Taxonomic Revision of the Pseudogekko Compresicorpus Complex (Reptilia: Squamata: Gekkonidae), with Descriptions of Three New Species

Herpetological Monographs, 28 2014, 110–139 E 2014 by The Herpetologists’ League, Inc.

TAXONOMIC REVISION OF THE PSEUDOGEKKO COMPRESICORPUS COMPLEX (REPTILIA: SQUAMATA: GEKKONIDAE), WITH DESCRIPTIONS OF THREE NEW SPECIES

1,6 2 3 1 1 CAMERON D. SILER ,LUKE J. WELTON ,DREW R. DAVIS ,JESSA L. WATTERS ,CONNER S. DAVEY , 4 5 6 ARVIN C. DIESMOS ,MAE L. DIESMOS , AND RAFE M. BROWN 1 Department of Biology and Sam Noble Oklahoma Museum of Natural History, University of Oklahoma, 2401 Chautauqua Avenue, Norman, OK 73072-7029, USA 2 Department of Biology, Brigham Young University, 401 WIDB, Provo, UT 84602, USA 3 Department of Biology, University of South Dakota, 414 E. Clark Street, Vermillion, SD 57069, USA 4 Herpetology Section, Zoology Division, Philippine National Museum, Rizal Park, Burgos Street, Manila, Philippines 5 University of Santo Tomas, Espana Boulevard, Manila, Philippines 6 Biodiversity Institute and Department of Ecology and Evolutionary Biology, University of Kansas, 1345 Jayhawk Boulevard, Lawrence, KS 66045-7593, USA

ABSTRACT: Recent phylogenetic analysis of false geckos, genus Pseudogekko, revealed unrecognized diversity within these exceedingly rare and enigmatic Philippine forest geckos. Newly available genetic datasets revealed that two of the four currently recognized species are complexes of multiple, deeply divergent evolutionary lineages. In this paper we evaluate species diversity in the Pseudogekko compresicorpus Complex and describe three new species in this unique clade of endemic Philippine geckos. For nearly a century, P. compresicorpus has been recognized as a single, ‘‘widespread’’ species with a geographic range spanning three major faunal regions and several isolated islands. This perception of the species’ wide geographic range has persisted due to the rarity of this species. We evaluate morphological data, in light of a recent phylogenetic study on the genus, to define species limits in P. compresicorpus, finding character-based evidence that unambiguously supports the recognition of four unique evolutionary lineages within the complex, three of which we describe as new species. These evolutionary species correspond to monophyletic lineages supported in recent molecular studies. We also address the historically controversial generic affiliation of Pseudogekko labialis and conclude that this poorly known species is a member of the genus Lepidodactylus. All species recognized in this study possess allopatric geographic ranges and differ from congeners by numerous diagnostic characters of external morphology and, therefore, should be recognized as full species in accordance with any lineage-based species concept. This study nearly doubles the known diversity of Philippine false geckos. Key words: Biodiversity; Conservation; Endemism; False geckos; Philippines; Species complex; Taxonomy

PHILIPPINE gecko diversity represents an gliding locomotion with highly specialized impressive array of diversification in morphol- cutaneousstructures(e.g.,Ptychozoon and ogy, behavior, and ecology (Brown and Alcala, Luperosaurus; Brown et al., 1997, 2012a; 1978). From ancient, micro-endemic lineages Dudley et al., 2007). with small ranges on larger islands (Ro¨sler Other than the work of Taylor (1922a), et al., 2006; Linkem et al., 2010) to several Brown and Alcala (1978) published the only widespread species groups (Siler et al., 2010), comprehensive systematic review of Philip- and to species limited to tiny isolated islets pine geckos. Their work summarized taxo- (Brown and Alcala, 2000; Brown et al., 2011a; nomic diversity, provided an identification Siler et al., 2012a), Philippine geckos are guide, and recognized 31 species. Many of quite ecologically variable, considering that these had been observed rarely in natural only 57 species are currently recognized conditions and were known only on the basis (PhilBREO, 2014). The archipelago’s gekko- of one or two specimens in museum collec- nids also range from morphologically conser- tions. Although Brown and Alcala’s (1978) vative gecko generalists (Brown and Alcala, foundational work has remained the only 1978) to delicate forest vegetation specialists synopsis of the archipelago’s geckos, species and to several lineages capable of derived diversity has now nearly doubled since its original publication. Remarkably, of the 6 CORRESPONDENCE: e-mail, [email protected] country’s 57 species, 47 (82%) are Philippine

110 2014] HERPETOLOGICAL MONOGRAPHS 111

FIG. 1.—Maximum clade credibility topology of Pseudogekko derived from Bayesian analyses in the recent phylogenetic study of Siler et al. (2014a). Numbers below nodes indicate maximum likelihood bootstrap values (left) and Bayesian posterior probabilities (right). Boxed numbers correspond to numbered sampling localities shown on the associated topographic map of the Philippines. endemics (Brown et al., 2008, 2009, 2011a,b). 2014a) for species of Pseudogekko. Yet even Despite this dramatic improvement in our in the absence of dense population genetic understanding of this predominantly endemic sampling, Siler et al. (2014a) revealed a fauna, a few genera are very poorly known considerable degree of cryptic genetic diver- (i.e., Luperosaurus; Brown et al., 2007, sity and high levels of genetic divergence 2011b, 2012a). A case in point is the between clades. As currently recognized, the extremely rare, endemic genus Pseudogekko distribution of each of these species spans (Taylor, 1915, 1922a), a group of four small, multiple recognized faunal regions or Pleisto- delicate, distinctly elongate, highly secretive, cene Aggregate Island Complexes (PAICs; and entirely arboreal forest geckos (Brown Brown and Guttman, 2002; Brown and and Alcala, 1978). Diesmos, 2009). In fact, as currently defined In the last two decades, our comprehensive (Brown and Alcala, 1978; Siler et al., 2012b), biodiversity surveys throughout the Philip- the species Pseudogekko compresicorpus is pines (e.g., Siler et al., 2012b; Brown et al., distributed across three distinct faunal regions 2013a,b) resulted in only a handful of (Luzon, Mindanao, and Visayan PAICs) and vouchered genetic samples (Siler et al., an isolated island group (Romblon Island 112 HERPETOLOGICAL MONOGRAPHS [No. 28

