 2(1); 2(3) 2(3) 2(1) 2(1) 2(3) 2(3) 2(3) 2(1) 3(3) 3(3) 3(3/5) Subocular supralabial Fifth and Fourth Fifth and Fourth Fourth Fourth Fourth Fifth Fifth Fifth Fourth and Fourth Fifth Fifth sixth and sixth and and and and and and and fifth or and and and fifth fifth fifth fifth fifth sixth sixth sixth fifth and fifth sixth sixth sixth Differentiated nuchals 2 22222 + 22 2 2 222 Uniform body color 2222++22222222 Continuous, light +++2 ++22or ++, poorly +, poorly 222+ dorsolateral stripes defined defined Continuous dark + 2 +++2 or + 22or ++ + 222or + 2 middorsal stripesb Dark lateral stripesb + +++++2 ++ + 222or ++ Dark ventral + 222 ++22 + 2222+ pigmentation 24 No. a Parentheses show the number of small ventral scale rows separating each enlarged pair of chin shields. b Character refers to longitudinal rows of dark scale pigmentation, often in the form of spots, aligned into rows. 2010] HERPETOLOGICAL MONOGRAPHS 15

Variation.—Variation in mensural charac- areas. Individuals have been observed under ters is summarized in Table 6. Among the 19 piles of rotting coconut husks, in the humus specimens examined for the degree of contact material within rotting logs, and in loose soil between parietal scales, nine specimens pos- and leaf litter surrounding the root networks sessed parietals separated by the interparietal of trees. This species is quite common at the (KU 307750, 307758, 320060, 322315–20), type locality, which has been virtually com- and 10 specimens possessed parietals in pletely converted to coconut plantations. moderate-to-broad medial contact (KU When disturbed, individuals immediately 307438–9, 307751–4, 307757, 320058–9, move in a rapid serpentine manner, attempt- 322314) behind the interparietal. ing to burrow into loose soil or humus. Scale counts were observed to vary among Sympatric lizard species observed on Lu- the measured series. With the exception of a zon, Polillo, and Marinduque islands include single specimen with five supraciliaries (KU the following: (Agamidae) Bronchocela crista- 307752), all specimens examined had six tella, Draco spilopterus, Gonocephalus so- supraciliaries. The number of supralabials phiae, Hydrosaurus pustulatus; (Gekkonidae) varied between six (CAS 61297, 62272–3, Cyrtodactylus philippinicus, Gehyra mutilata, 62276–7, KU 307438–9, 307752–4, 307757–8) Gekko gecko, Gekko mindorensis, Hemidacty- and seven (CAS 61096, KU 307751). Speci- lus frenatus, H. garnoti, H. luzonensis, H. mens were observed to have midbody scale platyurus, Pseudogekko compressicorpus, P. row counts of 26 (CAS 61096, 31297, 62273, smaragdina; (Scincidae) B. bonitae, B. bicolor, 62276–7, KU 307439, 307751–4, 307758) and B. elerae, B. lukbani, B. makusog, B. mun- 27 (CAS 62272, KU 307750, 307757); axilla– tingkamay, B. samarensis, B. kadwa, B. groin scale row counts of 42 (KU 307439, wrighti, Emoia atrocostata, Eutropis bonto- 307758), 43 (KU 307750–4, 307757), 44 (CAS censis, E. multicarinata, E. multifasciata, 61096, 62272–3, 62276–7), and 46 (CAS Lamprolepis smaragdina, Lipinia pulchella, 61297); and paravertebral scale row counts Sphenomorphus cumingi, S. decipiens, S. of 63 (KU 307439, 307750, 307752, 307754, jagori, S. leucospilos, S. luzonensis, S. steerei, 307757–8), 64 (CAS 61096, 62272–3, 62277, S. stejnegeri, Tropidophorus grayi; and (Var- KU 307751, 307753), 65 (CAS 62276), and 66 anidae) Varanus marmoratus and V. olivaceus. (CAS 61297). We also observed lamellae counts to vary Brachymeles boholensis Brown and Rabor among the measured series. With the excep- 1967 tion of a single specimen with six Finger-III Figs. 3, 6, 11C lamellae (KU 307750), all specimens examined Brachymeles gracilis boholensis, Brown and had five. Two specimens examined had 10 Toe- Rabor, 1967. Type locality: 6 km southeast IV lamellae (KU 307750, 307757), whereas the of Sierra Bullones, Teacher’s Park, Bohol remaining specimens examined had nine. Island, Philippines, 400–433-m elevation, Distribution.—Brachymeles boulengeri oc- 9u 479 32.530 N, 124u 189 14.40 E (holotype: curs in central and southern Luzon and on CAS-SU24528). Marinduque and Polillo islands (Fig. 3). The Brachymeles gracilis (part), Brown and Alcala, species has been collected in the Camarines 1970. Norte Province of the Bicol Peninsula and Brachymeles boulengeri boholensis (part), may eventually be found to occur further Brown and Alcala, 1980. south on the Bicol Peninsula and even on Diagnosis.—Brachymeles boholensis can be Catanduanes Island. distinguished from congeners by the following Ecology and natural history.—Brachymeles combination of characters: (1) body size boulengeri occurs in disturbed and secondary moderate (SVL 83.8–93.6 mm); (2) limbs growth forest. Little or no original, low- pentadactyl; (3) Finger-III lamellae six; (4) elevation forest remains throughout the range Toe-IV lamellae nine or 10; (5) limb length of B. boulengeri, but we assume the species moderate; (6) supralabials seven; (7) infra- once also occurred in first-growth forest when labials seven; (8) pineal eye spot present; (9) this forest type extended into low-elevation supranasals not contacting on midline; (10) 6HREOOIA OORPS[ MONOGRAPHS HERPETOLOGICAL 16

TABLE 6.—Summary of univariate morphological variation among mensural characters in series of Brachymeles boholensis, B. boulengeri, B. mindorensis, B. taylori, B. talinis, B. kadwa, B. tungaoi, B. orientalis, B. schadenbergi,andB. vindumi. Character abbreviations are as follows: SVL 5 snout–vent length; AGD 5 axilla–groin distance; TotL 5 total length; MBW 5 midbody width; MBH 5 midbody height; TL 5 tail length; TW 5 tail width; TH 5 tail height; HL 5 head length; HW 5 head width; HH 5 head height; SnFa 5 snout–forearm length; ED 5 eye diameter; END 5 eye–narial distance; SNL 5 snout length; IND 5 internarial distance; FLL 5 forelimb length; HLL 5 hind limb length.

boholensis boulengeri mindorensis taylori talinis kadwa tungaoi orientalis schadenbergi vindumi (5 male; (7 male; (6 male; (8 male; (11 male; (12 male; (2 male; (21 male; (14 male; (1 male; 14 female) 8 female) 12 female) 13 female) 10 female) 15 female) 6 female) 19 female) 10 female) 1 female) SVL (male) 84.1–93.6 72.3–93.1 93.9–104.2 83.1–99.2 103.1–123.1 97.0–128.2 89.2, 104.8 84.7–112.3 94.4–115.8 113.6 (89.1 6 4.1) (82.5 6 6.7) (100.2 6 4.1) (87.0 6 5.2) (113.6 6 7.1) (109.3 6 9.1) (99.0 6 8.4) (104.5 6 6.1) SVL (female) 83.8–94.0 60.5–95.5 90.0–106.8 65.8–93.2 103.8–126.7 90.6–135.7 78.2–106.2 97.6–115.2 93.1–113.5 104.9 (88.4 6 3.1) (84.0 6 11.2) (98.8 6 5.3) (83.9 6 7.4) (116.5 6 6.8) (109.7 6 11.7) (95.6 6 10.7) (104.6 6 5.9) (104.8 6 6.7) AGD (male) 54.7–61.9 46.3–60.7 59.5–67.6 52.8–66.0 64.2–90.3 54.1–84.5 56.8, 69.3 51.0–75.3 60.3–76.8 72.7 (58.1 6 2.8) (53.0 6 5.0) (63.3 6 3.3) (55.9 6 4.2) (74.6 6 7.5) (70.4 6 7.3) (62.7 6 7.0) (67.7 6 4.9) AGD 53.0–61.2 36.2–61.3 56.3–74.9 46.6–60.6 66.5–81.4 48.0–93.7 49.8–69.6 57.9–75.9 64.8–92.3 67.4 (female) (57.0 6 2.7) (54.3 6 8.3) (64.8 6 5.5) (54.7 6 4.6) (74.7 6 4.9) (71.8 6 10.7) (61.9 6 7.7) (66.6 6 4.8) (72.0 6 7.7) TotL (male) 154.5–166.2 124.3–173.1 165.3–197.0 149.6–176.7 191.7–238.4 181.9–255.2 178.3, 203.1 154.6–221.1 179.2–219.7 — (160.7 6 5.9) (151.4 6 19.4) (184.9 6 11.5) (164.3 6 11.3) (209.0 6 12.4) (208.3 6 20.2) (184.8 6 22.3) (203.0 6 14.4) TotL 129.6–174.8 129.7–167.4 162.5–206.7 130.3–168.5 187.5–236.2 170.4–221.0 168.9–206.5 159.7–213.0 180.8–217.4 — (female) (154.1 6 14.7) (159.3 6 13.1) (180.2 6 14.2) (149.9 6 13.0) (209.4 6 18.0) (196.3 6 18.7) (187.2 6 15.9) (194.3 6 17.1) (202.4 6 17.6) MBW (male) 12.2–13.9 10.1–14.7 12.8–16.0 11.6–16.8 15.9–20.1 14.3–19.6 13.9, 15.2 11.0–17.7 13.5–17.2 14.2 (13.0 6 0.7) (12.1 6 1.9) (14.7 6 1.1) (13.3 6 1.7) (17.7 6 1.3) (16.5 6 1.6) (14.4 6 1.6) (14.8 6 1.3) MBW 11.9–15.0 9.9–14.6 14.5–20.8 11.0–16.6 17.3–20.9 14.3–21.0 11.9–17.0 12.3–18.9 13.9–19.1 14.8 (female) (13.6 6 1.1) (12.7 6 1.7) (16.7 6 1.8) (14.0 6 1.8) (19.3 6 1.1) (17.2 6 2.0 (14.4 6 1.9) (15.8 6 1.8) (15.9 6 2.4) MBH (male) 11.7–12.5 7.9–10.4 10.2–12.9 9.4–14.6 10.4–15.9 9.2–14.9 8.6, 14.6 9.6–15.2 12.1–14.8 11.6 (12.1 6 0.4) (9.0 6 1.0) (12.0 6 1.0) (11.8 6 1.7) (13.5 6 1.7) (11.5 6 1.6) (12.6 6 1.3) (13.3 6 0.9) MBH 10.4–14.2 6.2–10.2 11.4–16.6 10.4–15.2 12.7–18.6 8.8–14.7 7.7–13.1 10.8–16.7 10.2–16.2 11.5 (female) (11.8 6 1.0) (8.9 6 1.3) (14.2 6 1.8) (12.2 6 1.5) (16.2 6 2.1) (11.1 6 1.7) (10.9 6 2.4) (13.9 6 1.6) (13.0 6 2.8) TL (male) 70.4–74.8 52.0–88.6 62.2–94.2 63.0–85.7 86.8–115.3 76.1–126.9 89.1, 98.3 63.8–108.8 70.3–112.5 — (72.9 6 2.3) (69.2 6 14.7) (84.6 6 12.0) (76.4 6 9.2) (96.1 6 8.5) (99.0 6 14.4) (85.4 6 15.1) (99.5 6 11.3) TL (female) 45.3–82.3 69.0–86.3 65.5–99.9 55.4–76.1 70.7–115.2 70.5–109.2 79.5–105.4 61.2–106.0 87.7–107.9 — (65.6 6 13.0) (76.0 6 6.7) (82.6 6 11.6) (67.0 6 7.3) (94.2 6 17.6) (88.9 6 13.1) (91.6 6 9.2) (89.2 6 13.7) (97.9 6 9.0) TW (male) 8.5–10.0 7.2–10.3 10.2–11.9 7.4–12.2 11.8–15.9 11.0–15.1 8.8, 12.1 8.6–11.1 9.6–11.1 11.8 (9.4 6 0.5) (8.7 6 1.0) (10.9 6 0.6) (9.5 6 1.4) (13.5 6 1.2) (13.0 6 1.3) (10.0 6 0.8) (10.6 6 0.6) TW (female) 8.4–10.4 7.0–9.9 10.0–12.5 7.4–12.1 12.9–15.6 10.3–15.7 7.3–12.5 9.2–12.5 9.0–11.0 11.6 (9.4 6 0.5) (8.6 6 1.2) (11.0 6 0.7) (9.4 6 1.3) (14.2 6 0.9) (12.4 6 1.5) (10.0 6 1.7) (10.9 6 0.9) (9.9 6 0.9) TH (male) 7.6–8.1 5.4–8.3 8.0–9.5 6.3–10.0 9.7–11.4 8.7–12.1 8.0, 10.5 7.6–9.9 7.7–9.5 8.4 (7.8 6 0.2) (6.8 6 0.9) (8.5 6 0.6) (8.0 6 1.0) (10.7 6 0.6) (10.2 6 1.0) (8.5 6 0.7) (8.6 6 0.7) TH (female) 6.9–8.5 5.1–8.0 7.3–10.0 6.9–10.3 10.6–13.2 8.2–12.7 6.0–10.1 7.3–10.5 7.8–9.7 9.3

(7.6 6 0.5) (6.5 6 0.9) (8.5 6 0.7) (8.0 6 1.0) (11.7 6 0.9) (9.9 6 1.3) (8.3 6 1.4) (9.1 6 0.8) (8.7 6 0.8) 24 No. HL (male) 8.3–9.0 7.3–8.9 8.6–10.5 8.4–9.0 9.2–12.2 8.6–12.1 8.7, 10.1 9.3–11.5 9.8–11.5 9.6 (8.7 6 0.4) (8.3 6 0.7) (9.6 6 0.7) (8.7 6 0.2) (10.7 6 0.9) (10.3 6 1.0) (11.2 6 0.6) (10.7 6 0.7) 2010

TABLE 6.—Continued. 17 MONOGRAPHS HERPETOLOGICAL ]

boholensis boulengeri mindorensis taylori talinis kadwa tungaoi orientalis schadenbergi vindumi (5 male; (7 male; (6 male; (8 male; (11 male; (12 male; (2 male; (21 male; (14 male; (1 male; 14 female) 8 female) 12 female) 13 female) 10 female) 15 female) 6 female) 19 female) 10 female) 1 female) HL (female) 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 6 0.4) (8.3 6 0.5) (9.9 6 0.7) (8.7 6 0.8) (11.3 6 0.9) (9.8 6 0.8) (8.8 6 0.7) (10.9 6 1.0) (10.6 6 1.0) HW (male) 9.3–10.9 9.1–10.3 10.3–12.1 9.6–11.6 11.9–14.5 11.2–14.3 10.1, 11.8 9.9–12.0 10.7–13.2 10.8 (10.2 6 0.6) (9.5 6 0.5) (11.2 6 0.6) (10.4 6 0.8) (13.2 6 0.8) (12.4 6 1.0) (11.1 6 0.6) (11.8 6 0.9) HW (female) 9.3–10.5 7.5–9.9 10.7–12.4 9.3–12.0 12.8–15.8 10.4–13.3 9.1–12.6 10.6–13.7 9.5–12.3 10.6 (10.0 6 0.4) (9.1 6 0.8) (11.4 6 0.5) (10.5 6 0.9) (14.0 6 0.9) (11.6 6 1.0) (10.8 6 1.4) (11.7 6 0.7) (11.4 6 1.3) HH (male) 7.2–7.9 6.2–7.5 8.2–10.1 7.3–8.8 8.8–12.7 7.9–10.9 7.2, 10.4 6.9–9.6 7.6–10.1 7.9 (7.7 6 0.3) (6.9 6 0.4) (9.0 6 0.6) (7.9 6 0.5) (9.9 6 1.1) (9.3 6 0.9) (8.4 6 0.6) (8.8 6 0.8) HH (female) 7.2–8.0 5.7–7.2 8.1–9.9 7.3–9.7 9.3–12.5 7.3–10.0 6.2–10.4 7.9–10.8 7.9–9.2 7.8 (7.5 6 0.3) (6.6 6 0.5) (8.9 6 0.5) (8.1 6 0.7) (10.7 6 1.0) (8.4 6 0.9) (8.3 6 1.6) (9.2 6 0.9) (8.7 6 0.6) SnFa (male) 22.9–25.1 19.1–24.0 25.9–28.1 20.7–23.3 26.2–34.1 26.5–32.7 22.8, 26.2 23.9–31.1 25.8–29.9 28.5 (23.7 6 1.0) (22.4 6 1.7) (27.0 6 0.7) (22.1 6 1.0) (30.6 6 2.3) (28.9 6 1.7) (27.4 6 2.1) (27.9 6 1.7) SnFa 21.3–24.2 17.9–24.4 24.3–27.5 20.8–25.1 29.1–34.7 25.4–31.9 20.9–28.6 25.6–32.3 24.8–30.3 26.8 (female) (22.7 6 0.8) (22.7 6 2.1) (25.9 6 1.0) (22.2 6 1.1) (31.4 6 1.9) (28.2 6 1.9) (25.0 6 2.8) (28.8 6 2.0) (27.9 6 2.3) ED (male) 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 6 0.2) (2.0 6 0.2) (2.2 6 0.2) (1.8 6 0.2) (2.2 6 0.1) (1.9 6 0.2) (1.8 6 1.0) (2.1 6 0.2) ED (female) 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 6 0.1) (1.9 6 0.3) (2.2 6 0.2) (1.8 6 0.2) (2.2 6 0.1) (1.9 6 0.2) (1.7 6 0.2) (1.9 6 0.2) (2.2 6 0.2) END (male) 2.4–2.6 2.7–3.0 2.5–2.9 2.8–3.4 3.7–5.1 3.7–4.3 3.3, 3.8 3.5–4.5 4.0–4.2 3.9 (3.5 6 0.1) (2.8 6 0.1) (2.7 6 0.2) (3.1 6 0.2) (4.3 6 0.4) (3.9 6 0.2) (4.0 6 0.3) (4.1 6 0.1) END 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 (female) (3.4 6 0.2) (2.8 6 0.3) (3.6 6 0.2) (3.0 6 0.2) (4.5 6 0.4) (3.8 6 0.3) (3.2 6 0.4) (4.2 6 0.3) (4.0 6 0.4) SNL (male) 5.0–5.3 4.2–4.6 5.4–6.0 4.1–4.8 5.7–7.5 5.3–6.8 5.2, 5.4 5.1–6.5 5.3–6.1 5.7 (5.2 6 0.1) (4.4 6 0.1) (5.7 6 0.2) (4.4 6 0.3) (6.4 6 0.6) (6.0 6 0.4) (5.9 6 0.4) (5.8 6 0.3) SNL (female) 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 6 0.2) (4.2 6 0.4) (5.5 6 0.4) (4.6 6 0.3) (6.6 6 0.5) (5.6 6 0.8) (5.3 6 0.4) (6.2 6 0.4) (5.9 6 0.5) IND (male) 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 6 0.1) (2.9 6 0.1) (3.4 6 0.1) (2.9 6 0.2) (3.9 6 0.3) (3.7 6 0.3) (3.3 6 0.2) (3.4 6 0.2) IND (female) 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 6 0.1) (2.8 6 0.3) (3.4 6 0.2) (2.9 6 0.2) (3.9 6 0.2) (3.6 6 0.2) (3.2 6 0.4) (3.5 6 0.3) (3.3 6 0.4) FLL (male) 9.6–11.1 10.0–11.7 10.7–12.8 9.3–10.4 11.3–17.7 10.9–14.8 12.5, 12.8 10.4–15.6 11.8–13.4 13.2 (10.5 6 0.7) (10.5 6 0.6) (11.4 6 0.7) (9.9 6 0.4) (13.9 6 1.8) (13.1 6 1.2) (12.8 6 1.6) (12.6 6 0.6) FLL (female) 9.0–11.2 8.2–11.3 10.0–13.0 9.0–10.3 13.5–18.9 10.7–15.0 11.0–13.8 11.6–15.6 11.1–13.5 13.3 (9.9 6 0.7) (10.4 6 1.0) (11.4 6 0.9) (9.7 6 0.4) (14.6 6 0.9) (12.9 6 1.3) (12.0 6 1.0) (13.9 6 1.2) (12.2 6 1.1) HLL (male) 16.2–17.5 16.6–18.5 20.1–22.6 16.1–17.9 20.5–27.9 20.5–23.7 20.1, 21.0 18.6–24.3 19.4–21.9 22.7 (16.8 6 0.5) (17.4 6 0.6) (20.8 6 0.9) (16.9 6 0.7) (23.0 6 2.4) (22.2 6 1.1) (21.2 6 2.0) (20.9 6 0.9) HLL 15.4–18.7 14.3–18.7 18.8–23.1 15.6–18.7 20.5–26.1 17.9–24.1 17.0–22.4 18.7–25.3 18.5–21.9 22.7 (female) (17.3–1.2) (17.0 6 1.4) (20.6 6 1.3) (17.1 6 1.1) (24.4 6 1.7) (21.5 6 1.8) (19.9 6 2.1) (22.8 6 2.1) (20.0 6 1.6) 18 HERPETOLOGICAL MONOGRAPHS [No. 24

