Herpetological Monographs, 25, 2011, 76–112 E 2011 by The Herpetologists’ League, Inc.

PHYLOGENY-BASED SPECIES DELIMITATION IN PHILIPPINE SLENDER (REPTILIA: : SCINCIDAE) II: TAXONOMIC REVISION OF SAMARENSIS AND DESCRIPTION OF FIVE NEW SPECIES

1,3 1 1 2 CAMERON D. SILER ,ALLISON M. FUITEN ,ROBIN M. JONES ,ANGEL C. ALCALA , 1 AND RAFE M. BROWN 1Biodiversity Institute and Department of Ecology and Evolutionary Biology, University of Kansas, 1345 Jayhawk Boulevard, Lawrence, KS 66045-7593, USA 2SUAKCREM Marine Laboratory, Silliman University, Bantayan, Dumaguete City, Philippines 6200

ABSTRACT: With robust new datasets from morphology and DNA sequences, we review the limbed, nonpentadactyl species of the Brachymeles samarensis complex (now known to include B. cebuensis, B. minimus, and B. lukbani), and describe five new species in this highly limb-reduced, endemic Philippine clade of scincid lizards. For more than four decades, B. samarensis has been recognized as a single ‘‘widespread’’ species. This perception of the species’ peculiar geographic range has persisted as a result of weak sampling and similar gross morphology (body sizes, scale pigmentation) among populations. However, previous authors have noted morphological variation between different island populations, and our new data build on these observations and extend them to delimit new proposed species boundaries. Our data indicate that the ‘‘widespread’’ species B. samarensis is actually a complex of six distinct lineages, some of which are not each others’ closest relatives, and each of which is genetically unique. The taxa we define possess allopatric geographic ranges and differ from their congeners by numerous diagnostic characters of external morphology, and therefore should be recognized as full species in accordance with any lineage-based species concept. Species diversity in the genus has doubled in the last 3 yr, with these six taxa increasing the total known number of species of Brachymeles to 30. Key words: Biodiversity; Endemism; Faunal region; Fossorial lizards; Limb reduction; Species delimitation;

FEW GENERA of scincid lizards are known to B. tridactylus, and B. wrighti), and five are possess species representing a full spectrum of entirely limbless (B. apus, B. minimus, B. body forms, from fully limbed, pentadactyl miriamae, B. lukbani, and B. vermis). species to limbless species (see Siler et al., Among the nonpentadactyl species, numer- 2011b; Siler and Brown, 2011). In the genus ous studies have documented a wide range of Brachymeles, all but two of the 26 recognized limb- and digit-reduced states, from minute species are endemic to the Philippines, with limbs with 0–3 digits (B. bonitae, B. cebuensis, the exceptions being a single species (B. apus) B. muntingkamay, B. samarensis, B. tridacty- from northern Borneo and another (B. lus), to moderately developed limbs with miriamae) from Thailand (Brown and Alcala, four to five digits on the hands and feet (B. 1980; Hikida, 1982; Siler, 2010; Siler et al., elerae, B. pathfinderi, B. wright;Dume´ril and 2009, 2010a,b, 2011a,b,c; Siler and Brown, Bibron, 1839; Brown, 1956; Brown and Rabor, 2010, 2011). Thirteen species are pentadactyl 1967; Brown and Alcala, 1980; Taylor, 1917, (B. bicolor, B. boholensis, B. boulengeri, B. 1918, 1925; Siler, 2010; Siler et al., 2009, gracilis, B. kadwa, B. makusog, B. mind- 2010a, 2011a,b,c; Siler and Brown, 2011). All orensis, B. orientalis, B. schadenbergi, B. species are semifossorial and typically found talinis, B. taylori, B. tungaoi, and B. vindumi), in dry, rotting material inside or underneath eight are nonpentadactyl with incompletely decaying logs or in loose soil, forest floor developed limbs and reduced numbers of detritus, and leaf litter. digits (B. bonitae, B. cebuensis, B. elerae, B. Although the genus was named well over muntingkamay, B. pathfinderi, B. samarensis, 150 yr ago (Dume´ril and Bibron, 1839), the rate of Brachymeles species descriptions 3 CORRESPONDENCE: e-mail, [email protected] reached an apparent asymptotic maximum in

76 2011] HERPETOLOGICAL MONOGRAPHS 77

1980 (Brown and Alcala, 1980). The one diversity in the genus for 30 yr, until Taylor exception is B. minimus, a legless species published a series of herpetofaunal descriptions described in 1995 (Brown and Alcala, 1995). in the early 1900s (Taylor, 1917, 1918, 1922, For more than a century, limited numbers of 1925). It would be 50 yr before Brown (1956) specimens in museum collections, combined described B. samarensis from a single juvenile with the similar body plans and external specimen (FMNH 44472) collected in Guiuan, morphological features among species of Samar Island, Philippines in 1945. At the time of Brachymeles, limited assessments of species- description, Brown (1956) hypothesized the level diversity (Brown, 1956; Brown and species was most closely related to B. elerae Alcala, 1980; Brown and Rabor, 1967; Taylor, due to similarities in the number of paraverte- 1917). Recent studies have revealed the bral scale rows. This single juvenile would species-level diversity within Brachymeles to remain the only vouchered specimen of this be drastically underestimated, and have iden- unique bidactyl species from Samar Island (Siler tified numerous nonmonophyletic species and Brown, 2010). complexes within the Philippines (Siler and By the time Brown and Rabor (1967) Brown, 2010, 2011; Siler et al., 2011a). revised the genus Brachymeles, samples of Additionally, several rare, mid-to-high eleva- specimens morphologically similar to B. sa- tion species long represented by only a few marensis had been collected from the islands specimens (e.g., B. bicolor, B. elerae, B. of and Leyte. Additionally, Brown and wrighti, B. pathfinderi) have recently been Rabor (1967) reported on a second specimen rediscovered (Siler, 2010) and redescribed as from Samar Island; however, no information valid taxa (Siler et al., 2011a,c). Together, on where the specimen was deposited, or its these studies coupled with increased sampling museum catalog number, was provided. Al- throughout the Philippines and a new, robust though Brown and Rabor (1967) treated B. molecular dataset, allow us to begin evaluating samarensis as a single widespread species, variation across the isolated populations of they referred to the species as a ‘‘complex,’’ widespread species in the Philippines. suggesting that they suspected it contained In a recent study, Siler and Brown (2010) multiple species. They also noted several revised two polytypic species (B. boulengeri distinct morphological differences between and B. schadenbergi) and one widespread island populations, including differences in species (B. talinis), and inferred the presence fore- and hind-limb digit number and head of 10 genetically and morphologically distinct scale patterns. allopatric evolutionary lineages (species). Sev- Additional island populations of B. samar- eral other species are still recognized as ensis were subsequently sampled by the time having widespread distributions that span Brown and Alcala (1980) revised the genus, historical faunal demarcations in the Philip- including populations from the Lapinig Group pines (Brown and Diesmos, 2002, 2009; islands off the northeast coast of Bohol Island Brown and Guttman, 2002; Heaney, 1985), (Fig. 1). Ross and Gonzales (1992) would later including B. samarensis and B. bonitae report on observations of B. samarensis from (Brown, 1956; Brown and Rabor, 1967; Brown Catanduan˜ es Island off the northeast coast of and Alcala, 1980). One of these species (B. the Bicol Peninsula (Fig. 1) and in 2001, RMB samarensis) is the focus of this study. (unpublished data) recorded B. samarensis on the southern tip of the Bicol Peninsula in the TAXONOMIC HISTORY foothills of Mt. Bulusan. The genus Brachymeles was first described by To date, Brachymeles samarensis remains a Dumeril and Bibron (1839) for the small, limb- widespread species spanning islands of the reduced species B. bonitae. Three additional Luzon and Mindanao Pleistocene Aggregate species (Senira bicolor Gray, 1845; Eumeces Island Complexes (PAICs; Brown and Dies- (Riopa) gracilis Fischer, 1885; E. (R.) schaden- mos, 2002; Brown and Guttman, 2002; bergi Fischer, 1885) were transferred to the Fig. 1). Widespread distributions such as this genus by Boettger (1886) and Boulenger (1887). have been the focus of many recent studies These four species represented the known (Brown et al., 2000; Siler and Brown, 2010; 78 HERPETOLOGICAL MONOGRAPHS [No. 25

distinct evolutionary lineages (species). In this paper we provide a phylogenetic analysis of all of these taxa, fully describe each species, clarify taxonomic boundaries, and provide the first illustrations of all included species. We also provide information on each species’ natural history, ecology, and geographic distribution.

MATERIALS AND METHODS Field Work, Sample Collection, and Specimen Preservation Fieldwork was conducted on Catanduan˜ es, Lapinig Grande, Leyte, Luzon, and Samar islands, all in the Philippines (Fig. 1) between 2001 and 2009. Specimens were collected between 900 and 1600 h, euthanized in aqueous chloretone, dissected for tissue sam- ples (liver preserved in 95% ethanol or flash frozen in liquid nitrogen), fixed in 10% formalin, and eventually (,2 mo) transferred to 70% ethanol. Newly sequenced specimens are deposited in US and Philippine museum collections (see Acknowledgments and Ap- pendix I, Additional Specimens Examined); voucher information corresponding to data FIG. 1.—Map of the Philippine islands, with island from GenBank sequences is included in labels provided for islands with representative samples used for this study. The five recognized major Pleistocene Table 1. Museum abbreviations for specimens Aggregate Island Complexes (PAICs), major island examined follow Leviton et al. (1985). groups, and additional deep-water islands are labeled for reference. Islands of the Romblon Island Group are Taxon Sampling and Outgroup Selection for designated by the first letter of the island name (T, Tablas Phylogenetic Analyses Island; R, Romblon Island; S, ). Current islands in the Philippines are shown in medium grey; light Because our primary goal was to estimate gray areas enclosed in black 120-m bathymetric contours phylogenetic relationships among the various indicate the hypothesized maximum extent of land during populations of Brachymeles samarensis we the mid- to late Pleistocene. sequenced only a few exemplars (2–4) per sampled population. We included samples of Siler et al., 2010a,b; Welton et al., 2009, Lygosoma bowringi as an outgroup based on 2010a,b), which have revealed that few relationships presented in recent phylogenetic endemic Philippine actually possess studies of the genus Brachymeles (Siler et al., broad distributions spanning these regional 2011b; Siler and Brown, 2011). Additionally, we faunistic boundaries (reviewed by Brown and included samples of B.apus,B.bonitae,B. Diesmos, 2009). cebuensis, B. lukbani, and B. minimus to explore The goal of the present study is to revise the the sister group relationships within the B. taxonomy of the B. samarensis complex such samarensis complex (Siler et al., 2011b; Siler that individual units (species) represent inde- and Brown, 2011). A total of 28 ingroup samples pendently evolving, cohesive lineage segments were used in the phylogenetic analyses. (sensu de Queiroz, 1998, 1999; Frost and Hillis, 1990; Simpson, 1961; Wiley 1978). DNA Extraction, Purification, Comprehensive examination of all recently and Amplification collected specimens from throughout the We extracted total genomic DNA from known range of B. samarensis results in the tissues (Table 1) using the modified guanidine reorganization of the species complex into six thiocyanate extraction method of Esselstyn et al. 2011 EPTLGCLMNGAH 79 MONOGRAPHS HERPETOLOGICAL ] TABLE 1.—Summary of specimens corresponding to genetic samples included in the study, general locality, and GenBank accession number. SP 5 Sabah Parks Reference Collection, KU 5 University of Kansas Natural History Museum, LSUHC 5 La Sierra University Herpetological Collections, TNHC 5 Texas Natural History Collections of the Texas Memorial Museum of the University of Texas at Austin.

GenBank accession numbers

Species Voucher Locality ND1 ND2 a-enolase PTGER4 Lygosoma bowringii LSUHC 6970 West Malaysia JN981975 JN981989 JN981962 JN982001 Lygosoma bowringii LSUHC 6998 West Malaysia HQ907328 HQ907430 HQ906969 — Brachymeles apus SP 06915 Malaysia, Borneo, Sabah, Mt. Kinabalu National Park HQ907331 HQ907433 HQ906972 HQ907533 Brachymeles bonitae KU 307747 Philippines, , Municipality of Quezon JN981976 JN981990 JN981963 JN982002 Brachymeles bonitae KU 326080 Philippines, Polillo Island, Municipality of Quezon JN981977 JN981991 JN981964 JN982003 Brachymeles minimus KU 308131 Philippines, Catanduanes Island, Municipality of Gigmoto HQ907404 HQ907508 HQ907039 HQ907608 Brachymeles minimus KU 308129 Philippines, Catanduanes Island, Municipality of Gigmoto HQ907405 HQ907509 HQ907040 HQ907609 Brachymele lukbani KU 313602 Philippines, Luzon Island, Municipality of Labo HQ907407 HQ907511 HQ907042 HQ907611 Brachymele lukbani KU 313596 Philippines, Luzon Island, Municipality of Labo HQ907408 HQ907512 HQ907043 HQ907612 Brachymeles cebuensis KU 320419 Philippines, Cebu Island, Municipality of Carcar HQ907410 HQ907514 HQ907045 HQ907614 Brachymeles cebuensis KU 320421 Philippines, Cebu Island, Municipality of Carcar HQ907411 HQ907515 HQ907046 HQ907615 Brachymeles libayani2 KU 320430 Philippines, Lapinig Grande Island, Municipality of Carlos P. Garcia JN981978 — JN981965 — Brachymeles libayani2 KU 320458 Philippines, Lapinig Grande Island, Municipality of Carlos P. Garcia JN981979 JN981992 — JN982004 Brachymeles libayani1 KU 320466 Philippines, Lapinig Grande Island, Municipality of Carlos P. Garcia JN981980 JN981993 JN981966 JN982005 Brachymeles libayani2 KU 320445 Philippines, Lapinig Grande Island, Municipality of Carlos P. Garcia JN981981 — JN981967 — Brachymeles bicolandia1 KU 324003 Philippines, Luzon Island, Municipality of Tobaco JN981982 JN981994 JN981968 JN982006 Brachymeles bicolandia2 KU 324016 Philippines, Luzon Island, Municipality of Tobaco JN981983 JN981995 JN981969 JN982007 Brachymeles bicolandia2 KU 324010 Philippines, Luzon Island, Municipality of Tobaco JN981984 JN981996 JN981970 JN982008 Brachymeles bicolandia2 KU 324004 Philippines, Luzon Island, Municipality of Tobaco JN981985 JN981997 JN981971 JN982009 Brachymeles brevidactylus3 TNHC 62469 Philippines, Luzon Island, Municipality of Sorsogon JN981986 JN981998 JN981972 JN982010 Brachymeles brevidactylus1 KU 324017 Philippines, Luzon Island, Municipality of Sorsogon JN981987 JN981999 JN981973 JN982011 Brachymeles cobos2 KU 324025 Philippines, Catanduanes Island, Municipality of San Miguel JN981988 JN982000 JN981974 JN982012 Brachymeles cobos2 KU 324020 Philippines, Catanduanes Island, Municipality of San Miguel HQ907349 HQ907450 HQ906989 HQ907550 Brachymeles cobos2 KU 324022 Philippines, Catanduanes Island, Municipality of San Miguel HQ907350 HQ907451 HQ906990 HQ907551 Brachymeles samarensis KU 310850 Philippines, Samar Island, Municipality of Taft HQ907416 HQ907520 HQ907051 HQ907620 Brachymeles samarensis KU 310851 Philippines, Samar Island, Municipality of Taft HQ907417 HQ907521 HQ907052 HQ907621 Brachymeles samarensis KU 310849 Philippines, Samar Island, Municipality of Taft HQ907413 HQ907517 HQ907048 HQ907617 Brachymeles paeforum2 KU 311225 Philippines, Leyte Island, Municipality of Baybay HQ907414 HQ907518 HQ907049 HQ907618 Brachymeles paeforum2 KU 311226 Philippines, Leyte Island, Municipality of Baybay HQ907418 HQ907522 HQ907053 HQ907622 Brachymeles paeforum2 KU 311227 Philippines, Leyte Island, Municipality of Baybay HQ907419 HQ907523 HQ907054 HQ907623

1 Holotype. 2 Paratopotype. 3 Paratype. 80 HERPETOLOGICAL MONOGRAPHS [No. 25

TABLE 2.—Models of evolution selected by Akaike Information Criterion (AIC) and applied for partitioned, Bayesian phylogenetic analyses.1

Partition AIC Model Model applied Number of characters ND1, 1st codon position GTR + I + G GTR + G 322 ND1, 2nd codon position GTR + I + G GTR + G 322 ND1, 3rd codon position GTR + I + G GTR + G 322 ND2, 1st codon position TVM + I + G GTR + G 287 ND2, 2nd codon position GTR + I + G GTR + G 287 ND2, 3rd codon position TVM + I + G GTR + G 287 a-enolase TVMef + G GTR + G 261 PTGER4 HKY + I + G HKY + G 490 1 The model GTR + G was used for partitioned RAxMLHPC analyses.