Group; Fig. 1). Currently, there are few In this study, we investigate species diver- examples of seemingly widespread species of sity within the Pseudogekko compresicorpus Philippine vertebrates that truly defy these Complex, with the understanding that several regional biogeographic boundaries (Brown of the divergent populations identified here and Diesmos, 2002, 2009; Brown et al., may represent threatened, and possibly en- 2002, 2013a). Phylogenetic studies support dangered, unique evolutionary lineages (Siler many species as more range-restricted, with et al., 2014a), worthy of both formal taxonom- patterns generally consistent with inferred ic recognition and prioritization for immediate PAIC formation (Siler et al., 2012c,d); Brown conservation action. The taxonomic revisions et al., 2013a). In fact, phylogenetic analyses herein are guided by the results of the recent reveal unique lineages of P. compresicorpus phylogenetic study on geckos of the genus with apparent distributions corresponding to Pseudogekko (Siler et al., 2014a). circumscribed biogeographic subregions of the archipelago (Brown and Diesmos, 2009; TAXONOMIC HISTORY Brown et al., 2013a; Siler et al., 2014a). Taylor (1915) described Luperosaurus com- Species of the genus Pseudogekko represent presicorpus on the basis of one specimen a critical conservation urgency (Alcala et al., collected from Limay, Bataan Province, Lu- 2004; Brown and Diesmos, 2009; Brown et al., zon Island. In this description, Taylor (1915) 2012a); the named taxa are nearly all micro- expressed uncertainty about the placement of endemics threatened by habitat destruction in this species in Luperosaurus (as opposed to the form of anthropogenic forest removal. erecting a new genus to accommodate the one First, all species are arboreal and considered specimen), noting that the new species had an obligate primary forest taxa or Pandanus spp. elongate, compressed body form (a character- (screw pines) plant microhabitat specialists istic generally shared with the other known (Brown and Alcala, 1978). Second, based on species of Luperosaurus; see Brown et al., the little information known about population 2000). Later, Taylor (1922a) transferred this health and microhabitat preferences, popula- species to a novel genus, Pseudogekko, where it tions apparently have decreased over the last remained a monotypic genus until the descrip- 100 yr (IUCN, 2013; Siler et al., 2014a). To tion of a new species, Pseudogekko shebae, make matters worse, the preferred habitat from the Soloman Islands (Brown and Tanner, (lowland and coastal forests) of these species 1949). Unfortunately, the type specimen of P. has been near completely removed from compresicorpus (Philippine Bureau of Science, throughout the Philippines (Catibog-Sinha No. 1781) was destroyed during World War II and Heaney, 2006; Brown and Diesmos, (Brown and Alcala, 1978), which limited 2009; Siler et al., 2014b). Finally, the genus comparisons of this species with other newly has had a complex taxonomic history (Brown described gekkonids from this region. Howev- and Alcala, 1978). Members of Pseudogekko er, the collection of additional specimens of P. previously have been assigned alternatively and compresicorpus from Mindanao and Bohol with little confidence (Taylor, 1922a; Brown islands allowed Brown (1964) to examine and Alcala, 1978) to the genera Luperosaurus shared characteristics between Pseudogekko (Taylor, 1915) and Lepidodactylus (Brown and and other phenotypically similar genera Tanner, 1949; Brown, 1964; Kluge, 1968; (Gekko, Hemiphyllodactylus, Lepidodactylus, Brown and Alcala, 1978) and even to Gekko Luperosaurus, Pseudothecadactylus). Brown (Taylor, 1922b). The combination of these (1964) noted similarities between P. shebae factors highlights the urgent need to assess and Lepidodactylus, and P. shebae was later species boundaries within the genus in order to placed in this genus by Kluge (1968). understand unique evolutionary lineages bet- Additionally, Kluge (1968) transferred two ter, as conservation targets, and to implement species to Pseudogekko: Pseudogekko bre- conservation strategies more efficiently for vipes—a taxon that was originally described protecting these threatened and charismatic as a Lepidodactylus (Boettger, 1897), and species (Sanguila et al., 2011; Siler et al., Pseudogekko smaragdinus, originally described 2014c). as a member of the genus Gekko (Taylor, 2014] HERPETOLOGICAL MONOGRAPHS 113

1922b). Prior to its placement in Pseudogekko, MATERIALS AND METHODS Brown (1964) questioned the assignment of P. Field Work, Sample Collection, and brevipes to the genus Lepidodactylus on the Specimen Preservation basis of body proportions (e.g., ‘‘breadth of head to SVL’’) that appeared more similar to We conducted fieldwork on Bohol, Leyte, members of Pseudogekko than to species of Luzon, Negros, Mindanao, and Polillo islands in the Philippines (Fig. 1). We collected Lepidodactylus. Finally, Peters (1867) original- specimens between 900 and 1600 h, which ly described Pseudogekko labialis as Gecko were euthanized via cardiac injection of labialis, which Boulenger (1885) later rede- nembutal or immersion in aqueous chlore- scribed as a Lepidodactylus. However, it was tone, dissected for genetic samples (liver or not until additional specimens became avail- muscle preserved in 95% ethanol or flash able that Brown and Alcala (1978) transferred frozen in liquid nitrogen), fixed in 10% this species to Pseudogekko. Other than Brown buffered formalin, and eventually transferred and Alcala’s samples (collected in 1971), this to 70% ethanol (,2 mo later). Museum exceedingly rare species had not been collected abbreviations for specimens examined or since its original description. A recent (2012) 5- sequenced in this study are those of Sabaj wk targeted survey effort in the neotype locality Perez (2013; CAS and CAS-SU: California (Brown and Alcala, 1978; Mt. Hilong-Hilong, Academy of Sciences, San Francisco, Califor- northeastern Mindanao Island) by a large nia; KU: The University of Kansas Biodiversity group of experienced herpetologists produced Institute, Lawrence, Kansas; PNM: National no specimen records (R. Brown, personal Museum of the Philippines [formerly Philip- observation). pine National Museum], Manila, Luzon). The genus Pseudogekko contains four species (P. brevipes, P. compresicorpus, P. Morphological Data labialis, P. smaragdinus), and no new species We examined fluid-preserved specimens (see have been described since the late 1970s Appendix) for variation in qualitative, meristic, (Brown and Alcala, 1978). Recently, Siler and mensural characters using the phylogenetic et al. (2014a) estimated phylogenetic relation- results of Siler et al. (2014a) conservatively as a ships and elucidated multiple highly divergent guide for the identification of possibly unique genetic lineages within P. compresicorpus, evolutionary lineages. We determined sex by which the authors interpreted as probable the presence in males of precloacal or pre- evidence for the existence of additional, as of cloacal-femoral pores, or as necessary (imma- yet undefined species contained within P. tures, females) by gonadal inspection. We brevipes and P. compresicorpus. (DRD and CDS) took measurements to the In this paper, we re-analyze new data from nearest 0.1 mm with digital calipers. all available P. compresicorpus specimens, Whenever possible, we scored meristic and including both our own collections from the mensural characters (based on Brown et al., last 20 yr (see Siler et al., 2014a) and older 2008, 2009, 2011a,b, with some modifications) museum specimens (Brown and Alcala, 1970, on the left side of the body. Characters include: 1978; Brown et al., 2013a). We use these data snout–vent length (SVL, distance from tip of and reliable diagnostic differences to revise snout to vent); tail length (TL, distance from Pseudogekko taxonomy and describe four posterior margin of vent to tip of tail); total distinct species. To relate our findings to length (TotL, distance from tip of snout to tip ongoing conservation efforts, we address of tail); tail width (TW, measured at widest conservation threats and priorities and pro- section of tail posterior to hemipene bulge); tail vide new or re-evaluated formal conservation height (TH, measured from ventral to dorsal status assessments (IUCN, 2013) for each surface of tail at the same point as TW); head species described in this study. Finally, we length (HL, from tip of snout to posterior tip of also reassessed the taxonomic affinities of P. mandible); head width (HW, widest measure of labialis and refer this species to the genus head width at jaw articulations); head height Lepidodactylus, in agreement with the con- (HH, measured from ventral to dorsal surface clusions of Boulenger (1885). of head at jaw articulations); midbody width 114 HERPETOLOGICAL MONOGRAPHS [No. 28