FIG. 6.—Illustration of head of adult female Brachymeles boholensis (KU 323972) and adult male neotype of Brachymeles boulengeri (PNM 9720; formerly KU 307756) in dorsal, lateral, and ventral views. Taxonomically diagnostic head 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 numerals indicate scales in the supraocular series, with Arabic numbers indicating scales in the supraciliary series. Illustrations by CDS. prefrontals not contacting on midline; (11) Comparisons.—Characters distinguishing enlarged chin shields in three pairs; (12) nuchal B. boholensis from all pentadactyl species of scales undifferentiated; (13) fifth and sixth Brachymeles are summarized in Tables 4 and supralabial below eye; (14) auricular opening 5. Brachymeles boholensis most closely re- present; (15) continuous, light, dorsolateral sembles B. boulengeri, B. mindorensis,andB. stripes present; and (16) continuous, dark taylori, but it differs from them by having six middorsal stripes present (Tables 4 and 5). lamellae on Finger-III and by the presence of 2010] HERPETOLOGICAL MONOGRAPHS 19 three pairs of enlarged chin shields (Tables 4 rounded in lateral profile, SNL 54.6–64.6% and 5). Brachymeles boholensis differs further (59.3 6 2.8) HL [57.4]; auricular opening from B. boulengeri by having seven suprala- present, moderate; eyes small, ED 1.8–2.2% bials, six supraciliaries, and the fifth and sixth (2.0 6 0.1) SVL [2.1], 17.6–23.9% (20.3 6 supralabial below the eye, and by the presence 1.6) HL [19.5], 45.1–61.8% (52.5 6 3.7) END of continuous, dark middorsal stripes (Ta- [52.9], pupil subcircular; body slightly de- bles 4 and 5); from B. mindorensis by having a pressed, MBW 98.1–136.2% (113.0 6 10.3) smaller body size, shorter hind limbs, nine or MBH [131.0]; scales smooth, glossy, imbri- 10 Toe-IV lamellae, seven infralabials, and by cate; longitudinal scale rows at midbody 26–28 the absence of contact between supranasals [28]; paravertebral scale rows 63–66 [64]; and the absence of contact between prefron- axilla–groin scale rows 42–46 [44]; limbs well tals (Tables 4 and 5); and from B. taylori by developed, pentadactyl, digits small; FinIII- having seven supralabials, the fifth and sixth lam 6 [6]; ToeIVlam 7–10 [7]; FLL 15.6– supralabial below the eye, and the presence of 20.5% (17.6 6 1.4) AGD [20.5], 10.2–12.9% continuous, light dorsolateral stripes (Tables 4 (11.4 6 0.8) SVL [12.4]; HLL 27.6–34.6% and 5). (30.0 6 2.3) AGD [32.9], 17.8–21.5% (19.4 6 From all nonpentadactyl species of Brachy- 1.2) SVL [20.0]; order of digits from shortest meles (B. apus, B. bonitae, B. cebuensis, B. to longest for hand: I 5 V , II 5 III 5 IV, for elerae, B. lukbani, B. minimus, B. muntingka- foot: V , I , II , III 5 IV; tail not as wide as may, B. pathfinderi, B. samarensis, B. tridac- body, sharply tapered toward the end, TW tylus, B. vermis,andB. wrighti), B. boholensis 61.6–78.4% (70.3 6 5.0) MBW [63.5], TL differs by having pentadactyl (vs. nonpenta- 52.7–90.0% (75.7 6 13.0) SVL [86.8]. dactyl) limbs, greater forelimb lengths Rostral projecting dorsoposteriorly to point (.9.0 mm vs. ,6.9 mm), greater hind limb in line with center of nasal; broader than high, lengths (.15.4 mm vs. ,12.9 mm), Toe-IV in contact with frontonasal; frontonasal wider lamellae nine or 10 (vs. eight or fewer), fifth than long; nostril ovoid, centered in a single and sixth supralabial below the eye (vs. fourth rectangular nasal; nasals well separated; supra- and fifth), and by presence of a postnasal scale nasals present, large, moderately separated by (vs. absence). In addition, B. boholensis differs frontonasal; postnasals present; prefrontals from all nonpentadactyl species except B. broadly separated by frontal; frontal nearly wrighti by having 26–28 midbody scale rows diamond shaped, in moderate contact with (vs. ,24); from all nonpentadactyl species frontonasal, first two anterior supraoculars, 43 except B. pathfinderi by having 63–66 para- wider than anterior supraocular; supraoculars vertebrals (vs. .84) and by the presence of five; frontoparietals moderate in size, in contact, auricular openings (vs. absence); and from B. each frontoparietal in contact with posterior apus, B. lukbani, B. minimus, and B. vermis three or four supraoculars; interparietal mod- by the presence of limbs (vs. absence). erate in size, quadrilaterally shaped, longer than Description (based on holotype and 38 wide, its length slightly greater than midline referred specimens, including 12 paratypes length of frontoparietals; parietal eyespot pres- at CAS).—Details of the head scalation of an ent in posterior half of scale; parietals separated adult female are shown in Fig. 6. Measure- by interparietal; nuchals undifferentiated; lor- ments of the holotype are provided below in eals two, anterior loreal largest, in contact with brackets. Body moderate relative to other prefrontal, postnasal, supranasal, second supra- Brachymeles; maximum SVL 93.6 mm for labial, posterior loreal and frontonasal; preocu- males, 94.0 mm for females [89.5, female] lar single, nearly two thirds as high as posterior (Tables 4 and 5); head weakly differentiated loreal; supraciliaries six, anterior supraciliary from neck, nearly as wide as body, HW 10.0– contacting prefrontal and separating posterior 12.4% (11.3 6 0.7) SVL [11.8], 104.2–131.9% loreal from first supraocular; subocular row (112.6 6 6.7) HL [110.7]; HL 35.2–43.6% complete; lower eyelid with one row of scales, (38.9 6 2.3) SnFa [39.3]; SnFa 24.1–28.0% lacking an enlarged oval window, largely (26.0 6 1.0) SVL [27.0]; snout moderately transparent; supralabials seven, fifth and sixth long, broadly rounded in dorsal profile, beneath center of eye; infralabials seven. 20 HERPETOLOGICAL MONOGRAPHS [No. 24

Mental wider than long, in contact with degree of contact between head scales. Six first infralabials; postmental single, enlarged, specimens were observed to have parietals slightly wider than mental, followed by three separated by the interparietal (KU 323944, pairs of enlarged chin shields, first pair in 323949, 323953, 323962, 323975–6), and 14 broad medial contact, second pair equal in specimens possessed parietals in moderate to width to first pair, broadly separated by single broad medial contact (KU 323948, 323952, medial scale, third pair separated by three 323954–6, 323960, 323963, 323966, 323970, medial scales. 323972, 323981, 323982, 323990, 324001) Scales on limbs smaller than body scales; behind the interparietal. In addition, seven scales on dorsal surfaces of digits large, specimens do not have the first pair of wrapping around lateral edges of digits; enlarged chin shields in medial contact lamellae undivided; palmar surfaces of hands (KU 323954–5, 323970, 323975, 323981–2, and plantar surfaces of feet covered by small, 324001), and 13 specimens with the first pair variably shaped scales, each bearing variably of enlarged chin shields in moderate to broad raised anterior edges; scales on dorsal surface medial contact (KU 323944, 323948–9, 323952– of hands and feet smaller than limb scales, 3, 323956, 323960, 323962–3, 323966, 323972, lacking raised edges. 323976, 323990). Coloration in preservative.—Ground color Scale counts were observed to vary among of body medium brown, lateral surface the measured series. Specimens were ob- lacking dark pigment, dorsal surface bearing served to have midbody scale row counts of continuous, longitudinal rows of olive green 26 (CAS-SU 18709, 18717, 24502, 24523–5, to brown pigment, spanning six full and two 24541, 24543, 25443–4), 27 (CAS-SU 24503– half rows of scales at midbody, narrowing to 4, 24522, 24867), and 28 (CAS-SU 24518, four full and two half rows of scales posterior 24520–21, 24528, 25447); axilla–groin scale to parietals, extending from posterior edge of row counts of 42 (CAS-SU 18717, 24502, parietals to tail tip; dark pigmentation cover- 24520, 24524), 43 (CAS-SU 24523, 24541, ing one half to three fourths of dorsal scales; 24543), 44 (CAS-SU 24504, 24518, 24525, lateral surface of body with six continuous 24528, 24867, 25447), 45 (CAS-SU 24503, to discontinuous longitudinal rows of olive 24521, 25443), and 46 (CAS-SU 18709, 24522, green to brown stripes, extending from post- 25444); and paravertebral scale row counts of erior edge of eye to base of tail; dorsolateral 63 (CAS-SU 18717, 24502, 24520, 24524, stripes present, lacking dark pigmentation, 24541, 24543), 64 (CAS-SU 24504, 24523, spanning two half rows of scales, extending 24525, 24528, 24867, 25447), 65 (CAS-SU from posterior edge of supraoculars to base of 24518), and 66 (CAS-SU 18709, 24503, tail. Ventral scales with or without dark spots. 24521–2, 25443–4). Head scales uniform medium brown, darker We also observed Toe-IV lamellae counts to brown than ventral scales; rostral, nasal, vary among the measured series. With the postnasal, supranasal, first supralabial, men- exception of a single specimen with seven tal, and first infralabial dark gray; pineal Two-IV lamellae (CAS-SU 24528, holotype), eyespot poorly defined, small, and light to all specimens examined had either nine (CAS- dark brown. Limbs mottled light and medium SU 18709, 24504, 24523–4, 24867, 25444) or brown dorsally, yellowish brown ventrally; 10 (CAS-SU 18717, 24502–3, 24518, 24520–2, dorsal and ventral surfaces of digits dark 24525, 24541, 24543, 25443, 25447). We brown. observed the holotype to have malformed Coloration in life.—Ground color of body digits that we believe resulted in fewer Toe-IV medium brown to tan (Fig. 11C); longitudinal lamellae than are observed for all other rows of darker brown pigmentation; dorsolat- individuals of this species. eral stripes light brown to tan; limbs bearing Distribution.—Brachymeles boholensis is dark brown mottling dorsally. known only from Bohol Island (Fig. 3). Variation.—Morphometric variation is Ecology and natural history.—Brachymeles summarized in Table 6. We observed varia- boholensis occurs in agricultural habitats, as tion among the 20 specimens examined for the well as in disturbed and secondary growth 2010] HERPETOLOGICAL MONOGRAPHS 21 forests. No original, low-elevation forest re- Brachymeles mindorensis Brown and Rabor mains on Bohol Island, but we assume the 1967 species once also occurred in primary forest at Figs. 3, 7 low elevations. Individuals have been ob- Brachymeles gracilis mindorensis, Brown and served under piles of rotting coconut husks, Rabor, 1967. Type locality: Bank of Tarogin in the humus material within rotting logs, and River, 30 km southeast of Calapan, Mind- in loose soil and leaf litter surrounding the oro Oriental Province, Mindoro Island, Phi- root networks of trees. Interestingly, this lippines, 0–33-m elevation, 13u 119 25.440 N, species seems to be a ubiquitous habitat 121u 89 52.80 E (holotype: CAS-SU 24487). generalist on Bohol, whereas its congener B. Brachymeles gracilis (part), Brown and Alcala, orientalis seems to be restricted to fallen and 1970. rotting logs in secondary growth forest on the Brachymeles boulengeri mindorensis (part), same island (CDS, personal observation). As is Brown and Alcala, 1980. typical for species in the genus, individuals Diagnosis.—Brachymeles mindorensis can immediately attempt to evade capture by be distinguished from congeners by the moving to quickly burrow into loose soil or following combination of characters: (1) body humus. size moderate (SVL 90.0–104.2 mm); (2) Although only two species of Brachymeles pentadactyl; (3) Finger-III lamellae five or have been confirmed to occur on Bohol Island six; (4) Toe-IV lamellae eight or nine; (5) (B. boholensis and B. orientalis), populations moderate limb length; (6) supralabials seven; of B. samarensis are known to occur on (7) infralabials six; (8) pineal eye spot present; Lapinig Grande and Lapinig Chico islands (9) enlarged chin shields in two pairs; (10) just off the northeast coast of Bohol Island nuchal scales undifferentiated; (11) fifth and (Brown and Alcala, 1980). No individuals of sixth supralabial below eye; (12) auricular this species lacking fully formed digits occur opening present; (13) continuous, light, dorso- on Bohol; however, given proximity of these lateral stripes present; and (14) continuous, small islands to mainland Bohol, it seems dark middorsal stripes present (Tables 4 and 5). likely that this neighboring, limb-reduced Comparisons.—Characters distinguishing species of Brachymeles eventually may be B. mindorensis from all pentadactyl species discovered on Bohol. of Brachymeles are summarized in Tables 4 Sympatric lizard species observed on Bohol and 5. Brachymeles mindorensis most closely Island include the following: (Agamidae) resembles B. boholensis, B. boulengeri, and B. Bronchocela cristatella, Draco bimaculatus, taylori, but it differs from these three taxa by D. ornatus, D. reticulates, Gonocephalus having a larger body size, longer hind limbs, semperi, Hydrosaurus pustulatus;(Gekkoni- eight or nine Toe-IV lamellae, and six infra- dae) Cyrtodactylus annulatus, Gehyra muti- labials (Tables 4 and 5). Brachymeles mind- lata, Gekko gecko, Hemidactylus frenatus, H. orensis further differs from B. boholensis by platyurus, Hemiphyllodactylus typus, Lepi- having five or six Finger-III lamellae and by dodactylus aureolineatus, L. planicaudus, the absence of a third pair of enlarged chin Pseudogekko compressicorpus, P. brevipes; shields (Tables 4 and 5); from B. boulengeri (Scincidae) Brachymeles schadenbergi orien- by having seven supralabials, six supraciliaries, talis, Emoia atrocostata, Eutropis multicar- and the fifth and sixth supralabial below the inata, E. multifasciata, Lamprolepis smarag- eye, and by the presence of continuous, dark dina, Lipinia pulchella, L. quadrivittata, middorsal stripes (Tables 4 and 5); and from Sphenomorphus acutus, S. cumingi, S. fas- B. taylori by having seven supralabials and the ciatus, S. jagori, S. minanensis, S. steerei, S. fifth and sixth supralabial below the eye, and variegatus; and (Varanidae) Varanus cumingi. by the presence of continuous, light dorsolat- Also, Brachymeles samarensis (Scincidae) is eral stripes (Tables 4 and 5). known to occur on Lipinig Grande and From all nonpentadactyl species of Brachy- Lipinig Chico islands just off the northeast meles (B. apus, B. bonitae, B. cebuensis, B. coast of Bohol Island (Brown and Alcala, elerae, B. lukbani, B. minimus, B. muntingka- 1980). may, B. pathfinderi, B. samarensis, B. tridac- 22 HERPETOLOGICAL MONOGRAPHS [No. 24

FIG. 7.—Illustration of head of adult male Brachymeles mindorensis (KU 304343) and adult female Brachymeles taylori (KU 324049) in dorsal, lateral, and ventral views. Labels for taxonomically diagnostic head scales follow those shown in Figure 6. Illustrations by CDS. tylus, B. vermis, and B. wrighti), B. mind- scale row count 26–28 (vs. ,24); from all orensis differs by having a pentadactyl body nonpentadactyl species except B. pathfinderi form (vs. nonpentadactyl), longer forelimb by having a paravertebral count 63–65 (vs. lengths (.10.0 mm vs. ,6.9 mm), greater .84) and by the presence of auricular hind limb lengths (.18.8 mm vs. ,12.9 mm), openings (vs. absence); and from B. apus, B. and the fifth supralabial below the eye (vs. lukbani, B. minimus, and B. vermis by the fourth), and by the presence of a postnasal presence of limbs (vs. absence). scale (vs. absence). In addition, Brachymeles Description (based on holotype and 33 mindorensis differs from all nonpentadactyl referred paratypes at CAS).—Details of the species except B. wrighti by having a midbody head scalation of an adult male are shown in 2010] HERPETOLOGICAL MONOGRAPHS 23