(2008). The mitochondrial NADH dehydroge- random addition-sequence replicates and tree nase subunit 1 (ND1), NADH dehydrogenase bisection and reconnection (TBR) branch subunit 2 (ND2), and the protein-coding nuclear swapping. To assess heuristic support, non- loci, a-enolase and PTGER4, were completely parametric bootstrapping was conducted us- sequenced for nearly all samples using the ing 1000 replicates, each with 100 random primers and protocols provided in Siler et al. addition-sequence replicates and TBR branch (2011b) and Siler and Brown (2011). We swapping. purified polymerase chain reaction templates Partitioned maximum likelihood (ML) anal- with 1 mLofa20%solutionofExoSAP-IT yses were conducted in RAxMLHPC v7.04 (US78201, Amersham Biosciences, Piscataway, (Stamatakis, 2006). The alignment was parti- NJ). Cycle-sequencing reactions were complet- tioned into eight regions consisting of the ed with the same primers and ABI Prism codon positions of ND1 and ND2, and the BigDye Terminator chemistry (Ver. 3.1; Applied two nuclear loci, a-enolase and PTGER4, Biosystems, Foster City, CA). Cycle-sequenc- following the methods of Siler et al. (2011b) ing products were purified with Sephadex Me- and Siler and Brown (2011). Analyses that dium (NC9406038, Amersham Biosciences) partition protein-coding genes by codon posi- in Centri-Sep 96 spin plates (CS-961, Prince- tion have been shown to improve resulting ton Separations, Princeton, NJ). We analyzed inferences (Brandley et al., 2005). The partitions purified products using an ABI Prism 3130xl were run under the same generalized time- Genetic Analyzer (Applied Biosystems), and reversible model (GTR + C) with 100 replicate gene sequences were assembled with Sequen- best-tree inferences. Each inference was per- cher 4.8 (Gene Codes Corp., Ann Arbor, MI). formed with a random starting tree and relied on the rapid hill-climbing algorithm (Stamatakis, Alignment and Phylogenetic Analysis 2006). Clade support was assessed with 1000 Initial alignments of the gene regions were bootstrap pseudoreplicates. We considered produced in Muscle v3.7 (Edgar, 2004) and branches receiving $70% bootstrap support to manual adjustments were made in MacClade be well supported (Wilcox et al., 2002). 4.08 (Maddison and Maddison, 2005). No The Akaike Information Criterion, as im- instances of insertions or deletions, or ambig- plemented in jModeltest v0.1.1 (Guindon and uously aligned regions, were observed in the Gascuel, 2003; Posada, 2008), was used to data, and all data were used for analyses. The select the best model of nucleotide substitu- final alignment consisted of 2570 nucleotide tion for each partition (Table 2). The best- positions. fit model for each of the eight partitions Phylogenetic analyses were conducted us- (Table 2) was used for Bayesian analyses ing parsimony and likelihood optimality crite- performed in MrBayes 3.1 (Ronquist and ria, as well as Bayesian methods. Parsimony Huelsenbeck, 2003). The same partitioning (MP) analyses were conducted in PAUP* 4.0 strategy used for ML analyses was used for (Swofford, 2002) with all characters weighted Bayesian inferences. Searches over tree space equally. Most-parsimonious trees were esti- were conducted with four runs, each with four mated using heuristic searches with 1000 chains, and were run for 2 3 107 generations. 2011] HERPETOLOGICAL MONOGRAPHS 81

Trees were sampled every 1000 generations, 1978). We consider as distinct lineages those with 4000 samples discarded as burn-in; this populations that are morphologically and left 16,001 post–burn-in trees from each run genetically distinct, especially if allopatric. included in the posterior distribution of Lineage-based species concepts have been topologies. Visual inspection for chain statio- employed in the recognition of Philippine narity and high effective sample size (ESS) biodiversity (Brown et al., 2000, 2008b, 2009; values (equal to or greater than 800) was Brown and Diesmos, 2002; Brown and Gutt- conducted within the program Tracer v1.4 man, 2002; Gaulke et al., 2007; Siler and (Rambaut and Drummond, 2007). Additional- Brown, 2010; Welton et al., 2009, 2010a,b) ly, correlations of split frequencies and cumu- due to the highly partitioned nature of the lative split frequencies were examined using archipelago (Brown and Diesmos, 2009), and the program AWTY (Nylander et al., 2008). We because the geological history of the islands considered topologies with posterior probabil- has been so well documented (Hall, 2002; ities $0.95 to be well supported (Leache´ and Voris, 2000; Yumul et al., 2009). In this study Reeder, 2002; Wilcox et al., 2002). we use an estimate of phylogenetic relation- ships as a guide for delimiting species but Morphological Data restrict our diagnoses of new species to those We examined fluid-preserved specimens populations that are clearly identified by (Appendix I) for variation in qualitative and diagnostic differences in nonoverlapping mor- mensural characters. Sex was determined by phological character states. gonadal inspection, and measurements were taken to the nearest 0.1 mm with digital RESULTS calipers by C. D. Siler. X-rays were taken with a company cabinet X-ray on Kodak MIN-R Phylogeny 2000 film exposed at 5 mA and 30 V for 1 min Of 2570 mitochondrial and nuclear charac- 15 s. ters, 848 were parsimony-informative. The Meristic and mensural characters were maximum parsimony analysis inferred 10 most chosen based on Siler et al. (2009, 2010a,b): parsimonious trees (tree length 5 2084; snout–vent length (SVL), axilla–groin distance topology not shown; bootstrap support sum- (AGD), total length, midbody width (MBW), marized in Fig. 2). The resulting 100 inferenc- midbody height (MBH), tail length (TL), tail es from the partitioned RAxML ML analysis width (TW), tail height (TH), head length show an average likelihood score 2lnL (HL), head width (HW), head height, snout– 12011.371112, with a single inference having forearm length (SnFa), eye diameter (ED), the highest likelihood score of 2lnL eye–narial distance (END), snout length 12011.367644. Trees recovered from ML, (SNL), internarial distance (IND), forelimb MP, and Bayesian analyses are topologically length (FLL), hind limb length (HLL), mid- identical. No inferences support the monophy- body scale-row count, paravertebral scale-row ly of Brachymeles samarensis. All analyses count, axilla–groin scale-row count, Finger-III recover two reciprocally monophyletic clades lamellae count, Toe-IV lamellae count, supra- within the B. samarensis complex, each char- labial count, infralabial count, supraciliary acterized by the presence of multiple highly count, and supraocular count. Additionally, divergent, genetically distinct lineages (Fig. 2). we counted the number of presacral vertebrae The Leyte Island and Lapinig Group from X-ray images of specimens. In the Islands populations were recovered as a clade, description, ranges are followed by mean 6 sister to B. cebuensis from Cebu Island SD in parentheses. (Fig. 2). Topotypic B. samarensis from Samar Island were recovered as sister to a clade of Species Concept two limbless species of Brachymeles (B. We follow the general lineage concept of minimus and B. lukbani) and the Luzon and species (de Queiroz, 1998, 1999) as a logical Catanduan˜ es island populations of B. samar- extension of the evolutionary species concept ensis (Fig. 2). Within Luzon Island, two se- (Frost and Hillis, 1990; Simpson, 1961; Wiley, parate lineages were recovered from the Bicol 82 HERPETOLOGICAL MONOGRAPHS [No. 25

FIG. 2.—Maximum likelihood (ML) estimate of combined mitochondrial and nuclear data for samples of Brachymeles used for this study (preferred ML tree, 2lnL 12011.367644; ND1, ND2, a-enolase, PTGER4). Nodes are shown with numerical values corresponding to maximum parsimony bootstrap proportions, maximum likelihood bootstrap proportions, and Bayesian posterior probabilities, respectively. Terminals are labeled with taxonomic names, fore- and hind limb digit states, and number of presacral vertebrae.

Peninsula, with no support for their mono- both limbless species, nested within a didactyl phyly (Fig. 2). clade of species formerly part of B. samarensis In addition to the finding of paraphyly of B. (topotypic B. samarensis, B. sp. nov. [Catan- samarensis complex members, several other duan˜ es Island], B. sp. nov. [Southern Bicol recognized species were recovered as part of Peninsula, Luzon Island], B. sp. nov. (Central this clade. Brachymeles cebuensis, one of only Bicol Peninsula, Luzon Island]; Fig. 2). All two recognized species to have unequal three previously recognized species (B. ce- numbers of fingers and toes (Brown and buensis, B. lukbani, B. minimus) have geo- Alcala, 1980), was supported as part of a clade graphical distributions that overlap, or are in of species (B. cebuensis + B. sp. nov. [Leyte close proximity to, the known ranges of other Island] + B. sp. nov. [Lapinig Group Islands]) species in the B. samarensis complex (Fig. 3). with three fingers and two or three toes All analyses result in the strong support (Fig. 2). Sister to this three-finger clade, all of six genetically distinct lineages within analyses recovered B. minimus and B. lukbani, the Brachymeles samarensis species complex 2011] HERPETOLOGICAL MONOGRAPHS 83

FIG. 3.—(Left) Map of the Philippine islands showing previously recognized distribution of Brachymeles samarensis (indicated by black shaded islands), and recognized distributions of other members of the B. samarensis complex (indicated by dark gray shapes). (Right) Hypothesized distributions of B. bicolandia, B. brevidactylus, B. cobos, B. libayani, B. paeforum, and B. samarensis in the eastern-central Philippines. The sampling localities are indicated by black shapes, and the hypothesized geographic range of each species is indicated by shaded islands and dashed lines, with shapes and shades of islands corresponding to the map’s key.

(Fig. 2). Uncorrected pairwise sequence di- Mitochondrial sequence divergences among vergences are low within the lineages defined the other three lineages within the B. samar- here as species and high between these ensis species complex (B. samarensis [Samar lineages (Table 3). Percentage of divergences Island], B. sp. nov. [Lapinig Group islands], B. for the mitochondrial and for the nuclear data, sp. nov. (Leyte Island]) are greater than 9.2% respectively, show that the monophyletic (Table 3; Fig. 2). Intraspecific sequence diver- lineages defined by our phylogenetic analyses gences are low (0.0–1.9%) in comparison to (B. samarensis, B. sp. nov. [Leyte Island], B. divergences among monophyletic lineages. sp. nov. [Lapinig Group Islands], B. sp. nov. Additionally, moderate levels of sequence diver- [Catanduan˜ es Island], B. sp. nov. [Southern gence are observed for nuclear sequence data Bicol Peninsula, Luzon Island], B. sp. nov. (Table 3). (Central Bicol Peninsula, Luzon Island]) are distinguished from congeners by levels of Morphology genetic divergence similar to, or greater than, Variation in morphological characters those between previously defined species— (Tables 4–6) mirrors the results observed in viz., B. bonitae, B. cebuensis, B. minimus,and phylogenetic analyses, and supports the rec- B. lukbani (Table 3; Fig. 2). The two most ognition of six B. samarensis group lineages. closely related lineages, B. sp. nov. (Catan- Characters differing among these six lineages duan˜ es Island) and B. sp. nov. (Southern Bicol include digit number, presacral vertebrae Peninsula, Luzon Island), are separated by number, degree of digit development, head 4.1–4.7% mitochondrial sequence divergence. and body scale counts and patterns, and 84 HERPETOLOGICAL MONOGRAPHS [No. 25

pigmentation patterns (Tables 4–6; species accounts below), all of which are typical morphological diagnostic characters employed 0.0

0.7–0.8 historically by taxonomists working with this genus (review: Brown and Alcala, 1980). We observed no intraspecific mensural or meristic differences between the sexes of any of the six

0.1 species. 2.8–3.0 3.2 Brachymeles samarensis, B. Superficially, the six lineages within the B. samarensis complex appear morphologically similar in overall body size and general shape; however, upon closer inspection, three dis- 2.2 2.6–3.0 3.2 0.0 tinct body forms are observed. Among the six lineages, two are tridactyl (B. sp. nov. [Leyte

(Fig. 2). Percentages on the diagonal represent Island] and B. sp. nov. [Lapinig Group Islands]), three are bidactyl (B. samarensis,

1.4 B. sp. nov. [Catanduan˜ es Island] and B. sp. 2.6–2.8 1.4–1.6 2.7–3.1 3.4–3.5 nov. [Central Bicol Peninsula, Luzon Island]),

B. minimus and one is bidactyl, but with small, highly

and reduced, and nearly imperceptible claws (B. sp. nov. [Southern Bicol Peninsula, Luzon

1.9 Island]). Additionally, numerous nonoverlap- 0.7–1.2 2.7 1.5 2.8–3.1 3.5 ping differences were detected in meristic, mensural, osteological, and color pattern characters for each complex member, readily defining six distinct lineages within the com- 0.5–0.9 0.4–1.1 2.3–2.7 1.1–1.6 2.6–3.2 3.2–3.6 0.0–0.2 plex (Tables 4–6). In summary, each lineage (most of which are allopatric) possesses unique and nonoverlapping suites of diagnostic char- acter states of morphology, perfectly cor- responding to the six clades defined in the 1.4–2.8 1.6–3.6 1.2–3.3 2.2–4.8 1.4–2.8 3.0–3.6 3.2–4.4 0.4–0.7 B. bonitae, B. cebuensis, B. lukbani,

, phylogenetic analyses of DNA sequence data. Taxonomic Conclusions Our estimate of phylogeny (Fig. 2); biogeo- graphically separate ranges of island or region 0.2–2.8 1.8–2.2 2.0 1.6–2.0 2.6 1.3 3.0 3.3 0.0–0.1

B. brevidactylus endemic species; diagnostic, nonoverlapping intraspecific genetic diversity for mitochondrial data (bolded for emphasis). , morphological character states; and genetic distances between the taxa (Table 3) indicate the distinctiveness of a new species from B. cobos 2.2 1.8–2.8 1.1–1.5 1.6 1.5 2.4 1.2 2.7–3.1 3.4 ,

0.0–0.7 Catanduan˜ es Island, two new species from the Bicol Peninsula of Luzon Island (one from the central Bicol region and one from the extreme southern tip of the peninsula), a new species

0.0 from the Lapinig Group Islands, and a new 14.3 16.2 11.6–11.7 12.0–12.4 14.6–14.8 14.7 14.8–14.9 16.5–16.7 14.114.1 14.1–14.4 14.0–14.2 14.8–15.3 13.8–14.3 11.5–11.9 11.6–12.0 11.2 11.0–11.2 11.6 11.8–11.9 15.0–15.1 15.0–15.1 13.4 13.7 4.9–5.0 14.4–14.514.9–15.016.9–17.3 14.4–14.6 14.6–14.9 15.0–15.6 14.5–14.8 14.7–15.4 15.5–16.5 4.1–4.7 9.4–9.6 15.8–16.5 15.7–16.2 9.3–9.5 15.7–16.3 14.6–15.214.5–14.7 9.2–10.5 14.1–14.5 14.9–15.8 B. samarensis B. paeforum B. libayani B. bicolandia B. cobos B. brevidactylus B. bonitaespecies B. cebuensis from B. lukbani Leyte B. minimus Island (Table 3; Fig. 2). Each of the six species of the B. samarensis complex is morphologically distinct from each other and all other known species in the

3.—Uncorrected pairwise sequence divergence (%) for mitochondrial data (below diagonal) and nuclear data (above diagonal), for genus, and the 11 species of Brachymeles included in phylogenetic analyses also are ABLE T B. samarensis B. paeforum B. cobos B. brevidactylus B. bonitae B. cebuensis paeforum, B. libayani, B. bicolandia B. libayani B. bicolandia B. lukbani B. minimus genetically distinct. All monophyletic lineages, 2011] HERPETOLOGICAL MONOGRAPHS 85 with the exception of the two occurring on (Tables 4 and 5); from B. brevidactylus by the Bicol Peninsula of Luzon Island, are having fewer presacral vertebrae and fewer endemic to single islands within two isolated paravertebral scale rows (Tables 4 and 5); and PAICs, thereby providing additional support from B. bonitae by having longer relative hind for the distinctiveness of each clade’s evo- limb lengths, fewer presacral vertebrae, fewer lutionary history and lineage integrity. Ac- paravertebral scale rows, six supraciliaries, five cordingly, we recognize B. samarensis as a supraoculars, and the presence of contact species that occurs only on Samar Island in between frontoparietals (Tables 4 and 5). the eastern Visayan (central) Philippine islands Brachymeles samarensis can be distin- (e.g., Mindanao PAIC; Fig. 3), and hereby guished from all limbless species of Brachy- recognize the five additional lineages within meles (B. apus, B. lukbani, B. minimus, B. the B. samarensis species complex as new miriamae, B. vermis) by having limbs; and species. from all pentadactyl species of Brachymeles (B. boholensis, B. boulengeri, B. bicolor, B. TAXONOMIC ACCOUNTS gracilis, B. kadwa, B. makusog, B. mind- Brachymeles samarensis Brown 1956: 6 orensis, B. orientalis, B. schadenbergi, B. Figs. 3–4 talinis, B. taylori, B. tungaoi, B. vindumi)by Brachymeles samarensis, Brown, 1956, having nonpentadactyl limbs, shorter adult Type locality: Guinuan, Samar Island, Philip- forelimb lengths (less than 2.6 mm vs. greater pines (holotype: FMNH 44472); Brown and than 5.9 mm), shorter adult hind limb lengths Alcala, 1970, 1980; Brown and Rabor, 1967. (less than 3.1 mm vs. greater than 10.3 mm), Diagnosis.—Brachymeles samarensis can and a narrower body (less than 6.4 mm vs. be distinguished from congeners by the greater than 7.9 mm), and by the absence of a following combination of characters: (1) body postnasal scale and auricular opening (vs. size small (SVL 57.9–66.1 mm), (2) limbs presence). bidactyl, (3) limb length short, (4) supralabials Description (based on holotype description six, (5) infralabials six, (6) supraciliaries six, (7) and six referred specimens).—Details of the supraoculars five, (8) midbody scale rows 19– head scalation of an adult female are shown in 22, (9) axilla–groin scale rows 66–69, (10) Fig. 5. Measurements and character states of paravertebral scale rows 86–88, (11) pineal the holotype are provided below in square eye spot present, (12) prefrontals not contact- brackets. Body small, slender; SVL 57.9 mm ing on midline, (13) frontoparietals contact, for males, maximum SVL 66.1 mm for (14) mental/first infralabial fusion absent, (15) females, [43.5 mm, juvenile] (Tables 4 and postnasals absent, (16) enlarged chin shields 5); head weakly differentiated from neck, in three pairs, (17) nuchal scales differentiat- nearly as wide as body, HW 7.3–9.2% (8.3 6 ed, (18) fourth supralabial below eye midline, 0.7) SVL, 91.4–117.8% (102.7 6 10.8) HL; (19) auricular opening absent, (20) presacral HL 36.6–42.5% (38.8 6 2.1) SnFa; SnFa vertebrae 45, and (21) uniform body color 18.8–23.5% (20.9 6 1.6) SVL; snout short, (Tables 4 and 5). bluntly rounded in dorsal and lateral profile, Comparisons.—Characters distinguishing SNL 50.9–55.3% (53.3 6 1.8) HL; ear Brachymeles samarensis from all nonpenta- completely hidden by scales; eyes small, ED dactyl, limbed species of Brachymeles are 1.3–1.6% (1.4 6 0.1) SVL, 17.0–18.7% (17.6 summarized in Tables 4 and 5. Brachymeles 6 0.6) HL, 42.6–48.0% (45.8 6 2.1) END, samarensis most closely resembles B. bicolan- pupil subcircular; body slightly depressed, dia, B. cobos, B. brevidactylus, and popula- nearly uniform in thickness, MBW 109.1– tions of B. bonitae, the only other bidactyl 150.6% (130.4 6 14.9) MBH; scales smooth, species. However, B. samarensis differs from glossy, imbricate; longitudinal scale rows at these four taxa by having midbody scale rows midbody 19–22 [22]; paravertebral scale rows as few as 19 and axilla–groin scale rows as few 86–88 [86]; axilla–groin scale rows 66–69; as 66 (Table 5). Brachymeles samarensis limbs short, poorly developed, with digits further differs from B. bicolandia by having reduced to two claws on both forelimbs and fewer presacral vertebrae and six infralabials hind limbs, finger and toe lamellae absent; 86 HERPETOLOGICAL MONOGRAPHS [No. 25

TABLE 4.—Summary of meristic and mensural characters in all known limbed, nonpentadactyl species of Brachymeles. Sample size, body length, and total length (TotL) among males and females, and general geographical distribution (Pleistocene Aggregate Island Complexes [PAIC], sensu Brown and Diesmos, 2002) are included for reference. SVL 5 snout–vent length, TotL 5 total length, TL 5 tail length, FLL 5 forelimb length, HLL 5 hind limb length, ToeIVlam 5 toe-IV lamellae count. (SVL, TotL, MBW, FLL, and HLL given as range over mean 6 SD; all body proportions given as percentage over mean 6 SD).