(MBW, measured from lateral surface to Evolutionary Species Concept (Simpson, 1961; opposing lateral surface at midpoint of axilla– Wiley, 1978; Frost and Hillis, 1990). We groin region); snout length (SNL, distance diagnose lineages as distinct species based on from anterior border of orbit to tip of snout); a suite of diagnostic morphological features, eye diameter (ED, at widest point); eye–nares genetic divergence, and allopatric distributions distance (END, distance from anterior margin in separate biogeographic subregions of the of eye to posterior margin of nares); internarial archipelago (Brown and Diesmos, 2009; Brown distance (IND, from dorsal aspect between et al., 2013a). Lineage-based species concepts most-laterally distal edges of nares); interorbit- have been shown to be particularly appropriate al distance (IOD, distance between midline of when applied to Philippine land vertebrate orbits from dorsal aspect); axilla–groin distance biodiversity (for review, see Davis et al., in (AGD, distance between posterior edge of arm press). In this study we use a morphological insertion and anterior edge of leg insertion); dataset for all available specimens in museum femur length (FL); tibia length (TBL); supra- collections of the focal lineages, conservatively labials (SUL, number of enlarged supralabials, guided by phylogenetic estimates of relation- from first supralabial in contact with rostral to ships (Siler et al., 2014a), to diagnose distinct posteriormost enlarged supralabial retaining lineages in this complex of false geckos, distinct, square to rectangular shape); infra- recognizing both previously described and labials (IFL, number of infralabials); circumor- new species on the basis of nonoverlapping bitals (CO, number of visible, small circumor- morphological character states. bital scales encircling the eye); pore-bearing precloacal scales (PPS, number of differentiat- RESULTS ed, enlarged, pore-bearing scales in series Morphology anterior to the cloaca); pore-bearing precloacal-femoral scales (PFPS, number of differen- Although sample sizes are low for many tiated, enlarged, pore-bearing scales in series lineages described in this study, we have anterior to the cloaca and, in some specimens, evaluated and examined all known specimens extending into the femoral region on the in museum collections of each putative species. ventral surface of the thigh); Finger III Despite the small sample sizes, multiple adult scansors (FinIII, scan, number of enlarged, specimens for each focal species are available, undivided scansors beneath Finger III, start- and each of the four identified lineages of the ing just distal to point where skin between Pseudogekko compresicorpus Complex are digits ends); Toe IV scansors (ToeIVscan, readily diagnosed on the basis of numerous, number of undivided scansors beneath Toe nonoverlapping differences in meristic, men- IV, starting just distal to point where skin sural, and color pattern characters (Tables 1, between digits ends); paravertebral scales 2). Variation in morphological characters (PVS, number of scales along dorsal surface (Tables 1, 2) mirrors the results observed in of body between midpoints of limb inser- phylogenetic analyses (Fig. 1; Siler et al., tions); ventral scales (VS, number of scales 2014a) and supports the recognition of four along ventral surface of body between P. compresicorpus Complex lineages. Char- midpoints of limb insertions); and interorbital acters differing among these lineages include scales (IOS, total number of scales in straight body, head, and snout length, body and digit line distance across interorbital region from scale counts, pore-bearing scale counts, and center of each eye, across both eyelids). In coloration and pigmentation patterns (Ta- the descriptions, ranges are followed by mean bles 1, 2; species accounts below), many of 6 standard deviation in parentheses. which are commonly employed diagnostic morphological characters in taxonomic stud- Species Concept ies of Philippine gekkonid lizards (Brown and As with many recent taxonomic revisions of Alcala, 1978; Brown et al., 2011a,b). With the organisms endemic to island systems, we exception of the presence (males) or absence embrace the General Lineage Concept (de (females) of pores in the precloacal or Queiroz, 1998, 1999) as an extension of the precloacal-femoral region of the body, we 2014] HERPETOLOGICAL MONOGRAPHS 115

did not find additional sexually dimorphic 6.0) 2.2) 2.5) 3.5) traits in any of these four species. 0.4) 0.6) 0.7) 1.0) 6 3) 1) 1) 6 6 6 6 6 6 6 6 6 6 Genetic Divergence Uncorrected pairwise sequence divergenc-

P. smaragdinus es are quite variable within the lineages (16 male, 17 female) defined here as species (0.0–19.0% mtDNA

Peninsula (Luzon Island) divergence; Siler et al., 2014a) as compared to Polillo Island and Bicol many recent observations of endemic vertebrate diversity in the Philippines (Siler and 11.0) 103.6–129.7(118.9 3.9) 26.2–35.5(31.5 2.1) 54.2–64.3(59.4 7.6) 50.2–61.7(57.4 0.7) 4.9–6.5(5.7 1.1) 7.1–9.8(8.6 1.2) 7.8–10.7(9.5 1.5) 6.2–10.4(8.1 6 5) 51–68(60 0) 14–16(15 1) 15–17(16 Brown, 2010; Welton et al., 2010; Siler et al., 6 6 6 6 6 6 6 6 6 6 2012c,d). However, genetic divergences between lineages are significantly higher, with

P. brevipes the exception of interpopulation divergences (4 male, 2 female) standard deviation follows ranges. among Luzon populations of P. compresicor- PAICs 6 Visayan and Mindanao pus (.26% mtDNA divergence; Siler et al., 2014a). The monophyletic lineages defined by Siler et al. (2014a; P. compresicorpus, Pseu- sp. nov. dogekko pungkaypinit sp. nov., Pseudogekko ditoy sp. nov., and Pseudogekko chavacano sp. Island Mindanao 95.8, — 89.4–113.5(105.6 nov., the latter three of which are first Western (1 male, 1 female) P. chavacano described herein) are distinguished from each

. In parentheses, mean other by levels of genetic divergence greater than those observed between species of most b sp. nov. other Philippine geckos (Siler et al., 2010; Welton et al., 2010; Brown et al., 2011b). P. ditoy islands (1 male, 1 female) Pseudogekko Given the higher observed intraspecific genetic diversity, we suspect that several of the 8.6) — 0.9) 7.7, 7.9 7.8, 8.5 5.3–8.3(7.4 0.9) 9.3, 9.6 10.4, 10.4 6.2–9.4(8.4 1.8) 25.1, 29.7 26.7, 30.0 17.2–28.4(24.4 5.1) 49.4 55.9 47.6–52.5(50.5 strongly supported clades (Fig. 1; Clades C, 0.4) 5.4, 5.7 5.8, 6.0 3.9–5.8(5.1 0.6) 6.3, 7.3 6.4, 6.7 3.6–7.7(6.3 6 3) 57, 59 55, 58 56–69(61 0) 15, 16 14, 15 15–16(16 0) 18, 19 19, 19 16–19(18 6 6 6 6 6 6

sp. nov. D) actually represent independent evolution- 6 6 6 ary lineages (Siler et al., 2014a). However, in two cases (P. compresicorpus, P. pungkaypinit sp. nov.), divergent populations are represent- (4 male, 2 female) Mindanao PAIC Samar and Leyte

P. pungkaypinit ed by single vouchered specimens, at times juvenile individuals, which prevents us from confidently evaluating these putatively unique, 3.8) 125.3–141.2(135.2 1.9) 37.2–41.2(39.6 2.6)1.5) 66.6–76.8(71.8 75.2, 75.3 52.6 54.7and genetically 34.0–44.8(39.4 divergent, populations at this 0.2) 6.7–7.5(7.2 0.3) 9.3–11.2(10.6 0.3) 11.4–13.6(12.6 0.5) 7.7–9.1(8.2 a 6 3) 54–60(58 1) 14–15(15 1) 17–18(17 6 6 6 time. 6 6 6 6 6 6 6 Status of Pseudogekko labialis One of the persistent taxonomic issues with 13–15(14 16–18(17 P. compresicorpus (3 male, 4 female) the diversity of Pseudogekko is whether P. labialis is appropriately placed in the genus rather than with morphologically more-similar species in the genus Lepidodactylus. Not only 1.—Summary of mensural characters in all known species of have researchers historically considered P.