Fig. 7. Measurements of the holotype are eyespot present in posterior one half of scale; included below in brackets. Body moderate parietals moderately separated behind inter- relative to other Brachymeles, elongate with parietal; nuchals undifferentiated from adja- respect to other lizards; maximum SVL cent dorsal scales; loreals two, anterior loreal 104.2 mm for males, 106.8 mm for females in contact with prefrontal, postnasal, suprana- [106.8, female] (Tables 4 and 5); head weakly sal, second supralabial, posterior loreal and differentiated from neck, nearly as wide as frontonasal; preocular, single, nearly one third body, HW 10.5–12.9% (11.4 6 0.7) SVL as high as posterior loreal; presubocular [10.9], 104.2–130.1% (116.1 6 6.0) HL single; supraciliaries six, the anteriormost [114.6]; HL 31.6–42.9% (37.3 6 3.1) SnFa contacting prefrontal and separating posterior [39.2]; SnFa 24.1–29.9% (26.5 6 1.3) SVL loreal from first supraocular; subocular row [24.1]; snout moderately long, rounded in complete; lower eyelid with one row of scales, dorsal and lateral profile, SNL 50.6–68.2% lacking an enlarged oval window, largely (57.3 6 4.9) HL [52.2]; auricular opening transparent; supralabials seven, fifth and sixth present, small; eyes moderate, ED 2.0–2.6% below eye; infralabials six. (2.2 6 0.2) SVL [2.0], 19.4–27.4% (22.7 6 Mental wider than long, in contact with first 2.0) HL [21.1], 51.8–72.2% (60.7 6 5.1) END infralabials; single enlarged postmental, equal [58.5], pupil nearly round; body slightly in width to mental followed by two pairs of depressed, MBW 105.4–156.0% (120.2 6 enlarged chin shields; first pair in moderate 14.0) MBH [125.0]; scales smooth, glossy, contact, second pair wider than first, moder- imbricate; longitudinal scale rows at midbody ately separated by single medial scale. 26–28 [26]; paravertebral scale rows 63–65 Scales on limbs smaller than body scales; [65]; axilla–groin scale rows 42–45 [45]; limbs scales on dorsal surfaces of digits large, well developed, pentadactyl, digits small; wrapping around lateral edges of digits; FinIIIlam 5–6 [5]; ToeIVlam 8–9 [9]; FLL lamellae undivided; palmar surfaces of hands 14.5–21.3% (17.8 6 1.9) AGD [15.5], 10.1– and plantar surfaces of feet covered by small, 13.1% (11.5 6 0.9) SVL [10.9]; HLL 16.7– variably shaped scales, each with variably 37.9% (31.5 6 5.0) AGD [25.1], 10.5–24.0% raised anterior edges; scales on dorsal surface (20.3 6 2.9) SVL [17.6]; order of digits from of hands and feet smaller than limb scales, shortest to longest for hand: I 5 V , II 5 IV lacking raised edges. , III, for foot: I , V , II , III 5 IV; tail not Coloration in preservative.—Ground color as wide as body, sharply tapered toward the of body medium brown; dorsal surface of body end, TW 54.4–80.1% (69.1 6 8.1) MBW with eight longitudinal rows of dark brown [54.4], TL 60.4–99.3% (84.6 6 11.5) SVL stripes, six continuous medial rows, two [93.5]. discontinuous lateral rows, spanning eight full Rostral projecting dorsoposteriorly to point rows of scales at midbody, narrowing to six full in line with anterior edge of nasal, broader rows of scales posterior to parietals, extending than high, in narrow contact with frontonasal; from posterior edge of parietals to tail tip; frontonasal wider than long; nostril ovoid, pigmentation covering middle one third of centered in a single rectangular nasal; nasals dorsal scales; dorsolateral stripes present, well separated; supranasals present, large, lacking dark pigmentation, spanning one narrowly separated by frontonasal; postnasals whole and two half rows of scales, extending present; prefrontals narrowly separated by from posterior edge of supraoculars to base of frontal; frontal octagonal, narrowly contacting tail. Lateral surface of body light brown frontonasal and first two supraoculars anteri- ground color with three or four rows of nearly orly, 43 wider than anteriormost supraocular; continuous spots of dark brown pigmentation, supraoculars five; frontoparietals moderate, in extending from auricular opening to tail tip. broad medial contact, each frontoparietal in Tail striped with longitudinal rows of dark contact with supraoculars 2–4; interparietal pigmentation. Ventral scales lacking dark large, quadrilaterally shaped, slightly longer spots. Head scales uniform medium brown, than wide, its length slightly greater than darker brown than ventral scales; posterior- midline length of frontoparietal; parietal most supraocular lacking pigmentation; dark 24 HERPETOLOGICAL MONOGRAPHS [No. 24 pigmentation surrounding auricular opening, 24487, 24549–54, 24566, 24570, 24574, connected to dark pigmentation on head 24577–9). scales; rostral, nasal, postnasal, supranasal, We also observed lamellae counts to vary and first supralabial dark gray; pineal eyespot among the series. With the exception of two poorly defined, small and light cream. Limbs specimens with six Finger-III lamellae (CAS- mottled dark brown dorsally, yellowish brown SU 24561, 24574), all specimens examined ventrally; dorsal and ventral surface of digits had five. Specimens were observed to have dark brown. Toe-IV lamellae counts of eight (CAS-SU Coloration in life.—Ground color of body 24549–54, 24561–2, 24564, 24566, 24570, dark to medium brown; continuous, dark 24573, 24577–8) or nine (CAS-SU 24487, middorsal stripes dark brown to black; dorso- 24568, 24574, 24579). lateral stripes light brown to tan, gradually Distribution.—Brachymeles mindorensis is become predominately tan on tail; limbs dark known only from Mindoro Island (Fig. 3). brown dorsally. Ecology and natural history.—Brachymeles Variation.—Morphometric variation of the mindorensis occurs in disturbed and second- series is summarized in Table 6. We observed ary growth forest. Individuals have been variation among the 16 specimens examined observed under piles of rotting coconut husks, for the degree of contact between head scales. in the humus material within rotting logs, and Five specimens have parietals separated by in loose soil and leaf litter surrounding the the interparietal (KU 304352, 304354, 304488, root networks of trees. The species has been 308447–8), and 11 specimens have parietals observed to be quite common in certain in moderate to broad medial contact (KU habitats on Mindoro Island (CDS and RMB, 304351, 304353, 304355, 304412–3, 307739– personal observations). Brachymeles mindor- 42, 308404, 308534) behind the interparietal; ensis occurs sympatrically with B. bonitae four specimens have supranasals narrowly (Brown, 1956; Brown and Alcala, 1980; Brown separated by the frontonasal (KU 307740–1, and Rabor, 1967), and it is the largest species 308404, 308448), six specimens have supra- of the B. boulengeri complex. When individ- nasals in medial point contact (KU 304351–4, uals were disturbed, they attempted to quickly 307742, 308447), and six specimens have burrow back into loose soil or humus. supranasals in moderate medial contact (KU Sympatric lizard species observed on Mind- 304355, 304412–3, 304488, 307739, 308534); oro Island include the following: (Agamidae) 13 specimens have prefrontals moderately Bronchocela marmoratus, Draco quadrasi, separated by the frontal (KU 304351–4, Gonocephalus interruptus, Hydrosaurus pustu- 304412–3, 304488, 307739–41, 308447–8, latus; (Gekkonidae) Cyrtodactylus philippini- 308534), one specimen has prefrontals nar- cus, Gehyra mutilata, Gekko gecko, G. mind- rowly separated by the frontal ( KU 308404), orensis, Hemidactylus frenatus, H. garnoti, H. and two specimens have prefrontals in medial platyurus, Hemiphyllodactylus typus, Lepido- point contact (KU 304355, 307742). dactylus naujanensis;(Scincidae)Brachymeles Scale counts were observed to vary among bonitae, Dasia olivaceum, Emoia atrocostata, the measured series. Specimens were ob- Eutropis multicarinata, E. multifasciata, Lam- served to have midbody scale row counts of prolepis smaragdina, Lipinia auriculatum, 26 (CAS-SU 24487, 24551, 24561, 24566, Sphenomorphus cumingi, S. jagori, S. steerei; 24574, 24577, 24579), 27 (CAS-SU 24549, and (Varanidae) Varanus marmoratus. 24573), and 28 (CAS-SU 24550, 24552–4, 24562, 24564, 24568, 24570, 24578); axilla– Brachymeles taylori Brown 1956 groin scale row counts of 42 (CAS-SU 24564), Figs. 3, 7, 11B 43 (CAS-SU 24573), 44 (CAS-SU 24553–4, Brachymeles gracilis taylori, Brown, 1956. 24561–2, 24568, 24570, 24578), and 45 Type locality: Sitio Lunga, 13 km West (CAS-SU 24487, 24549–52, 24566, 24574, Dumaguete, 3 km West Valencia, on low 24577, 24579); and paravertebral scale row ridge on the north side of the Maite River, counts of 63 (CAS-SU 24564, 24573), 64 Negros Oriental Province, Negros Island, (CAS-SU 24561–2, 24568), and 65 (CAS-SU Philippines, 600-m elevation, 9u 179 32.960 2010] HERPETOLOGICAL MONOGRAPHS 25

N, 123u 149 2.40 E (holotype: CAS-SU ,6.9 mm), longer hind limbs (.15.6 mm vs. 24487). ,12.9 mm), and Toe-IV lamellae nine or 10 Brachymeles gracilis (part), Brown and Alcala, (vs. eight or fewer), and by the presence of a 1970. postnasal scale (vs. absence). In addition, Brachymeles boulengeri taylori (part), Brown Brachymeles taylori differs from all nonpen- and Alcala, 1980. tadactyl species except B. wrighti by having a Diagnosis.—Brachymeles taylori can be midbody scale row count 26–28 (vs. ,24); distinguished from congeners by the following from all nonpentadactyl species except B. combination of characters: (1) body size pathfinderi by having a paravertebral count moderate (SVL 65.8–99.2 mm); (2) pentadac- 62–69 (vs. .84) and by the presence of tyl; (3) Finger-III lamellae five or six; (4) Toe- auricular openings (vs. absence); and from B. IV lamellae nine or 10; (5) moderate limb apus, B. lukbani, B. minimus,andB. vermis length; (6) supralabials six; (7) infralabials by the presence of limbs (vs. absence). seven; (8) pineal eye spot present; (9) Description (based on holotype and 33 supranasals not contacting on midline; (10) referred specimens, including five paratypes prefrontals not contacting on midline; (11) at CAS).—Details of the head scalation of an enlarged chin shields in two pairs; (12) nuchal adult female are shown in Fig. 7. Measure- scales undifferentiated; (13) fourth and fifth ments of the holotype are provided below in supralabial below eye; (14) auricular opening brackets. Body moderate relative to other present; (15) dorsolateral stripes absent; and Brachymeles, elongate with respect to other (16) continuous, dark middorsal stripes pres- lizards; maximum SVL 99.2 mm for males, ent (Tables 4 and 5). 93.2 mm for females [65.8, female] (Tables 4 Comparisons.—Characters distinguishing and 5); head weakly differentiated from neck, Brachymeles taylori from all pentadactyl nearly as wide as body, HW 11.3–15.3% (12.3 species of Brachymeles are summarized in 6 1.0) SVL [15.3], 104.4–139.5% (120.5 6 Tables 4 and 5. Brachymeles taylori most 10.2) HL [132.8]; HL 34.8–44.6% (39.2 6 2.5) closely resembles B. boholensis, B. boulengeri, SnFa [36.4]; SnFa 23.4–31.7% (26.2 6 1.7) and B. mindorensis, but it differs from these SVL [31.7]; snout moderately long, broadly three taxa by the absence of continuous, light rounded in dorsal profile, depressed in lateral dorsolateral stripes (Tables 4 and 5). Brachy- profile, SNL 44.1–62.5% (52.2 6 4.6) HL meles taylori can further be distinguished [62.5]; auricular opening present, moderate; from B. boholensis by having five or six eyes moderate, ED 1.6–2.8% (2.1 6 0.2) SVL Finger-III lamellae, six supralabials, and the [2.8], 15.9–24.4% (20.8 6 2.4) HL [24.4], 48.8– fourth and fifth supralabial below the eye, and 69.5% (59.6 6 6.1) END [62.7], pupil nearly by the absence of a third pair of enlarged chin round; body slightly depressed, MBW 89.1– shields (Tables 4 and 5); from B. boulengeri 148.4% (115.1 6 14.7) MBH [112.9]; scales by having six supralabials and six supracili- smooth, glossy, imbricate; longitudinal scale aries, and the presence of continuous, dark rows at midbody 26–28 [27]; paravertebral middorsal stripes (Tables 4 and 5); and from scale rows 62–69 [64]; axilla–groin scale rows B. mindorensis by having a smaller body size, 42–47 [43]; limbs well developed, pentadactyl, shorter hind limbs, nine or 10 Toe-IV digits small; FinIIIlam 5–6 [6]; ToeIVlam 8–10 lamellae, six supralabials, seven infralabials, [9]; FLL 15.7–20.4% (17.8 6 1.2) AGD [19.9], the fourth and fifth supralabial below the eye, 10.4–14.1% (11.5 6 0.9) SVL [14.1]; HLL and no contact between supranasals and 26.7–36.8% (31.1 6 2.5) AGD [35.6], 17.8– prefrontals (Tables 4 and 5). 25.2% (20.1 6 1.7) SVL [25.2]; order of digits From all nonpentadactyl species of Brachy- from shortest to longest for hand: I 5 V , IV , meles (B. apus, B. bonitae, B. cebuensis, B. II 5 III, for foot: I , V , II , III 5 IV; tail not elerae, B. lukbani, B. minimus, B. muntingka- as wide as body, sharply tapered posteriorly, may, B. pathfinderi, B. samarensis, B. tridac- TW 54.0–80.3% (68.9 6 6.7) MBW [75.0], TL tylus, B. vermis,andB. wrighti), B. taylori 69.2–103.1% (83.3 6 10.4) SVL [98.2]. differs by having pentadactyl limbs (vs. non- Rostral projecting dorsoposteriorly to point pentadactyl), longer forelimbs (.9.0 mm vs. in line with anterior edge of nasal, broader 26 HERPETOLOGICAL MONOGRAPHS [No. 24 than high, in moderate contact with frontona- brown ventral surface of body, ventral surface sal; frontonasal wider than long; nostril ovoid, without dark pigmentation; dark dorsal pig- in center of single rectangular nasal; nasals mentation in nearly continuous block across well separated; supranasals present, large, dorsal surface or at times forming 10 contin- narrowly separated by frontonasal; postnasals uous longitudinal rows of dark brown pig- present; prefrontals moderately separated by ment, spanning six full rows of scales across frontal; frontal nearly octagonal, its anterior dorsal surface, extending from posterior edge margin in moderate contact with frontonasal, of parietals to tail tip, additional three to four in contact with first two anterior supraoculars, scale rows on each lateral surface with dark 4.53 wider than anteriormost supraocular; coloration, covering one half to two thirds of supraoculars five; frontoparietals moderate, in dorsal and lateral scales; dorsolateral stripes broad medial contact, each frontoparietal in absent. Tail coloration matches body colora- contact with supraoculars 2–4; interparietal tion. Head scales uniform dark brown, darker moderate, quadrilaterally shaped, longer than brown than ventral scales; rostral, nasal, wide, its length nearly equal to midline length postnasal, supranasal, and first supralabial of frontoparietal; parietal eyespot present dark gray; pineal eyespot poorly defined, small in posterior one half of scale; parietals in and light cream. Limbs mottled medium moderate contact behind interparietal or brown dorsally, yellowish brown ventrally; narrowly separated; nuchals undifferentiated; dorsal and ventral surface of digits dark loreals two, decreasing in size from anterior to brown. posterior, subequal, anterior loreal in contact Coloration in life.—Ground color of body with prefrontal, postnasal, supranasal, second light to medium brown (Fig. 11B); medium to supralabial, posterior loreal and frontonasal; dark brown dorsal pigmentation gradually preocular single, nearly two thirds as high as fading into medium brown lateral pigmenta- posterior loreal; single presubocular; supraci- tion; limbs mottled medium to dark brown liaries six, the anteriormost contacting pre- dorsally. frontal and separating posterior loreal from Variation.—Morphometric variation of the first supraocular; subocular row complete; series is summarized in Table 6. We observed lower eyelid with one row of scales, lacking variation among the 14 specimens examined an enlarged oval window, largely transparent; for the degree of contact between head scales. supralabials six, fourth and fifth below the eye; Six specimens have parietals moderately infralabials seven. separated by the interparietal (KU 324048, Mental wider than long, in contact with first 324050, 324052–5), one specimen has parie- infralabial; postmental single, enlarged, slight- tals narrowly separated by the interparietal ly wider than mental, followed by two pairs of (KU 324051), and seven specimens have enlarged chin shields, scales of first pair parietals in moderate medial contact (KU separated or in moderate contact, second pair 306651, 324044–7, 324049, 324056) behind slightly wider than first, broadly separated by the interparietal; eight specimens do not have a single medial scale. the first pair of enlarged chin shields in medial Scales on limbs smaller than body scales; contact (KU 324045, 324048–50, 324053–6), scales on dorsal surfaces of digits large, one specimen has the first pair of enlarged wrapping around lateral edges of digits; chin shields in point medial contact (KU lamellae undivided; palmar surfaces of hands 324046), and five specimens have the first pair and plantar surfaces of feet covered by small, of enlarged chin shields in moderate medial irregular scales, each with irregular raised contact (KU 306651, 324044, 324047, anterior edges; scales on dorsal surface of 324051–2). hands and feet smaller than limb scales, Scale counts were observed to vary among lacking raised edges. the measured series. Specimens were ob- Coloration in preservative.—Ground color served to have midbody scale row counts of of body medium olive brown; dorsal pigmen- 26 (CAS-SU 18641, 18656–7, 18748, 21873, tation nearly all dark brown, gradually fading 21877, 21880, 21884, 22355, CAS 154971, into olive brown lateral surface and yellowish 154673, 154680–2, 154686), 27 (CAS-SU 2010] HERPETOLOGICAL MONOGRAPHS 27

18615, 22356, CAS 154679), and 28 (CAS-SU rotting logs, and in loose soil and leaf litter 18649, 21883, CAS 154678); axilla–groin scale surrounding the root networks of trees. The row counts of 42 (CAS 154680, 154686), 43 species is quite common throughout its range (CAS-SU 18615, 18656, CAS 154682), 44 (CDS, personal observation) and occurs sym- (CAS-SU 18641, 18748, 21873, 21877, 21880, patrically with three other species of Brachy- 22356, CAS 154678, 154679, 154681), 45 meles (B. cebuensis, B. talinis,andB. tridac- (CAS-SU 21884, 22355, CAS 154671), 46 tylus; Brown, 1956; Brown and Alcala, 1980; (CAS-SU 18649, 21883, CAS 154673), and 47 Brown and Rabor, 1967). Similar to B. (CAS-SU 18657); and paravertebral scale row boulengeri, B. taylori seems to have a wider counts of 62 (CAS 154680, 154686), 64 (CAS- geographic distribution that spans multiple SU 18615, 18656, CAS 154682), 66 (CAS-SU Philippine islands. This is in contrast to the 18641, 18748, 21873, 21877, 21880, 21884, island endemic species B. boholensis and B. 22356, CAS 154678–9, 154681), 67 (CAS-SU mindorensis that are known from just Bohol 18649, CAS 154671, 154673), 68 (CAS-SU and Mindoro islands, respectively. As with all 21883, 22355), and 69 (CAS-SU 18657). members of the genus, when disturbed, We also observed lamellae counts to vary individuals move rapidly attempting to burrow among the measured series. Specimens were into the loose soil or humus. observed to have Finger-III lamellae counts of Sympatric lizard species observed within five (CAS-SU 18641, 18649, 18656, 18748, the range of B. taylori include the following: 21883–4, 22356, CAS 154671, 154673, (Agamidae) Bronchocela marmoratus, Draco 154678–80) or six (CAS-SU 18615, 18657, spilopterus, Gonocephalus sophiae, Hydro- 21873, 21877, 21880, 22355, CAS 154681–2, saurus pustulatus; (Dibamidae) Dibamus ar- 154686); Toe-IV lamellae counts of eight genteus; (Gekkonidae) Cyrtodactylus philip- (CAS 154678), nine (CAS-SU 18615, 18649, pinicus, Gehyra mutilata, Gekko gecko, G. 18656, 21880, 22356, CAS 154671, 154673, mindorensis, G. enrstkelleri, Luperosaurus 154682, 154686), and 10 (CAS-SU 18641, corfieldi, Pseudogekko brevipes, Hemidactylus 18657, 18748, 21873, 21877, 21883–4, 22355, frenatus, H. platyurus, Hemiphyllodactylus CAS 15479–81). typus, Lepidodactylus christiani, L. herrei, There is some color variation in the L. lugubris; (Scincidae) Brachymeles cebuen- examined series, with the degree and defini- sis, B. talinis, B. cebuensis, Emoia atrocostata, tion of continuous, dark middorsal stripes. All Eutropis multicarinata, E. multifasciata, Lam- specimens have continuous, dark lines run- prolepis smaragdina, Lipinia auriculata, L. ning down the middorsal surface of the body; pulchella, L. quadrivittata, L. rabori, Spheno- however, the continuous lines in some spec- morphus arborens, S. coxi, S. jagori, S. steerei, imens are present without a dark, middorsal Tropidophorus grayi; and (Varanidae) Vara- background coloration (KU 324045–6, 324049– nus nuchalis. 51, 324053–4). The dark lines in other specimens overlay a dark, middorsal region covered by a Brachymeles orientalis Brown and Rabor long streak of dark background pigmentation 1967 (KU 324044, 324047–8, 324052, 324055–6). Figs. 4, 8, 11F,G Distribution.—Brachymeles taylori is Brachymeles schadenbergi orientalis, Brown known from Negros, Cebu, Inampulugan, and Rabor, 1967. Type locality: Bario Pan de Azucar, Danjugan, Ponson, and Poro Dusita, 11 km southeast of Sierra Bullones, islands (Fig. 3). Bohol Province, Bohol Island, Philippines, Ecology and natural history.—Brachymeles 533-m elevation, 9u 469 57.50 N, 124u 189 taylori occurs in agricultural areas as well as 10.80 E (holotype: CAS-SU 24436). disturbed and secondary growth forest. Little Brachymeles schadenbergi (part), Brown and or no original, low-elevation forest remains in Alcala, 1970. the Visayas, but we assume the species once Brachymeles schadenbergi (part), Brown and also occurred in primary forest. Individuals Alcala, 1980. have been observed under piles of rotting Diagnosis.—Brachymeles orientalis can be coconut husks, in the humus material within distinguished from congeners by the following 28 HERPETOLOGICAL MONOGRAPHS [No. 24