B. samarensis (1 B. paeforum (3 males, B. libayani (10 males, B. bicolandia (6 males, B. cobos B. brevidactylus (1 male, 5 females) 9 females) 25 females) 10 females) (9 females) male, 2 females)

Lapinig Group Central Bicol Southern Bicol Range Samar Island Leyte PAIC Islands Peninsula Catanduan˜ es Island Peninsula SVL (female) 62.4–66.1 47.2–61.4 52.8–66.1 46.4–67.4 54.0–64.4 54.0, 60.0 (63.4 6 1.5) (56.5 6 4.2) (58.6 6 3.3) (59.0 6 6.7) (58.7 6 3.5) SVL (male) 57.9 62.4–66.1 52.7–57.4 56.4–66.1 — 56.8 (63.4 6 1.5) (56.0 6 1.6) (61.7 6 3.5) TotL (female) 97.7–112.9 99.5–108.5 91.4–111.2 94.1–112.7 102.2–109.4 92.3, 95.2 (107.3 6 8.3) (102.6 6 5.1) (102.2 6 6.4) (102.1 6 8.8) (106.2 6 2.7) TotL (male) 93.0 106.7–114.6 92.1–103.4 99.6–107.9 — 102.0 (110.6 6 5.6) (99.4 6 4.5) (104.1 6 4.2) TL/SVL 57–81 69–79 63–84 58–93 76–96 59–80 (68 6 12) (75 6 4) (77 6 6) (75 6 12) (88 6 8) (70 6 11) FLL 1.1–2.6 1.3–1.7 1.1–1.8 1.1–1.9 1.4–2.1 1.1–1.5 (1.8 6 0.5) (1.5 6 0.1) (1.3 6 0.2) (1.4 6 0.3) (1.7 6 0.2) (1.5 6 0.2) FLL/SVL 2–4 2–4 2–3 2–4 3 2–3 (3 6 1) (3 6 0) (2 6 0) (2 6 0) (3 6 0) (3 6 0) HLL 2.5–3.1 2.3–3.0 2.0–2.7 1.9–3.1 2.5–3.6 2.1– 2.7 (2.9 6 0.2) (2.6 6 0.3) (2.4 6 0.2) (2.6 6 0.3) (3.0 6 0.3) (2.5 6 0.3) HLL/SVL 4–5 4–5 3–5 3–5 4–6 4–5 (5 6 0) (4 6 0) (4 6 0) (4 6 1) (5 6 1) (4 6 0) ToeIVlam 0 0 0 0 0 0

FLL 2.4–5.7% (3.9 6 1.3) AGD, 1.8–3.9% anterior loreal about as long as and slightly (2.9 6 0.9) SVL; HLL 5.3–7.2% (6.2 6 0.7) higher than posterior loreal; preocular one; AGD, 4.0–5.0% (4.6 6 0.4) SVL [6.9]; tail not presubocular one; supraciliaries six, the ante- as wide as body, gradually tapered towards riormost contacting prefrontal and separating end, TW 70.2–82.6% (76.7 6 5.0) MBW, TL posterior loreal from first supraocular, poste- 56.5–80.6% (68.4 6 11.6) SVL. riormost extending to posterior edge of fifth Rostral projecting onto dorsal snout to level supraocular; subocular scale row single, com- in line with middle of nasal, broader than plete, in contact with supralabials; lower high, in contact with frontonasal; frontonasal eyelid with one row of scales; supralabials wider than long; nostril ovoid, in center of six, first twice the anteroposterior length of single trapezoidal nasal, longer axis directed others, fourth below eye midline; infralabials anteroventrally and posterodorsally; suprana- six (Fig. 4). sals present, large, broadly separated; postna- Mental wider than long, in contact with first sals absent; prefrontals moderately separated; infralabials; postmental single, enlarged, its frontal octagonal-shaped, its anterior margin width equal to width of mental; followed by in moderate contact with frontonasal, in con- three pairs of enlarged chin shields, first pair tact with first two anterior supraoculars, 33 in broad medial contact, second pair wider wider than anterior supraocular; supraoculars than first, broadly separated by single medial five; frontoparietals moderate, in broad medi- scale, third pair separated by three medial al contact, each frontoparietal in contact with scales (Fig. 4). supraoculars 2–4; interparietal moderate, Scales on limbs smaller than body scales; its length roughly equal to midline length of scales on dorsal surfaces of digits wrapping frontoparietal, longer than wide, diamond- around lateral edges of digits; lamellae absent; shaped, wider anteriorly; parietals broader palmar surfaces of hands and plantar surfaces than frontoparietals, in broad contact behind of feet with several small, irregular scales, interparietal; nuchals enlarged; loreals two, each with irregular raised anterior edges; 2011] HERPETOLOGICAL MONOGRAPHS 87

TABLE 4.—Extended

B. muntingkamay B. tridactylus (9 males, B. bonitae (6 males, B. cebuensis B. elerae (2 males, B. pathfinderi B. wrighti (1 (12 females) 11 females) 7 females) (8 females) 1 female) (14 m, 23 females) male, 1 female)

Mindoro & Luzon Mindanao Luzon Island Visayan PAIC PAICs Cebu Island Luzon Island Island Luzon Island 61.8–81.3 45.5–59.1 49.7–59.8 51.5–67.9 68.2, 71.9 55.8–68.3 113.0 1 (73.6 6 5.9) (52.1 6 5.0) (56.4 6 3.9) (62.0 6 3.4) N/A 55.7–78.3 65.1–80.0 (61.8 6 5.3) 71.5 54.5–65.1 25.8 (68.5 6 7.4) (73.5 6 6.4) N/A (59.4 6 3.8) 107.4–136.0 102.6–154.1 93.4–150.4 104.3–128.0 109.9, 131.9 111.1–133.2 205.6 (124.0 6 8.6) (132.6 6 14.0) (126.7 6 19.9) (119.0 6 8.5) (119.7 6 8.2) N/A 105.3–133.67 102.6–144.5 N/A 101.4–107.0 216.4 (115.9 6 15.4) (121.3 6 15.6) N/A (104.2 6 4.0) 50–79 69–112 35–93 78–115 61–84 69–95 72, 82 (65 6 10) (92 6 12) (69 6 18) (92 6 13) (72 6 16) (84 6 10) 2.4–3.0 1.5–2.5 1.0–1.5 1.1–1.8 3.3–3.5 4.4–6.9 7.5, 7.5 (2.7 6 0.2) (2.0 6 0.3) (1.3 6 0.1) (1.5 6 0.3) (3.4 6 0.1) (5.8 6 0.5) 3–4 2–3 1–2 2–3 5–5 8–11 6, 7 (4 6 0) (3 6 0) (2 6 0) (2 6 0) (5 6 0) (10 6 1) 5.3–6.0 2.6–3.6 1.3–2.0 2.3–3.0 4.3–5.4 8.4–12.9 10.9, 13.9 (5.7 6 0.2) (3.1 6 0.3) (1.6 6 0.2) (2.7 6 0.3) (5.0 6 0.6) (10.8 6 1.0) 7–9 3–6 2–3 3–5 6–8 15–21 10, 11 (8 6 1) (5 6 1) (2 6 0) (4 6 0) (7 6 1) (18 6 1) 0 0 0 0 3 5–8 4, 5 fingers equal in size; toes unequal in size, minor differences, including a dark brown middle digit greatest in length, first and third body color and dark brown to black streaks of digits equal in length. pigmentation. Coloration in preservative.—Body ground Variation.—Morphometric variation of the color medium brown, each dorsal scale light series is summarized in Table 6. We observed a brown posteriorly, with a dark auburn streak single instance of digit variation, where one on the anterior two-thirds to half of the scale. specimen (KU 310849) has no fingers and two Dark streaks on each scale consist of four to toes. All specimens have two loreals with the seven thin longitudinal lines with smudges exception of a single specimen (KU 310852), between lines. Streaks on scales present which has a single loreal on the right side of the around entire body, more distinct on venter, body resulting from the fusion of the two scales where posterior ends of scales are cream, in this position. Additionally, the first and giving greater contrast. Posterior edge of all second pairs of enlarged chin shields are equal body scales transparent. Forelimb and hind in width among all specimens with the exception limb scales same color as body scales. of a single specimen (KU 310850), in which the Precloacal scale coloration matches surround- width of the second pair of enlarged chin shields ing ventral scale coloration. Head scales is greater than the width of the first pair. mottled light and dark brown, matching dorsal Distribution.—Brachymeles samarensis is background coloration. Supraocular, rostral, known only from Samar Island (Fig. 3). nasal, supranasal, and supralabials scales gray- Ecology and natural history.—Brachymeles cream. Mental, infralabial, postmental, and samarensis occurs in primary- and secondary- chin shields scales cream with slight brown growth forest habitats. In contrast to the other mottling, lighter than bordering ventral scales. members of the B. samarensis complex, this Coloration in life.—Coloration in life close- species appears to be a forest obligate, and ly matches the coloration in preservative with was only observed within rotting logs in 88 HERPETOLOGICAL MONOGRAPHS [No. 25

TABLE 5.—Summary of qualitative diagnostic characters (present, absent) in all known limbed, nonpentadactyl species of Brachymeles. The pairs of enlarged scales posterior to the postmental scale are abbreviated as chin shield pairs with reference to the first, second, and third pairs (when present). In cases of scale count variation within species, numbers of individuals showing specific counts are given in parentheses. PSV 5 presacral vertebrae, MBSR 5 midbody scale-row count, AGSR 5 axilla–groin scale-row count, PVSR 5 paravertebral scale-row count, SL 5 supralabial count, IFL 5 infralabial count, SC 5 supraciliary count, SO 5 supraocular count.

B. samarensis (1 male, B. paeforum B. libayani (10 B. bicolandia (6 B. cobos B. brevidactylus 5females) (3 males, 9 females) males, 25 females) males, 10 females) (9 females) (1 male, 2 females) Number of digits 2/2 3/3 3/3 2/2 2/2 2/2 (fore/hind) PSV 45 47 47 46–49 45 47–48 MBSR 19–22 21–22 22–23 20–22 21–22 20 AGSR 66–69 71–74 72–75 68–73 68–72 73–77 PVSR 86–88 93–96 90–92 85–90 85–89 90–94 SL 6 (6) 6 (12) 6 (35) 6 (16) 6 (9) 6 (3) IFL 6 (6) 5 (4) 6 (8) 5 (35) 5 (11) 6 (5) 6 (9) 6 (3) SC 6 (6) 6 (12) 6 (35) 6 (16) 6 (9) 6 (3) SO 5 (6) 5 (12) 5 (35) 5 (16) 5 (9) 5 (3) Pineal eyespot Present Present Present Present Present Present Prefrontal contact Absent Absent Point contact Absent Absent Absent or Absent Frontoparietal Present Present Present Present or Present Present contact Absent First chin shield Present Present or Present or Present or Present Absent pair contact Absent Absent Absent Thirdrd chin shield pair Present Present Present Present Present Present Chin shield pair size 3 , 1 # 23, 1 # 23, 1 , 23, 1 , 23, 1 # 23, 1 , 2 Chin shield pair 1(0); 2(1); 1(0/1); 2(1); 1(0/1); 2(1); 1(0/1); 2(1); 1(0); 2(1); 1(0); 2(1); separation1 3(3) 3(3) 3(3) 3(3) 3(3) 3(3) First/second loreal Present or Present or Present or Present or Absent Absent fusion Absent Absent Absent Absent Mental/first IFL fusion Absent Present or Present Present or Absent Absent Absent Absent Differentiated nuchals Present Present Present Present Present Present Continuous subocular Present Present Present Present Present Present scale row Auricular opening Absent Absent Absent Absent Absent Absent Dorsolateral stripes Absent Absent Absent Absent Absent Absent Longitudinal rows Absent Absent Absent Absent Absent Absent of dark spots

1Parentheses show the number of small ventral scale rows separating each enlarged pair of chin shields. 2Due to head damage in the nuchal region for both known specimens of B. wrighti, the presence of differentiated nuchals remains tentative. secondary-growth forest. Three species of Other sympatric lizard species observed on Brachymeles have been confirmed to occur Samar Island include the following: (Agamidae) on Samar Island: B. gracilis hilong, B. orien- Bronchocela cristatella, Draco bimacula- talis, and B. samarensis (Brown and Alcala, tus, D. ornatus, D. reticulates, Gonocephalus 1980; Siler and Brown, 2010). semperi, Hydrosaurus pustulatus; (Gekkoni- We have evaluated this species against the dae) Cyrtodactylus annulatus, C. sumoroi, International Union for Conservation of Gehyra mutilata, Gekko gecko, Gek. mind- Nature (IUCN) criteria for classification, and orensis, Hemidactylus frenatus, H. platyurus, find that it does not qualify for Critically Hemiphyllodactylus typus, Lepidodactylus aureo- Endangered, Endangered, Vulnerable, or lineatus, L. planicaudus, Pseudogekko compressi- Near Threatened status. Brachymeles samar- corpus; (Scincidae) Emoia atrocostata, Eutropis ensis has been documented to have a broad multicarinata, Eu. multifasciata, Lamprolepis geographic distribution across southern Samar smaragdina, Lipinia pulchella, Li. quadrivittata, Island. We therefore classify this species as Sphenomorphus acutus, S. cumingi, S. fasciatus, having Least Concern status (IUCN, 2011). S. jagori, S. cf. mindanensis, S. steerei, S. 2011] HERPETOLOGICAL MONOGRAPHS 89

TABLE 5.—Extended

B. muntingkamay B. tridactylus B. bonitae B. cebuensis B. elerae (2 males, B. pathfinderi B. wrighti (1 male, (12 females) (9 males, 11 females) (6 males, 7 females) (8 females) 1 females) (14 males, 23 females) 1 female) 3/3 3/3 0–2/0–2 3/2 4/4 5/4 4/4

42, 44 47 47–57 45 43 34 — 22–24 22–24 21–23 22–24 22–24 23–25 28, 28 65–70 70–79 73–90 65–69 63–67 44–48 85, 85 85–90 88–98 90–109 84–88 84–87 64–67 106, 108 6 (12) 6 (12) 7 (8) 6 (12) 7 (1) 6 (8) 6 (3) 6 (37) 6 (1) 7 (1) 6 (12) 6 (12) 7 (8) 5 (1) 6 (10) 7 (2) 6 (5) 7 (3) 6 (3) 6 (37) 7 (2) 6 (10) 7 (2) 5 (20) 5 (12) 6 (1) 6 (8) 5 (2) 6 (1) 5 (17) 6 (19) 6 5 (11) 6 (1) 4 (20) 4 (13) 5 (8) 4 (2) 5 (1) 5 (37) 5 Absent Present Present Present Absent Present Present Present Absent Absent Present or Present Absent Present or Absent Absent Absent Absent Absent Present Present Present or Present Absent Absent Present or Absent Present Absent Absent Absent Absent Present Present Present Present Present Absent Present 3 , 1 , 23, 1 , 23, 2 , 115 3 , 21, 3 , 21, 22, 1 1(1); 2(1); 1(0/1); 2(1); 1(1); 2(1); 1(0); 2(1); 1(1); 2(1); 1(1); 2(1) 1(1); 2(3) 3(3) 3(3) 3(3) 3(3) 3(3) Absent Absent Absent Absent Absent Absent Absent Absent Absent Present or Absent Absent Absent Absent Absent Absent Present Present Present Absent Absent Present2

Absent Absent Present Present Present Present Present

Absent Absent Absent Absent Absent Present Absent Absent Absent Absent Absent Absent Present Absent Present, around Present, vague Absent Absent Present, around Present, 6 Present body to indistinct body variegatus, Tropidophorus misaminus;(Varani- collected between 29 October and 8 Novem- dae) Varanus cumingi samarensis. ber 2007. Other paratypes.—One adult male (CAS- Brachymeles paeforum sp. nov. SU 26120), four adult females (CAS-SU Figs. 3, 6, 7 26110, 26112, 26121–22), and two juveniles Holotype.—PNM 9746 (CDS Field (CAS-SU 26115, 26123) collected between 1 No. 3418, formerly KU 311228), adult female, May and 4 June 1964, in Barrio Tambis, collected under rotting logs in secondary-growth Municipality of Burauen, Leyte Province, Leyte forest (1000 to 1230 h) on 8 November 2007 in Island, Philippines (11u009370N, 124u529190E; the Sitio San Vicente Tree Nursery, Barangay WGS-84), by D. S. Rabor; one adult male Pilim, Baybay City, Leyte Province, Leyte (CAS-SU 26771), two adult females (CAS-SU Island, Philippines (10u439350N, 124u499050E; 26770, 26772), and one juvenile (CAS-SU WGS-84), by CDS and J. Fernandez. 26773), collected between 10 June and 17 July Paratopotypes.—One adult male (KU 1964, in the Municipality of Mahaplag, Leyte 311225), one adult female (KU 311229), and Province, Leyte Island, Philippines (10u359420N, three juveniles (KU 311224, PNM 9747–48) 124u599130E; WGS-84), by D. S. Rabor. 0HREOOIA OORPS[ MONOGRAPHS HERPETOLOGICAL 90 TABLE 6.—Summary of univariate morphological variation among mensural characters in series of Brachymeles samarensis, B. paeforum, B. libayani, B. bicolandia, B. cobos, and B. brevidactylus.SVL5 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.