ABLE labialis as a species of the genus Lepidodactylus T based on morphological similarity (Boulenger,

Pleistocene Aggregate Island Complexes. 1885; Wermuth, 1965; Kluge, 1968) but, also, 5 Data unavailable due to small sampleover sizes. the years key morphological differences 5 — length length PAIC have been highlighted between these two a b Snout length/head length 57–65(60 Head width/snout–vent Snout length 5.4–6.0(5.8 Head length/snout–vent Head width 7.5–8.6(8.0 Total length 111.9–117.3(114.6 Axilla–groin distance 27.0–32.6(30.4 Head length 9.2–10.2(9.7 Character Range Luzon PAIC Snout–vent length (female)Snout–vent length (male) 54.9–59.7(57.1 55.9–58.8(57.6 Midbody width 5.4–6.7(6.0 genera. Kluge (1968) described the genus 1 EPTLGCLMNGAH [ MONOGRAPHS HERPETOLOGICAL 116

TABLE 2.—Summary of qualitative diagnostic characters in species of Pseudogekko. In cases of scale count variation within species, numbers of individuals showing specific counts are given in parentheses.

P. compresicorpus P. pungkaypinit sp. nov. P. ditoy sp. nov. P. chavacano sp. nov. P. brevipes P. smaragdinus Character (3 male, 4 female) (4 male, 2 female) (1 male, 1 female) (1 male, 1 female) (4 male, 2 female) (16 male, 17 female) Finger III scansor count 15 (4) 15 (3) 14 (1) 15 (1) 12 (2) 15 (1) 16 (2) 16 (1) 15 (1) 16 (1) 13 (3) 16 (4) 17 (1) 17 (2) 14 (1) 17 (9) 18 (19) Toe IV scansor count 18 (5) 17 (1) 16 (1) 17 (1) 13 (2) 16 (1) 19 (2) 18 (2) 17 (1) 20 (1) 14 (2) 18 (6) 19 (2) 15 (2) 19 (8) 21 (1) 20 (12) 21 (5) 22 (1) Supralabial count 16 (1) 16 (1) 17 (1) 15 (1) 14 (1) 16 (8) 17 (1) 18 (2) 20 (1) 16 (1) 15 (3) 17 (10) 18 (3) 19 (2) 16 (2) 18 (11) 19 (1) 20 (1) 19 (4) 20 (1) Infralabial count 13 (1) 17 (4) 16 (1) 16 (1) 12 (1) 14 (6) 15 (2) 18 (1) 17 (1) 17 (1) 13 (2) 15 (12) 16 (4) 19 (1) 14 (3) 16 (10) 17 (5) Circumorbital count 39–45 50–55 40, 43 46, 46 35–37 33–35 Paravertebral scale 226–234 265–280 180–185 195–197 228–240 241–252 count Ventral scale count 127–130 125–155 111–118 122–123 119–128 124–130 Enlarged pore series 10–14 (precloacal) 17–20 (precloacal) 18, — (precloacal) 16, — (precloacal) 13–15 (precloacal) 32–41 (precloacal-femoral) count Femoral pores Absent Absent Absent Absent Absent Present Dominant body Dark brown to tan Grayish brown Light brown Light brown Dark brown Bright neon yellow to coloration orange (undisturbed) to neon green (disturbed) Conspicuous head Dense, neon green Absent Absent Dense, neon green Sparse, cream Dense black, sparse white spotting Conspicuous Faint neon green Absent Absent Neon green Cream Large black, small white dorsolateral spotting Conspicuous limb Faint neon green Absent Absent Dense, neon green Absent Sparse black and white spotting

Tail banding Absent Absent Absent Present Absent Neon yellow, white, and 28 No. neon orange 2014] HERPETOLOGICAL MONOGRAPHS 117

Pseudogekko as differing from the genera Lepidodactylus and Hemiphyllodactylus based on the shape of the enlarged, precloacal pore- bearing scale series (series greatly arched anteromedially in Pseudogekko vs. not arched;

P. smaragdinus Fig. 2). Boettger (1897) described Lepidodac-

(16 male, 17 female) tylus brevipes (eventually recognized as a member of the genus Pseudogekko by Kluge [1993]), noting it could be separated from Lepidodactylus labialis Peters and Lepidodac- tylus pulcher Boulenger on the basis of having a more-slender body, fewer precloacal pores,

P. brevipes and less-distinctive series of submental scales.

(4 male, 2 female) Even in their redescription of P. labialis and recognition of the species as a member of the genus Pseudogekko, Brown and Alcala (1978) recognized that the general body coloration sp. nov. and pigmentation patterns of P. labialis did not match the patterns of any other species in the genus Pseudogekko. They note that all individ- (1 male, 1 female) P. chavacano uals of P. labialis have distinct dorsal markings that range from a vertebral row of bars or spots to irregular transverse bars or even dark dorsolateral stripes (Brown and Alcala, 1978); these markings are absent in the remaining sp. nov. species of Pseudogekko (Brown and Alcala, Absent Absent Absent1978; C. Siler, Absent personal observation). 2.—Continued. P. ditoy (1 male, 1 female) Based on measurements and comparisons

ABLE of individuals of P. labialis available in T museum collections (see Appendix; Speci- mens Examined) and on published accounts documenting character differences between sp. nov. P. labialis and all other members of the genus Pseudogekko, we now formally recognize this species as a member of the genus Lepido- (4 male, 2 female) slender, anterodorsal– posteroventral, light brown

P. pungkaypinit dactylus. This decision is supported by a suite of diagnostic character differences between L. labialis and all other recognized species of Pseudogekko. Species of the genus Pseudo- gekko differ from L. labialis by having enlarged, pore-bearing scale series (precloacal or precloacal-femoral scales) that are P. compresicorpus (3 male, 4 female) greatly arched anteromedially (vs. not distinctly arched anteromedially; Fig. 2), having markedly narrower bodies (HW/SVL 15–16% vs. .18%), longer relative snout lengths (SNL/HW 63–71% vs. ,57%), by the presence of small, juxtaposed postmental scales (vs. distinctively enlarged, strongly imbricate postmentals; Fig. 2), and the absence (vs. presence) of darkly pigmented, and often

Character Body stripingInterorbital bandingCiliary ring coloration Absent Absent Light blue Lateral body surface, striped, Absent Absent body coloration. Absent Absent Absent Absent Absent Light brown Absent Absent 118 HERPETOLOGICAL MONOGRAPHS [No. 28