FIG. 8.—Illustration of head of adult male Brachymeles orientalis (KU 311241) and adult male Brachymeles schadenbergi (KU 314992) in dorsal, lateral, and ventral views. Labels for taxonomically diagnostic head scales follow those shown in Fig. 6. Illustrations by CDS. combination of characters: (1) body large undifferentiated; (13) fourth and fifth supra- (SVL 97.6–112.3 mm); (2) pentadactyl; (3) labial below eye; (14) auricular opening Finger-III lamellae six or seven; (4) Toe-IV present; (15) dorsolateral stripes absent; and lamellae 8–10; (5) limbs relatively long; (6) (16) middorsal stripes absent (Tables 4 and 5). supralabials six or seven; (7) infralabials six or Comparisons.—Characters distinguishing seven; (8) pineal eye spot present; (9) Brachymeles orientalis from all pentadactyl supranasal contact absent; (10) prefrontals species of Brachymeles are summarized in not contacting on midline; (11) enlarged chin Tables 4 and 5. Brachymeles orientalis most shields in two pairs; (12) nuchal scales closely resembles B. makusog and B. scha- 2010] HERPETOLOGICAL MONOGRAPHS 29 denbergi, but it differs from both taxa by HL [62.5]; auricular opening present, moder- having six or seven Finger-III lamellae, 8–10 ate; eyes moderate, ED 1.5–2.3% (1.8 6 0.2) Toe-IV lamellae, and the fourth and fifth SVL [1.8], 14.8–19.6% (17.3 6 1.3) HL [18.2], supralabial below the eye (Tables 4 and 5), 38.2–56.7% (44.8 6 3.7) END [45.9], pupil and by the presence (vs. absence) of reddish subcircular; body slightly depressed, MBW orange to salmon-colored scales on the lateral 93.1–138.2% (114.5 6 11.4) MBH [124.9]; surfaces of the body. Brachymeles orientalis scales smooth, glossy, imbricate; longitudinal can further be distinguished from B. makusog scale rows at midbody 26–30 [28]; paraverte- by having a greater maximum axilla–groin bral scale rows 69–73 [72]; axilla–groin scale scale row count and a greater maximum rows 46–49 [48]; limbs well developed, paravertebral scale row count (Table 5) and pentadactyl, digits small; FinIIIlam 6–7 [6]; from B. schadenbergi by having no contact ToeIVlam 8–10 [8]; FLL 17.0–24.8% (20.7 6 between supranasals, and by the absence of 2.0) AGD [18.8], 10.7–15.4% (13.1 6 1.1) continuous, dark middorsal stripes and dark SVL [12.3]; HLL 27.6–42.7% (34.2 6 3.4) lateral stripes (Table 5). AGD [32.2], 18.4–24.2% (21.6 6 1.7) SVL From all nonpentadactyl species of Brachy- [21.2]; order of digits from shortest to longest meles (B. apus, B. bonitae, B. cebuensis, B. for hand: I , V , II 5 IV , III, for foot: I , elerae, B. lukbani, B. minimus, B. muntingka- V , II , III 5 IV; tail nearly as wide as body, may, B. pathfinderi, B. samarensis, B. tridac- sharply tapered toward the end, TW 55.6– tylus, B. vermis,andB. wrighti), B. orientalis 86.9% (69.7 6 6.4) MBW [70.9], TL 62.2– differs by having pentadactyl limbs (vs. 106.0% (85.2 6 11.6) SVL [106.0]. nonpentadactyl), longer forelimb lengths Rostral projecting dorsoposteriorly to point (.10.4 mm vs. ,6.9 mm), greater hind limb in line with anterior edge of nasal, broader lengths (.18.6 mm vs. ,12.9 mm), and by the than high, in narrow contact with frontonasal; presence (vs. absence) of a postnasal scale. In frontonasal wider than long; nostril ovoid, addition, B. orientalis differs from all non- centered in a single rectangular nasal; supra- pentadactyl species except B. pathfinderi by nasals present, large, narrowly separated by having Toe-IV lamellae 8–10 (vs. four or frontonasal; postnasals present; prefrontals fewer), a paravertebral scale row count 69– moderately separated by frontal; frontal nearly 72 (vs. .84), and by the presence (vs. diamond shaped, its anterior margin in absence) of auricular openings; from all moderate contact with frontonasal, in contact nonpentadactyl species except B. wrighti by with first two anterior supraoculars, 43 wider having a midbody scale row count 26–28 (vs. than anteriormost supraocular; supraoculars ,24); and from B. apus, B. lukbani, B. five; frontoparietals moderate, in broad medi- minimus, and B. vermis by the presence of al contact, each frontoparietal in contact with limbs (vs. absence). supraoculars 2–4; interparietal moderate, Description (based on holotype and 52 quadrilaterally shaped, longer than wide, its referred specimens, including 13 paratypes length slightly greater than midline length of from CAS).—Details of the head scalation of frontoparietal; parietal eyespot present in an adult male are shown in Fig. 8. Measure- posterior half of scale; parietals in narrow ments of the holotype are provided below in contact or separated behind interparietal; brackets. Body large relative to other Brachy- nuchals nonenlarged, undifferentiated; loreals meles, elongate with respect to other lizards; two, decreasing in size from anterior to maximum SVL 112.3 mm for males, 115.2 mm posterior, anterior loreal about as long as for females [99.9, female] (Tables 4 and 5); and 1.83 higher than posterior loreal, in head weakly differentiated from neck, nearly contact with prefrontal, postnasal, supranasal, as wide as body, HW 10.2–12.7% (11.2 6 0.7) second supralabial, posterior loreal and fron- SVL [11.9] 87.5–120.0% (108.4 6 7.3) HL tonasal, and occasionally with first supralabial; [120.0]; HL 33.7–45.0% (37.6 6 2.6) SnFa preocular single, nearly two thirds as high as [38.0]; SnFa 25.3–30.2% (27.6 6 1.1) SVL posterior loreal; presubocular single; supraci- [26.1]; snout long, rounded in dorsal and liaries six, the anteriormost contacting pre- lateral profile, SNL 46.6–65.2% (57.5 6 4.2) frontal and separating posterior loreal from 30 HERPETOLOGICAL MONOGRAPHS [No. 24 first supraocular; subocular row complete; variation among the 19 specimens examined lower eyelid with one row of scales, lacking for the degree of contact between head scales. an enlarged oval window, largely transparent; Twelve specimens were observed to have supralabials six or seven (six), first 23 size of parietals moderately separated by the inter- other supralabials, fourth and fifth below the parietal (KU 305470, 310734–6, 310942, eye; infralabials six or seven (seven). 310944, 310949, 310951, 311232–5), one Mental wider than long, in contact with first specimen has parietals in point medial contact infralabial; single enlarged postmental, wider (KU 311231), and six specimens have parietals than mental; followed by two pairs of enlarged in moderate medial contact (KU 310739, chin shields, first pair narrowly separated by 310943, 310945–6, 310955, 311241) behind single row of undifferentiated scales or in the interparietal; nine specimens do not have moderate medial contact, scales of second pair the first pair of enlarged chin shields in medial narrower than first, broadly separated by contact (KU 210736, 310942, 310945–6, three medial scales. 310949, 310951, 311231, 311234–5), and 10 Scales on limbs smaller than body scales; specimens have the first pair of enlarged scales on dorsal surfaces of digits large, chin shields in moderate medial contact wrapping around lateral edges of digits; (KU 305470, 310734–5, 310739, 310943–4, lamellae undivided; palmar surfaces of hands 310955, 311232–3, 311241). and plantar surfaces of feet covered by small, Scale counts were observed to vary among irregular scales, each with irregular raised the measured series. The number of suprala- anterior edges; scales on dorsal surface of bials varied between six (CAS-SU 18702, hands and feet smaller than limb scales, 24428, 24434, 24436–7, 24442, 24446–9, lacking raised edges. 24451, 24458, 25452, 25460, 28332, CAS Coloration in preservative.—Ground color 102404, 110978–81, 133301, 133616, 133749, of body cream; lateral and ventral surfaces of 133752, 133754, KU 310734, 310736, 310942, body lacking dark pigment; dorsum of body, 311231–2, 311234) and seven (CAS-SU from posterior edge of supranasals to tail tip, 24450, 28320–1, 28338, 28370, CAS 110976– uniformly dark brown with dark pigmentation 7, 110982–3); infralabials varied between six spanning six and two half rows of scales at (CAS-SU 24446, KU 310734, 310736, 310942, midbody and narrowing to cover four and two 311231–2, 311234) and seven (CAS-SU half rows of scales posterior to parietals; body 18702, 24428, 24434, 24436–7, 24442, dark brown dorsally abruptly changing to 24447–51, 24458, 25452, 25460, 28320–1, cream laterally and ventrally; head scales 28332, 28338, 28370, CAS 102404, 110976– uniform dark brown; rostral, nasal, postnasal, 83, 133301, 133616, 133749, 133752, 133754). supranasal, and first supralabial light gray; Specimens were observed to have midbody pineal eyespot charcoal; small dark brown scale row counts of 26 (CAS-SU 24428, blotch dorsal to auricular openings. Limbs 24446), 27 (CAS-SU 24458, 25460), 28 mottled light and medium brown dorsally, (CAS-SU 18702, 24434, 24436–7, 24442, cream colored ventrally; dorsal and ventral 24447–52, CAS 102404, 133301, 133752, surface of digits light brown. 133754), 29 (CAS-SU 28320, 28338, CAS Coloration in life.—Dorsal ground color 110976, 110981, 133616, 133749), and 30 homogeneous medium brown (Fig. 11F,G); (CAS-SU 28231–2, 28370, CAS 110977–80, sharp lateral demarcation between dorsal and 110982–3); axilla–groin scale row counts of 46 lateral and ventral coloration; lateral and ventral (CAS-SU 25452, CAS 110983, 133616), 47 surfaces of body bright burnt orange, orange- (CAS-SU 24450, 24458, 25460, CAS 110979– brown, or salmon colored; dark brown spots 81, 133301), 48 (CAS-SU 18702, 24428, and longitudinal lines of spots absent from 24436, 24442, 24446, 24449, 28332, 28338, lateral surfaces. Limbs medium brown dorsally, CAS 110976, 110978, 110982, 133749), and 49 burnt orange to orange-brown ventrally. Dorsal (CAS-SU 24434, 24437, 24447–8, 24451, head scales uniform medium brown. 28320–1, 28370, CAS 102404, 110977, Variation.—Morphometric variation of the 133752, 133754); and paravertebral scale row series is summarized in Table 6. We observed counts of 69 (CAS-SU 25452, CAS 133616), 2010] HERPETOLOGICAL MONOGRAPHS 31

70 (CAS-SU 24450, 24458, 25460, CAS observation). Similar to B. boulengeri, B. 133301), 71 (CAS-SU 18702, 24428, 24442, orientalis seems to have a wider geographic 24446, 24449, 28332, 28338, CAS 110980, distribution that spans multiple Philippine 110983, 133749), 72 (CAS-SU 24434, 24436– islands. This is in contrast to the pentadactyl, 7, 24447–8, 24451, 28320–1, CAS 102404, island endemic species B. boholensis, B. 110976–9, 110981–2, 133752, 133754), and 73 tungaoi, and B. mindorensis that are known (CAS-SU 28370). from just Bohol, Masbate, and Mindoro We also observed lamellae counts to vary islands, respectively. As do all members of among the measured series. Specimens were the genus, disturbed individuals move in a observed to have Finger-III lamellae counts of rapid serpentine manner and always attempt six (CAS-SU 18702, 24428, 24434, 24436–7, to burrow back into loose soil or humus. 24442, 24446–7, 24449–51, 24458, 25460, Sympatric lizard species observed within 28320–1, 28332, 28338, 28370, CAS 102404, the range of Brachymeles orientalis include 110976–83, 133301, 133616, 133749, 133754, the following: (Agamidae) Bronchocela crista- KU 310734, 311231–2, 311234) or seven tella, Draco bimactulatus, D. cyanopterus, D. (CAS-SU 24448, CAS 133752, KU 310736, mindanensis, D. ornatus, Gonocephalus inter- 310942); Toe-IV lamellae counts of eight ruptus, G. semperi, Hydrosaurus pustulatus; (CAS-SU 24436, 25452, 28320, CAS 102404), (Gekkonidae) Cyrtodactylus agusanensis, C. nine (CAS-SU 18702, 24428, 24434, 24437, annulatus, C. jambangan, Gehyra mutilata, 24442, 24446–51, 24458, 25460, 28321, 28332, Gekko gecko, Gekko mindorensis, Hemidacty- 28370, CAS 110976–7, 110979, 110981–3, lus frenatus, H. platyurus, Pseudogekko com- 133301, 133616, 133754, KU 310734, 310942, pressicorpus; (Scincidae) Brachymeles grac- 311232, 311234), or 10 (CAS-SU 28338, CAS ilis hilong, B. cf. g. gracilis, B. samarensis, 110978, 110980, 133749, 133752, KU 310736, Eutropis indeprensa, E. multifasciata, Lam- 311231). prolepis smaragdina, L. pulchella, L. quad- Distribution.—Brachymeles orientalis is rivittata, Sphenomorphus abdictus abdictus, known from Bohol, Samar, Leyte, Dinagat, S. acutus, S. cumingi, S. cf. mindanensis, S. Camiguin Sur islands and the eastern and coxi, S. fasciatus, S. jagori, S. llanosi, S. central portions of Mindanao Island (Fig. 4). steerei, S. variegatus, Tropidophorus misami- Ecology and natural history.—Brachymeles nus; and (Varanidae) Varanus cumingi. orientalis occurs in agricultural areas as well as disturbed and secondary growth forest. On Brachymeles schadenbergi (Fischer 1885) Samar, Leyte, Mindanao, and Camiguin Sur Figs. 4, 8, 11E islands, we have collected this species in Senira bicolor (part), Gray, 1845. primary forest, and on Bohol Island it is Eumeces (Riopa) schadenbergi, Fischer, 1885. present in mature secondary growth. Individ- Type locality: ‘‘Southern Mindanao Island, uals have been observed under piles of rotting Philippines’’ (reported by Fischer [1885] as coconut husks, in the humus material within No. 845 housed in the Dresden Museum). rotting logs, and in loose soil and leaf litter Brachymeles schadenbergi (part), Boettger, surrounding the root networks of trees. The 1886; Boulenger, 1887; Boettger, 1893; species is quite common throughout its range Taylor, 1917, 1922a,b; Brown and Alcala, with the exception of Bohol Island (CDS, 1970. personal observation), and it occurs sympatri- Brachymeles schadenbergi schadenbergi (part), cally with four other species of Brachymeles in Brown, 1956; Brown and Rabor, 1967. different parts of its range (Brown, 1956; Diagnosis.—Brachymeles schadenbergi can Brown and Alcala, 1980; Brown and Rabor, be distinguished from congeners by the 1967). Brachymeles orientalis occurs sympat- following combination of characters: (1) body rically with B. boholensis on Bohol Island, B. large (SVL 93.1–115.8 mm); (2) pentadactyl; gracilis hilong and B. samarensis on Samar (3) Finger-III lamellae five or six; (4) Toe-IV and Leyte islands, B. gracilis hilong on lamellae eight or nine; (5) limbs relatively Mindanao Island, and B. cf. gracilis hilong long; (6) supralabials six or seven; (7) infra- on Camiguin Sur Island (CDS, personal labials six or seven; (8) pineal eye spot present; 32 HERPETOLOGICAL MONOGRAPHS [No. 24

(9) supranasals in contact; (10) prefrontals not lizards; maximum SVL 115.8 mm for males, contacting on midline; (11) enlarged chin 113.5 mm for females [85] (Tables 4 and 5); shields in two pairs; (12) nuchal scales head weakly differentiated from neck, nearly undifferentiated; (13) fifth and sixth suprala- as wide as body, HW 10.2–11.7% (11.2 6 0.5) bial below eye; (14) auricular opening present; SVL, 102.0–116.8% (108.7 6 4.2) HL; HL and (15) continuous, light dorsolateral stripes 37.0–40.2% (38.4 6 0.9) SnFa; SnFa 25.7– absent (Tables 4 and 5). 27.4% (26.8 6 0.5) SVL; snout moderately Comparisons.—Characters distinguishing long, rounded in dorsal and lateral profile, Brachymeles schadenbergi from all pentadac- SNL 48.8–58.8% (54.3 6 2.9) HL; auricular tyl species of Brachymeles are summarized in opening present, moderate; eyes moderate, Tables 4 and 5. Brachymeles schadenbergi ED 1.8–2.2% (2.1 6 0.1) SVL, 18.4–21.8% most closely resembles B. makusog and B. (20.0 6 1.0) HL, 44.7–57.5% (52.3 6 4.3) orientalis, but it differs from both taxa by END, pupil nearly round; body slightly having eight or nine Toe-IV lamellae, and the depressed, MBW 94.8–135.4% (115.8 6 fifth and sixth supralabial below the eye, and 13.0) MBH; scales smooth, glossy, imbricate; by contact between supranasals (Tables 4 and longitudinal scale rows at midbody 26–28 [28 5). Brachymeles schadenbergi can further be fide Fischer, 1885]; paravertebral scale rows distinguished from B. makusog by having a 67–72; axilla–groin scale rows 45–50 [46 fide greater maximum axilla–groin scale row count Fischer, 1885]; limbs well developed, penta- and a greater maximum paravertebral scale dactyl, digits small; FinIIIlam 5–6; ToeIVlam row count (Table 5) and from B. orientalis by 8–9; FLL 12.5–21.6% (18.0 6 2.4) AGD, the absence (vs. presence) of reddish orange 10.3–13.7% (12.0 6 0.9) SVL [12.9 fide to salmon-colored scales on the lateral surfac- Fischer, 1885]; HLL 20.1–24.4% (29.7 6 es of the body. 3.9) AGD, 17.4–22.0% (19.8 6 1.4) SVL From all nonpentadactyl species of Brachy- [22.4 fide Fischer, 1885]; order of digits from meles (B. apus, B. bonitae, B. cebuensis, B. shortest to longest for hand: V 5 I , II 5 IV elerae, B. lukbani, B. minimus, B. muntingka- , III, for foot: I , V , II , III , IV; tail not may, B. pathfinderi, B. samarensis, B. tridac- as wide as body, gradually tapered toward the tylus, B. vermis,andB. wrighti), B. schaden- end, TW 57.4–76.8% (68.5 6 5.7) MBW, TL bergi differs by having a pentadactyl body 64.6–102.6% (91.6 6 10.8) SVL. form (vs. nonpentadactyl), longer forelimb Rostral projecting dorsoposteriorly to point lengths (.11.1 mm vs. ,6.9 mm), greater in line with anterior edge of nasal, broader hind limb lengths (.18.5 mm vs. ,12.9 mm), than high, separated from frontonasal; fronto- and by the presence of a postnasal scale (vs. nasal wider than long; nostril ovoid, centered absence). In addition, B. schadenbergi differs in a single rectangular nasal; supranasals from all nonpentadactyl species except B. present, large, in broad medial contact; pathfinderi by having Toe-IV lamellae eight or postnasals present; prefrontals moderately nine (vs. four or fewer), 67–72 paravertebrals separated by frontal; frontal nearly diamond (vs. .84), and by the presence (vs. absence) of shaped, its anterior margin in moderate auricular openings; from all nonpentadactyl contact with frontonasal, in contact with first species except B. wrighti by having a midbody two anterior supraoculars, 53 wider than scale row count 26–28 (vs. ,24); and from B. anteriormost supraocular; supraoculars five; apus, B. lukbani, B. minimus, and B. vermis frontoparietals moderate, broad contact me- by the presence (vs. absence) of limbs. dially, each frontoparietal in contact with Description (based on holotype description supraoculars 2–4; interparietal moderate, and 34 referred specimens).—Details of the quadrilaterally shaped, longer than wide, its head scalation of an adult male are shown in length greater than midline length of fronto- Fig. 8. The holotype was not examined by us; parietal; parietal eyespot present in posterior however, measurements of the holotype taken half of scale; parietals in moderate to broad from the original description are provided contact behind interparietal or moderately below in brackets. Body large relative to other separated; nuchals undifferentiated; loreals Brachymeles, elongate with respect to other two, decreasing in size from anterior to 2010] HERPETOLOGICAL MONOGRAPHS 33 posterior, subequal, in contact with prefrontal, medium brown; lateral surfaces with irregu- postnasal, supranasal, second supralabial, pos- larly shaped rows of dark brown spots. Limbs terior loreal and frontonasal; preocular single, dark brown dorsally, medium brown ventrally. nearly two thirds as high as posterior loreal; Dorsal head scales blotched dark and medium presubocular single; supraciliaries six, the brown. anteriormost contacting prefrontal and sepa- Variation.—Morphometric variation of the rating posterior loreal from first supraocular; series is summarized in Table 6. We observed subocular row complete; lower eyelid with variation among the 36 specimens examined one row of scales, lacking an enlarged oval for the degree of contact between head scales. window, largely transparent; supralabials six Twenty-one specimens were observed to have or seven, fifth and sixth beneath center of eye; parietals moderately separated by the inter- infralabials six or seven. parietal (KU 314969, 314976, 314984–5, Mental wider than long, in contact with first 314988–9, 314992, 314997; MCZ 26552–3, infralabials; single enlarged postmental, wider 26556–8, 26561, 26563, 26566, 26568, 26571– than mental, followed by two pairs of enlarged 2, 26574), one specimen has parietals narrow- chin shields; first pair in slight contact or ly separated by the interparietal (KU 314996), narrowly separated by single undifferentiated and 14 specimens have parietals in moderate scale, second pair narrower than first, broadly medial contact (KU 314967, 314970–5, separated by undifferentiated scales. 314977–8, 314980, 314990–1, 314994; MCZ Scales on limbs smaller than body scales; 26555) behind the interparietal; seven speci- scales on dorsal surfaces of digits large, mens do not have the first pair of enlarged wrapping around lateral edges of digits; chin shields in medial contact (KU 314971, lamellae undivided; palmar surfaces of hands 314973, 314976, 314990, 314992, 314994; and plantar surfaces of feet covered by small, MCZ 26552, 26554, 26563), one specimen irregular scales, each with irregular raised has the first pair of enlarged chin shields in anterior edges; scales on dorsal surface of point medial contact (KU 314997), and 26 hands and feet smaller than limb scales, specimens have the first pair of enlarged chin lacking raised edges. shields in moderate medial contact (KU Coloration in preservative.—Ground color 314967, 314969, 314970, 314972, 314974–5, of body medium brown; dorsal surfaces nearly 314977–8, 314980, 314984–5, 314988–9, all dark brown, gradually fading into medium 314991, 314996; MCZ 26553, 26555–8, brown lateral and ventral surfaces of body; 26561, 26566, 26568, 26571–2, 26574). dark dorsal pigmentation in nearly continuous Scale counts were observed to vary among block across dorsal surface, spanning six full the measured series. The number of suprala- and two half rows of scales at midbody and bials varied between six (CAS 23495, KU narrowing to cover four full and two half rows 314991) and seven (CAS 23468–9, 23471, of scales posterior to parietals; lateral surfaces 23479–81, 23484–5, 23494, 23496, 60493, KU with one to two irregular dark brown lines on 314967, 314969, 314974–5, 314977–8, posterior half of axilla–groin region; head 314980, 314984–5, 314994, 314996); infrala- scales uniform dark brown; rostral, nasal, bials varied between six (KU 314967, 314969, postnasal, supranasal, first supralabial, mental, 314974–5, 314977, 314980, 314984–5, and first infralabial dark gray; pineal eyespot 314991, 314996) and seven (CAS 23468–9, poorly defined, surrounded by light cream 23471, 23479–81, 23484–5, 23494–6, 60493). border. Tail coloration matches body colora- Specimens were observed to have midbody tion. Limbs mottled dark brown dorsally, scale row counts of 26 (CAS 23468, 23479–81, medium brown ventrally; dorsal and ventral 23494–6, 60493), 27 (CAS 23469, 23484), and surface of digits dark brown. 28 (CAS 23471, 23485); axilla–groin scale row Coloration in life.—Dorsal ground color counts of 45 (CAS 23495), 46 (CAS 23469, homogeneous dark brown (Fig. 11E); 23494, 23496), 47 (CAS 23468, 23484), 48 blotched, irregular, lateral demarcation be- (CAS 23485, 60493), 49 (CAS 23471, 23480), tween dorsal and lighter lateral and ventral and 50 (CAS 23479, 23481); and paravertebral coloration; lateral and ventral surfaces of body scale row counts of 67 (CAS 23495), 68 (CAS 34 HERPETOLOGICAL MONOGRAPHS [No. 24