B. brevidactylus (1 B. samarensis (1 male, 5 females) B. paeforum (3 males, 9 females) B. libayani (10 males, 25 females) B. bicolandia (6 males, 10 females) B. cobos (9 females) male, 2 females) SVL (male) 57.9 59.7–64.1 (61.8 6 2.2) 52.7–57.4 (56.0 6 1.6) 56.4–66.1 (61.7 6 3.5) — 56.8 SVL (female) 62.4–66.1 (63.4 6 1.5) 47.2–61.4 (56.5 6 4.2) 52.8–66.1 (58.6 6 3.3) 46.4–67.4 (59.0 6 6.7) 54.0–64.4 (58.7 6 3.5) 54.0, 60.0 AGD (male) 40.3 44.1–48.2 (46.0 6 2.1) 39.5–43.6 (42.1 6 1.2) 42.9–52.0 (47.4 6 3.4) — 43.8 AGD (female) 45.3–50.3 (47.6 6 1.8) 35.6–46.5 (43.2 6 3.1) 39.4–50.6 (44.7 6 2.8) 32.9–52.3 (44.8 6 5.8) 41.0–49.1 (44.6 6 2.3) 45.5, 45.8 TotL (male) 93.0 106.7–114.6 (110.6 6 5.6) 92.1–103.4 (99.4 6 4.5) 99.6–107.9 (104.1 6 4.2) — 102.0 TotL (female) 97.7–112.9 (107.3 6 8.3) 99.5–108.5 (102.7 6 5.1) 91.4–111.2 (102.2 6 6.4) 94.1–112.7 (102.1 6 8.8) 102.2–109.4 (106.2 6 2.7) 92.3, 95.2 MBW (male) 5.7 5.0–5.3 (5.1 6 0.2) 4.0–4.7 (4.4 6 0.3) 4.2–5.1 (4.5 6 0.3) — 4.0 MBW (female) 5.2–6.4 (5.7 6 0.5) 3.7–5.0 (4.2 6 0.4) 3.4–5.9 (4.5 6 0.6) 3.8–4.8 (4.4 6 0.3) 4.2–5.4 (4.8 6 0.4) 3.2, 4.8 MBH (male) 4.0 2.9–4.5 (3.8 6 0.8) 3.2–4.7 (3.6 6 0.5) 3.0–5.1 (3.6 6 0.8) — 3.2 MBH (female) 4.3–4.8 (4.5 6 0.2) 3.0–4.7 (3.9 6 0.5) 2.9–5.3 (3.7 6 0.8) 2.9–5.0 (3.7 6 0.8) 3.4–4.4 (3.8 6 0.4) 2.5, 4.3 TL (male) 35.1 45.0–50.5 (47.8 6 3.9) 39.3–47.1 (43.6 6 3.4) 38.7–51.5 (43.2 6 7.2) — 45.2 TL (female) 35.3–50.4 (44.5 6 8.1) 41.0–47.1 (43.9 6 3.1) 38.3–50.1 (44.2 6 4.1) 32.3–52.1 (42.5 6 7.4) 45.6–53.5 (49.5 6 3.1) 38.3, 35.2 TW (male) 4.1 3.8–4.1 (4.0 6 0.2) 3.1–3.9 (3.4 6 0.2) 3.4–4.3 (3.7 6 0.3) — 3.7 TW (female) 4.1–4.6 (4.4 6 0.2) 3.3–3.8 (3.4 6 0.2) 2.8–4.6 (3.5 6 0.5) 2.8–4.2 (3.5 6 0.4) 3.6–4.5 (4.0 6 0.3) 2.4, 3.6 TH (male) 3.5 3.2–3.4 (3.3 6 0.1) 2.8–3.5 (3.1 6 0.2) 2.6–3.6 (3.0 6 0.4) — 2.6 TH (female) 3.5–3.9 (3.8 6 0.2) 2.9–3.4 (3.1 6 0.2) 2.5–4.5 (3.1 6 0.5) 2.3–3.4 (2.8 6 0.4) 2.7–3.9 (3.1 6 0.4) 2.3, 3.2 HL (male) 5.3 4.5–4.8 (4.6 6 0.2) 3.7–4.7 (4.1 6 0.4) 3.6–4.2 (3.8 6 0.2) — 3.9 HL (female) 4.8–5.2 (5.0 6 0.2) 3.5–4.4 (4.1 6 0.3) 3.5–4.5 (4.0 6 0.3) 3.5–5.3 (4.1 6 0.6) 3.8–5.5 (4.3 6 0.6) 3.1, 4.2 HW (male) 5.1 4.6–4.7 (4.6 6 0.1) 3.9–4.6 (4.2 6 0.2) 3.9–4.9 (4.2 6 0.4) — 4.0 HW (female) 4.6–5.8 (5.2 6 0.5) 3.8–4.6 (4.1 6 0.2) 3.6–5.2 (4.2 6 0.4) 3.7–5.4 (4.2 6 0.5) 4.0–5.3 (4.4 6 0.4) 3.5, 4.2 HH (male) 3.5 3.0–3.8 (3.4 6 0.4) 2.7–3.1 (2.9 6 0.2) 2.8–3.6 (3.0 6 0.3) — 2.9 HH (female) 3.4–4.2 (3.9 6 0.3) 2.9–3.4 (3.1 6 0.2) 2.5–3.9 (3.0 6 0.4) 2.5–3.9 (3.0 6 0.5) 2.9–3.9 (3.3 6 0.3) 2.5, 3.1 SnFa (male) 13.6 11.6–12.8 (12.0 6 0.6) 10.9–12.2 (11.6 6 0.4) 11.3–12.3 (11.8 6 0.3) — 11.9 SnFa (female) 11.7–14.1 (12.9 6 0.9) 10.1–11.9 (11.2 6 0.6) 10.9–12.5 (11.6 6 0.5) 10.6–13.0 (11.7 6 0.8) 11.3–13.2 (11.8 6 0.6) 11.5, 12.1 ED (male) 0.9 1.0–1.1 (1.0 6 0.1) 1.0–1.1 (1.0 6 0.0) 0.8–0.9 (0.9 6 0.0) — 0.9 ED (female) 0.8–1.0 (0.9 6 0.1) 0.8–1.1 (1.0 6 0.1) 0.9–1.1 (1.0 6 0.1) 0.8–0.9 (0.9 6 0.1) 0.9–1.1 (1.0 6 0.1) 1.0, 1.0 END (male) 1.9 1.1–1.7 (1.5 6 0.3) 1.5–1.9 (1.7 6 0.1) 1.6–1.9 (1.7 6 0.1) — 1.6 END (female) 1.8–2.2 (1.9 6 0.1) 1.6–1.8 (1.7 6 0.1) 1.5–1.9 (1.7 6 0.1) 1.4–1.9 (1.7 6 0.2) 1.6–1.9 (1.8 6 0.1) 1.7, 1.9 SNL (male) 2.7 1.8–2.5 (2.2 6 0.4) 2.1–2.5 (2.4 6 0.1) 2.2–2.5 (2.4 6 0.1) — 2.3 SNL (female) 2.6–2.9 (2.7 6 0.1) 2.2–2.6 (2.3 6 0.1) 2.2–2.6 (2.4 6 0.1) 2.1–2.7 (2.4 6 0.2) 2.3–2.6 (2.4 6 0.1) 2.4, 2.6 IND (male) 1.4 1.2–1.4 (1.3 6 0.1) 1.2–1.4 (1.3 6 0.1) 1.1–1.5 (1.3 6 0.1) — 1.1 IND (female) 1.1–1.4 (1.3 6 0.1) 1.2–1.4 (1.3 6 0.1) 1.1–1.5 (1.3 6 0.1) 1.0–1.4 (1.3 6 0.1) 1.1–1.5 (1.3 6 0.1) 1.1, 1.3 FLL (male) 2.3 1.4–1.7 (1.6 6 0.1) 1.1–1.5 (1.3 6 0.1) 1.1–1.6 (1.3 6 0.2) — 1.4 FLL (female) 1.1–2.6 (1.7 6 0.5) 1.2–1.7 (1.4 6 0.1) 1.1–1.8 (1.3 6 0.2) 1.1–1.9 (1.5 6 0.3) 1.4–2.1 (1.7 6 0.2) 1.1, 1.5 o 25 No. HLL (male) 2.9 2.3–3.0 (2.6 6 0.4) 2.1–2.4 (2.3 6 0.1) 2.4–2.8 (2.6 6 0.2) — 2.6 HLL (female) 2.5–3.1 (2.8 6 0.2) 2.3–2.9 (2.6 6 0.2) 2.0–2.7 (2.4 6 0.2) 1.9–3.1 (2.7 6 0.4) 2.5–3.6 (3.0 6 0.3) 2.1, 2.7 2011] HERPETOLOGICAL MONOGRAPHS 91

FIG. 4.—Illustration of head of adult female Brachymeles samarensis (KU 310849) and adult female B. brevidactylus (PNM 4856; formerly KU 324004) 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; Nu, nuchal; 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, AMF, and RMJ.

Diagnosis.—Brachymeles paeforum can be paravertebral scale rows 93–96, (11) pineal distinguished from congeners by the following eye spot present, (12) prefrontals not contact- combination of characters: (1) body size small ing on midline, (13) frontoparietals contact, (SVL 47.2–66.1 mm), (2) limbs tridactyl, (14) enlarged chin shields in three pairs, (15) (3) limb length short, (4) supralabials six, nuchals enlarged, (16) fourth supralabial (5) infralabials five or six, (6) supraciliaries six, below eye midline, (17) auricular opening (7) supraoculars five, (8) midbody scale rows absent, (18) presacral vertebrae 47, and (19) 21–22, (9) axilla–groin scale rows 71–74, (10) uniform body color (Tables 4 and 5). 92 HERPETOLOGICAL MONOGRAPHS [No. 25

FIG. 5.—Illustration of head of adult male holotype Brachymeles bicolandia (PNM 9756; formerly KU 324003) and adult female holotype Brachymeles cobos (PNM 9761; formerly KU 324023) in dorsal, lateral, and ventral views. Labels for taxonomically diagnostic head scales follow those shown in Fig. 4. Illustrations by CDS, AMF, and RMJ.

Comparisons.—Characters distinguishing lengths among males, longer relative forelimb Brachymeles paeforum from all nonpentadac- lengths, a greater number of paravertebral tyl, limbed species of Brachymeles are sum- scale rows, and a uniform body color (Tables 4 marized in Tables 4 and 5. Brachymeles and 5); from B. muntingkamay by having a paeforum most closely resembles B. libayani, shorter maximum body length, shorter fore- B. muntingkamay, and B. tridactylus, the only limb lengths, shorter hind limb lengths, a other tridactyl species, but differs from these greater number of axilla–groin scale rows, a three taxa by having five or six infralabials greater number of paravertebral scale rows, (Table 5). Brachymeles paeforum further dif- six supraciliaries, five supraoculars, the pres- fers from B. libayani by having longer body ence of a pineal eyespot, contact between 2011] HERPETOLOGICAL MONOGRAPHS 93

FIG. 6.—Illustration of head of adult female holotype Brachymeles paeforum (PNM 9746; formerly KU 311228) and adult male holotype Brachymeles libayani (PNM 9749; formerly KU 320466) in dorsal, lateral, and ventral views. Labels for taxonomically diagnostic head scales follow those shown in Fig. 4. Illustrations by CDS, AMF, and RMJ. frontoparietals, differentiated nuchals, and a frontoparietals, the presence of a continuous continuous subocular scale row, and the subocular scale row, and the absence of absence of contact between prefrontals and longitudinal rows of spots around the body longitudinal rows of spots around the body (Tables 4 and 5). (Tables 4 and 5); and from B. tridactylus by Brachymeles paeforum can be distinguished having a shorter maximum body length, from all limbless species of Brachymeles (B. shorter relative tail length, shorter forelimb apus, B. lukbani, B. minimus, B. miriamae, B. length, a greater number of presacral verte- vermis) by having limbs; and from all penta- brae, six supralabials, six supraciliaries, five dactyl species of Brachymeles (B. boholensis, supraoculars, the presence of contact between B. boulengeri, B. bicolor, B. gracilis, B. kadwa, 94 HERPETOLOGICAL MONOGRAPHS [No. 25

B. makusog, B. mindorensis, B. orientalis, B. preocular one; presubocular one; supracili- schadenbergi, B. talinis, B. taylori, B. tungaoi, aries six, the anteriormost contacting prefron- B. vindumi) by having nonpentadactyl limbs, tal and separating posterior loreal from first shorter forelimb lengths (less than 1.7 mm vs. supraocular, posteriormost extending to mid- greater than 5.9 mm), shorter hind limb lengths dle of fifth supraocular; subocular scale row (less than 3.0 mm vs. greater than 10.3 mm), a single, complete, in contact with supralabials; narrower body (less than 5.3 mm vs. greater lower eyelid with one row of scales; suprala- than 7.9 mm), and by the absence of a postnasal bials six, first twice the anteroposterior length scale and auricular opening (vs. presence). of others, fourth below eye midline; infra- Description of holotype.—Details of the labials six (Fig. 6). head scalation are shown in Fig. 7. Adult Mental wider than long, in contact with first female, body small, slender, SVL 59.1 mm; infralabials; postmental single, enlarged, its head weakly differentiated from neck, nearly width greater than width of mental; followed as wide as body, HW 6.5% SVL, 105.5% HL; by three pairs of enlarged chin shields, first HL 31.5% SnFa; SnFa 19.5% SVL; snout pair in broad medial contact, equal in width to short, bluntly rounded in dorsal and lateral third pair, second pair wider than first and profile, SNL 71.2% HL; ear completely third, broadly separated by single medial hidden by scales; eyes small, ED 1.7% SVL, scale, third pair separated by three medial 27.5% HL, 56.2% END, pupil subcircular; scales (Fig. 6). body slightly depressed, nearly uniform in Scales on limbs smaller than body scales; thickness, MBW 126.7% MBH; scales smooth, scales on dorsal surfaces of digits wrapping glossy, imbricate; longitudinal scale rows at around lateral edges of digits; lamellae absent; midbody 22; paravertebral scale rows 93; palmar surfaces of hands and plantar surfaces axilla–groin scale rows 71; limbs short, poorly of feet with several small, irregular scales, developed, with digits reduced to three claws each with irregular raised anterior edges; on both forelimbs and hind limbs, finger and fingers equal in size; toes unequal in size, toe lamellae absent; FLL 2.7% AGD, 2.1% middle digit greatest in length, first and third SVL; HLL 5.2% AGD, 4.1% SVL; tail not as digits equal in length. wide as body, gradually tapered towards end, Coloration in preservative.—Body ground TW 73.7% MBW, TL 69.4% SVL. color medium brown, each dorsal scale light Rostral projecting onto dorsal snout to level brown posteriorly, with a dark auburn streak on in line with posterior edge of nasal, broader the anterior two-thirds to half of the scale. Dark than high, in contact with frontonasal; fronto- streaks on each scale consist of four to six thin nasal wider than long; nostril ovoid, in center longitudinal lines with smudges between lines. of single trapezoidal nasal, longer axis directed Streaks on scales present around entire body, anteroventrally and posterodorsally; suprana- more distinct on venter, where posterior ends sals present, large, broadly separated; post- of scales are cream, giving greater contrast. nasals absent; prefrontals broadly separated; Subcaudal scales with reduced dark pigmenta- frontal nearly diamond-shaped, its anterior tion. Posterior edge of all body scales trans- margin in moderate contact with frontonasal, parent. Forelimb and hind limb scales dark in contact with first two anterior supraoculars, brown. Forelimb scales with weakly defined 43 wider than anterior supraocular; suprao- scale boundaries. Precloacal scale coloration culars five; frontoparietals moderate, in nar- lighter than surrounding ventral scale colora- row medial contact, each frontoparietal in tion. Head scales mottled light and dark brown, contact with supraoculars 2–4; interparietal matching dorsal background coloration. Ros- moderate, its length roughly equal to midline tral, nasal, supranasal, and first supralabial length of frontoparietal, longer than wide, scales cream, lighter than supraocular scales. diamond-shaped, wider anteriorly; parietals as Supraocular scales light gray-umber. Suprala- broad as frontoparietals, in broad contact bial, infralabial, postmental, and chin shields behind interparietal; enlarged nuchals pres- scales beige with slight light brown mottling. ent; loreals two, anterior loreal about as long Coloration in life.—Coloration in life (Fig. 7) as and slightly higher than posterior loreal; closely matches coloration in preservative with 2011] HERPETOLOGICAL MONOGRAPHS 95

it does not qualify for Critically Endangered, Endangered, Vulnerable, or Near Threatened status. Brachymeles paeforum has been doc- umented to have a broad geographic distribu- tion on Leyte Island, and has been observed to be moderately abundant in a variety of habitats. We therefore classify this species as one of Least Concern (IUCN, 2011). Other lizard species observed in sympatry on Leyte Island include the following: (Agami- dae) Bronchocela cristatella, Draco bimaculatus, FIG. 7.—Photograph in life of Brachymeles paeforum D. ornatus, D. reticulates, Gonocephalus sem- (PNM 9746; formerly KU 311228), SVL 5 59.1 mm. Photograph by CDS. peri, Hydrosaurus pustulatus; (Gekkonidae) Cyrtodactylus annulatus, C. gubaot, Gehyra minor differences, including a dark brown body mutilata, Gekko gecko, Gek. mindorensis, Hemi- color and dark brown to black streaks of dactylus frenatus, H. platyurus, Hemiphyllodac- pigmentation. tylus typus, Lepidodactylus aureolineatus, L. Variation.—Morphometric variation of the planicaudus, Pseudogekko compressicorpus; series is summarized in Table 6. We observed a (Scincidae) Emoia atrocostata, Eutropis multi- single instance of digit variation in which one carinata, Eu. multifasciata, Lamprolepis smar- specimen (KU 311225) has two toes. The pineal agdina, Lipinia pulchella, Li. quadrivittata, eyespot was absent in a single specimen (CAS Sphenomorphus acutus, S. cumingi, S. fasciatus, 26120) and present in all others. The number of S. jagori, S. cf. mindanensis, S. steerei, S. infralabials ranged from nearly 70% (11/16) of variegatus, Tropidophorus misaminus;(Varani- the specimens possessing six infralabials (CAS- dae) Varanus cumingi samarensis. SU 26110, 26112, 26123, 26772, 26120, 26771, Etymology.—CDS is pleased to name this KU 311224–8), to five specimens possessing five new species in honor of the Philippine- infralabials (CAS-SU 26121–2, 26770, 26773, American Education Foundation (PAEF), in KU 311229). Five specimens have enlarged honor of their continued support and contri- mental scales resulting from fusion with the first butions to our Philippine biodiversity research infralabial (CAS-SU 26121–2, 26770, 26773, KU program. As the Fulbright Commission in the 311229). All specimens have two loreals on each Philippines, PAEF is responsible for leading side of the head, with the exception of four the advancement of international exchange specimens, which have single, enlarged loreals programs between the United States and the on both sides of the head (KU 311227) or on Philippines, with the mission of promoting only the left side (KU 311224–5, 311229). mutual understanding between citizens of Distribution.—Brachymeles paeforum is both countries. Suggested common name: known only from Leyte Island (Fig. 3). The PAEF Slender . Ecology and natural history.—Brachymeles Brachymeles libayani sp. nov. paeforum occurs in agricultural habitats, as Figs. 3, 6, 8 well as in disturbed and secondary-growth forest. Little lower-elevation original forest Holotype.—PNM 9749 (CDS Field No. remains on Leyte Island, but we assume the 3700, formerly KU 320466), adult male, species once also occurred in first-growth collected under rotting coconut husks in forest at low elevations. Individuals have been secondary-growth forest (1000 to 1230 h) on observed in the rotting material within fallen 21 March 2009, in Barangay Villa Milagrosa, logs and in leaf litter surrounding the root Municipality of President Carlos P. Garcia, networks of trees. Similar to B. samarensis, Bohol Province, Lapinig Grande Island, Phi- this species is found in sympatry with B. lippines (10u079160N, 124u349300E; WGS-84), orientalis and B. gracilis hilong. by CDS and J. Fernandez. We have evaluated this species against the Paratopotypes.—Nine adult males (KU IUCN criteria for classification, and find that 320435, 320444–6, 320451, 320462, 320466, 96 HERPETOLOGICAL MONOGRAPHS [No. 25