Taxonomic Conclusions Pseudogekko compresicorpus (Taylor): Brown With the removal of Lepidodactylus labialis and Tanner, 1949:41; Kluge, 1993:30. from the genus Pseudogekko, the remaining Pseudogekko compressicorpus (Taylor) [Mis- four focal lineages of this study each possess spelled]: Taylor, 1922a:103; Underwood, unique, nonoverlapping suites of diagnostic 1954:479; Wermuth, 1965:151; Kluge, 1967: character states of morphology (Tables 1, 2), 30, 1968:333; Brown and Alcala, 1970:112, and all correspond to clades defined in 1978:119; Brown et al., 2012a:920, 2012b: multilocus phylogenetic analyses of DNA 355, 2013b:54; Siler et al., 2012b:454, sequence data (Fig. 1; Siler et al., 2014a). 2014a:205. Combined with biogeographic evidence and Diagnosis.—Pseudogekko compresicorpus can allopatric distributions, our data support the be distinguished from congeners by the interpretation of four distinct evolutionary following combination of characters: (1) body lineages (full evolutionary species) within the size large (SVL 54.9–59.7 mm); (2) axilla– P. compresicorpus Complex. groin distance 49.3–55.2% SVL; (3) head The type of the genus, Pseudogekko com- length 16.1–18.0% SVL; (4) snout long 56.9– presicorpus (Taylor, 1915), was described on 64.6% head length; (5) Toe IV scansors 18 or the basis of a specimen from the Bataan 19; (6) paravertebral scales 226–234; (7) Peninsula of Luzon Island (Fig. 1). Our new ventral scales 127–130; (8) supralabials 16– specimens from localities close to Taylor’s 20; (9) infralabials 13–16; (10) circumorbitals (1915) type locality match the holotype 39–45; (11) precloacal pores 13 or 14; (12) description in all regards. Additionally, several femoral pores absent; (13) dominant body distinct morphological characters not empha- coloration dark brown to tan; (14) conspicu- sized by Taylor (1915) closely ally with ous head spotting present, neon green; (15) sampled populations from the northern Phi- conspicuous dorsolateral spotting present, lippines (Fig. 1, Clade D; Siler et al., 2014a) faint, neon green; (16) conspicuous limb and with all previously published accounts and spotting present, faint, neon green; (17) tail references to the holotype (Taylor, 1915, banding absent; (18) body striping absent; 1922a): (1) infralabials 13–16, (2) precloacal (19) interorbital banding absent; and (20) pores 13–14, and (3) body coloration homog- ciliary ring coloration present, light blue enous brown and unpatterned. Accordingly, (Fig. 3; Tables 1 and 2). we recognize P. compresicorpus as a species Comparisons.—Characters distinguishing that occurs in the northern Philippines, Pseudogekko compresicorpus from all other restricting the species distribution to the species of Pseudogekko are summarized in Luzon PAIC, Romblon Island Group, and Tables 1 and 2. Pseudogekko compresicorpus Masbate Island (Visayan PAIC). Furthermore, most closely resembles P. pungkaypinit sp. we recognize large-bodied populations from nov. However, it differs from this species by Mindanao Island (Fig. 1; Clade C), small- having a shorter total body length (TotL , bodied populations from Leyte Island (Fig. 1; 117.3 mm vs. .125.3), narrower body (MBW Clade B), and small-bodied populations from ,6.7 mm vs. .7.7), fewer infralabials (13–16 the Zamboanga Peninsula of Mindanao Island vs. 17–19), fewer circumorbitals (39–45 vs. (Fig. 1; Clade A) as members of three, unique 50–55), fewer precloacal pores (13 or 14 vs. evolutionary lineages which we describe 17–20), fewer paravertebral scales (226–234 below as new species. vs. 265–280), a dark brown to tan (vs. grayish brown) body coloration, and by the presence (vs. absence) of conspicuous neon green spots TAXONOMIC ACCOUNTS on the head, dorsolateral region of the body, Pseudogekko compresicorpus (Taylor, 1915) and limbs, absence (vs. presence) of striped (Figs. 1–8) pigmentation patterns on the body, and Luperosaurus compresicorpus Taylor, 1915:96, presence (vs. absence) of a light blue ciliary holotype female (Philippine Bureau of ring. Science 1781) from ‘‘Limay, Bataan Prov- Pseudogekko compresicorpus can be distin- ince, Luzon,’’ Brown and Tanner, 1949:41. guished from P. ditoy sp. nov. and P. 2014] HERPETOLOGICAL MONOGRAPHS 119

FIG. 2.—Ventral surface of head of (A) Lepidodactylus labialis (CAS 133396) and (B) Pseudogekko compresicorpus (KU 331657) and precloacal pore-bearing scale series of (C) adult male L. labialis (CAS 133318) and (D) adult male P. compresicorpus (KU 331657). Scale bar 5 2 mm. Illustrations by CDS. chavacano sp. nov. by having a greater length (SVL . 54.9 mm vs. ,52.6), a greater number of paravertebral scales (.226 vs. number of Toe IV scansors (18 or 19 vs. 16 or ,197) and ventral scales (.127 vs. ,123); 17), fewer precloacal pores (13–14 vs. 18), by from P. ditoy by having a longer snout–vent the presence (vs. absence) of neon green spots on the head, dorsolateral region of the body and limbs, and presence (vs. absence) of a light blue ciliary ring; from P. chavacano by having a longer total body length (TotL . 111.9 mm vs. 95.8), a shorter relative head length (HL 16–18% SVL vs. 19%), fewer circumorbitals (39–45 vs. 46), fewer precloacal pores (13 or 14 vs. 16), and by the absence (vs. presence) of tail banding and presence (vs. absence) of a light blue ciliary ring; from P. brevipes by having a longer snout–vent length FIG. 3.—Lateral view of head of Pseudogekko compresicorpus (KU 326436). Scale bar 5 2 mm. Illustration (SVL . 54.9 mm vs. ,52.5), a greater number by CDS. of Finger III scansors (15–17 vs. 12–14), a 120 HERPETOLOGICAL MONOGRAPHS [No. 28

FIG. 4.—Photographic plates at 315 magnification of lateral and ventral views of the head and dorsal views of the trunk of preserved specimens for (A) Pseudogekko compresicorpus (KU 326436), (B) Pseudogekko pungkaypinit sp. nov. (Holotype, PNM 9810, formerly KU 326435), (C) Pseudogekko ditoy sp. nov. (Paratype, KU 326437), and (D) Pseudogekko chavacano sp. nov. (Holotype, PNM 9812, formerly KU 314963). Scale bar 5 2 mm. Photographs by JLW and CDS. 2014] HERPETOLOGICAL MONOGRAPHS 121

coloration, absence (vs. presence) of femoral pores, presence of dense, neon green spots on the head (vs. dense black and sparse white), presence of neon green (vs. large black and small white) dorsolateral spotting, presence of neon green limb spotting (vs. sparse black and white), absence of tail banding (vs. presence, neon yellow, white, and neon orange), and presence (vs. absence) of a light blue ciliary ring. Description(basedondescriptionofholotype [Taylor, 1915] and 7 referred specimens).— Details of the head scalation are shown in Figures 3 and 4A. Measurements and meristic data scored from the holotype are provided below in brackets. Body small, slender, SVL 54.9–59.7 mm (females), 55.9–58.8 mm (males) [62.0]; limbs well developed, moderately slender; tail slender; margins of limbs smooth, lacking cutaneous flaps or dermal folds; trunk lacking ventrolateral cutaneous fold. Head size moderate, slightly differentiated from neck, characterized by only slightly hypertrophied temporal and adductor muscu- lature; snout rounded in dorsal and lateral aspect (Fig. 4A); HW 118.2–146.8% MBW [120%], 77.2–89.2% HL; HL 16.1–18.0% SVL; SNL 67.9–78.8% HW, 56.9–64.6% HL;