23494, 23496), 70 (CAS 23484), 71 (CAS of the genus, disturbed individuals move in a 23468–9, 23471, 23480, 23485, 60493), and 72 rapid serpentine manner and always attempt (CAS 23479, 23481). to burrow back into loose soil or humus. We also observed lamellae counts to vary Sympatric lizard species observed within the among the measured series. Specimens were range of Brachymeles schadenbergi include the observed to have Finger-III lamellae counts of following: (Agamidae) Bronchocela cristatella, five (CAS 23469, 23495, 60493, KU 314967, Draco bimaculatus, D. cyanopterus, D. mind- 314969, 314974, 314977–8, 314984–5, anensis, Gonocephalus interruptus, Hydro- 314991, 314994, 314996) or six (CAS 23468, saurus amboinensis; (Gekkonidae) Cyrtodac- 23471, 23479–81, 23484–5, 23494, 23496, KU tylus jambangan, Gehyra mutilata, Gekko 314975, 314980); Toe-IV lamellae counts of gecko, Hemidactylus frenatus, H. platyurus, eight (CAS 23468, 23494–5, 60493, KU Hemiphyllodactylus typus, Lepidodactylus sp., 314967, 314969, 314974–5, 314977, 314984– L. quadrivittata, Luperosaurus joloensis, Pseu- 5, 314991, 314994, 314996) or nine (CAS dogekko compressicorpus; (Scincidae) Brachy- 23469, 23471, 23479–81, 23484–5, 23496, KU meles gracilis gracilis, Eutropis indeprensa, E. 314978, 314980). multicarinata, E. multifasciata, E. englei, Lam- There is a small degree of color variation in prolepis smaragdina, Sphenomorphus atrigu- the examined series, with the degree and laris, S. fasciatus, S. jagori, S. steerei, S. definition of continuous, dark middorsal variegatus, Tropidophorus misaminus, T. par- pigmentation. Most of the examined speci- telloi; and (Varanidae) Varanus cumingi. mens show patterns consistent with a contin- uous, dark streak of pigmentation covering the Brachymeles talinis Brown 1956 middorsal region of the body (KU 314969, Figs. 5, 9, 11H 314974–5, 314977–8, 314980, 314984–5, Brachymeles schadenbergi talinis, Brown, 314991, 314994, 314996). In several speci- 1956. Type locality: ‘‘On the low ridge mens, continuous, dark middorsal stripes are north side of the Maite River, 5 to 6 km evident overlaying the dark ground coloration west of Valencia,’’ Negros Oriental Prov- (KU 314967). ince, Negros Island, Philippines, 933-m Distribution.—Brachymeles schadenbergi elevation, 9u 179 19.250 N, 123u 119 56.40 is known from Basilan and western Mindanao E (holotype: CAS-SU 18358). islands (Fig. 4). Brachymeles talinis, Brown and Rabor, 1967. Ecology and natural history.—Brachymeles Brachymeles talinis, Brown and Alcala, 1980. schadenbergi occurs in a variety of habitats Diagnosis.—Brachymeles talinis can be from disturbed and secondary growth to distinguished from congeners by the following primary forest and intact climax forest. combination of characters: (1) body size large Individuals have been observed in the humus (SVL 103.8–123.1 mm); (2) pentadactyl; (3) material within rotting logs and in loose soil Finger-III lamellae five or six; (4) Toe-IV and leaf litter surrounding the root networks lamellae 8–10; (5) limbs relatively long; (6) of trees. Individuals are moderately common paravertebral scale rows 67–72; (7) suprala- in populations sampled (CDS and RMB, bials seven; (8) infralabials seven; (9) pineal personal observations) and occur sympatrical- eye spot present; (10) supranasals in contact; ly with B. gracilis gracilis in western Mind- (11) prefrontals not contacting on midline; anao Island (Brown, 1956; Brown and Alcala, (12) enlarged chin shields in two pairs; (13) 1980; Brown and Rabor, 1967). We collected nuchal scales undifferentiated; (14) fifth and numerous specimens in pitfall traps, indicat- sixth supralabial below eye; (15) auricular ing some level of surface activity. Although B. opening present; (16) dark lateral stripes schadenbergi occurs on multiple islands in the present; and (17) venter devoid of dark southern Philippines, the species seems to pigmentation (Tables 4 and 5). have a more restricted geographic distribution Comparisons.—Characters distinguishing compared with more widespread pentadactyl Brachymeles talinis from all pentadactyl species, such as B. boulengeri, B. talinis, B. species of Brachymeles are summarized in orientalis, and B. kadwa. As in other members Tables 4 and 5. Brachymeles talinis most 2010] HERPETOLOGICAL MONOGRAPHS 35

FIG. 9.—Illustration of head of adult male Brachymeles talinis (KU 306769) and adult male holotype of Brachymeles kadwa (PNM 9721; formerly KU 323091) in dorsal, lateral, and ventral views. Labels for taxonomically diagnostic head scales follow those shown in Fig. 6. Illustrations by CDS. closely resembles B. kadwa, B. makusog, B. than the second, frontoparietals in contact, tungaoi, and B. vindumi, but it differs from and by the absence of dark ventral pigmenta- these four taxa by having the range of tion (Tables 4 and 5); from B. makusog by paravertebral scale rows reaching .70 but having seven supralabials the fifth and sixth ,74, and seven infralabials (Table 5). Bra- supralabial below the eye, supranasals in chymeles talinis can further be distinguished contact, and by the presence of dark lateral from B. kadwa by having 8–10 Toe-IV stripes (Tables 4 and 5); from B. tungaoi by lamellae, the first enlarged chin shield wider having a larger body size, shorter relative tail 36 HERPETOLOGICAL MONOGRAPHS [No. 24 length, 8–10 Toe-IV lamellae, and the first midbody 26–30 [29]; paravertebral scale rows enlarged chin shield wider than the second 67–72 [72]; axilla–groin scale rows 43–48 [48]; (Tables 4 and 5); and from B. vindumi by limbs well developed, pentadactyl, digits having fewer axilla–groin scale rows, fewer moderate; FinIIIlam 5–6 [6]; ToeIVlam 8–10 paravertebral scale rows, and by the absence [10]; FLL 15.1–23.9% (19.1 6 1.8) AGD of dark ventral pigmentation (Table 5). [19.6], 10.1–15.3% (12.4 6 1.2) SVL [14.9]; From all nonpentadactyl species of Brachy- HLL 26.8–38.9% (31.8 6 3.0) AGD [30.9], meles (B. apus, B. bonitae, B. cebuensis, B. 18.0–24.9% (20.6 6 1.8) SVL [23.5]; order of elerae, B. lukbani, B. minimus, B. muntingka- digits from shortest to longest for hand: V , I may, B. pathfinderi, B. samarensis, B. tridac- , IV , II , III, for foot: I 5 V , II , III 5 tylus, B. vermis, and B. wrighti), B. talinis IV; tail nearly as wide as body at base, sharply differs by having a pentadactyl body form (vs. tapered toward the end, TW 63.4–94.0% (75.1 nonpentadactyl), longer forelimb lengths 6 6.2) MBW [94.0], TL 60.6–107.2% (83.9 6 (.11.3 mm vs. ,6.9 mm), and greater hind 12.0) SVL [73.2]. limb lengths (.20.5 mm vs. ,12.9 mm), and Rostral projecting dorsoposteriorly to point by the presence of a postnasal scale (vs. in line with anterior edge of nasal, broader absence). In addition, B. talinis differs from all than high, completely separated from fronto- nonpentadactyl species except B. wrighti by nasal by broad supranasal contact; frontonasal having a midbody scale row count 26–30 (vs. wider than long; nostril ovoid, in center of ,24); from all nonpentadactyl species except single trapezoidal nasal; supranasals present, B. pathfinderi by having a paravertebral scale large, in broad medial contact; postnasals row count 68–70 (vs. .84), and by the present; prefrontals moderately separated by presence of auricular openings (vs. absence); frontal; frontal nearly octagonal shaped, its from all nonpentadactyl species except B. anterior margin in moderate contact with apus and B. wrighti by having a larger body frontonasal, in contact with first two anterior size (SVL .103.1 mm vs. ,81.3 mm); and supraoculars, 43 wider than anteriormost from B. apus, B. lukbani, B. minimus, and B. supraocular; supraoculars five; frontoparietals vermis by the presence (vs. absence) of limbs. moderate, in moderate medial contact, each Description (based on holotype and 30 frontoparietal in contact with supraoculars referred specimens, including two paratypes 2–4; interparietal moderate, quadrilaterally from CAS).—Details of the head scalation of shaped, width nearly equal to length, its an adult male are shown in Fig. 9. Measure- length nearly equal to midline length of ments of the holotype are provided below in frontoparietal; parietal eyespot present in brackets. Body large relative to other Brachy- posterior one third of scale; parietals in point meles, elongate with respect to other lizards; to moderate contact behind interparietal or maximum SVL 123.1 mm for males, 116.5 mm narrowly separated; nuchals undifferentiated; for females [118.7, male] (Tables 4 and 5); loreals two, decreasing in size from anterior to head weakly differentiated from neck, nearly posterior, subequal, in contact with prefrontal, as wide as body, HW 10.6–13.2% (11.8 6 0.5) postnasal, supranasal, second supralabial, pos- SVL [12.1], 111.7–136.2% (124.1 6 6.8) HL terior loreal and frontonasal; preocular single, [130.3]; HL 30.6–40.7% (35.4 6 2.7) SnFa nearly three fourths as high as posterior loreal; [33.5]; SnFa 25.1–29.8% (27.0 6 1.3) SVL single presubocular; supraciliaries six, the [27.6]; snout moderately long, broadly round- anteriormost contacting prefrontal and sepa- ed in dorsal and lateral profile, SNL 51.5– rating posterior loreal from first supraocular; 65.8% (58.9 6 4.0) HL [65.8]; auricular subocular row complete; lower eyelid with opening present, moderate; eyes moderate, one row of scales, lacking an enlarged oval ED 1.7–2.2% (1.9 6 0.1) SVL [1.7], 17.4– window, largely transparent; supralabials sev- 24.8% (20.0 6 1.9) HL [18.2], 40.9–62.1% en, fifth and sixth below the eye; infralabials (50.3 6 5.7) END [41.5], pupil nearly round; seven. body slightly depressed, MBW 109.3–153.8% Mental wider than long, in contact with first (126.7 6 14.4) MBH [109.6]; scales smooth, infralabials; single enlarged postmental, wider glossy, imbricate; longitudinal scale rows at than mental; followed by two pairs of enlarged 2010] HERPETOLOGICAL MONOGRAPHS 37 chin shields; first pair in moderate contact or brown, bordered middorsally by rows of dark moderately separated by a single medial scale, spots; lateral surface ground color light brown wider than second pair; second pair separated to tan; ventral surfaces of body light brown to by three undifferentiated scales. tan. Dorsal surfaces of limbs dark to medium Scales on limbs smaller than body scales; brown, ventral surfaces light brown. Dorsal scales on dorsal surfaces of digits large, head scales blotched dark and medium brown. wrapping around lateral edges of digits; Variation.—Morphometric variation of the lamellae undivided; palmar surfaces of hands series is summarized in Table 6. We observed and plantar surfaces of feet covered by small, variation among the 19 specimens examined irregular scales, each with irregular raised for the degree of contact between head scales. anterior edges; scales on dorsal surface of Four specimens were observed to have hands and feet smaller than limb scales, parietals moderately separated by the inter- lacking raised edges. parietal (KU 306757, 306763, 306767, Coloration in preservative.—Ground color 306786), one specimen has parietals narrowly of body medium brown; longitudinal stripes separated by the interparietal (KU 306758), on dorsal surface of body present or absent; and 14 specimens have parietals in moderate when present a total of eight longitudinal dark medial contact (KU 306756, 306759–60, brown spot rows, extending from posterior 306762, 306764–6, 306769–71, 306773–6) edge of parietals to base of tail: six continuous behind the interparietal; 13 specimens do medial rows and two discontinuous postero- not have the first pair of enlarged chin shields lateral rows, together spanning eight full rows in medial contact (KU 304756–7, 306759–60, of scales at midbody, narrowing to six full rows 306762, 306764–6, 306767, 306769–70, of scales posterior to parietals; when dark spot 306774–5) and six specimens have the first rows are absent, pigmentation forms nearly pair of enlarged chin shields in moderate continuous dark dorsal surface, covering one medial contact (KU 306758, 306763, 306771, half to entire surface of dorsal scales; dorso- 306773, 306776, 306786). lateral stripes present or absent, when pres- Scale counts were observed to vary among ent, well defined, continuous, lacking dark the measured series. Specimens were ob- pigmentation, spanning two whole and two served to have midbody scale row counts of half row of scales from auricular opening to 26 (KU 306766), 28 (CAS-SU 22311, 22317, base of tail. Lateral and ventral surface of 37996, KU 306765), 29 (CAS-SU 12225, body medium brown. Lateral surface with 18358, 22323, 27972, 89813, CAS 133871, three to six discontinuous longitudinal rows of KU 306758, 306774), and 30 (CAS-SU 22312, dark brown spots, rows often extending to 27997, KU 306756, 306760, 306769, 306772–3, edge of ventral surface. Ventral surface 306786); axilla–groin scale row counts of 43 without dark pigmentation. Tail coloration (KU 306786), 44 (CAS-SU 12225, 22311, equal to body coloration, dorsal surface 27996–7), 45 (CAS-SU 22323, KU 306756, covered with dark brown blotches, ventral 306758, 306760, 306765–6, 306772), 46 (CAS- surface covered with scattered dark brown SU 22312, 22317, 27972, 89813, CAS 133871, spots, fewer than dorsal surface. Head scales KU 306773), 47 (KU 306774), and 48 (CAS-SU homogeneous dark brown; rostral, nasal, 18358, KU 306769); and paravertebral scale postnasal, supranasal, first supralabial, mental row counts of 67 (KU 306786), 68 (CAS-SU and first infralabial dark gray; pigment sur- 12225, 22311, 27996–7, KU 306756, 306758), rounding pineal eyespot reduced to indistinct, 69 (CAS-SU 22312, 22323, KU 306760, small and medium brown. Limbs mottled 306765–6, 306772–3), 70 (CAS-SU 22317, medium brown dorsally, yellowish brown 37972, 89813, CAS 133871, KU 306769), 71 ventrally; dorsal and ventral surface of digits (KU 306774), and 72 (CAS-SU 18358). dark brown. We also observed lamellae counts to vary Coloration in life.—Dorsal ground color among the measured series. With the excep- medium brown (Fig. 11H); when present, tion of two specimens observed to have six longitudinal rows of spots dark brown to Finger-III lamellae (CAS-SU 18358, CAS black; dorsolateral stripes light to medium 133871), all other examined specimens were 38 HERPETOLOGICAL MONOGRAPHS [No. 24 observed to have five. We also observed Toe- Sympatric lizard species observed within IV lamellae counts of eight (KU 306769, the range of Brachymeles talinis include the 306786), nine (CAS-SU 22311, 22317, following: (Agamidae) Bronchocela marmor- 89813, KU 306756, 306758, 306760, 306765– atus, Draco spilopterus, Hydrosaurus pustu- 6, 306772–4), and 10 (CAS-SU 12225, 18358, latus; (Dibamidae) Dibamus argenteus; (Gek- 22312, 22323, 27972, 27996–7, CAS 133871). konidae) Cyrtodactylus philippinicus, Gehyra Color variation exists in the degree and mutilata, Gekko gecko, Gekko mindorensis, definition of continuous, dark middorsal Gonocephalus sophiae, Hemidactylus frena- stripes. Many specimens show patterns con- tus, H. platyurus, Hemiphyllodactylus typus, sistent with continuous, middorsal dark lines Lepidodactylus christiani, L. herrei, L. lugu- (KU 306651, 306756–7, 306759, 30676,2 bris, Luperosaurus corfieldi, Pseudogekko 306765–7, 306769–72, 306776, 306786, brevipes; (Scincidae) Brachymeles tridactylus, 306763). The dark lines are obscured in some B. taylori, Emoia atrocostata, Eutropis multi- and irregular in others, where the middorsal carinata, E. multifasciata, Lamprolepis smar- region is covered by a long streak of dark agdina, Lipinia auriculata, L. pulchella, L. pigmentation, with little to moderate line quadrivittata, L. rabori, Sphenomorphus ar- definition (KU 306759–60, 306764, 306774). borens, S. coxi, S. jagori, S. steerei, Tropido- Distribution.—Brachymeles talinis is phorus grayi; and (Varanidae) Varanus nu- known from Negros, Panay, Romblon, Sibu- chalis. yan, and Tablas islands (Fig. 5). It is also likely to occur on Guimaras Island. Brachymeles kadwa sp. nov. Ecology and natural history.—Brachymeles Figs. 5, 9, 11D talinis occurs in a variety of habitats from Holotype.—PNM 9721 (RMB Field agricultural areas, to disturbed and second- No. 12466, formerly KU 323091), adult male, ary growth forest. Little or no original, collected under rotting logs in secondary- lowland forest remains in the Visayas, but growth forest (1000 to 1230 h) on 4 June 2009, we assume the species originally occurred in on the campus of Aurora State College of primary forest. Individuals have been ob- Technology, Barangay Zabali, Municipality of served under piles of rotting coconut husks, Baler, Aurora Province, Luzon Island, Philip- in the humus material within rotting logs, pines (15u 449 310 N, 121u 349 340 E; WGS- and in loose soil and leaf litter surrounding 84), by CDS, RMB, J. Fernandez, L. Welton, the root networks of trees. The species is J. Brown, J. Siler, Y. Vicente, and M. Vicente. moderately common throughout its range Paratopotypes.—Three adult males (KU (CDS, personal observation) and occurs 323092, 323095, 323096) and four adult fe- sympatrically with three other species (B. males (KU 323106, 323094, 323104, 323100), bonitae, B. talinis,andB. tridactylous; collected between 4 and 7 June 2009. Brown, 1956; Brown and Alcala, 1980; Paratypes.—Four adult males (KU 304875, Brown and Rabor, 1967). Individuals were 304900, 304915, 304941) and six adult females often encountered in pitfall traps, indicating (KU 304897, 304902–3, 304905–6, 304929) some level of activity outside of fossorial collected between 15 and 22 March 2006 (19u microhabitats. Similar to B. boulengeri, B. 179 380 N, 121u 249 320 E; WGS-84; 245 m talinis seems to have a wider geographic above sea level) Barangay Magsidel, Munici- distribution that spans multiple Philippine pality of Calayan, Cagayan Province, Calayan islands. This is in contrast to the island Island, Philippines, by RMB, C. Oliveros, and endemic species B. boholensis, B. tungaoi, J. Fernandez; four adult males (KU 304575, and B. mindorensis that are known from just 307984, 307996, 307998) and five adult Bohol, Masbate, and Mindoro islands, re- females (KU 304559, 304593, 304708, spectively. As do all members of the genus, 304754, 308011), four adult males and five when disturbed, individuals attempt to adult females collected between 3 and 11 escape by moving in a rapid serpentine March 2006 from 300-m elevation (18u 559 450 manner and attempting to burrow back into N, 121u 539 560 E; WGS-84) Barangay loose soil or humus. Balatubat, Municipality of Calayan, Cagayan 2010] HERPETOLOGICAL MONOGRAPHS 39