(SVL 52.7–66.1 mm), (2) limbs tridactyl, (3) limb length short, (4) supralabials six, (5) infralabials five, (6) supraciliaries six, (7) supraoculars five, (8) midbody scale rows 22–23, (9) axilla–groin scale rows 72–75, (10) paravertebral scale rows 90–92, (11) pineal eye spot present, (12) frontoparietals contact, (13) enlarged chin shields in three pairs, (14) nuchal scales differentiated, (15) fourth su- pralabial below eye midline, (16) auricular opening covered with scales, (17) presacral vertebrae 47, (18) fusion of mental and first infralabials, and (19) nonuniform body color (Tables 4 and 5). Comparisons.—Characters distinguishing Brachymeles libayani from all nonpentadactyl, limbed species of Brachymeles are summa- rized in Tables 4 and 5. Brachymeles libayani most closely resembles B. paeforum, B. muntingkamay, and B. tridactylus, the only other tridactyl species, but differs from these FIG. 8.—Photographs in life of Brachymeles libayani three taxa by having five infralabials and (PNM 9749; formerly KU 320466), SVL 5 56.3 mm. fusion of the mental and first infralabials Photographs by CDS. (Table 5). Brachymeles libayani further dif- fers from B. paeforum by having shorter body PNM 9754–55), 21 adult females (KU 320428– lengths among males, shorter relative forelimb 30, 320438–40, 320442, 320447, 320449–50, lengths, fewer paravertebral scale rows, and a 320452–5, 320457–61, 320463, 320467), and nonuniform body color (Tables 4 and 5); from 10 juveniles (KU 320436–7, 320441, 320443, B. muntingkamay by having a shorter maxi- 320448, 320456, PNM 9750–53) collected mum body length, shorter forelimb lengths, between 19 and 21 March 2009. shorter hind limb lengths, a greater number of Other paratypes.—One adult male (CAS- axilla–groin scale rows, six supraciliaries, five SU 28453) collected on 11 April 1967, under supraoculars, the presence of a pineal eyespot, rotting coconut tree, 0.5 km SW of Barrio contact between frontoparietals, differentiat- Pitogo, in the Municipality of Ubay, Bohol ed nuchals, and a continuous subocular scale Province, Lapinig Grande Island, Philippines row, and the absence of longitudinal rows (10u079050N, 124u339040E; WGS-84), by A. C. of spots around the body (Tables 4 and 5); Alcala; one adult female (CAS-SU 27554) and from B. tridactylus by having a shorter collected on 15 April 1967, under rotting log maximum body length, shorter relative tail in secondary-growth forest, in the Municipal- length, shorter forelimb length, shorter hind ity of Ubay, Bohol Province, Polong Dako limb lengths, a greater number of presacral Island, Philippines (10u049110N, 124u309140E; vertebrae, six supralabials, five infralabials, six WGS-84), by A. C. Alcala; three adult females supraciliaries, five supraoculars, the presence (CAS-SU 27556, 28454–5) collected on 20 of contact between frontoparietals, the pres- April 1967, under rotting logs and leaves in ence of a continuous subocular scale row, and a patch of secondary-growth trees, in the the absence of longitudinal rows of spots Municipality of Ubay, Bohol Province, Lapi- around the body (Tables 4 and 5). nig Chico Island, Philippines (10u059220N, Brachymeles libayani can be distinguished 124u309320E; WGS-84), by A. C. Alcala. from all limbless species of Brachymeles (B. Diagnosis.—Brachymeles libayani can be apus, B. lukbani, B. minimus, B. miriamae, B. distinguished from congeners by the following vermis) by having limbs; and from all penta- combination of characters: (1) body size small dactyl species of Brachymeles (B. boholensis, 2011] HERPETOLOGICAL MONOGRAPHS 97

B. boulengeri, B. bicolor, B. gracilis, B. kadwa, contact behind interparietal; enlarged nuchals B. makusog, B. mindorensis, B. orientalis, B. present; loreals two, anterior loreal about as schadenbergi, B. talinis, B. taylori, B. tungaoi, long as and slightly higher than posterior B. vindumi) by having nonpentadactyl limbs, loreal; preocular one; presubocular one; shorter forelimb lengths (less than 1.8 mm vs. supraciliaries six, the anteriormost contacting greater than 5.9 mm), shorter hind limb prefrontal and separating posterior loreal from lengths (less than 2.7 mm vs. greater than first supraocular, posteriormost extending to 10.3 mm), a narrower body (less than 5.9 mm middle of fifth supraocular; subocular scale vs. greater than 7.9 mm), and by the absence row single, complete, in contact with suprala- of a postnasal scale and auricular opening (vs. bials; lower eyelid with one row of scales; presence). supralabials six, first twice the anteroposterior Description of holotype.—Details of the length of others, fourth below eye midline; head scalation are shown in Fig. 7. Adult infralabials five (Fig. 6). male, hemipenes everted; body small, slender, Mental wider than long, fused with first SVL 56.3 mm; head weakly differentiated infralabials on both sides; postmental single, from neck, nearly as wide as body, HW 6.5% enlarged, its width greater than width of SVL, 112.7% HL; HL 31.5% SnFa; SnFa mental; followed by three pairs of enlarged 20.9% SVL; snout short, bluntly rounded in chin shields, first pair in broad medial contact, dorsal and lateral profile, SNL 65.0% HL; ear equal in width to third pair, second pair wider completely hidden by scales; eyes small, ED than first and third, separated by single medial 1.8% SVL, 28.0% HL, 60.8% END, pupil scale, third pair separated by three medial subcircular; body slightly depressed, nearly scales (Fig. 6). uniform in thickness, MBW 120.3% MBH; Scales on limbs smaller than body scales; scales smooth, glossy, imbricate; longitudinal scales on dorsal surfaces of digits wrapping scale rows at midbody 22; paravertebral scale around lateral edges of digits; lamellae absent; rows 90; axilla–groin scale rows 73; limbs palmar surfaces of hands and plantar surfaces short, poorly developed, with digits reduced to of feet with several small, irregular scales, three claws on both forelimbs and hind limbs, each with irregular raised anterior edges; finger and toe lamellae absent; FLL 3.3% fingers equal in size; toes unequal in size, AGD, 2.5% SVL; HLL 5.3% AGD, 3.9% SVL; middle digit greatest in length, first and third tail not as wide as body, gradually tapered digits equal in length. towards end, TW 81.5% MBW, TL 83.6% Coloration in preservative.—Body ground SVL. color dark brown, each dorsal scale light brown Rostral projecting onto dorsal snout to level posteriorly, with a dark chocolate-brown streak in line with posterior edge of nasal, broader on the anterior one-third to half of the scale. than high, in contact with frontonasal; fronto- Dark streaks on each scale consist of four to nasal wider than long; nostril ovoid, in center seven thin longitudinal lines with smudges of single trapezoidal nasal, longer axis directed between lines. Streaks on scales present around anteroventrally and posterodorsally; suprana- entire body, less dominant on venter. Subcau- sals present, large, broadly separated; post- dal scales darker brown than venter. Posterior nasals absent; prefrontals moderately separat- edge of all body scales transparent. Forelimb ed; frontal nearly diamond-shaped, its anterior and hind limb scales dark brown. Precloacal margin in moderate contact with frontonasal, scale coloration matches surrounding ventral in contact with first two anterior supraoculars, scale coloration. Head scales mottled light and 43 wider than anterior supraocular; suprao- dark brown, matching dorsal background col- culars five; frontoparietals moderate, in mod- oration. Rostral, nasal, supranasal, and first erate medial contact, each frontoparietal in supralabial scales light gray. Supraocular and contact with interior two supraoculars; inter- remaining supralabial scales dark brown. Men- parietal moderate, its length roughly equal to tal scale cream. Chin shields and postmental midline length of frontoparietal, longer than scales match bordering ventral scales. wide, kite-shaped, wider anteriorly; parietals Coloration in life.—Coloration in life (Fig. 8) as broad as frontoparietals, in moderate closely matches coloration in preservative with 98 HERPETOLOGICAL MONOGRAPHS [No. 25 minor differences. Head scales mottled medium Brachymeles (C. D. Siler, personal observa- brown to dark brown or black. Ground color of tion). Individuals were regularly observed body medium to dark brown. Streaks of under piles of rotting coconuts, in loose soil pigmentation on each scale dark brown to black. around trees and root systems, and in loose leaf Variation.—Morphometric variation of the litter. Interestingly, this species seems to be a series is summarized in Table 6. We observed ubiquitous habitat generalist on the Lapinig variation in the degree of head scale contact: Group Islands, but has not been recorded from (1) prefrontals were observed in point contact the nearby island of Bohol, whereas its medially in seven specimens (KU 320444, congeners, B. orientalis and B. boholensis, are 320449, 320458–60, 320462, 320465), and known. It remains possible that populations of separated for the remaining specimens (CAS- this species will eventually be discovered on SU 27554, 27556, 28453–5, KU 320428–43, the northeast coast of Bohol Island. 320445–8, 320450–57, 320461, 320463–4, We have evaluated this species against the 320466–7); (2) frontoparietals were observed IUCN criteria for classification, and find that it in point contact for a single specimen (KU does not qualify for Critically Endangered, 320467), and separated for the remaining Endangered, Vulnerable, or Near Threatened specimens (CAS-SU 27554, 27556, 28453–5, status. Brachymeles libayani has been docu- KU 320428–66); (3) first pair of enlarged chin mented to occur throughout the islands of the shields were observed separated in a single Lapinig Island Group, and is abundant at all specimen (KU 320451), and in contact medi- sampled localities. We therefore classify this ally for the remaining specimens (CAS-SU species as one of Least Concern (IUCN, 2011). 27554, 27556, 28453–5, KU 320428–50, Sympatric lizard species observed in the 320452–67). Lapinig Group islands include (Gekkonidae) Additionally, the degree of fusion between Hemidactylus frenatus and H. platyurus. loreals on each side of the head was observed Etymology.—We are pleased to name this to vary among the type series. The majority of new species in honor of Carlos Polestico specimens have two loreals and no fused Garcia. Born on Bohol Island, Carlos P. Garcia scales (CAS-SU 27554, 27556, 28453–55, KU later became the eighth President of the 320431–7, 320439–42, 320445–46, 320448, Philippines. He was the first Philippine pres- 320450–51, 320455, 320457–60, 320462, ident to be laid to rest in the Libingan ng mga 320464, 320466), five specimens have single, Bayani, or Cemetery of the Heroes, located enlarged loreals on both sides of the head (KU within Fort Bonifacio in Manila, Philippines. 320444, 320449, 320452, 320456, 320467), The municipality of Carlos P. Garcia, and type five specimens have single, enlarged loreals on locality for Brachymeles libayani, was named the left side of the head (KU 320430, 320453– after this Philippine hero. The word ‘‘libayani’’ 54, 320463, 320465), and six specimens have is derived from the phrase Libingan ng mga single, enlarged loreals on the right side of the Bayani. Suggested common name: Lapinig head (KU 320428–29, 320438, 320443, Islands’ Slender Skink. 320447, 320461). Distribution.—Brachymeles libayani is Brachymeles bicolandia sp. nov. known from Lapinig Chico, Lapinig Grande, Figs. 3, 5 Tilmubo, and Tintiman islands off the north- Holotype.—PNM 9756 (CDS Field No. east coast of Bohol Island (Fig. 3). 4050, formerly KU 324003), adult male, Ecology and natural history.—Brachymeles collected under rotting coconut husks in libayani occurs in agricultural habitats, and secondary-growth forest (1000 to 1230 h) on disturbed forest habitat. No original forest 1 June 2009, in Barangay Common, Munici- remains on any of the Lapinig Group Islands, pality of Tabaco City, Albay Province, Luzon and common habitat consists of grassland, rice Island, Philippines (13u149N, 123u389E; fields, agricultural habitats, and human habita- WGS-84), by CDS and J. Fernandez. tions. Surprisingly, B. libayani on Lapinig Paratopotypes.—Four adult males (KU Grande Island was observed to be more 324015–6, PNM 9759–60), six adult females common than any other known species of (KU 323087, 324005–7, 324009–10), and four 2011] HERPETOLOGICAL MONOGRAPHS 99 juveniles (KU 324008, 324011, PNM 9757– scale rows (Tables 4 and 5); and from B. 58), collected between 1 and 23 June 2009. bonitae by having longer relative fore- and Other paratypes.—One adult male (CAS hind limb lengths, six supralabials, five or six 152025) and two adult females (CAS 140065, infralabials, six supraciliaries, five supraocu- 152026) collected on 16 December 1991, in lars, and a tendency towards fewer presacral an Abaca plantation, Barangay Labnig, Mu- vertebrae, axilla–groin, and paravertebral nicipality of Malinao, Albay Province, Luzon scale rows (Tables 4 and 5). Island, Philippines (13u229380N, 123u409590E; Brachymeles bicolandia can be distin- WGS-84), by C. A. Ross; two adult females guished from all limbless species of Brachy- (CAS-SU 24173, 24413) collected between 26 meles (B. apus, B. lukbani, B. minimus, B. March and 22 April 1961, on Mt. Isarog, Bario miriamae, B. vermis) by having limbs; and Curry, Municipality of Pili, Camarines Sur from all pentadactyl species of Brachymeles Province, Luzon Island, Philippines (13u389350N, (B. boholensis, B. boulengeri, B. bicolor, B. 123u21940E; WGS-84), by D. S. Rabor. gracilis, B. kadwa, B. makusog, B. mind- Diagnosis.—Brachymeles bicolandia can be orensis, B. orientalis, B. schadenbergi, B. distinguished from congeners by the following talinis, B. taylori, B. tungaoi, B. vindumi)by combination of characters: (1) body size small having nonpentadactyl limbs, shorter forelimb (SVL 46.4–66.1 mm), (2) limbs bidactyl, (3) lengths (less than 1.9 mm vs. greater than limb length short, (4) supralabials six, (5) 5.9 mm), shorter hind limb lengths (less than infralabials five or six, (6) supraciliaries six, (7) 3.1 mm vs. greater than 10.3 mm), a narrower supraoculars five, (8) midbody scale rows 20– body (less than 5.1 mm vs. greater than 22, (9) axilla–groin scale rows 68–73, (10) 7.9 mm), and by the absence of a postnasal paravertebral scale rows 85–90, (11) pineal scale and auricular opening (vs. presence). eye spot present, (12) prefrontals not contact- Description of holotype.—Details of the ing on midline, (13) postnasals absent, (14) head scalation are shown in Fig. 6. Adult enlarged chin shields in three pairs, (15) male, body small, slender, SVL 60.2 mm; head nuchals enlarged, (16) fourth supralabial below weakly differentiated from neck, nearly as eye midline, (17) mental and first infralabials wide as body, HW 6.7% SVL, 104.9% HL; HL fused or separated, (18) auricular opening 33.1% SnFa; SnFa 19.3% SVL; snout short, absent, (19) presacral vertebrae 46–49, and bluntly rounded in dorsal and lateral profile, (20) uniform body color (Tables 4 and 5). SNL 58.2% HL; ear completely hidden by Comparisons.—Characters distinguishing scales; eyes small, ED 1.4% SVL, 22.6% HL, Brachymeles bicolandia from all nonpenta- 55.4% END, pupil subcircular; body slightly dactyl, limbed species of Brachymeles are depressed, nearly uniform in thickness, MBW summarized in Tables 4 and 5. Brachymeles 124.0% MBH; scales smooth, glossy, imbri- bicolandia most closely resembles B. samar- cate; longitudinal scale rows at midbody 22; ensis, B. cobos, B. brevidactylus, and popula- paravertebral scale rows 85; axilla–groin scale tions of B. bonitae, the only other species to rows 68; limbs short, poorly developed, with be bidactyl or have bidactyl populations. digits reduced to two claws on both forelimbs However, B. bicolandia differs from these and hind limbs, finger and toe lamellae four taxa by having five or six infralabials absent; FLL 2.7% AGD, 2.1% SVL; HLL (Table 5). Brachymeles bicolandia further 6.1% AGD, 4.7% SVL [6.9]; tail nearly as wide differs from B. samarensis by having a greater as body, gradually tapered at end, TW 88.9% number of presacral vertebrae, and a tenden- MBW, TL 65.6% SVL. cy toward a greater number of midbody, Rostral projecting onto dorsal snout to level axilla–groin, and paravertebral scale rows just past posterior edge of nasal, broader than (Table 5); from B. cobos by having a smaller high, in broad contact with frontonasal; relative forelimb length and a greater number frontonasal wider than long; nostril ovoid, in of presacral vertebrae (Tables 4 and 5); from center of single trapezoidal nasal, longer axis B. brevidactylus by having a smaller relative directed anteroventrally and posterodorsally; forelimb length, and a tendency towards supranasals present, large, broadly separated; having fewer axilla–groin and paravertebral postnasals absent; prefrontals moderately 100 HERPETOLOGICAL MONOGRAPHS [No. 25 separated; frontal octagonal, its anterior mar- body scales transparent. Forelimb and hind gin in moderate contact with frontonasal, in limb scales dark brown, with weakly defined contact with first two anterior supraoculars, scale boundaries. Precloacal scale coloration 53 wider than anterior supraocular; suprao- darker than surrounding ventral scale colora- culars five; frontoparietals moderate, just tion. Head scales mottled light and dark brown, barely separated by anterior point of interpa- matching dorsal background coloration. Rostral, rietal in contact with frontal, each frontopari- nasal, supranasal, and first supralabial scales etal in contact with supraoculars 2–4; inter- cream, lighter than supraocular scales. Suprao- parietal large, its length roughly 1.53 midline cular scales dark brown. Mental, infralabial, length of frontoparietal, longer than wide, postmental, and chin shields scales cream with diamond-shaped, wider anteriorly; parietals as light brown mottling. broad as frontoparietals, in broad contact Coloration in life.—Coloration in life unre- behind interparietal; enlarged nuchals pres- corded; however, because Brachymeles spec- ent; anterior and posterior loreals fused into imens do not change significantly during single, enlarged loreal (Fig. 5), or distinct; preservation (C. D. Siler and R. M. Brown, preocular one; presubocular one; supracili- personal observation), we suspect that the aries six, the anteriormost contacting prefron- preserved coloration and patterns are much tal and separating posterior loreal from first like those in life. supraocular, posteriormost extending to mid- Variation.—Morphometric variation of the dle of fifth supraocular; subocular scale row series is summarized in Table 6. A single single, complete, in contact with supralabials; instance of digit variation was observed, in lower eyelid with one row of scales; suprala- which one specimen (KU 324006) has three bials six, first 1.53 size of other supralabials, forelimb claws and two hind limb claws. We fourth below eye midline; infralabials five observed variation in the degree of head scale (Fig. 5). contact and the number of infralabials: (1) Mental wider than long, fused with first frontoparietals were observed in point contact infralabials; postmental single, enlarged, its medially for a single specimen (KU 324003), width less than width of mental; followed by in moderate to broad contact medially for 11 three pairs of enlarged chin shields, first pair in specimens (CAS-SU 24173, 24413, CAS broad medial contact, greater in width than third 140065, KU 323087, 324006–8, 324011–3, pair, narrower than second pair, second pair 324018), and separated for eight specimens broadly separated by single medial scale, third (CAS 152025–6, KU 324005, 324014–6, pair separated by three medial scales (Fig. 5). 324009–10); (2) parietals were observed in Scales on limbs smaller than body scales; point contact medially for a single specimen scales on dorsal surfaces of digits wrapping (CAS-SU 24413), in moderate to broad around lateral edges of digits; lamellae absent; contact medially for 17 specimens (CAS-SU palmar surfaces of hands and plantar surfaces 24173, CAS 140065, CAS 152025–6, KU of feet with several small, irregular scales, 323087, 324003, 324005–9, 324011, 324013– each with irregular raised anterior edges; 6, 234018), and separated for two specimens fingers equal in size; toes unequal in size, (KU 324010, 324012); (3) first pair of enlarged second digit greatest in length. chin shields were observed in point contact Coloration in preservative.—Body ground medially for a single specimen (CAS 152025), color medium brown, each dorsal scale light in moderate to broad contact medially for 17 brown posteriorly, with a dark auburn streak specimens (CAS-SU 24173, 24413, CAS on the anterior two-thirds to half of the scale. 140065, 152026, KU 324003, 324005–13, Dark streaks on each scale consist of four to 324016, 324018), and separated in three five thin longitudinal lines with smudges specimens (KU 324014–5, 323087); (4) the between lines. Streaks on scales present number of infralabials varied among the type around entire body, more distinct on venter, series, with eight specimens observed to have where posterior ends of scales are sandy brown, six infralabials (CAS-SU 24413, KU 324009– giving greater contrast. Subcaudal scales with 14, 324018), and 12 observed to have five reduced dark pigmentation. Posterior edge of all infralabials (CAS-SU 24173, CAS 140065, 2011] HERPETOLOGICAL MONOGRAPHS 101