FIG. 5.—Illustration of Star of David configuration dorsal surfaces of head relatively homoge- formed by interstitial granules surrounding body scales, neous, with only slightly pronounced concave visible under high magnification. Magnification 325. postnasal, internasal, prefrontal, and interor- Illustration by CDS. bital concavities; auricular opening moderate, ovoid, angled slightly anteroventrally and posterodorsally from beneath temporal swell- greater number of Toe IV scansors (18 or 19 ings on either side of head; tympanum deeply vs. 13–15), a greater number of infralabials sunken; eye large; pupil vertical, margin wavy (13–16 vs. 12–14), a greater number of (Fig. 4A); limbs and digits relatively short and circumorbitals (39–45 vs. 35–37), and by the moderately slender; thighs moderately thicker presence of dense, neon green spots on the compared to brachium; tibia length 7.2–9.6% head (vs. sparse and cream colored), presence SVL, 53.5–64.6% femur length. of neon green (vs. cream colored) dorsolateral Rostral rectangular in anterior view, 33 as spotting, presence (vs. absence) of neon broad as high, sutured anterolaterally with green limb spotting, absence (vs. presence) of anteriormost enlarged supranasals, projecting interorbital banding, and presence (vs. absence) onto dorsal surface of head to point in line of a light blue ciliary ring; and from P. with midline of nasal; nostril surrounded by smaragdinus by having a greater number of first labial, or first and second labials, rostral, circumorbitals (39–45 vs. 33–35), fewer en- one or two enlarged postnasals, and one or larged pores (13 or 14 precloacal pores vs. 32– two enlarged supranasals; supranasals sepa- 41 precloacal-femoral pores), fewer paraverte- rated by 2–5 small median scales, or touching bral scales (226–234 vs. 241–252), dark brown at midline; enlarged supranasals equal in size to tan (vs. bright neon yellow to orange to enlarged postnasals or greatly enlarged [undisturbed] to neon green [disturbed]) body compared to postnasals. 122 HERPETOLOGICAL MONOGRAPHS [No. 28

FIG. 6.—Illustrations of precloacal pore-bearing scale series of adult males for (A) Pseudogekko compresicorpus (KU 331657), (B) Pseudogekko pungkaypinit sp. nov. (Holotype, PNM 9810, formerly KU 326435), (C) Pseudogekko ditoy sp. nov. (Paratype, KU 326438), and (D) Pseudogekko chavacano sp. nov. (Holotype, PNM 9812, formerly KU 314963). For comparison, the precloacal-femoral pore-bearing scale series of (E) P. smaragdinus (KU 302824) is provided for reference. Scale bar 5 2 mm. Illustrations by CDS.

Total number of differentiated supralabials scales; postrictal scales undifferentiated; re- 16–20, bordered dorsally by one row of maining undifferentiated gulars very small, differentiated, slightly enlarged snout scales; round, nonimbricate, juxtaposed (Fig. 4A), each total number of differentiated infralabials 13–16 scale surrounded by six interstitial granules, [16], bordered ventrally by 3–5 rows of slightly giving the appearance of a Star of David enlarged scales; undifferentiated chin and gular configuration under high magnification (Fig. 5). 2014] HERPETOLOGICAL MONOGRAPHS 123

convex, juxtaposed, relatively homogeneous in size; each dorsal scale surrounded by six interstitial granules; dorsals sharply transition to imbricate ventrals along lateral body surface; paravertebrals between midpoints of limb insertions 226–234; ventrals between midpoints of limb insertions 127–130; scales on dorsal surfaces of limbs more imbricate than dorsals; scales on dorsal surfaces of hands and feet similar to dorsal limb scales, heavily imbricate; ventral body scales flat, cycloid, strongly imbricate, much larger than lateral or dorsal body scales, relatively homogeneous in size. Ten to 14 pores [14], in continuous precloacal pore-bearing series, arranged in a widely obtuse, W-formation (Fig. 6); patch of slightly enlarged scales posterior to precloacal series, roughly three scale rows in size, forming an oval patch; precloacals situated atop a substantial precloacal bulge. Digits moderately expanded and covered on palmar and plantar surfaces by bowed, unnotched, undivided scansors (Fig. 7); digits with minute vestiges of interdigital webbing; subdigital scansors of Finger III 15–17, Toe IV 18–19; subdigital scansors of hands and feet bordered proximally (on palmar and plantar surfaces) by 1–4 slightly enlarged scales that form a near-continuous series with enlarged scansors; all digits clawed, but first claw greatly reduced (Fig. 7); remaining terminal phalanges compressed, with large recurved claws (Fig. 7). Tail short, 52.9–58.5 mm [48], 89.7–99.6% FIG. 7.—Illustration of left hand and foot of Pseudo- SVL [77.4%]; round, not heavily depressed; gekko compresicorpus (KU 326436). Scale bar 5 2 mm. TH 76.1–91.9% TW; caudals similar in size to Illustrations by CDS. dorsals, subcaudals similar in size to ventrals. Coloration in preservative (based on Dorsal cephalic scales fairly homogeneous seven referred specimens).—Background dor- in size, shape, disposition, and distribution; sal body coloration light tan with intermittent cephalic scalation slightly convex, round to small cream and dark brown speckles; pattern oval scales; postnasal, prefrontal, internasal, continued down tail but speckled areas are and interorbital depressions; undifferentiated concentrated occasionally into larger blotches; posterior head scales granular, slightly convex; dorsal region of head with same color pattern, throat and chin scales small, juxtaposed, except for darker brown interorbital region; and nonimbricate, making a moderately sharp dorsal surfaces of limbs with same color transition to gular and pectoral region scala- pattern; one individual with sparse, orange- tion, with enlarged cycloid, imbricate scales; tan spots on arms and legs (KU 331657); circumorbitals 39–45. lateral region of body with same coloring Axilla–groin distance 49.3–55.2% SVL [58.1]; pattern as dorsal region; lateral region of head undifferentiated dorsal body scales round, with same color patterns as body, except for 124 HERPETOLOGICAL MONOGRAPHS [No. 28