Province, Camiguin Norte Island, Philippines, tinuous, dark middorsal stripes, dark lateral by RMB, C. Oliveros, and J. Fernandez. stripes, and dark ventral pigmentation (Ta- Referred specimens.—CALAYAN IS- ble 5); from B. tungaoi by having a greater LAND: CAGAYAN PROVINCE: Municipality of midbody width, shorter relative tail length, Calayan: Barangay Magsidel: KU 304908, paravertebrals 68–70, and the presence of 304899, 304907, 304909, 304921, 304941; dark ventral pigmentation (Tables 4 and 5); CAMIGUIN NORTE ISLAND: CAGAYAN from B. talinis by having 28 or fewer midbody PROVINCE: Municipality of Calayan: Barangay scale rows, 70 or fewer paravertebrals, infra- Balatubat: KU 304558, 304562–65, 304569, labials six, and by the presence dark ventral 304571–74, 304627–30, 304643, 304647, pigmentation (Table 5); and from B. vindumi 304696–99, 304704–07, 304709–12, 304714, by having five or six Finger-III lamellae, 26– 304753, 304755–59, 307965–66, 307985–86, 28 midbody scale rows, paravertebrals 68–70, 307997, 307999–8003, 308006–10, 308012– and by the presence of continuous, dark 15, 308017–18; LUZON ISLAND: AURORA middorsal stripes (Tables 4 and 5). PROVINCE: Municipality of Baler: Barangay From all nonpentadactyl species of Brachy- Zabali, ASCOT: KU 323090–91, 323093, meles (B. apus, B. bonitae, B. cebuensis, B. 323097–99, 323101–03, 323105, 323107; Mu- elerae, B. lukbani, B. minimus, B. muntingka- nicipality of Casiguran, IDC property: KU may, B. pathfinderi, B. samarensis, B. tridac- 323108–48; Municipality of San Luis: Baran- tylus, B. vermis, and B. wrighti), B. kadwa gay Real, Sitio Minoli: KU 322320. differs by having a pentadactyl body form (vs. Diagnosis.—Brachymeles kadwa can be nonpentadactyl), longer forelimb lengths distinguished from congeners by the following (.10.7 mm vs. ,6.9 mm), and greater hind combination of characters: (1) body size large limb lengths (.17.9 mm vs. ,12.9 mm), and (SVL 90.6–128.2 mm); (2) pentadactyl; (3) by the presence of a postnasal scale (vs. Finger-III lamellae five or six; (4) Toe-IV absence). In addition, B. kadwa differs from lamellae 7–10; (5) limbs relatively long; (6) all nonpentadactyl species except B. wrighti paravertebrals 68–70; (7) supralabials seven; by having 26–28 midbody scales (vs. ,24); (8) infralabials six; (9) pineal eye spot present, from all nonpentadactyl species except B. small; (10) supranasals in contact; (11) pre- pathfinderi by having 68–70 paravertebrals frontals not contacting on midline; (12) (vs. .84), and by the presence of auricular enlarged chin shields in two pairs; (13) nuchal openings (vs. absence); from all nonpentadac- scales undifferentiated; (14) fifth and sixth tyl species except B. apus and B. wrighti by supralabial below eye; (15) auricular opening having a larger body size (SVL .90.6 mm vs. present; (16) continuous, light dorsolateral ,81.3 mm); and from B. apus, B. lukbani, B. stripes present, indistinct; (17) continuous, minimus, and B. vermis by the presence of dark middorsal stripes present; (18) dark limbs (vs. absence). lateral stripes present; and (19) dark ventral Description of holotype.—Mature male, pigmentation present (Tables 4 and 5). hemipenes everted (Fig. 10); SVL 106.2 mm; Comparisons.—Characters distinguishing body moderately large relative to other the new species from all pentadactyl species Brachymeles, elongate with respect to other of Brachymeles are summarized in Tables 4 lizards; head weakly differentiated from neck, and 5. Brachymeles kadwa most closely nearly as wide as body, HW 11.0% SVL, resembles B. makusog, B. tungaoi, B. talinis, 111.1% HL; HL 38.1% SnFa; SnFa 26.0% and B. vindumi, but it differs from these four SVL; snout moderately long, rounded in taxa by having 7–10 Toe-IV lamellae and the dorsal and lateral profile, SNL 56.1% HL; second enlarged chin shield wider than the auricular opening present, small; eyes moder- first (Tables 4 and 5). Brachymeles kadwa can ate, ED 1.9% SVL, 19.6% HL, 54.8% END, further be distinguished from B. makusog by pupil nearly round; body slightly depressed, having seven supralabials, the fifth and sixth MBW 157.3% MBH; body scales smooth, supralabial below the eye, six infralabials, the glossy, imbricate; longitudinal scale rows at presence of supranasal contact, the presence midbody 28; paravertebral scale rows 68; of continuous, light dorsolateral stripes, con- axilla–groin scale rows 47; limbs well devel- 40 HERPETOLOGICAL MONOGRAPHS [No. 24

FIG. 10.—Illustration of head of adult male holotype of Brachymeles tungaoi (PNM 9722; formerly KU 323933) and adult male holotype of Brachymeles vindumi (CAS 60724) in dorsal, lateral, and ventral views. Labels for taxonomically diagnostic head scales follow those shown in Fig. 6. Illustrations by CDS. oped, pentadactyl, digits moderate; FinIIIlam Rostral projecting dorsoposteriorly to point 5; ToeIVlam 10; FLL 20.3% AGD, 13.0% in line with anterior edge of nostril, broader SVL; HLL 32.0% AGD, 20.6% SVL; order of than high, separated from frontonasal by digits from shortest to longest for hand: I 5 V moderate contact of supranasals; frontonasal , II 5 IV , III, for foot: V , I , II , III 5 wider than long; nostril ovoid, centered in a IV; tail nearly as wide as body at base, single rectangular nasal; supranasals large, in gradually tapered toward the end, TW 73.5% moderate medial contact; postnasals present; MBW, TL 101.6% SVL. prefrontals moderately separated by frontal; 2010] HERPETOLOGICAL MONOGRAPHS 41

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 (KU 314973), female, SVL 5 107.0 mm; (F) B. orientalis (KU 311240), juvenile, SVL 5 51.0 mm; (G) B. orientalis (KU 324029), 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. 42 HERPETOLOGICAL MONOGRAPHS [No. 24 frontal nearly diamond shaped, its anterior covering middle three fourths of dorsal scales; margin in moderate contact with frontonasal, dorsolateral stripes somewhat indistinct, dis- in contact with first two anterior supraoculars, continuous, spanning two half rows of scales 43 wider than anteriormost supraocular; from auricular opening point just posterior to supraoculars five; frontoparietals moderate, forelimb insertion; dark dorsal coloration point contact medially or moderately separat- blends gradually into medium brown lateral ed, each frontoparietal in contact with suprao- and ventral surface of body. Lateral surface culars 2–4; interparietal moderate, quadrilat- with six discontinuous, dark brown spot rows, erally shaped, its length slightly greater than extending to edge of ventral surface. Ventral midline length of frontoparietal; parietal surface with scattered dark brown spots. Tail eyespot present in posterior one third of scale, coloration similar to body coloration, dorsal indistinct; parietals in moderate contact be- surface covered with dark brown blotches, hind interparietal; nuchals undifferentiated; ventral surface covered with few dark brown loreals two, decreasing in size from anterior to spots. Head scales homogeneous dark brown; posterior, subequal, in contact with prefrontal, rostral, nasal, postnasal, supranasal, first postnasal, supranasal, second supralabial, pos- supralabial, mental, and first infralabial dark terior loreal and frontonasal; preocular single, gray; pineal eyespot indistinct, small and light nearly two thirds as high as posterior loreal; brown. Limbs mottled dark brown dorsally, single presubocular; supraciliaries six, the yellowish brown ventrally; dorsal and ventral anteriormost contacting prefrontal and sepa- surface of digits dark brown. rating posterior loreal from first supraocular, Coloration in life.—Ground color of body posteriormost extending to midline of last light to medium brown (Fig. 11D); dorsal supraocular; subocular row complete; lower surfaces of limbs medium brown. eyelid with one row of scales, lacking an Measurements of holotype (in millime- enlarged oval window, largely transparent; ters).—SVL 106.2; AGD 68.2; TotL 214.0; supralabials seven, fifth and sixth below the MBW 15.9; MBH 10.1; TL 107.9; TW 11.7; eye; infralabials six. TH 9.2; HL 10.5; HW 11.7; HH 8.6; SnFa Mental wider than long, in contact with first 27.6; ED 2.1; END 3.8; SNL 5.9; IND 3.7; infralabials; single enlarged postmental, slight- FLL 13.8; HLL 21.8; MBSR 28; PVSR 68; ly wider than mental; followed by two pairs of AGSR 47; FinIIIlam 5; ToeIVlam 10; SL 7; enlarged chin shields, first pair in moderate IFL 6; SC 6; SO 5. medial contact, second pair slightly wider than Variation.—Morphometric variation of the first, separated by a single undifferentiated series is summarized in Table 6. We observed scale. variation among the 25 specimens examined Scales on limbs smaller than body scales; for the degree of contact between head scales. scales on dorsal surfaces of digits large, Fourteen specimens were observed to have wrapping around lateral edges of digits; parietals moderately separated by the inter- lamellae undivided; palmar surfaces of hands parietal (KU 304559, 304574–5, 304593, and plantar surfaces of feet covered by small, 304630, 304708, 304754–5, 304759, 304906, irregular scales, each with raised anterior 307984–5, 307996, 308007), one specimen has edges; scales on dorsal surface of hands and parietals in point medial contact (KU 308011), feet smaller than limb scales, lacking raised and 10 specimens have parietals in moderate edges. medial contact (KU 304875, 304897, 304900, Coloration in preservative.—Ground color 304902–3, 304905, 304915, 304929, 304941, of body dark brown; dorsal surface of body 307998) behind the interparietal; two speci- with eight longitudinal rows of dark brown mens have frontoparietals moderately sepa- spots spanning eight full rows of scales at rated by the frontal (KU 304559, 307984), one midbody and extending from posterior edge of specimen has frontoparietals narrowly sepa- parietals to base of tail: six rows in middorsal rated by the frontal (KU 304574), and 22 region, flanked by discontinuous dorsolateral specimens have frontoparietals in moderate rows; spot rows narrowing to six full rows of medial contact (KU 304575, 304593, 304630, scales posterior to parietals; dark coloration 304708, 304754–5, 304759, 304875, 304897, 2010] HERPETOLOGICAL MONOGRAPHS 43

304900, 304902–3, 304905–6, 304915, humus material within rotting logs, and in 304929, 304941, 307985, 307996, 307998, loose soil and leaf litter surrounding the root 308007, 308011). We observed the first pair networks of trees. This species is quite of enlarged chin shields narrowly separated in common in all sampling localities, and we a single specimen (KU 307996) and in have taken large series in pitfall traps, moderate contact for all other examined indicating some level of surface activity. When specimens. disturbed, individuals immediately moved in a Scale counts were observed to vary among rapid serpentine manner and attempted to the measured series. Specimens were ob- burrow back into loose soil or humus. served to have midbody scale row counts of 26 Sympatric lizard species observed on Lu- (KU 304559, 307996, 323104), 27 (KU zon, Camiguin Norte, and Calayan islands 304593, 307984, 307998), and 28 (KU include the following: (Agamidae) Broncho- 304575, 304708, 304754, 304875, 304897, cela cristatella, Draco spilopterus, Gonoce- 304900, 304902–3, 304905–6, 304915, phalus sophiae, Hydrosaurus pustulatus; 304929, 304941, 308011, 323091, 323091–2, (Gekkonidae) Cyrtodactylus philippinicus, 323094–6, 323100, 323106); axilla–groin scale Gehyra mutilata, Gekko gecko, Gekko mind- row counts of 47 (KU 304559, 304575, orensis, Hemidactylus frenatus, H. garnoti, H. 304593, 304875, 304902, 304929, 307996, luzonensis, H. platyurus, Luperosaurus cf. 307998, 308011, 323091, 323096), 48 (KU cumingi, L. kubli, Pseudogekko compressicor- 304708, 304754, 304897, 304900, 304915, pus, P. smaragdina; (Scincidae) Brachymeles 304941, 307984, 323092, 323094–5, 323100, bonitae, B. bicolor, B. elerae, B. lukbani, B. 323104, 323106), 49 (KU 304903, 304905–6); makusog, B. muntingkamay, B. samarensis, B. and paravertebral scale row counts of 68 (KU cf. talinis, B. wrighti, Emoia atrocostata, 304559, 304593, 304900, 307996, 323091, Eutropis bontocensis, E. multicarinata, E. 323096), 69 (KU 304575, 304708, 304754, multifasciata, Lamprolepis smaragdina, Lipi- 304875, 304929, 304941, 307984, 307998, nia pulchella, Sphenomorphus cumingi, S. 308011, 323092, 323094–5, 323100, 323104, decipiens, S. jagori, S. leucospilos, S. luzonen- 323106), and 70 (KU 304897, 304902–3, sis, S. steerei, S. stejnegeri, Tropidophorus 304905–6, 304915). grayi; and (Varanidae) Varanus marmoratus. We also observed lamellae counts to vary Etymology.—CDS is pleased to name this among the measured series. With the excep- new species for his loving wife Jessi M. Siler tion of two specimens observed to have six for her endless support that has made all of Finger-III lamellae (KU 304903, 304906), all this research possible. The name of the new other examined specimens were observed to species is derived from one of the local have five. We also observed Toe-IV lamellae dialects spoken in the Philippines. The word counts of seven (KU 304593), eight (KU ‘‘kadwa’’ is the Ilonggo term for friend and 304559, 304575, 304708, 304754, 304875, companion. Suggested common name: Jessi’s 304897, 304900, 304915, 304929, 304941, slender skink. 307984, 307996, 307998, 308011), nine (KU 304902–3, 304905–6, 323092, 323094–6, Brachymeles tungaoi sp. nov. 323100, 323104, 323106), or 10 (KU 323091). Figs. 5, 10 Distribution.—Brachymeles kadwa is Holotype.—PNM 9722 (CDS Field known from numerous localities on Luzon No. 5125, formerly KU 323933), adult male, Island as well as from Calayan and Camiguin collected in rotting stump in disturbed, Norte Islands of the Babuyan Island group residential habitat (1000 to 1230 h) 4 Sep- (Fig. 5). tember 2009, at 61-m elevation in Municipal- Ecology and natural history.—Brachymeles ity of Masbate City, Masbate Province, kadwa occurs in agricultural areas, disturbed Masbate Island, Philippines (12u 219 010 N, secondary growth forest, and first growth 123u 379 420 E; WGS-84), by CDS and J. forests of Luzon, Camiguin Norte, and Fernandez. Calayan. Individuals have been observed Paratopotypes.—KU 323934–36, three under piles of rotting coconut husks, in the adult females, collected between 3 and 7 44 HERPETOLOGICAL MONOGRAPHS [No. 24

September 2009, from 61–99-m elevation by continuous, light dorsolateral stripes, contin- CDS and J. Fernandez. uous, dark middorsal stripes, and dark lateral Paratypes.—One adult male (CAS 144313), stripes (Table 5); from B. talinis by having three adult females (CAS 144229–30, nine or 10 Toe-IV lamellae, 66–68 paraverte- 144341), and four juvenile specimens of brals, infralabials six (Tables 4 and 5); and unknown sex (CAS 144290, 144306–7, from B. vindumi by having five or six Finger- 144342), collected 2 June 1976 ‘‘in humus III lamellae, 26–28 midbody scale rows, and under rotting log,’’ in Barangay Tugbo, 66–68 paravertebrals (Tables 4 and 5). Municipality of Mobo, Masbate Province, From all nonpentadactyl species of Brachy- Masbate Island, Philippines (12u 209 11.040 meles (B. apus, B. bonitae, B. cebuensis, B. N, 123u 379 58.80 E; WGS-84; 400-m eleva- elerae, B. lukbani, B. minimus, B. muntingka- tion) by A. Alcala. may, B. pathfinderi, B. samarensis, B. tridac- Diagnosis.—Brachymeles tungaoi can be tylus, B. vermis, and B. wrighti), B. tungaoi distinguished from congeners by the following differs by having a pentadactyl body form (vs. combination of characters: (1) body size nonpentadactyl), longer forelimb lengths moderate (SVL 78.2–106.2 mm); (2) relative (.11.0 mm vs. ,6.9 mm), and greater hind tail length long; (3) pentadactyl; (3) Finger-III limb lengths (.17.0 mm vs. ,12.9 mm), and lamellae five or six; (4) Toe-IV lamellae nine by the presence of a postnasal scale or 10; (5) limb length moderate; (6) paraver- (vs. absence). In addition, Brachymeles tun- tebral scale rows 66–68; (7) supralabials seven; gaoi differs from all nonpentadactyl species (8) infralabials six; (9) pineal eye spot present, except B. wrighti by having a midbody scale large; (10) supranasals in contact; (11) pre- row count 26–28 (vs. ,24); from all non- frontals not contacting on midline; (12) pentadactyl species except B. pathfinderi by contact between first pair of chin shields; having a paravertebral scale row count 66 (vs. (13) enlarged chin shields in two pairs; (14) .84), and by the presence of auricular nuchal scales undifferentiated; (15) fifth and openings (vs. absence); and from B. apus, B. sixth supralabial below eye; (16) auricular lukbani, B. minimus, and B. vermis by the opening present; (17) continuous, light dorso- presence of limbs (vs. absence). lateral stripes present, indistinct; (18) contin- Description of holotype.—Mature male, uous, dark middorsal stripes present; (19) dark hemipenes everted (Fig. 10); SVL 89.2 mm; lateral stripes present; and (20) dark ventral body moderate relative to other Brachymeles, pigmentation absent (Tables 4 and 5). elongate with respect to other lizards; head Comparisons.—Characters distinguishing weakly differentiated from neck, nearly as the new species from all pentadactyl species wide as body, HW 11.3% SVL, 115.9% HL; of Brachymeles are summarized in Tables 4 HL 38.0% SnFa; SnFa 25.6% SVL; snout and 5. Brachymeles tungaoi most closely moderately long, bluntly rounded in dorsal resembles B. kadwa, B. makusog, B. talinis, profile, sharply rounded in lateral profile, and B. vindumi, but it differs from these four SNL 60.1% HL; auricular opening present, taxa by having a smaller body size, smaller moderate; eyes small, ED 1.9% SVL, 19.2% midbody width, greater relative tail length, the HL, 50.0% END, pupil nearly round; body first and second pairs of enlarged chin shields slightly depressed, MBW 162.0% MBH; body equal in width, and contact between the first scales smooth, glossy, imbricate; longitudinal pair of enlarged chin shields (Tables 4 and 5). scale rows at midbody 28; paravertebral scale Brachymeles tungaoi can be further distin- rows 66; axilla–groin scale rows 46; limbs well guished from B. kadwa by having nine or 10 developed, pentadactyl, digits moderate; Fin- Toe-IV lamellae, paravertebral scale rows 66– IIIlam 6; ToeIVlam 10; FLL 22.5% AGD, 68, frontoparietal in contact, and by the 14.4% SVL; HLL 35.4% AGD, 22.6% SVL; absence of dark ventral pigmentation (Ta- order of digits from shortest to longest for bles 4 and 5); from B. makusog by having hand: I 5 V , II 5 IV , III, for foot: I , V , seven supralabials, six infralabials, the fifth II , IV , III; tail not as wide as body, and sixth supralabial below the eye, suprana- gradually tapered toward the end, TW 63.7% sals in moderate contact, the presence of MBW, TL 99.9% SVL. 2010] HERPETOLOGICAL MONOGRAPHS 45