152025–6, KU 323087, 324003, 324005–8, steerei, Tropidophorus grayi; (Varanidae) Var- 324015–6). anus marmoratus, V. olivaceus. Additionally, the degree of fusion between Etymology.—The specific epithet is chosen loreals, and between the mental and first in reference to the biogeographically and infralabials, varies in the type series. All culturally distinct Bicol Region of southern specimens have two loreals on each side of the Luzon Island (Albay, Camarines Norte, Ca- head, with the exception of a single specimen marines Sur, Catanduan˜ es and Sorsogon (KU 324003) with single, enlarged loreals on provinces). Inhabited by peaceful and partic- both sides of the head. Eleven specimens have ularly hospitable Bicolanos, the unique central an enlarged mental scale resulting from fusion Bicol peninsula is home to many dozens of with the first infralabial on both sides of the head endemic vertebrates, delicious local cuisine, (CAS-SU 24173, CAS 152025–6, KU 323087, unique linguistic stock, and rich cultural 324003, 324005–8, 324015–6), a single specimen traditions. Suggested common name: Bicol has fused scales only on the right side of the head Slender Skink. (CAS 140065), and eight specimens have distinct mentals and infralabials, with no scale fusion Brachymeles cobos sp. nov. (CAS-SU 24413, KU 324009–14, 324018). Figs. 3, 5 Distribution.—Brachymeles bicolandia is Holotype.—PNM 9761 (CDS Field No. known only from the central Bicol Peninsula 5255, formerly KU 324023), adult female, of Luzon Island (Fig. 3). collected under rotting coconut husks in Ecology and natural history.—Brachymeles secondary-growth forest (1000 to 1230 h) on bicolandia occurs in agricultural habitats, as 8 October 2009, in Barangay Palta Small, well as in disturbed and secondary-growth Municipality of Virac, Catanduan˜ es Province, forest, and is found in sympatry with B. Catanduan˜ es Island, Philippines (13u349440N, boulengeri and B. makusog. Three additional 124u139520E; WGS-84), by J. Fernandez. species of Brachymeles have also been record- Paratopotypes.—Eight adult females (KU ed from the Bicol Peninsula of Luzon Island: B. 306311, 308077, 324019–20, 324025–26, brevidactylus, B. kadwa, and B. lukbani. PNM 9562–63) and one juvenile (KU We have evaluated this species against the 324021) collected between 4 and 7 June IUCN criteria for classification, and find that it 2009 by CDS and J. Fernandez. does not qualify for Critically Endangered, Diagnosis.—Brachymeles cobos can be dis- Endangered, Vulnerable, or Near Threatened tinguished from congeners by the following status. Brachymeles bicolandia has been doc- combination of characters: (1) body size small umented to occur throughout much of the (SVL 54.0–64.4 mm), (2) limbs bidactyl, (3) limb central Bicol Peninsular of Luzon Island, and length short, (4) supralabials six, (5) infralabials is abundant at all sampled localities. We six, (6) supraciliaries six, (7) supraoculars five, therefore classify this species as one of Least (8) midbody scale rows 21–22, (9) axilla–groin Concern (IUCN, 2011). scale rows 68–72, (10) paravertebral scale rows Sympatric lizard species occurring in the 85–89, (11) pineal eye spot present, (12) Bicol Peninsula include the following: (Agami- prefrontals not contacting on midline, (13) dae) Bronchocela cristatella, Draco spilopterus, frontoparietals contact, (14) postnasals absent, Gonocephalus sophiae, Hydrosaurus pustula- (15) enlarged chin shields in three pairs, (16) tus; (Gekkonidae) Cyrtodactylus philippinicus, nuchals enlarged, (17) fourth supralabial below Hemidactylus frenatus, H. platyurus, Gehyra eye midline, (18) auricular opening absent, (19) mutilata, Gekko gecko, Gek. mindorensis, presacral vertebrae 45, (20) nonfusion of mental Luperosaurus cumingii, Pseudogekko compres- and first infralabials, (21) nonfusion of loreals, sicorpus, P. smaragdina; (Scincidae) Emoia and (22) uniform body color (Tables 4 and 5). atrocostata, Eutropis multicarinata borealis, Comparisons.—Characters distinguishing Eu. multifasciata, Lamprolepis smaragdina, Brachymeles cobos from all nonpentadactyl, Lipinia pulchella pulchella, Sphenomorphus limbed species of Brachymeles are summa- abdictus abdictus, S. cumingi, S. decipiens, S. rized in Tables 4 and 5. Brachymeles cobos jagori, S. laterimaculatus, S. leucospilos, S. most closely resembles B. samarensis, B. 102 HERPETOLOGICAL MONOGRAPHS [No. 25 bicolandia, B. brevidactylus, and populations scale rows 68; limbs short, poorly developed, of B. bonitae, the only other species to be with digits reduced to two claws on both bidactyl or have bidactyl populations. Howev- forelimbs and hind limbs, finger and toe er, B. cobos can be distinguished from B. lamellae absent; FLL 3.4% AGD, 2.6% SVL; samarensis by having a tendency toward a HLL 6.2% AGD, 4.7% SVL; tail as wide as greater number of axilla–groin scale rows body, tail tip regenerated, sharply tapered at (Table 5); from B. bicolandia by having a end, TW 87.6% MBW, TL 75.6% SVL. greater relative forelimb length, fewer presa- Rostral projecting onto dorsal snout to level cral vertebrae, and six infralabials (Tables 4 in line with middle of nasal, broader than and 5); from B. brevidactylus by having fewer high, in contact with frontonasal; frontonasal presacral vertebrae, a greater number of wider than long; nostril ovoid, in center of midbody scale rows, fewer axilla–groin and single trapezoidal nasal, longer axis directed paravertebral scale rows, and the presence of anteroventrally and posterodorsally; suprana- contact between the first pair of enlarged chin sals present, large, broadly separated; post- shields (Table 5); and from B. bonitae by nasals absent; prefrontals moderately separat- having longer relative fore- and hind limb ed; frontal octagonal-shaped, its anterior lengths, fewer presacral vertebrae, fewer margin in moderate contact with frontonasal, axilla–groin and paravertebral scale rows, six in contact with first two anterior supraoculars, supralabials, six infralabials, six supraciliaries, 43 wider than anterior supraocular; suprao- five supraoculars, the presence of contact culars five; frontoparietals large, in broad between frontoparietals, and the presence of medial contact, each frontoparietal in contact contact between the first pair of enlarged chin with supraoculars 2–4; interparietal moderate, shields (Tables 4 and 5). its length equal to midline length of fronto- Brachymeles cobos can be distinguished parietal, longer than wide, diamond-shaped, from all limbless species of Brachymeles (B. wider anteriorly; parietals narrower than apus, B. lukbani, B. minimus, B. miriamae, B. frontoparietals, in broad contact behind inter- vermis) by having limbs; and from all penta- parietal; nuchals enlarged; loreals two, anteri- dactyl species of Brachymeles (B. boholensis, B. or loreal about as long as and slightly higher boulengeri, B. bicolor, B. gracilis, B. kadwa, B. than posterior loreal; preocular one; presubo- makusog, B. mindorensis, B. orientalis, B. cular one; supraciliaries six, the anteriormost schadenbergi, B. talinis, B. taylori, B. tungaoi, contacting prefrontal and separating posterior B. vindumi) by having nonpentadactyl limbs, loreal from first supraocular, posteriormost shorter forelimb lengths (less than 2.1 mm vs. extending nearly to middle of fifth supraocu- greater than 5.9 mm), shorter hind limb lengths lar; subocular scale row single, complete, in (less than 3.6 mm vs. greater than 10.3 mm), a contact with supralabials; lower eyelid with narrower body (less than 5.4 mm vs. greater one row of scales; supralabials six, first twice than 7.9 mm), and by the absence of a postnasal the anteroposterior length of others, fourth scale and auricular opening (vs. presence). below eye midline; infralabials six (Fig. 5). Description of holotype.—Details of the Mental wider than long, fused with first head scalation are shown in Fig. 6. Adult infralabials; postmental single, enlarged, its female, body small, slender, SVL 60.2 mm; width greater than width of mental; followed head weakly differentiated from neck, nearly by three pairs of enlarged chin shields, first as wide as body, HW 7.0% SVL, 107.1% HL; pair in broad medial contact, its width greater HL 33.8% SnFa; SnFa 19.2% SVL; snout than width of third pair, narrower than second short, bluntly rounded in dorsal and lateral pair, second pair broadly separated by single profile, SNL 63.0% HL; ear completely hidden medial scale, third pair separated by three by scales; eyes small, ED 1.7% SVL, 26.5% medial scales (Fig. 5). HL, 62.7% END, pupil subcircular; body Scales on limbs smaller than body scales; slightly depressed, nearly uniform in thickness, scales on dorsal surfaces of digits wrapping MBW 120.4% MBH; scales smooth, glossy, around lateral edges of digits; lamellae absent; imbricate; longitudinal scale rows at midbody palmar surfaces of hands and plantar surfaces 22; paravertebral scale rows 85; axilla–groin of feet with several small, irregular scales, 2011] HERPETOLOGICAL MONOGRAPHS 103 each with irregular raised anterior edges; We have evaluated this species against the fingers equal in size; toes unequal in size on IUCN criteria for classification, and find that right foot, second digit greatest in length, it does not qualify for Critically Endangered, digits absent on left foot. Endangered, Vulnerable, or Near Threatened Coloration in preservative.—Body ground status. Brachymeles cobos has been docu- color medium brown, each dorsal scale light mented to be widely distributed on the island brown posteriorly, with a dark auburn streak on of Catanduan˜ es, and is abundant at all the anterior two-thirds to half of the scale. Dark sampled localities. We therefore classify this streaks on each scale consist of four to seven species as one of Least Concern (IUCN, thin longitudinal lines with smudges between 2011). lines. Streaks on scales present around entire Sympatric lizard species occurring on Cat- body, more distinct on venter, where posterior anduan˜ es Island include the following: (Aga- ends of scales are cream, giving greater midae) Bronchocela cristatella, Draco spilop- contrast. Subcaudal scales match venteral scale terus, Gonocephalus sophiae, Hydrosaurus coloration. Posterior edge of all body scales pustulatus; (Gekkonidae) Cyrtodactylus philip- transparent. Forelimb and hind limb scales pinicus, Hemidactylus frenatus, H. platyurus, dark brown, with weakly defined scale bound- Gehyra mutilata, Gekko gecko, Gek. mind- aries. Precloacal scales match ventral scale orensis, Luperosaurus cumingii, Pseudogekko coloration. Head scales mottled light and dark smaragdina, P. compressicorpus; (Scincidae) brown, matching dorsal background coloration. Emoia atrocostata, Eutropis multicarinata bor- Rostral, nasal, supranasal, and first supralabial ealis, Eu. indeprensa, Eu. multifasciata, Lam- scales cream. Supraocular scales dark brown. prolepis smaragdina, Lipinia pulchella pul- Mental, postmental, and chin shields scales chella, Sphenomorphus abdictus, S. decipiens, mottled brown to light cream. S. cumingi, S. jagori, S. laterimaculatus, S. Coloration in life.—Ground color of body lawtoni, S. steerei, Tropidophorus grayi;(Var- medium to dark brown; streaks of darker anidae) Varanus marmoratus, V. olivaceus. pigmentation on body dark-brown. Etymology.—The specific epithet is chosen Variation.—Morphometric variation of the in recognition of the Catanduan˜ es indigenous series is summarized in Table 6. We observed people’s group for which the first adopted no variation among the type series in digit name for the island, ‘‘Isla de Cobos,’’ was number, head scale counts, or in the degree of coined. The name was adopted by the Spanish head scale contact. conquistadores who encountered the original Distribution.—Brachymeles cobos is known Catanduan˜ es tribes living in thatched huts only from Catanduan˜ es Island (Fig. 3). called ‘‘cobos.’’ Suggested common name: Ecology and natural history.—Brachymeles Catanduan˜ es Slender Skink. cobos occurs in residential and agricultural habitats, as well as in disturbed and second- Brachymeles brevidactylus sp. nov. ary-growth forest. No original, low-elevation Figs. 3–4 forest remains on Catanduan˜ es Island, but we Holotype.—PNM 9764 (CDS Field assume the species once also occurred in first- No. 4099, formerly KU 324017), adult male, growth forest at low elevations when these collected under pile of rotting coconut husks habitats persisted. Individuals have been in secondary-growth forest (1000 to 1230 h) observed under piles of rotting coconut husks, on 18 June 2009, in the Municipality of Irosin, in the rotting material within fallen logs, and Sorsogon Province, Luzon Island, Philippines in loose soil and leaf litter surrounding the (15u509 N, 123u559 E; WGS-84), by J. root networks of trees. This species occurs Fernandez. sympatrically with the pentadactyl species, B. Paratypes.—Adult female (TNHC 62469) makusog, and the limbless species, B. mini- collected in a rotting log on ridge above Lake mus. On Catanduan˜ es Island, both B. maku- Bulusan on 24 November 2001, 500–700 m sog and B. minimus have only been observed elevation, Mt. Balusan National Park, Baran- in disturbed, secondary-growth forest, where- gay San Roque, Municipality of Irosin, Sorso- as B. cobos appears to be a habitat generalist. gon Province, Luzon Island, Philippines, by 104 HERPETOLOGICAL MONOGRAPHS [No. 25