FIG. 8.—Photographs in life of (A) Pseudogekko compresicorpus (KU 326436) and the holotypes of (B) Pseudogekko pungkaypinit sp. nov. (Holotype, PNM 9810, formerly KU 326435), (C) Pseudogekko ditoy sp. nov. (Holotype, PNM 9811, formerly KU 326437), and (D) Pseudogekko chavacano sp. nov. (Holotype, PNM 9812, formerly KU 314963). Photographs by RMB. slightly lighter area just posterior to orbits and trunk, but with random placement light green along both sets of labial scales; circumorbital blotches. Tail colored as trunk, but with series scales with mixture of cream and medium of longitudinally elongate, paravertebral brown coloration; ventral side of body with cream blotches. Ventral coloration light gray background cream color, with speckling pat- ground color with minimal chocolate brown tern of lateral side of body wrapping around to mottling along the lateral margins. Taylor approximately halfway to midpoint of venter; (1915) reports simply that the holotype was ventral surfaces of head with same color cinnamon brown in life. pattern as body; ventral surfaces of limbs Distribution.—Pseudogekko compresicorpus cream with medium to dark brown speckles occurs on the Luzon PAIC (Luzon and Polillo scattered sparsely throughout; palmar and islands), Visayan PAIC (Masbate Island), and plantar surfaces solid cream, except medium the Romblon Island Group (Tablas Island; brown regions between scansors; ventral Fig. 1). Although currently not recorded from surface of tail solid cream with occasional other islands in the Luzon PAIC, we would not light brown speckles, speckles increase in be surprised if future surveys discover addi- intensity towards tail tip. tional island populations (i.e., Catanduanes Coloration in life (based on CDS and RMB Island). field notes and photographs in life; Fig. 8A).— Ecology and natural history.—Pseudogekko Dorsal ground color of head, trunk, and tail compresicorpus has been observed in first- chocolate brown. Head with conspicuous light and secondary-growth forest (Fig. 1) on leaves green mottling in canthal and interocular of shrubs and small trees 2–4 m above the regions as well as on lateral surface between ground. Taylor (1922a) notes that two, fully the eye and ear opening. Dorsolateral surface developed embryos were found in eggs that of body with series of eight faint, light green had been attached to the underside of a leaf at blotches running from nuchal region onto the top of a recently felled tree. This natural base of tail. Dorsal limb surfaces colored as history observation may indicate that species 2014] HERPETOLOGICAL MONOGRAPHS 125 of this genus may have a more arboreal A.C. Alcala; one adult male (KU 324426) lifestyle than currently is appreciated and collected on 6 August 2009 in Raja Sikatuna that, although encountered by us in forest Natural Park, Barangay Danicop, Municipality of lower strata, this species may also inhabit tree Sierra Bullones, Bohol Province, Bohol Island, canopies. Philippines (09u42919.8360N, 124u7924.3840E; Both Pseudogekko compresicorpus and P. WGS-84) by C.D. Siler; one adult female smaragdinus occur on Luzon and Polillo (KU 326243) collected on 17 April 2008 on islands (Fig. 1); however, there is insufficient the campus of Visayas State University, Baybay evidence to determine if populations of these City, Municipality of Baybay, Leyte Province, species occur in sympatry. Similar to P. Leyte Island, Philippines (10u44944.4660N, pungkaypinit sp. nov., numerous populations 124u47929.22720E; WGS-84) by R.M. Brown; of P. compresicorpus have been observed in one adult male (KU 334019) collected on 19 the wild as compared to other species in the July 2012 near Ginoog River, Mt. Lumot, Sitio genus. In fact, as currently recognized, P. Kibuko, Barangay Lawaan, Municipality of compresicorpus possesses the broadest geo- Gingoog City, Misamis Oriental Province, graphic distribution of any species in the Mindanao Island, Philippines (08u4990.33960N, genus. Unfortunately, even with this broad 125u5944.32920E; WGS-84) by R.M. Brown. distribution, few specimens exist in museum Diagnosis.—Pseudogekko pungkaypinit can collections. At this time we do not find this be distinguished from congeners by the species qualifies for Critically Endangered, following combination of characters: (1) body Endangered, Vulnerable, or Near Threatened size large (SVL 66.6–76.8 mm); (2) axilla– status under the IUCN criteria for classifica- groin distance 53.7–55.9% SVL; (3) head tion (IUCN, 2013). Therefore, we recommend length 17.1–17.7% SVL; (4) snout 53.6– that the species be considered Least Concern 60.2% head length; (5) Toe IV scansors 17– until additional information can be obtained 21; (6) paravertebral scales 265–280; (7) concerning the health and diversity of wild ventral scales 125–155; (8) supralabials 16– populations throughout the Luzon and Visa- 20; (9) infralabials 17–19; (10) circumorbitals yan PAICs and Romblon Island Group. 50–55; (11) precloacal pores 17–20; (12) femoral pores absent; (13) dominant body Pseudogekko pungkaypinit sp. nov. coloration grayish brown; (14) head, body and (Figs. 1, 4, 6, 8) tail immaculate; (15) body stripes present; (16) interorbital band absent; and (17) ciliary Holotype.—PNM 9810 (RMB Field ring coloration undifferentiated (Tables 1, 2). No. 4392, formerly KU 326435), adult male, Comparisons.—Characters distinguishing collected in secondary-growth forest on 3 Pseudogekko pungkaypinit from all other September 2002, in the Calbiga—a creek area species of Pseudogekko are summarized in on the Visayas State University Visca campus, Tables 1 and 2. Pseudogekko pungkaypinit Barangay Guadalupe, Municipality of Baybay, most closely resembles P. compresicorpus; Leyte Province, Leyte Island, Philippines however, P. pungkaypinit differs from P. (10u4590.270N, 124u47924.00E; WGS-84), by compresicorpus by having a longer total R.M. Brown. body length (TotL . 125.3 mm vs. ,117.3), Paratypes.—One adult female (CAS 131854) wider body (MBW . 7.7 mm vs. ,6.7), more collected from the bark of a rotten stump on 31 infralabials (17–19 vs. 13–16), circumorbitals March 1964 in Dusita Barrio, Municipality of (50–55 vs. 39–45), precloacal pores (17–20 vs. Sierra Bullones, Bohol Province, Bohol Island, 13 or 14), and paravertebral scales (265–280 Philippines (09u46959.340N, 124u18910.80E; vs. 226–234); a grayish brown (vs. dark brown WGS-84) by S. Magusara; one adult male to tan) body coloration, the absence (vs. (CAS-SU 23655) collected on floor of lowland presence) of neon green spots on the head, bamboo forest on 9 May 1962 in Dusita Barrio, dorsolateral region of the body, and limbs, the Municipality of Sierra Bullones Municipality, presence (vs. absence) of dark stripes on the Bohol Province, Bohol Island, Philippines body, and absence (vs. presence) of a light (09u46957.50N, 124u18910.80E; WGS-84) by blue ciliary ring. 126 HERPETOLOGICAL MONOGRAPHS [No. 28