Rostral projecting dorsoposteriorly to point tudinal rows of dark brown spots, extending in line with anterior edge of nasal, broader from posterior edge of parietals to base of tail; than high, moderately separated from fronto- spot rows span six full and two half rows of nasal; frontonasal wider than long; nostril scales at midbody, narrowing to four full and ovoid, centered in a single rectangular nasal; two half rows of scales posterior to parietals; supranasals present, large, in narrow medial pigmentation covering middle one third of contact; postnasals present; prefrontals broad- dorsal scales; dorsolateral stripes indistinct, ly separated by frontal; frontal nearly octago- discontinuous, spanning one whole and two nal, its anterior margin in broad contact with half row of scales from auricular opening to frontonasal, in contact with first two anterior midbody. Lateral and ventral surface of body supraoculars, 53 wider than anteriormost light brown. Lateral surface with three supraocular; supraoculars five; frontoparietals discontinuous rows of dark brown spots, moderate, in broad contact medially, each spanning posterior two thirds of axilla–groin frontoparietal in contact with supraoculars distance. Ventral surface without dark pig- 2–4; interparietal moderate, quadrilaterally mentation. Tail with dark dorsal blotches and shaped, its length nearly equal to midline spots; dark pigment reduced ventrally. Head length of frontoparietal; distinct parietal scales homogeneous dark brown; rostral, eyespot present, large, in posterior half of nasal, postnasal, supranasal, first supralabial, scale; parietals broadly separated by interpa- mental, and first infralabial light gray; pineal rietal; nuchals undifferentiated; loreals two, eyespot large distinct, light cream. Limbs decreasing in size from anterior to posterior, mottled medium to dark brown dorsally, subequal, in contact with prefrontal, postna- yellowish brown ventrally; dorsal and ventral sal, supranasal, second supralabial, posterior surface of digits dark brown. loreal and frontonasal; preocular single, nearly Coloration of holotype in life.—Coloration one half as high as posterior loreal; single in life is unrecorded; however, because presubocular; supraciliaries six, the anterior- Brachymeles specimens do not change signif- most contacting prefrontal and separating icantly during preservation (CDS and RMB, posterior loreal from first supraocular, poste- personal observations), we suspect that the riormost extending to midline of last suprao- preserved coloration and patterns are much cular; subocular row complete; lower eyelid like those in life. with one row of scales, lacking an enlarged Measurements of holotype (in millime- oval window, largely transparent; supralabials ters).—SVL 89.2; AGD 56.8; TotL 178.3; seven, fifth and sixth below the eye; infra- MBW 13.9; MBH 8.6; TL 89.1; TW 8.8; TH labials six. 8.0; HL 8.7; HW 10.1; HH 7.2; SnFa 22.8; ED Mental wider than long, in contact with first 1.7; END 3.3; SNL 5.2; IND 3.0; FLL 12.8; infralabials; single enlarged postmental, wider HLL 20.1; MBSR 28; PVSR 66; AGSR 46; than mental; followed by two pairs of enlarged FinIIIlam 6; ToeIVlam 10; SL 7; IFL 6; SC 6; chin shields; first pair in broad medial contact, SO 5. second pair separated by single undifferenti- Variation.—Morphometric variation of the ated scale. series is summarized in Table 6. Specimens Scales on limbs smaller than body scales; were observed to have parietals moderately scales on dorsal surfaces of digits large, separated by the interparietal (CAS 144229– wrapping around lateral edges of digits; 30, 144341, KU 323933, 323935–6) or in point lamellae undivided; palmar surfaces of hands medial contact (CAS 144313, KU 323934). and plantar surfaces of feet covered by small, Scale counts were observed to vary among irregular scales, each with raised anterior the measured series. Specimens were ob- edges; scales on dorsal surface of hands and served to have midbody scale row counts of 26 feet smaller than limb scales, lacking raised (KU 323935), 27 (CAS 144313), and 28 (CAS edges. 144229–30, 144341, KU 323933–4, 323936); Coloration of holotype in preservative.— axilla–groin scale row counts of 46 (KU Ground color of body medium brown; dorsal 323933, 323935–6), 47 (CAS 144229–30, surface of body with eight continuous, longi- 144341, KU 323934), and 49 (CAS 144313); 46 HERPETOLOGICAL MONOGRAPHS [No. 24 and paravertebral scale row counts of 66 (KU 60723), collected over the same dates and in 323933–6), 67 (CAS 144229), and 68 (CAS the same locality as holotype. 144230, 144313, 144341). Diagnosis.—Brachymeles vindumi can be We also observed lamellae counts to vary distinguished from congeners by the following among the measured series. Specimens were combination of characters: (1) body size observed to have Finger-III lamellae counts of moderate (SVL 104.9–113.6 mm); (2) penta- five (CAS 144229–30, 144341, KU 323934–5) dactyl; (3) Finger-III lamellae six; (4) Toe-IV or six (CAS 144313, KU 323933, 323936); lamellae nine or 10; (5) moderate limb length; Toe-IV lamellae counts of nine (CAS 144229, (6) midbody scale rows 30 or 31; (7) axilla– 144313, 144341, KU 323935–6) or 10 (CAS groin scale rows 49; (8) paravertebral scale 144230, KU 323933–4). rows 74; (9) supralabials seven; (10) infra- Distribution.—Brachymeles tungaoi is labials six; (11) pineal eye spot present, known only from Masbate Island (Fig. 5). indistinct; (12) supranasals in contact; (13) Ecology and natural history.—Brachymeles prefrontals separate; (14) parietal in contact; tungaoi occurs in agricultural areas as well as (15) enlarged chin shields in two pairs; (16) disturbed and secondary growth forest habi- first pair of chin shields separated; (17) tat. Little or no original, low-elevation forest nuchals undifferentiated; (18) fifth and sixth remains on Masbate Island, but we assume supralabials below the eye; (19) auricular the species once also occurred in primary opening present; (20) continuous, light dorso- forest. Individuals were collected in the lateral stripes present, distinct; (21) dark humus material within the rotting stumps of lateral stripes present; and (22) dark ventral trees. When disturbed, individuals immedi- pigmentation present (Tables 4 and 5). ately moved in a rapid serpentine manner and Comparisons.—Characters distinguishing attempted to burrow back into loose soil or the new species from all pentadactyl species humus. of Brachymeles are summarized in Tables 4 Sympatric lizard species observed on Mas- and 5. Brachymeles vindumi most closely bate Island include the following: (Agamidae) resembles B. kadwa, B. talinis,andB. tungaoi, Bronchocela cristatella, Draco spilopterus, but it differs from these three taxa by having Gonocephalus sophiae; (Gekkonidae) Cyrto- six Finger-III lamellae, six supralabials, mid- dactylus philippinicus, Gehyra mutilata, body scale rows 30 or 31, axilla–groin scale Gekko gecko, Hemidactylus frenatus, H. pla- rows 49, paravertebral scale rows 74, the first tyurus; (Scincidae) Brachymeles bonitae, pair of enlarged chin shields separated, and Emoia atrocostata, E. multicarinata, E. multi- the presence of continuous, light dorsolateral fasciata, Lamprolepis smaragdina, Lipinia stripes (Tables 4 and 5). Brachymeles vindumi pulchella, Sphenomorphus decipiens, S. ja- can further be distinguished from B. kadwa gori; and (Varanidae) Varanus marmoratus. and B. talinis by having nine or 10 Toe-IV Etymology.—We take pleasure in naming lamellae (Tables 4 and 5); from B. talinis by the new species after our friend and dedicated having six infralabials (Table 5); from B. field collaborator Jason B. ‘‘Tungao’’ Fernan- kadwa by contact between frontoparietals dez, with thanks for years of hard work toward (Table 5); and from B. kadwa and B. tungaoi the research of semifossorial lizards. Suggest- by contact between parietals (Table 5). ed common name: Tungao’s slender skink. From all nonpentadactyl species of Brachy- meles (B. apus, B. bonitae, B. cebuensis, B. Brachymeles vindumi sp. nov. elerae, B. lukbani, B. minimus, B. muntingka- Figs. 4, 10 may, B. pathfinderi, B. samarensis, B. tridac- Holotype.—CAS 60724 (EHT Field No. tylus, B. vermis, and B. wrighti), B. vindumi 1718), adult male, collected between 25 Octo- differs by having a pentadactyl body form (vs. ber and 17 November 1920, in Sulu Province, nonpentadactyl), longer forelimb lengths Jolo Island, Philippines, by Edward H. Taylor. (.13.2 mm vs. ,6.9 mm), greater hind limb Paratypes.—One adult female (CAS lengths (.22.7 mm vs. ,12.9 mm), and 60725), one juvenile female (MCZ 26577), greater number of midbody scale rows (30 and one juvenile of unknown sex (CAS or 31 vs. ,28), and by the presence of a 2010] HERPETOLOGICAL MONOGRAPHS 47 postnasal scale (vs. absence). In addition, B. posterior, subequal, in contact with prefrontal, vindumi differs from all nonpentadactyl spe- postnasal, supranasal, second supralabial, pos- cies except B. pathfinderi by having a terior loreal and frontonasal; preocular single, paravertebral scale row count 74 (vs. .84) nearly two thirds as high as posterior loreal; and by the presence of auricular openings (vs. single presubocular; supraciliaries six, the absence); and from B. apus, B. lukbani, B. anteriormost contacting prefrontal and sepa- minimus, and B. vermis by the presence of rating posterior loreal from first supraocular, limbs (vs. absence). posteriormost extending to midline of last Description of holotype.—Mature male, supraocular; single subocular row complete; hemipenes not everted (Fig. 9); SVL lower eyelid with one row of scales, lacking an 113.6 mm; body moderate relative to other enlarged oval window, largely transparent; Brachymeles, elongate with respect to other supralabials seven, fifth and sixth below the lizards; head weakly differentiated from neck, eye; infralabials six. nearly as wide as body, HW 9.5% SVL, Mental wider than long, in contact with first 112.5% HL; HL 33.7% SnFa; SnFa 25.1% infralabials; single enlarged postmental, slight- SVL; snout moderately long, rounded in ly wider than mental; followed by two pairs of dorsal and lateral profile, SNL 59.7% HL; enlarged chin shields, scales of first pair auricular opening present, moderate; eyes separated by a single undifferentiated scale, small, ED 2.0% SVL, 23.5% HL, 58.0% second pair separated by three undifferenti- END, pupil nearly round; body slightly ated scales. depressed, MBW 122.9% MBH; body scales Scales on limbs smaller than body scales; smooth, glossy, imbricate; longitudinal scale scales on dorsal surfaces of digits large, rows at midbody 31; paravertebral scale rows wrapping around lateral edges of digits; 74; axilla–groin scale rows 49; limbs well lamellae undivided; palmar surfaces of hands developed, pentadactyl, digits moderate; Fi- and plantar surfaces of feet covered by small, nIIIlam 6; ToeIVlam 9; FLL 18.1% AGD, irregular scales, each with raised anterior 11.6% SVL; HLL 31.2% AGD, 20.0% SVL; edges; scales on dorsal surface of hands and order of digits from shortest to longest for feet smaller than limb scales, lacking raised hand: I , V , II , IV , III, for foot: I 5 V , edges. II 5 III , IV; tail regenerated, not as wide as Coloration of holotype in preservative.— body, sharply tapered toward the end, TW Ground color of body medium to dark brown; 83.1% MBW. middorsal surface of body covered with dark Rostral projecting dorsoposteriorly to point pigmentation, extending from posterior edge in line with anterior edge of nasal, broader of supranasals to base of tail, made of eight than high, moderately separated from fronto- irregular, longitudinal middorsal rows of dark nasal by supranasal contact; frontonasal wider brown spots, spanning six full and two half than long; nostril ovoid, centered in a single rows of scales at midbody, narrowing to six full rectangular nasal; supranasals present, large, rows of scales posterior to parietals, pigmen- in moderate medial contact; postnasals pres- tation covering middle one third of dorsal ent; prefrontals narrowly separated by frontal; scales; dorsolateral stripes present, clearly frontal nearly octagonal, its anterior margin in defined, continuous, lacking dark pigmenta- narrow contact with frontonasal, in contact tion, spanning one whole and one half row of with first two anterior supraoculars, 43 wider scales from anterior-most supraocular to base than anteriormost supraocular; supraoculars of tail. Lateral and ventral surface of body five; frontoparietals moderate, in broad medi- medium to dark brown. Lateral surface with al contact, each frontoparietal in contact with six to eight irregular dark spot rows, gradually supraoculars 2–4; interparietal small, diamond becoming fainter on ventral surface. Ventral shaped, its length equal in size to midline surface with irregular dark spots and blotches. length of frontoparietal; parietal eyespot Tail with continuous dark blotches and spots absent; parietals in broad contact behind dorsally, dark pigment reduced ventrally. interparietal; nuchals undifferentiated; loreals Head scales homogeneous mottled medium two, decreasing in size from anterior to and dark brown dorsally; rostral, nasal, 48 HERPETOLOGICAL MONOGRAPHS [No. 24 postnasal, supranasal, first supralabial, mental, colleague Jens Vindum. The specific epithet is and first infralabial light brown to tan; pineal a patronym in the genitive singular, chosen in eyespot absent; dark brown blotch of pigmen- thanks for the many years of support and tation on lateral surfaces of head, spanning assistance he has provided during our re- from posterior edge of eye to posterior edge of search on Philippine amphibians and reptiles. auricular openings. Limbs mottled medium to Suggested common name: Jens’ slender skink. dark brown; dorsal surface of digits dark brown, ventral surface of digits medium DISCUSSION brown. Phylogenetic analyses of the mitochondrial Coloration of holotype in life.—Coloration ATP8 and ATP6 genes resulted in strong in life is unrecorded; however, because support for nine lineages of Brachymeles Brachymeles specimens do not change signif- (Fig. 2). The phylogeny, combined with mor- icantly during preservation (CDS and RMB, phological data, supports the elevation of all personal observations), we suspect that the subspecies of the polytypic species B. boulen- preserved coloration and patterns are much geri and B. schadenbergi to full species. like those in life. However, the inferred relationships between Measurements of holotype (in millime- several of the species sampled are weakly ters).—SVL 113.6; AGD 72.7; TotL N/A; supported. This may be indicative of rapid MBW 14.2; MBH 11.6; TL N/A; TW 11.8; diversification of Brachymeles or simply indi- TH 8.4; HL 9.6; HW 10.8; HH 7.9; SnFa 28.5; cate a lack of character support at some ED 2.3; END 3.9; SNL 5.7; IND 3.4; FLL internal nodes. Given the use of only mito- 13.2; HLL 22.7; MBSR 31; PVSR 74; AGSR chondrial data for our phylogenetic analyses, 49; FinIIIlam 6; ToeIVlam 9; SL 7; IFL 6; SC caution must be taken when interpreting 6; SO 5. interspecies relationships, because a single Variation.—Morphometric variation of the locus can be subject to random variation, deep series is summarized in Table 6. Specimens coalescence, lineage sorting, and natural were observed to have midbody scale row selection (Brown et al., 2010; Edwards and counts of 30 (CAS 60725) or 31 (CAS 60724), Beerli, 2002; Galtier et al., 2009). Regardless and Toe-IV lamellae counts of nine (CAS of the potential weaknesses of our single-locus 60724) or 10 (CAS 60725). approach, our results are strongly supported Distribution.—Brachymeles vindumi is by an independent, comprehensive dataset of known only from Jolo Island (Fig. 4). morphological characters. Ecology and natural history.—Brachymeles No analyses supported the monophyly of vindumi presumably occurs in disturbed species formerly part of Brachymeles boulen- habitat as well as secondary growth forest on geri (B. boholensis, B. boulengeri, B. mind- Jolo Island. Due to security concerns, no orensis, and B. taylori; Fig. 2). Another clade recent surveys have been conducted on Jolo including B. talinis, B. kadwa, and B. tungaoi Island; therefore, no information is available was estimated with strong support (Fig. 2). As on the ecology of this species. previously recognized, B. talinis spanned two Sympatric lizard species observed on Jolo distinct, recognized faunal regions (Luzon and Island include the following: (Agamidae) Visayas). Given this formerly wide geograph- Draco guentheri; (Gekkonidae) Cyrtodactylus ical distribution, it is not surprising that the annulatus, Gehyra mutilata, Gekko gecko, G. northern populations (Luzon and the Babuyan mindorensis, Hemidactylus frenatus, H. pla- islands) constitute a genetically distinct line- tyurus, Luperosaurus joloensis; (Scincidae) age that we describe here as B. kadwa.We Brachymeles vermis, Eutropis multifasciata, were surprised, however, to discover an E. rudis, Lamprolepis smaragdina, Lipinia additional genetically distinct lineage on quadrivittata, Lygosoma bowringi, Spheno- Masbate Island (Fig. 2). The fauna of Masbate morphus biparietalis, S. variegatus; and (Var- Island is recognized as part of the Visayan or anidae) Varanus cumingi. central Philippine islands and has been Etymology.—We take pleasure in naming hypothesized to have shared land bridge the new species for our close friend and connections with the central islands during 2010] HERPETOLOGICAL MONOGRAPHS 49 periods of glacial maxima (Dickerson, 1928; loose soil, or leaf litter, whereas others are Heaney, 1985; Inger, 1954; Voris, 2000). common beneath piles of rotting coconut Although we expected Masbate populations husks in disturbed, agricultural habitat. The to be more closely related to Visayan (Negros species now found in residential and agricul- + Panay) populations, all analyses strongly tural areas were once native to forested supported the sister relationship between B. habitats. Before recent, focused survey efforts, tunagoi (Masbate) and B. kadwa (Luzon), the relatively low numbers of specimens of providing additional biogeographic support Brachymeles in museum collections handi- for the distinctiveness of B. tungaoi. We are capped our efforts at delimiting species. The unaware of phylogeographic or phylogenetic rarity of Brachymeles in collections was due to studies including other vertebrate taxa from their secretive, semifossorial lifestyle. Masbate. Comparison of the systematic affin- This is the first, species-level phylogenetic ities of other Masbate species may provide study of Brachymeles. To date, taxonomic interesting exceptions to the prevailing PAIC- reviews of Brachymeles have focused solely on oriented perspective of Masbate as a faunistic morphological variation (Brown, 1956; Brown extension of the central Visayas (Heaney, and Alcala, 1980; Brown and Alcala, 1995; 1985). Brown and Rabor, 1967; Hikida, 1982). It is The species recognized in this paper apparent that species diversity in the genus increase the total number of known species has been considerably underestimated; ac- of Brachymeles to 25, all but one of which are cordingly, discovery of additional undocu- endemic to the Philippines. During the past mented (possibly cryptic) diversity is antici- 2 yr, our knowledge of the species diversity in pated in other species groups (e.g., Siler et al., the genus has expanded rapidly as the result of 2009, 2010a,b). Several studies have shown large-scale sampling efforts across the Philip- that the evolution of a burrowing lifestyle is pines and the detailed analyses of morpho- correlated with decreasing dispersal abilities logical variation among species and popula- (Nevo, 1979; Patton and Feder, 1978; Patton tions (Siler et al., 2009, 2010a,b). Before this and Yang, 1977; Selander et al., 1974; Wiens effort, estimates of Brachymeles species di- et al., 2006). Many Brachymeles lineages have versity remained nearly constant for .30 yr experienced reduction or loss of limbs, which (but see Brown and Alcala, 1995), which is a may further reduce vagility (Daniels et al., testament to the extent of morphological 2005; Mulvaney et al., 2005; Wiens et al., similarity among species within the genus 2006). Through time, reduced dispersal abil- and a lack of systematic studies of the group. ities may lead to increasingly patchy distribu- It comes as little surprise that allopatric tions, reduced gene flow between populations, populations of ‘‘B. boulengeri’’ from the and the accumulation of interpopulation Luzon, Mindanao, Mindoro, and Visayan differences (Nevo, 1979). However, the role PAICs have proven to be morphologically that geological history and complex geography diagnosable with increased sampling. To date, play on the dispersal abilities and diversifica- few studies have provided evidence of truly tion patterns of Brachymeles species remains ‘‘widespread’’ reptile species that have geo- unknown. Regardless of what processes pro- graphic distributions spanning recognized duce species diversity, we expect that addi- zoogeographic boundaries in the Philippines tional species await discovery. With several (but see Siler et al., 2010c) and as is quite species represented by only a few vouchered often the case, these species frequently turn specimens, and frequent morphological con- out to constitute multiple evolutionary lineag- vergence, it is clear that a comprehensive es (Brown and Diesmos, 2002; Brown et al., phylogenetic analysis of the genus will be 2009; Gaulke et al., 2007; McGuire and required to assess, with accuracy, the species Alcala, 2000; Welton et al., 2009, 2010). diversity within Brachymeles. All species of Brachymeles have a semi- Following the recognition of Brachymeles fossorial life style, specializing in dry rotting boholensis, B. boulengeri, B. tungaoi, B. material within rotten logs. Many are habitat kadwa, B. mindorensis, B. orientalis, B. specialists found exclusively in rotting logs, schadenbergi, B. taylori, and B. vindumi, 50 HERPETOLOGICAL MONOGRAPHS [No. 24 there are now 13 pentadactyl species of that continued efforts be made to conduct Brachymeles. Of these species, eight are surveys focused on rotting log and leaf litter large-bodied (B. bicolor, B. tungaoi, B. kadwa, microhabitats throughout the ranges of all B. makusog, B. orientalis, B. schadenbergi, B. species. Accurate data on the distributions talinis, and B. vindumi) and five (B. boholen- of these species will allow for a complete sis, B. boulengeri, B. gracilis, B. mindorensis, assessment of the geographic ranges of the and B. taylori) have moderately sized bodies. species and appropriate decision of conserva- The distribution of pentadactyl species in the tion status and actions can be made. At Philippines is relatively even across the major present, all 10 species are known or believed recognized faunal regions, with four species to be common throughout their ranges. known to occur in the Luzon Faunal Region, Although these species currently inhabit five in the Mindanao Faunal Region, three in highly disturbed, agricultural and residential the Visayan Faunal Region, one in the Mind- areas, no studies on the long-term effect of oro Faunal Region, and one in the Sulu deforestation on populations of Brachymeles archipelago (Brown and Alcala, 1980, Brown exist. Therefore, according to the Internation- and Alcala, 1995; Brown and Diesmos, 2002; al Union for Conservation of Nature catego- Siler et al., 2010a). In contrast, the distribu- ries and classification structure, we consider tion of total species diversity in the genus is the conservation status of these species as less uniform, with 11 species known from the ‘‘least concern,’’ pending the collection of Luzon Faunal Region versus six in the Mind- additional information that might suggest anao Faunal Region, six in the Visayan Faunal otherwise. Region, and only one and two in the Sulu Acknowledgments.—We thank the Protected Areas and archipelago and Mindoro Faunal Region, Wildlife Bureau of the Philippine Department of respectively (Brown and Alcala, 1980; Brown Environment and Natural Resources for facilitating and Alcala, 1995; Brown and Diesmos, 2002; collecting and export permits necessary for this and Siler et al., 2009, 2010a,b). New species related studies; we are particularly grateful to M. Lim, C. Custodio, and A. Tagtag. Financial support for fieldwork discoveries on Luzon Island have occurred for CDS was provided by a Panorama Fund grant from with consistency during the last two decades; The University of Kansas Biodiversity Institute, a Madison given the island’s complex mountain ranges and Lila Self Fellowship from the University of Kansas, a (Sierra Madres, Cordillera, Zambales, Bicol Fulbright-Hayes Fellowship, and a grant from the National Science Foundaton (NSF; DEB 0804115). Peninsula volcanoes) and geographic com- Financial support for RMB and ACD was provided by plexity (Defant et al., 1989; Yumul et al., NSF (EF-0334952 and DEB 0743491) funds to RMB. 2009), the increase in the region’s faunal For the loans of specimens, we thank J. Vindum and A. diversity is likely to continue (Brown et. al. Leviton (CAS), R. Sison and A. C. Diesmos (Philippine 1995a,b, 1999, 2000a,b, 2007; Ross and National Museum), J. Ferner (Cincinnati Museum of Natural History), A. Resetar and H. Voris (Field Museum Gonzales, 1992; Siler et al., 2009, 2010a,b). of Natural History), R. Crombie (United States Natural It is worth noting that efforts to survey History Museum), and T. LaDuc (Texas Memorial Mindanao have been less extensive than Natural History Museum). Critical reviews of the efforts on Luzon; this may account for some manuscript were provided by L. Trueb, D. McLeod, of the differences in diversity between the and J. Esselstyn, and C. Oliveros assisted with the Tagalog abstract. CDS thanks the CAS’s Stearns Fellowship and regions, which may be artifacts of sampling the Museum of Comparative Zoology’s Ernst Mayr biases. Fellowship for funding recent visits to examine compar- At present, there remains one polytypic ative material. CDS and RMB extend a special thanks to species (B. gracilis) and two ‘‘widespread’’ Arvin and Mae Diesmos for continued support. species (B. bonitae and B. samarensis), all with distributions spanning boundaries between LITERATURE CITED recognized faunal regions (Brown and Alcala, BOETTGER, O. 1886. Aufzahlung der von Philippinen 1980). Closer investigation of island popula- bekannten Reptilien und Batrachier. Bericht der tions within each of these species may result Senckenbergischen Naturforschenden Gesellschaft fur 1886, Frankfurt am Main. Germany. in the discovery of new diversity in the genus. BOETTGER, O. 1893. Katalog der Reptilien. Sammlung im As our understanding of the total diversity Museum de Senckenbergischer Naturforschenden Ge- within Brachymeles increases, it is important sellschaft in Frankfurt am Main, Frankfurt, Germany. 2010] HERPETOLOGICAL MONOGRAPHS 51