RMB and B. Fernandez; adult female (PNM Brachymeles brevidactylus can be distin- 4856) collected 1 July 1995, in Barangay guished from all limbless species of Brachy- Salvacion, Municipality of Santa Magdalena, meles (B. apus, B. lukbani, B. minimus, B. Sorsogon Province, Luzon Island, Philippines, miriamae, B. vermis) by having limbs; and by R. V. Sison. from all pentadactyl species of Brachymeles Diagnosis.—Brachymeles brevidactylus can (B. boholensis, B. boulengeri, B. bicolor, B. be distinguished from congeners by the gracilis, B. kadwa, B. makusog, B. mind- following combination of characters: (1) body orensis, B. orientalis, B. schadenbergi, B. size small (SVL 54.0–60.0 mm), (2) limbs talinis, B. taylori, B. tungaoi, B. vindumi)by bidactyl, (3) limb length short, (4) suprala- having nonpentadactyl limbs, shorter forelimb bials six, (5) infralabials six, (6) supraciliaries lengths (less than 2.1 mm vs. greater than six, (7) supraoculars five, (8) midbody scale 5.9 mm), shorter hind limb lengths (less than rows 20, (9) axilla–groin scale rows 73–77, 3.6 mm vs. greater than 10.3 mm), a narrower (10) paravertebral scale rows 90–94, (11) body (less than 5.4 mm vs. greater than pineal eye spot present, (12) prefrontals not 7.9 mm), and by the absence of a postnasal contacting on midline, (13) frontoparietals scale and auricular opening (vs. presence). contact, (14) postnasals absent, (15) enlarged Description of holotype.—Details of the chin shields in three pairs, (16) nuchals head scalation are shown in Fig. 5. Adult enlarged, (17) fourth supralabial below eye male, body small, slender, SVL 56.8 mm; head midline, (18) auricular opening absent, (19) weakly differentiated from neck, nearly as presacral vertebrae 47–48, (20) nonfusion of wide as body, HW 7.0% SVL, 103.6% HL; HL mental and first infralabials, (21) nonfusion 32.4% SnFa; SnFa 20.9% SVL; snout short, of loreals, and (22) uniform body color bluntly rounded in dorsal and lateral profile, (Tables 4 and 5). SNL 59.2% HL; ear completely hidden by Comparisons.—Characters distinguishing scales; eyes small, ED 1.6% SVL, 24.2% HL, Brachymeles brevidactylus from all nonpenta- 59.6% END, pupil subcircular; body slightly dactyl, limbed species of Brachymeles are depressed, nearly uniform in thickness, MBW summarized in Tables 4 and 5. Brachymeles 125.2% MBH; scales smooth, glossy, imbri- brevidactylus most closely resembles B. sa- cate; longitudinal scale rows at midbody 20; marensis, B. bicolandia, B. cobos, and popu- paravertebral scale rows 90; axilla–groin scale lations of B. bonitae, the only other species to rows 73; limbs short, poorly developed, with be bidactyl or have bidactyl populations. digits highly reduced to two small claws on However, B. brevidactylus can be distin- both forelimbs and hind limbs, finger and toe guished from B. samarensis by having a lamellae absent; FLL 3.3% AGD, 2.5% SVL; greater number of presacral vertebrae, and a HLL 5.9% AGD, 4.5% SVL; tail nearly as greater number of axilla–groin and paraverte- wide as body, gradually tapered at end, TW bral scale rows (Table 5); from B. bicolandia 91.5% MBW, TL 79.5% SVL. by having a greater relative forelimb length, Rostral projecting onto dorsal snout to level six infralabials, and a tendency toward a just past posterior edge of nasal, broader than greater number of axilla–groin and paraverte- high, in broad contact with frontonasal; bral scale rows (Tables 4 and 5); from B. cobos frontonasal wider than long; nostril ovoid, in by having a greater number of presacral center of single trapezoidal nasal, longer axis vertebrae, fewer midbody scale rows, and a directed anterodorsally and posteroventrally; greater number of axilla–groin and paraverte- supranasals present, large, broadly separated; bral scale rows (Table 5); and from B. bonitae postnasals absent; prefrontals broadly sepa- by having longer relative fore- and hind limb rated; frontal octagonal-shaped, its anterior lengths, fewer midbody scale rows, six supra- margin in broad contact with frontonasal, in labials, six infralabials, six supraciliaries, five contact with first two anterior supraoculars, supraoculars, the presence of contact between 53 wider than anterior supraocular; suprao- frontoparietals, and a tendency towards fewer culars five; frontoparietals large, in moderate presacral vertebrae, axilla–groin, and paraver- medial contact, each frontoparietal in contact tebral scale rows (Tables 4 and 5). with supraoculars 2–4; interparietal large, its 2011] HERPETOLOGICAL MONOGRAPHS 105 length greater than midline length of fronto- dorsal background coloration. Rostral, nasal, parietal, longer than wide, diamond-shaped, supranasal, and supralabial scales light beige. wider anteriorly; parietals as broad as fronto- Supraocular scales dark brown. Mental and parietals, in broad contact behind interpari- infralabial scales cream. Chin shields and etal; nuchals enlarged; loreals two, anterior postmental scales cream with slight brown loreal about as long as and slightly higher than mottling. posterior loreal; preocular one; presubocular Coloration in life.—Coloration in life unre- one; supraciliaries six, the anteriormost con- corded; however, because Brachymeles spec- tacting prefrontal and separating posterior imens do not change significantly during loreal from first supraocular, posteriormost preservation (C. D. Siler and R. M. Brown, extending to middle of fifth supraocular; personal observation), we suspect that the subocular scale row single, complete, in preserved coloration and patterns are much contact with supralabials; lower eyelid with like those in life. one row of scales; supralabials six, first twice Variation.—Morphometric variation of the the anteroposterior length of others, fourth series is summarized in Table 6. We observed below eye midline; infralabials six (Fig. 4). no variation among the type series in digit Mental wider than long, fused with first number, head scale counts, or in the degree of infralabials; postmental single, enlarged, its head scale contact. width narrower than width of mental; fol- Distribution.—Brachymeles brevidactylus lowed by three pairs of enlarged chin shields, is known only from the southern Bicol first pair in broad medial contact, its width Peninsula (Fig. 3). greater than width of third pair, narrower than Ecology and natural history.—Brachymeles second pair, second pair broadly separated by brevidactylus occurs in disturbed and second- single medial scale, third pair separated by ary-growth forest, and is found in sympatry three medial scales (Fig. 4). with B. boulengeri; however, B. brevidactylus, Scales on limbs smaller than body scales; B. kadwa, B. lukbani,andB. makusog are also scales on dorsal surfaces of digits wrapping recognized to occur on the Bicol Peninsula of around lateral edges of digits; lamellae absent; Luzon Island. We have evaluated this species palmar surfaces of hands and plantar surfaces against the IUCN criteria for classification, of feet with several small, irregular scales, and find that it qualifies for the status of each with irregular raised anterior edges; Vulnerable (VU) based on the following fingers absent on left hand, highly reduced criteria: VU B2ab(iii); D2 (IUCN, 2011). to two small claw tips on right hand; toes Sympatric lizard species occurring in the unequal in size, first digit highly reduced to Bicol Peninsula include the following: (Aga- small claw tip, second digit greatest in length. midae) Bronchocela cristatella, Draco spilop- Coloration in preservative.—Body ground terus, Gonocephalus sophiae, Hydrosaurus color medium brown, each dorsal scale light pustulatus; (Gekkonidae) Cyrtodactylus phi- brown posteriorly, with a dark auburn streak lippinicus, Hemidactylus frenatus, H. pla- on the anterior one-third to half of the scale. tyurus, Gehyra mutilata, Gekko gecko, Gek. Dark streaks on each scale consist of four to mindorensis, Luperosaurus cumingii, Pseudo- six thin longitudinal lines with smudges gekko compressicorpus, P. smaragdina; (Scin- between lines. Streaks on scales present cidae) Emoia atrocostata, Eutropis multicar- around entire body, more distinct on venter, inata borealis, Eu. multifasciata, Lamprolepis where posterior ends of scales are sandy smaragdina, Lipinia pulchella pulchella, brown, giving greater contrast. Subcaudal Sphenomorphus abdictus abdictus, S. cu- scales with reduced dark pigmentation. Pos- mingi, S. decipiens, S. jagori, S. laterimacula- terior edge of all body scales transparent. tus, S. leucospilos, S. steerei, Tropidophorus Forelimb and hind limb scales sandy brown. grayi; (Varanidae) Varanus marmoratus, V. Forelimb scales with weakly defined scale olivaceus. boundaries. Precloacal scale coloration match- Etymology.—The name of the new species es surrounding ventral scale coloration. Head is derived from the Latin root word ‘‘brevis,’’ scales mottled light and dark brown, matching meaning short, and ‘‘dactylus,’’ meaning digit, 106 HERPETOLOGICAL MONOGRAPHS [No. 25 to represent the species’ small, highly reduced Despite the past taxonomic assessments digits. Suggested common name: Southern (reviewed by Brown and Alcala, 1980), it Bicol Slender Skink. comes as little surprise that allopatric popu- lations of ‘‘B. samarensis’’ from the Luzon and DISCUSSION Mindanao PAICs have proven, with improved sampling, to be a series of morphologically The six species recognized in this paper are diagnosable, independent evolutionary lineag- inferred to be part of two clades with markedly different body plans (Fig. 2). One es. We now recognize nine species as part of of these clades (B. cebuensis, B. libayani, and the B. samarensis complex (Fig. 2). B. paeforum) consists of species with three This study also adds to a growing body of digits on their forelimbs and two or three literature suggesting a need for reevaluation of digits on their hind limbs (Fig. 2). In contrast, amphibian and species boundaries the remaining species (B. brevidactylus, B. within the Philippines (Brown et al., 2008a; bicolandia, B. cobos, B. samarensis) are part of Brown and Stuart, in press; Diesmos and a second clade consisting of limbless and Brown, 2011). Few examples exist of truly bidactyl body forms (Fig. 2). Two of the five ‘‘widespread’’ reptile species that have geo- recognized species of limbless Brachymeles graphic distributions spanning recognized (B. minimus and B. lukbani) are sister to a zoogeographic boundaries, and as is often clade of bidactyl species from the Bicol the case, these species frequently turn out Peninsula of Luzon Island and Catanduan˜ es to constitute multiple evolutionary lineages Island. As previously recognized, B. samar- (Brown et al., 2009; Gaulke et al., 2007; ensis spanned two distinct, recognized faunal McGuire and Alcala, 2000; Siler and Brown, regions: Luzon and Mindanao PAICs (Brown 2010; Welton et al., 2009, 2010a,b). The and Diesmos, 2002, 2009). Given this former- exceptions, in contrast, appear to be invasive ly wide geographical distribution, it is not species and human-mediated range expan- surprising that populations between the two sions (Brown et al., 2010; Diesmos et al., PAICs are inferred herein to be distinct; 2006). however, we were surprised to discover high All species of Brachymeles live a semifos- levels of intra-PAIC species diversity (Fig. 2). sorial existence, specializing in dry rotting The new species recognized in this paper material inside and underneath fallen decom- increase the total number of known species of posing logs, leaf litter, and other forest floor Brachymeles to 30, and all but two of these are detritus. Many are habitat specialists found endemic to the Philippines. Species diversity exclusively in rotting logs, loose soil, or leaf within the genus has doubled in the last 2 yr as litter, whereas others are common beneath the result of large-scale sampling efforts across piles of rotting coconut husks in disturbed, the Philippines and the detailed analyses of agricultural habitat. We assume that the morphological variation among species and species now found in residential and agricul- populations (Siler, 2010; Siler et al., 2009, tural areas were once native to forested 2010a,b, 2011a,b,c; Siler and Brown, 2010, habitats and were possibly forest edge spe- 2011). Brachymeles has long been considered a cialists. small clade of Southeast Asian lizards (Brown, Prior to recent, focused survey efforts, the 1956; Brown and Alcala, 1980; Brown and modest sample sizes of specimens of Brachy- Rabor, 1967; Taylor, 1917), and estimates of meles available in museum collections limited species diversity had remained nearly constant appropriate, lineage-based species delimita- for more than 30 yr (but see Brown and Alcala, tions. Although the rarity of Brachymeles in 1995). This vast underestimation of true collections may be due to their secretive, diversity within the genus is a testament to semifossorial lifestyle, focused survey efforts the extent of morphological similarity among that target the appropriate microhabitat have species and, until recently, a lack of systematic proven effective in sampling Brachymeles in studies of the group (Siler, 2010; Siler et al., their native environments (C. D. Siler, per- 2009, 2010a,b, 2011a,b,c; Siler and Brown, sonal observation; Siler and Brown, 2010; 2010, 2011). Siler et al., 2011a,c). 2011] HERPETOLOGICAL MONOGRAPHS 107

Recent fine- and broad-scale phylogenetic and one in the Faunal Region analyses of species of the genus Brachymeles (Brown and Alcala, 1980, Brown and Alcala, have made it apparent that species diversity in 1995; Brown and Diesmos, 2002; Siler et al., the clade has been considerably underesti- 2011a,c). mated; accordingly, discovery of additional Additionally, the distribution of total spe- undocumented (possibly cryptic) diversity is cies diversity in the genus is also uneven, with anticipated in other Brachymeles species 13 species known from the Luzon Faunal groups (e.g., Siler et al., 2009, 2010a,b, Region vs. eight in the Mindanao Faunal 2011a,b,c; Siler and Brown, 2010, 2011). A Region, six in the Visayan Faunal Region, and number of studies have shown that the only two and two in the Sulu archipelago and evolution of a burrowing lifestyle is correlated Mindoro Faunal Region, respectively (Brown with decreasing dispersal abilities (Nevo, and Alcala, 1980, 1995; Brown and Diesmos, 1979; Patton and Feder, 1978; Patton and 2002; Siler et al., 2009, 2010a,b, 2011a,c; Siler Yang, 1977; Selander et al., 1974; Siler et al., and Brown, 2010). New species discoveries on 2011b). Many Brachymeles lineages have Luzon Island have occurred with consistency experienced reduction or loss of limbs, which during the last two decades; given the island’s may further reduce vagility (Siler and Brown, complex mountain ranges (Sierra Madres, 2010, 2011; Siler et al., 2011b). Through time, Cordillera, Zambales, Bicol Peninsula volca- reduced dispersal abilities and semifossorial noes) and geographic complexity (Defant lifestyles may lead to increasingly patchy et al., 1989; Yumul et al., 2009), the increase distributions, reduced gene flow among pop- in the faunal region’s diversity is likely to ulations, and the accumulation of interpopu- continue (Brown et al. 1995a,b, 1999, 2000; lation differences (Nevo, 1979). Still, the role Linkem et al., 2010; Ross and Gonzales, 1992; that reduced dispersal abilities associated with Siler et al., 2009, 2010a,b,c; Welton et al., fossoriality play on the dispersal abilities and 2010a). It is worth noting that efforts to survey diversification patterns of Brachymeles spe- Mindanao have been less extensive than cies remains unknown. Regardless of which efforts on Luzon; this may account for some processes produce species diversity, we sus- of the differences in diversity between the pect that additional species await discovery. regions—which may be artifacts of sampling Following the recognition of B. bicolandia, biases. B. brevidactylus, B. cobos, B. libayani, B. At present there remains one polytypic paeforum, and Brachymeles samarensis there species (B. gracilis) and one ‘‘widespread’’ are now 13 nonpentadactyl, limbed species species (B. bonitae) recognized in the genus, of Brachymeles. Of these, four species are both with distributions spanning boundaries bidactyl (B. bicolandia, B. brevidactylus, B. between recognized faunal regions (Brown cobos, B. samarensis), four are tridactyl (B. and Alcala, 1980; Siler and Brown, 2010; Siler libayani, B. muntingkamay, B. tridactylus, B. et al., 2011a,b,c). Recent phylogenetic studies paeforum), and two are tetradactyl (B. elerae, of the genus Brachymeles have not supported B. wrighti). Additionally, two species have the monophyly of either of these species, an unequal fore- and hind limb digit numbers (B. indication that taxonomic revisions are needed cebuensis, 3/2; B. pathfinderi, 5/4), and (Siler and Brown, 2011; Siler et al., 2011b). As populations of B. bonitae have been observed our understanding of the total diversity within to have 0–2 fingers and toes. All nonpenta- Brachymeles increases, it is important that dactyl species have smaller body lengths with continued efforts be made to conduct surveys the exception of B. wrighti (Taylor, 1925; Siler focused on rotting log and leaf litter micro- et al., 2011c). Interestingly, the distribution of habitats throughout the ranges of all species. limbed, nonpentadactyl species in the Philip- Accurate data on the distributions of these pines is relatively uneven across the major species will allow for a complete assessment biogeographic regions of the Philippines, with of the geographic ranges of the species and seven species known to occur in the Luzon appropriate assessment of conservation status Faunal Region, four in the Mindanao Faunal can be made. At present, eight of the nine Region, two in the Visayan Faunal Region, species of the B. samarensis complex are 108 HERPETOLOGICAL MONOGRAPHS [No. 25 known or believed to be common throughout and Bornean stream frogs: Reconsideration of Huxley’s their ranges. Although these species currently modification of Wallace’s Line at the Oriental–Austra- lian faunal zone interface. Biological Journal of the inhabit highly disturbed, agricultural and Linnean Society 76:393–461. residential areas, no studies on the long-term BROWN,R.M.,AND B. L. STUART. 2011. Patterns of effect of deforestation and habitat modifica- biodiversity discovery through time: An historical tion on populations of Brachymeles exist. analysis of amphibian species discoveries in the South- east Asian mainland and island archipelagos. In D. J. Acknowledgments.—We thank the Protected Areas Gower, K. G. Johnson, J. E. Richardson, B. R. Rosen, L. and Wildlife Bureau of the Philippine Department of Ru¨ ber, and S. T. Williams (Eds.), Biotic Evolution and Environment and Natural Resources for facilitating Environmental Change in Southeast Asia. Cambridge collecting and export permits necessary for this and University Press, Cambridge, United Kingdom. related studies; we are particularly grateful to T. M. Lim, BROWN, R. M., J. W. FERNER, AND L. A. RUEDAS. 1995a.A C. Custodio, A. Tagtag, and J. L. De Leon for their new species of lygosomine lizard (Reptilia: Lacertilia: logistical support of this research. Financial support for Scincidae; Sphenomorphus) from Mt. Isarog, Luzon fieldwork was provided by a Panorama Fund grant from Island, Philippines. Proceedings of the Biological The University of Kansas Biodiversity Institute, travel Society of Washington 108:18–28. funds from The University of Kansas Ecology and BROWN, R. M., J. W. FERNER, AND R. V. SISON. 1995b. Evolutionary Biology department, a Madison and Lila Rediscovery and redescription of Sphenomorphus Self Fellowship from the University of Kansas, a Fulbright beyeri Taylor (Reptilia: Lacertilia: Scincidae) from the Fellowship, a Fulbright-Hayes Fellowship, NSF DEB Zambales mountains of Luzon, Philippines. Proceed- 0804115 to CDS, and DEB 0743491, and NSF EF- ings of the Biological Society of Washington 108:6–17. 0334952 to RMB. For the loans of specimens we thank J. BROWN, R. M., J. A. MCGUIRE,J.W.FERNER, AND A. C. Vindum and A. Leviton (California Academy of Sciences ALCALA. 1999. A new diminutive species of skink [CAS]), R. Sison and A. C. Diesmos (Philippine National (Squamata; Scincidae; Lygosominae: Sphenomorphus) Museum), J. Ferner (Cincinnati Museum of Natural History), A. Resetar and H. Voris (Field Museum of from Luzon Island, Republic of the Philippines. Copeia Natural History), R. Crombie (United States Natural 1999:362–370. History Museum), and T. LaDuc (Texas Memorial Natural BROWN, R. M., J. A. MCGUIRE, AND A. C. DIESMOS. 2000. History Museum). Critical reviews of the manuscript were Status of some Philippine frogs referred to Rana provided by J. Esselstyn, C. Oliveros, T. Reeder, and three everetti (Anura: Ranidae), description of a new species, anonymous reviewers. CDS thanks the CAS’s Stearns and resurrection of R. igorota Taylor 1922. Herpetolo- Fellowship and the Museum of Comparative Zoology’s gica 56:81–104. Ernst Mayr Fellowship for funding recent visits to examine BROWN,R.M.,A.C.DIESMOS, AND A. C. ALCALA. 2008a. comparative material. Both CDS and RMB extend a special Philippine amphibian biodiversity is increasing in thanks to Arvin and Mae Diesmos for their continued leaps and bounds. Pp. 82–83. In S. N. Stuart, M. enthusiastic support of our Philippine biodiversity research Hoffmann, J. S. Chanson, N. A. Cox, R. Berridge, P. program. Ramani, and B. E. Young (Eds.), Threatened Amphibians of the World. Lynx Ediciones, Barcelona, Spain; IUCN–The World Conservation Union, Gland, LITERATURE CITED Switzerland; and Conservation International, Arling- BOETTGER, O. 1886. Aufzahlung der von den Philippinen ton Virginia, USA. bekannten Reptilien und Batrachier. Berlin Sencken- BROWN, R. M., C. OLIVEROS,C.D.SILER, AND A. C. bergische Naturforschende Gesellschaft 1886:91–134. DIESMOS. 2008b.AnewGekko from the Babuyan Islands, BOULENGER, G. A. 1887. Catalogue of the Lizards in the northern Philippines. Herpetologica 64:305–320. British Museum (Natural History, Volume III, Lacer- BROWN, R. M., C. OLIVEROS,C.DSILER, AND A. C. tidae, Gerrhosauridae, Scincidae, Anelytropidae, Diba- DIESMOS. 2009. Phylogeny of Gekko from the northern midae, Chamaeleontidae). Taylor & Francis, London, Philippines, and description of a new species from UK. Calayan Island. Journal of Herpetology 43:620–635. BRANDLEY, M. C., A. SCHMITZ, AND T. W. REEDER. 2005. BROWN, R. M., C. W. LINKEM,C.D.SILER,J.SUKUMARAN, Partitioned Bayesian analyses, partition choice, and the J. A. ESSELSTYN,A.C.DIESMOS,D.T.ISKANDAR,D. phylogenetic relationships of scincid lizards. Systematic BICKFORD,B.J.EVANS,J.A.MCGUIRE,L.GRISMER,J. Biology 54:373–390. SUPRIATNA, AND N. ANDAYANI. 2010. Phylogeography and BROWN, R. M., AND A. C. DIESMOS. 2002. Application of historical demography of Polypedates leucomystax in lineage-based species concepts to oceanic island frog the islands of Indonesia and the Philippines: Evidence populations: The effects of differing taxonomic philos- for recent human-mediated range expansion? Molecu- ophies on the estimation of Philippine biodiversity. lar Phylogenetics and Evolution 57:598–619. Silliman Journal 42:133–162. BROWN, W. C. 1956. A revision of the genus Brachymeles BROWN, R. M., AND A. C. DIESMOS. 2009. Philippines, (Scincidae), with descriptions of new species and Biology. Pp. 723–732. In R. Gillespie and D. Clague subspecies. Breviora 54:1–19. (Eds.), Encyclopedia of Islands. University of California BROWN, W. C., AND A. C. ALCALA. 1970. The zoogeography Press, Berkeley, California, USA. of the herpetofauna of the Philippine Islands, a fringing BROWN, R. M., AND S. I. GUTTMAN. 2002. Phylogenetic archipelago. 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Phylogeny of Philippine slender skinks Numbers in parentheses indicate the number of (Scincidae: Brachymeles) reveals underestimated spe- specimens examined. With the exception of Brachymeles cies diversity, complex biogeographical relationships, apus and B. miriamae, all specimens examined are from and cryptic patterns of lineage diversification. Molec- the Philippines. Numbers in parentheses indicate the ular Phylogenetics and Evolution 59:53–65. number of specimens examined for each species. Several SILER, C. D., R. JONES,L.J.WELTON, AND R. M. BROWN. sample sizes are greater than those observed in the 2011c. Redescription of tetradactyl, Philippine slender description due to the examination of subadult specimens, skinks (genus Brachymeles). Herpetologica 67:300–317. which were excluded from morphometric analyses. SIMPSON, G. G. 1961. Principles of Taxonomy. Brachymeles apus (1).—BORNEO: MALAYSIA: Sabah: Columbia University Press, New York, New York, USA. Mt. Kinabalu National Park, Sayap Substation: SP 06915. STAMATAKIS, A. 2006. RAxML-VI-HPC: Maximum likeli- Brachymeles bicolandia (20).—See type description. hood-based phylogenetic analyses with thousands of Brachymeles bicolor (24).—LUZON ISLAND: Aurora taxa and mixed models. Bioinformatics 22:2688–2690. Province: Municipality of Maria Aurora: Barangay Villa SWOFFORD, D. L. 2002. PAUP*. Phylogenetic analysis Aurora, Sitio Dimani, Aurora Memorial National Park: using parsimony (*and other methods). Ver 4.0. Sinauer KU 323149–52; CAGAYAN PROVINCE: Municipality of Associates, Sunderland, Massachusetts, USA. Baggao: Sitio Hot Springs: CAS 186111, USNM 140847, TAYLOR, E. H. 1917. Brachymeles, a genus of Philippine 498829–30, 498833; Isabela Province, Sierra Madres lizards. Philippine Journal of Science 12:267–279. Mountain Range: KU 324097–99, PNM 5785, 9568–77; TAYLOR, E. H. 1918. Reptiles of Sulu archipelago. KALINGA PROVINCE: Balbalasang-Balbalan National Park: Philippine Journal of Science 13:233–267. FMNH 259438. TAYLOR, E. H. 1922. The lizards of the Philippine Islands. Brachymeles boholensis (19).—BOHOL ISLAND: BO- Department of Agriculture and Natural Resources, HOL PROVINCE: Municipality of Sierra Bullones, Barangay Bureau of Science, Manila, Publication 17:1–269. Danicop: KU 323944, 323948–9, 323952–6, 323960, 2011] HERPETOLOGICAL MONOGRAPHS 111

323962–3, 323966, 323970, 323972, 323975–6, 323981–2, NORTE ISLAND: CAGAYAN PROVINCE: 323990, 324001; BOHOL ISLAND: BOHOL PROVINCE: Municipality of Calayan: Barangay Balatubat: KU (Para- 6 km S of Municipality of Sierra Bullones: Teachers Park: types) 304559, 304575, 304593, 304708, 304754, 307984, CAS-SU (Holotype) 24528; 13 km SE Municipality of 307996, 307998, 308011, KU 304558, 304562–65, 304569, Sierra Bullones: Dusita Barrio: CAS-SU (Paratypes) 304571–74, 304627–30, 304643, 304647, 304696–99, 24502–04, 24518, 24520–25, 24541, 24543, CAS-SU 304704–07, 304709–12, 304714, 304753, 304755–59, 25443–44, 25447; 1 km E Dusita Barrio: Abacjanan: 307965–66, 307985–86, 307997, 307999–8003, 308006– CAS-SU 24867; Municipality of Sierra Bullones: San- 10, 308012–15, 308017–18. dayong: CAS-SU 18709, 18717. Brachymeles libayani (46).—See type description. Brachymeles bonitae (13).—MASBATE ISLAND: Brachymeles lukbani (14).—LUZON ISLAND: CAMAR- MASBATE PROVINCE: Municipality of Mobo: Tugbo Barrio: INES NORTE PROVINCE: Municipality of Labo: Barangay CAS 144223; Mapuyo Barrio: Palangkahoy: CAS 144270; Tulay Na Lupa, Mt. Labo: PNM (Holotype) 9567, MINDORO ISLAND: MINDORO ORIENTAL PROVINCE:Mt (Paratopotypes) 9589–92, KU (Paratopotypes) 313597– Halcon: SE slope Barawanan Peak: CAS-SU 25713, 99, 313601, 313603–04, 313606, 313608, FMNH (Para- 25793, 25886–88, 25904; Sumagui: CAS 62064 (Paratype); topotype) 270191. POLILLO ISLAND: QUEZON PROVINCE: Municipality of Brachymeles makusog (17).—CATANDUAN˜ ES IS- Polillo: Barangay Pinaglubayan: KU 307747–49, 307755. LAND: CATANDUAN˜ ES PROVINCE: Municipality of Gigmoto: Brachymeles boulengeri (26).—LUZON ISLAND: AU- Barangay San Pedro, Sitio Tungaw: PNM (Holotype) RORA PROVINCE: Municipality of Baler: KU 322314–20; 9565, (Paratopotypes) 9583–9584, KU (Paratopotypes) LUZON ISLAND: LAGUNA PROVINCE: Municipality of Los 308126, 308128, 308136, 308208; LUZON ISLAND: Banos, Barangay Batong Malake: KU 32058–60; Munic- CAMARINES NORTE PROVINCE: Municipality of Labo, ipality of Los Banos: CAS 61096; Mt. Maquiling: CAS Barangay Tulay Na Lupa, Mt. Labo: KU (Paratypes) 61297; POLILLO ISLAND: QUEZON PROVINCE: Munici- 313612–313614, 313616, 313617, PNM (Paratypes) 9585– pality of Polillo: CAS (Paratypes) 62272–73, 62276–77; 9588, FMNH (Paratype) 270200. Barangay Pinaglubayan: KU 307438–9, 307750–54, Brachymeles mindorensis (34).—MINDORO IS- 307756 (Neotype), 307757–58. LAND: MINDORO OCCIDENTAL PROVINCE: KU 304351–5, Brachymeles brevidactylus (3).—See type description. 304412–3, 304488, 307739–42, 308404, 308447–8, Brachymeles cebuensis (8).—CEBU ISLAND: 40 km 308534; MINDORO ISLAND: MINDORO ORIENTAL PROV- SW of Cebu City: Tapal Barrio, Sitio Mantalungon: CAS- INCE: 30 km SE Municipality of Calapan: Bank of Tarogin SU (Holotype) 24400, (Paratypes) 24396–97, 24399, River: CAS-SU (Holotype) 24487; SE slope Mt Halcon, 24401, 24403; Municipality of Carcar: Tapal Barrio: Tarogin Barrio: CAS-SU (Paratypes) 24549–54, 24561–62, CAS 102405 (Paratype); 3 km NW Cebu City, Buhisan 24564; 24566, 24568, 24573–74, 24577–79; Mt Halcon, Barrio, Buhisan Reforestation Project: CAS-SU 27537. SE slope Barawanan Peak: CAS-SU (Paratype) 24570. Brachymeles cobos (10).—See type description. Brachymeles minimus (6).—CATANDUAN˜ ES IS- Brachymeles elerae (5).—LUZON ISLAND: KALINGA LAND: CATANDUAN˜ ES PROVINCE: Municipality of Gigmoto: PROVINCE: Municipality of Balbalan: CAS 61499–500, CM Barangay San Pedro: KU 308129–31, 308210–12. (Paratype) 1717, PNM 9563–4. Brachymeles miriamae (2).—THAILAND: NAKHON Brachymeles gracilis gracilis (18).—MINDANAO IS- RATCHASIMA PROVINCE: Wang Nam Khieo District: Sa- LAND: DAVAO DEL SUR PROVINCE: Municipality of Malalag: Sitio Kibawalan: CAS-SU 24163, 24165, CAS kaerat Environmental Research Station: DSM 1293, 1363 124811, 139307–09; Davao City: Buhangin, Kabanti-an: (Currently uncataloged specimens housed at KU). CAS 124803–04, 139293–95, 139303–05; Digos City: Tres Brachymeles muntingkamay (17).—LUZON ISLAND: de Mayo Barrio: CAS 124806–08, 139300. NUEVA VIZCAYA PROVINCE: Municipality of Quezon: Bar- Brachymeles gracilis hilong (20).—MINDANAO IS- angay Maddiangat, Mt. Palali: PNM (Holotype) 9566, LAND: AGUSAN DEL NORTE PROVINCE: Municipality of (Paratopotypes) 9578–82, KU (Paratopotypes) 308865–66, Cabadbaran: Diuata Mountain Range: Mt. Hilonghilong: 308900–06, 308908, 308953. Balangbalang: CAS-SU (Holotype) 24407, (Paratype) Brachymeles orientalis (53).—BOHOL ISLAND: 102406, 133578, CAS-SU 24411, 133577, 133581–82, BOHOL PROVINCE: Municipality of Sierra Bullones: Dusita 133609, 133612, 133692–93, 133703–06, 133743, 133745– Barrio: CAS-SU (Holotype) 24436, CAS-SU (Paratypes) 47; SURIGAO DEL SUR PROVINCE: Municipality of Lanuza: 24428, 24434, 24437, CAS (Paratype) 102404, CAS-SU Diuata Mountain Range: Sibuhay Barrio: CAS-SU (Para- 25452; Dusita Barrio: Abacjanan: CAS-SU (Paratypes) type) 24315. 24446–51, CAS-SU 25460; Cantaub Barrio: CAS-SU Brachymeles kadwa (101).—LUZON ISLAND: AURORA (Paratypes) 18702, 24442, 24458; CAMIGUIN SUR PROVINCE: Municipality of Baler: Barangay Zabali, Aurora ISLAND: CAMIGUIN PROVINCE: Municipality of Catarman: State College of Technology campus: PNM (Holotype) Mt. Mambajao: Sitio Sangsangan: CAS 110976–83; 9721, KU (Paratopotypes) 232092, 323094–96, 323100, LEYTE ISLAND: Leyte PROVINCE: Municipality of 323104, 323106, KU 323090–91, 323093, 323097–99, Baybay: KU 311231–5, 311241; MINDANAO ISLAND: 323101–03, 323105, 323107; Municipality of Casiguran, AGUSAN DEL NORTE PROVINCE: Municipality of Cabad- IDC property: KU 323108–48; Municipality of San Luis, baran: Diuata Mountain Range: Mt. Hilonghilong: Barangay Real, Sitio Minoli: KU 322320; CALAYAN Kasinganan: CAS-SU 133301, 133616, 133749, 133752, ISLAND: CAGAYAN PROVINCE: Municipality of Calayan: 133754; SAMAR ISLAND: Eastern Samar PROVINCE: Barangay Magsidel: KU (Paratypes) 304875, 304897, Municipality of Taft: KU 305470, 310734–6, 310739, 304900, 304902–3, 304905–6, 304915, 304929, 304941, 310942–6, 310949, 310951, 310955. KU 304908, , 304899, 304907, 304909, 304921, 304941; Brachymeles paeforum (17).—See type description. 112 HERPETOLOGICAL MONOGRAPHS [No. 25

Brachymeles pathfinderi (40).—MINDANAO IS- NEGROS ORIENTAL PROVINCE: 3 km W Municipality of LAND: SARANGANI PROVINCE: Municipality of Glan: Valencia: Cuernos de Negros Mountain Range: Sitio Barangay Taluya: KU 324089–96; Barangay Tanibulad, Lunga: ridge on north side of Maiti River: CAS-SU Sitio Padido: KU 324057–88. (Holotype) 18615, CAS-SU 21873; ridge on south side of Brachymeles samarensis (7).—SAMAR ISLAND: EAST- Maiti River: CAS-SU (Paratype) 18641, 18656–57, 18748; ERN SAMAR PROVINCE: Municipality of Taft: Barangay San Cuernos de Negros Mountain Range: CAS-SU (Paratype) Rafael: KU 310849–52, 311294–6. 18649; top of Dayungan Ridge: CAS-SU 21877, 21880, Brachymeles schadenbergi (34).—BASILAN ISLAND: 21883–84; 24 km NW Bondo Barrio: Bantolinao: CAS-SU BASILAN PROVINCE: Port Holland: Sawmill: CAS 60493; 22355–56; CEBU ISLAND: CEBU PROVINCE: Municipality MINDANAO ISLAND: MISAMIS OCCIDENTAL PROVINCE: of Carcar: Tapal Barrio: Sitio Mantalongon: CAS 154671, 2 km NW of Masawan: CAS 23468–69; 4 km NW of 154673, 154678–82, 154686. Masawan: CAS 23471; 3 km NW Masawan: south bank of Brachymeles tridactylus (20).—NEGROS ISLAND: Dapitan River: CAS 23479–81, 23484–85; ZAMBOANGA DEL NEGROS OCCIDENTAL PROVINCE: 16 km E Municipality of NORTE PROVINCE: Dapitan River: CAS-SU 23494–96; ZAM- La Castellana: Barrio Cabagna-an: Southern Slope of Mt. BOANGA CITY PROVINCE: Municipality of Pasonanca: Barangay Canlaon: CAS-SU 19424, 19426–27, 19429, 19452, 19458; Baluno: Pasonanca Natural Park: KU 314967, 314969, 20 km E Municipality of La Castellana: Sitio Kalapnagan: 314970–8, 314980, 314984–85, 314988–92, 314994, 314996–7. CAS-SU 27082–83; NEGROS ORIENTAL PROVINCE: Hills Brachymeles talinis (31).—NEGROS ISLAND: NE- North and Northwest of Mayaposi: CAS-SU (Holotype) GROS ORIENTAL PROVINCE: 6 km W Municipality of 18354; PANAY ISLAND: ANTIQUE PROVINCE: Municipal- Valencia: Cuernos de Negros Mountain Range: ridge on ity of Culasi: Barangay Alojipan: KU 307726–36. north side of Maite River: CAS-SU (Holotype) 18358, Brachymeles tungaoi (12).—MASBATE ISLAND: MAS- (Paratype) 89813; Cuernos de Negros Mountain Range: BATE PROVINCE: Municipality of Masbate City: PNM Dayungan Ridge: CAS 133871; Dumaguete City: CAS-SU (Holotype) 9722, KU (Paratopotypes) 323934–36; Munic- (Paratype) 12225; Municipality of Siaton: 20 km N Bondo ipality of Mobo, Barangay Tugbo: CAS (Paratypes) Barrio: CAS-SU 22311–12; 22317, 22323; INAMPULA- 144229–30, 144290, 144306–7, 144313, 144341–2. GAN ISLAND: GUIMARAS PROVINCE: Municipality of Brachymeles vermis (5).—JOLO ISLAND: SULU PROV- Sibunag: 8 km W Pulupandan Town: CAS-SU 27972, INCE: CAS-SU (Paratype) 62489, CAS-SU 60720–22, 60857. 27996–97; PANAY ISLAND: ANTIQUE PROVINCE: Munic- Brachymeles vindumi (4).—JOLO ISLAND: SULU ipality of San Remigio: KU 306756–60, 306762–7, 306769, PROVINCE: CAS (Holotype) 60724, CAS (Paratypes) 306770–6, 306786. 60723, 60725, MCZ (Paratype) 26577). Brachymeles taylori (34).—NEGROS ISLAND: NE- Brachymeles wrighti (2).—LUZON ISLAND: BENGUET GROS OCCIDENTAL PROVINCE: Municipality of Silay City, PROVINCE: Municipality of La Trinidad: MCZ (Holotype) Barangay Patag: KU 324044–56; NEGROS ISLAND: 26589, USNM 140756.