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M. bonitae.—(13) MASBATE ISLAND: MASBATE PROVINCE: BROWN. 2009. A new bent-toed Gekko (Genus Cyrto- Municipality of Mobo: Tugbo Barrio: CAS 144223; dactylus) from southern Palawan Island, Philippines Mapuyo Barrio: Palangkahoy: CAS 144270; MINDORO and clarification of the taxonomic status of C. ISLAND: MINDORO ORIENTAL PROVINCE: Mt. Halcon: SE annulatus. Herpetologica 65:328–343. slope Barawanan Peak: CAS-SU 25713, 25793, 25886–88, WELTON, L. J., C. D. SILER,A.C.DIESMOS, AND R. M. 25904; Sumagui: CAS 62064 (paratype); POLILLO BROWN. 2010. Phylogeny-based species delimitation of ISLAND: QUEZON PROVINCE: Municipality of Polillo: southern Philippines bent-toed geckos and a new Barangay Pinaglubayan: KU 307747–49, 307755. Brachy- species of Cyrtodactylus (Squamata; Gekkonidae) from meles boulengeri.—(26) LUZON ISLAND: AURORA western Mindanao and the Sulu Archipelago. Zootaxa PROVINCE: Municipality of Baler: KU 322314–20; LUZON 2390:49–68. ISLAND: LAGUNA PROVINCE: Municipality of Los Banos, WIENS, J. J., AND J. L. SLINGLUFF. 2001. How lizards turn Barangay Batong Malake: KU 32058–60; Municipality of into snakes: A phylogenetic analysis of body-form Los Banos: CAS 61096; Mt. Maquiling: CAS 61297; evolution in anguid lizards. Evolution 55:2303–2318. POLILLO ISLAND: QUEZON PROVINCE: Municipality of WIENS, J. J., M. C. BRANDLEY, AND T. W. REEDER. 2006. Polillo: CAS (paratypes) 62272–73, 62276–77; Barangay Why does a trait evolve multiple times within a clade? Pinaglubayan: KU 307438–9, 307750–54, 307756 (neo- Repeated evolution of snake-like body form in squa- type), 307757–58. Brachymeles cebuensis.—(8) CEBU mate reptiles. Evolution 60:123–141. ISLAND: 40 km SW of Cebu City: Tapal Barrio, Sitio WILCOX, T. P., D. J. ZWICKL,T.A.HEATH, AND D. M. Mantalungon: CAS-SU (holotype) 24400, (paratypes) HILLIS. 2002. Phylogenetic relationships the dwarf boas 24396–97, 24399, 24401, 24403; Municipality of Carcar: and a comparison of Bayesian and bootstrap measures Tapal Barrio: CAS 102405 (paratype); 3 km NW Cebu of phylogenetic support. Molecular Phylogenetics and City, Buhisan Barrio, Buhisan Reforestation Project: Evolution 25:361–371. CAS-SU 27537. Brachymeles elerae.—(4) LUZON WILEY, E. O. 1978. The evolutionary species concept ISLAND: KALINGA PROVINCE: Municipality of Balbalan: reconsidered. Systematic Zoology 21:17–26. CAS 61499–500, PNM 9563–4. Brachymeles gracilis YUMUL, G. P., JR., C. B. DIMALANTA,K.QUEAN˜ O, AND E. gracilis.—(18) MINDANAO ISLAND: DAVAO DEL SUR MARQUEZ. 2009. Philippines, geology. Pp. 732–738. In PROVINCE: Municipality of Malalag: Sitio Kibawalan: CAS- R. Gillespie and D. Calgue (Eds.), Encyclopedia of SU 24163, 24165, CAS 124811, 139307–09; Davao City: Islands. University of California Press, Berkeley, Buhangin, Kabanti-an: CAS 124803–04, 139293–95, California, USA. 139303–05; Digos City: Tres de Mayo Barrio: CAS 124806–08, 139300. Brachymeles gracilis hilong.— (20) MINDANAO ISLAND: AGUSAN DEL NORTE PROV- APPENDIX INCE: Municipality of Cabadbaran: Diuata Mountain Range: Mt. Hilonghilong: Balangbalang: CAS-SU (holo- type) 24407, paratype) 102406, 133578, CAS-SU 24411, Additional Specimens Examined 133577, 133581–82, 133609, 133612, 133692–93, 133703– Numbers in parentheses indicate the number of 06, 133743, 133745–47; SURIGAO DEL SUR PROVINCE: specimens examined. With the exception of B. apus, all Municipality of Lanuza: Diuata Mountain Range: Sibuhay specimens examined are from the Philippines. Several Barrio: CAS-SU (paratype) 24315. Brachymeles luk- sample sizes are greater than those observed in the bani.—(14) LUZON ISLAND: CAMARINES NORTE PROV- description due to the examination of subadult specimens INCE: Municipality of Labo: Barangay Tulay Na Lupa, Mt. that were excluded from morphometric analyses. Bra- Labo: PNM (holotype) 9567, (paratopotypes) 9589–92, chymeles apus.—(1) BORNEO: MALAYSIA: Sabah: Mt. KU (paratopotypes) 313597–99, 313601, 313603–04, Kinabalu National Park, Sayap Substation: SP 06915. 313606, 313608, FMNH (paratopotype) 270191. Brachy- Brachymeles bicolor.—(24) LUZON ISLAND: Aurora meles makusog.—(17) CATANDUANES ISLAND: CAT- Province: Municipality of Maria Aurora: Barangay Villa ANDUANES PROVINCE: Municipality of Gigmoto: Barangay Aurora, Sitio Dimani, Aurora Memorial National Park: San Pedro, Sitio Tungaw: PNM (holotype) 9565, (para- KU 323149–52; CAGAYAN PROVINCE: Municipality of topotypes) 9583–9584, KU (paratopotypes) 308126, Baggao: Sitio Hot Springs: CAS 186111, USNM 140847, 308128, 308136, 308208; LUZON ISLAND: CAMARINES 498829–30, 498833; Isabela Province, Sierra Madres NORTE PROVINCE: Municipality of Labo, Barangay Tulay Mountain Range: KU 324097–99, PNM 5785, 9568–77; Na Lupa, Mt. Labo: KU (paratypes) 313612–313614, KALINGA PROVINCE: Balbalasang-Balbalan National Park: 313616, 313617, PNM (paratypes) 9585–9588, FMNH FMNH 259438. Brachymeles boholensis.—(19) BO- (paratype) 270200. Brachymeles mindorensis.—(34) HOL ISLAND: BOHOL PROVINCE: Municipality of Sierra MINDORO ISLAND: MINDORO OCCIDENTAL PROVINCE: Bullones, Barangay Danicop: KU 323944, 323948–9, KU 304351–5, 304412–3, 304488, 307739–42, 308404, 54 HERPETOLOGICAL MONOGRAPHS [No. 24

308447–8, 308534; MINDORO ISLAND: MINDORO PROVINCE: Municipality of Tatayan: MCZ 26553, 26555– ORIENTAL PROVINCE: 30 km SE Municipality of Calapan: 8, 26561, 26566, 26568, 26571–2, 26574; ZAMBOANGA DEL Bank of Tarogin River: CAS-SU (holotype) 24487; SE NORTE PROVINCE: Dapitan River: CAS-SU 23494–96; slope Mt. Halcon, Tarogin Barrio: CAS-SU (paratypes) ZAMBOANGA CITY PROVINCE: Municipality of Pasonanca: 24549–54, 24561–62, 24564; 24566, 24568, 24573–74, Barangay Baluno: Pasonanca Natural Park: KU 314967, 24577–79; Mt. Halcon, SE slope Barawanan Peak: CAS- 314969, 314970–8, 314980, 314984–85, 314988–92, SU (paratype) 24570. Brachymeles minimus.—(6) CAT- 314994, 314996–7. Brachymeles talinis.—(31) NE- ANDUANES ISLAND: CATANDUANES PROVINCE: Munic- GROS ISLAND: NEGROS ORIENTAL PROVINCE: 6kmW ipality of Gigmoto: Barangay San Pedro: KU 308129–31, Municipality of Valencia: Cuernos de Negros Mountain 308210–12. Brachymeles muntingkamay.—(17) LU- Range: ridge on north side of Maite River: CAS-SU ZON ISLAND: NUEVA VIZCAYA PROVINCE: Municipality (holotype) 18358, (paratype) 89813; Cuernos de Negros of Quezon: Barangay Maddiangat, Mt. Palali: PNM Mountain Range: Dayungan Ridge: CAS 133871; Duma- (holotype)9566, (paratopotypes) 9578–82, KU (paratopo- guete City: CAS-SU (paratype) 12225; Municipality of types) 308865–66, 308900–06, 308908, 308953. Brachy- Siaton: 20 km N Bondo Barrio: CAS-SU 22311–12; 22317, AR- meles pathfinderi.—(40) MINDANAO ISLAND: S 22323; INAMPULAGAN ISLAND: GUIMARAS PROVINCE: ANGANI PROVINCE: Municipality of Glan: Barangay Taluya: Municipality of Sibunag: 8 km W Pulupandan Town: CDS 5235–42; Barangay Tanibulad, Sitio Padido: CDS CAS-SU 27972, 27996–97; PANAY ISLAND: ANTIQUE 5192–5206, 5210–20, 5222–27. Brachymeles samaren- PROVINCE: Municipality of San Remigio: KU 306756–60, sis.—(7) SAMAR ISLAND: EASTERN SAMAR PROVINCE: 306762–7, 306769, 306770–6, 306786. Brachymeles Municipality of Taft: Barangay San Rafael: KU 310849– taylori.—(34) NEGROS ISLAND: NEGROS OCCIDENTAL 50, 310852, 311294–6; LEYTE ISLAND: LEYTE PROV- PROVINCE: Municipality of Silay City, Barangay Patag: KU INCE: Municipality of Baybay: Barangay Pilim: Sitio San 324044–56; NEGROS ISLAND: NEGROS ORIENTAL PROV- Vicente: KU 311225. Brachymeles orientalis.—(53) INCE: 3 km W Municipality of Valencia: Cuernos de BOHOL ISLAND: BOHOL PROVINCE: Municipality of Negros Mountain Range: Sitio Lunga: ridge on north side Sierra Bullones: Dusita Barrio: CAS-SU (holotype) 24436, CAS-SU (paratypes) 24428, 24434, 24437, CAS of Maiti River: CAS-SU (holotype) 18615, CAS-SU 21873; (paratype) 102404, CAS-SU 25452; Dusita Barrio: Abac- ridge on south side of Maiti River: CAS-SU (paratype) janan: CAS-SU (paratypes) 24446–51, CAS-SU 25460; 18641, 18656–57, 18748; Cuernos de Negros Mountain Cantaub Barrio: CAS-SU (paratypes) 18702, 24442, Range: CAS-SU (paratype) 18649; top of Dayungan Ridge: CAS-SU 21877, 21880, 21883–84; 24 km NW 24458; CAMIGUIN SUR ISLAND: CAMIGUIN PROVINCE: Municipality of Catarman: Mt. Mambajao: Sitio Sangsan- Bondo Barrio: Bantolinao: CAS-SU 22355–56; CEBU gan: CAS 110976–83; LEYTE ISLAND: Leyte PROVINCE: ISLAND: CEBU PROVINCE: Municipality of Carcar: Tapal Municipality of Baybay: KU 311231–5, 311241; MIND- Barrio: Sitio Mantalongon: CAS 154671, 154673, 154678– ANAO ISLAND: AGUSAN DEL NORTE PROVINCE: Munici- 82, 154686. Brachymeles tridactylus.—(20) NEGROS pality of Cabadbaran: Diuata Mountain Range: Mt. ISLAND: NEGROS OCCIDENTAL PROVINCE: 16 km E Hilonghilong: Kasinganan: CAS-SU 133301, 133616, Municipality of La Castellana: Barrio Cabagna-an: 133749, 133752, 133754; SAMAR ISLAND: Eastern southern slope of Mt. Canlaon: CAS-SU 19424, 19426– Samar PROVINCE: Municipality of Taft: KU 305470, 27, 19429, 19452, 19458; 20 km E Municipality of La 310734–6, 310739, 310942–6, 310949, 310951, 310955. Castellana: Sitio Kalapnagan: CAS-SU 27082–83; NEGROS Brachymeles schadenbergi.—(45) BASILAN ISLAND: ORIENTAL PROVINCE: Hills North and Northwest of BASILAN PROVINCE: Port Holland: Sawmill: CAS 60493; Mayaposi: CAS-SU (holotype) 18354; PANAY ISLAND: MINDANAO ISLAND: MISAMIS OCCIDENTAL PROVINCE: ANTIQUE PROVINCE: Municipality of Culasi: Barangay 2 km NW of Masawan: CAS 23468–69; 4 km NW of Alojipan: KU 307726–36. Brachymeles vermis.—(5) Masawan: CAS 23471; 3 km NW Masawan: south bank of JOLO ISLAND: SULU PROVINCE: CAS-SU (paratype) Dapitan River: CAS 23479–81, 23484–85; COTABATO 62489, CAS-SU 60720–22, 60857.