Pseudogekko pungkaypinit can be distin- HL; HL 17.3% SVL; SNL 69.6% HW, 60.2% guished from P. ditoy sp. nov., P. chavacano HL; dorsal surfaces of head relatively homo- sp. nov., P. brevipes, and P. smaragdinus by geneous, with only slightly pronounced con- having a longer body (SVL . 66.6 mm vs. cave postnasal, internasal, prefrontal, and ,64.3), a longer trunk (AGD . 37.2 mm vs. interorbital concavities; auricular opening ,35.5), more circumorbitals (50–55 vs. ,46) moderate, ovoid, angled slightly anteroven- and paravertebral scales (.265 vs. ,252), and trally and posterodorsally from beneath by the presence (vs. absence) of dark dorso- temporal swellings on either side of head; lateral body stripes; from P. ditoy and P. tympanum deeply sunken; eye large; pupil chavacano by having a wider body (MBW . vertical, margin wavy (Fig. 4B); limbs and 7.7 mm vs. ,7.3); from P. chavacano by digits relatively short and moderately slender; having a shorter relative head length (HL 17– thighs moderately thicker compared to bra- 18% SVL vs. 19%), a greater number of chium; tibia length 10.1% SVL, 63.5% femur precloacal pores (17–20 vs. 16), and by the length. absence (vs. presence) of neon green spots on Rostral rectangular in anterior view, 33 as the head, dorsolateral region of the body, broad as high, sutured anterolaterally with and limbs, and absence (vs. presence) of tail anteriormost enlarged supranasals; nostril sur- bands; from P. brevipes by having greater rounded by first labial, rostral, one enlarged numbers of Finger III scansors (15–17 vs. 12– postnasal, and two enlarged supranasals; su- 14), Toe IV scansors (17–21 vs. 13–15), pranasals separated by five small median infralabials (17–19 vs. 12–14), and precloacal scales. pores (17–20 vs. 13–15), and by the absence Total number of differentiated supralabials (vs. presence) of cream spots on the head and 18/19 (left/right [L/R]), bordered dorsally by dorsolateral region of the body, and absence 1–2 rows of differentiated, slightly enlarged (vs. presence) of interorbital band; and from snout scales; total number of differentiated P. smaragdinus by having fewer enlarged infralabials 17/17 (L/R), bordered ventrally by pores (17–20 precloacal pores vs. 32–41 7–9 rows of slightly enlarged scales; undiffer- precloacal-femoral pores), its grayish brown entiated chin and gular scales; postrictal scales (vs. bright neon yellow to orange [undis- undifferentiated; remaining undifferentiated turbed] to neon green [disturbed]) body gulars very small, round, nonimbricate to coloration, and by the absence (vs. presence) slightly imbricate, juxtaposed (Fig. 4B). of femoral pores, absence (vs. presence) of Dorsal cephalic scales fairly homogeneous in black and white spots on the head, dorsolat- size, shape, disposition, and distribution; ce- eral surfaces of the body and limbs, and phalic scalation slightly convex, round to oval absence (vs. presence) of transverse tail scales; postnasal, prefrontal, internasal, and bands. interorbital depressions; undifferentiated pos- Description of holotype.—Details of the terior head scales granular, slightly convex; head scalation are shown in Figure 4B. Adult throat and chin scales small, juxtaposed, and male in excellent condition, hemipenes evert- nonimbricate, making a moderately sharp ed, hemipenal bulge present; small incision in transition to gular and pectoral region scalation, the sternal region (portion of liver removed with enlarged cycloid, imbricate scales; circu- for genetic sample). Body small, slender, SVL morbitals 51/53 (L/R); interorbital scales 74. 71.9 mm; limbs well developed, moderately Axilla–groin distance 54.3% SVL; undiffer- slender; tail original, slender; margins of limbs entiated dorsal body scales round, convex, smooth, lacking cutaneous flaps or dermal juxtaposed, relatively homogeneous in size; folds; trunk lacking ventrolateral cutaneous dorsal scales surrounded by interstitial gran- fold. ules, however, granules do not give clear Head moderate in size, slightly differenti- appearance of a Star of David configuration ated from neck, characterized by only slightly under high magnification; dorsals sharply hypertrophied temporal and adductor muscu- transition to imbricate ventrals along lateral lature; snout rounded in dorsal and lateral body surface; paravertebrals between mid- aspect (Fig. 4B); HW 132.3% MBW, 86.5% points of limb insertions 280; ventrals between 2014] HERPETOLOGICAL MONOGRAPHS 127 midpoints of limb insertions 140; scales on to approximately halfway to midpoint of dorsal surfaces of limbs more imbricate than venter; ventral region of head follows same dorsals; scales on dorsal surfaces of hands and pattern as trunk; ventral regions of limbs, feet similar to dorsal limb scales, strongly hands, and feet are cream with medium imbricate; ventral body scales flat, cycloid, to dark brown speckles scattered sparsely strongly imbricate, much larger than lateral or throughout; ventral surface of tail is solid dorsal body scales, relatively homogeneous in cream with occasional light brown speckles size. that increase in intensity towards tip. Twenty enlarged pore-bearing scales pres- Coloration in life (based on field notes and ent, in continuous precloacal pore-bearing photographs in life; Fig. 8B).—Dorsal ground series, arranged in a widely obtuse, W- color of head, trunk and tail olive-brown; formation (Fig. 6); precloacals 10/10 (L/R); lateral surfaces of head and trunk with series patch of slightly enlarged scales posterior to of light brown stripes, from eye to leg precloacal series, roughly four scale rows in insertion, stripes oriented posteroventrally; size, forming an oval patch; precloacals dorsal limb color similar to trunk, but lacking situated atop a substantial precloacal bulge. any discernable pattern; dorsal tail color Digits moderately expanded and covered on similar to trunk, but with occasional small, palmar/plantar surfaces by bowed, unnotched, cream spots; ventrum light cream ground undivided scansors; digits with minute vestig- color with minimal light olive-brown mottling es of interdigital webbing; subdigital scansors along the lateral margins. of hand: 11/10, 12/12, 15/14, 17/17, and 11/11 Measurements and scale counts of holotype (L/R) on Finger I–Finger V, respectively; foot: in millimeters.—Snout–vent length 71.9; tail 12/12, 14/14, 16/17, 19/19, and 12/12 (L/R) on length 67.2; total length 139.1; axilla–groin Toe I–Toe V, respectively; subdigital scansors distance 39.1; tail width 3.6; tail height 3.2; of hands and feet bordered proximally (on head length 12.4; head width 10.7; head palmar and plantar surfaces) by 1–4 slightly height 7.0; midbody width 8.1; snout length enlarged scales that form a near-continuous 7.5; eye diameter 4.8; eye–nares distance 5.9; series with enlarged scansors; all digits clawed internarial distance 2.8; interorbital distance but first claw greatly reduced; remaining 5.8; femur length 11.4; tibia length 7.3; Finger terminal phalanges compressed, with large III scansors 17; Toe IV scansors 19; suprala- recurved claws. bials 18; infralabials 17; circumorbitals 51; Tail short, 67.2 mm, 93.5% SVL; round, not preanofemoral pores 20; paravertebral scales heavily depressed; TH 88.4% TW; caudals 280; ventral scales 140. similar in size to dorsals, subcaudals similar in Variation.—Among the six specimens ex- size to ventrals. amined, we observed variation in the numbers Coloration of holotype in preservative.— of precloacal pores, supralabials, infralabials, Background dorsal color medium brown with and digital scansors. The number of suprala- small dark brown speckles throughout, pattern bials varied between 16 (KU 334019), 18 (KU continued down tail; dorsal nuchal region 326243, 326435), 19 (CAS-SU 23655, KU slightly lighter brown than remainder of body; 324426), and 20 (CAS 131854); infralabials dorsal regions of head and limbs have same varied between 17 (CAS 131854, KU 324426, color pattern as body; lateral regions of trunk 326243, 326435), 18 (CAS-SU 23655), and 19 have same coloration pattern as dorsum; (KU 334019). Precloacal pore counts in adult lateral side of head has thin cream circumor- males were observed to vary between 17 (KU bital ring of scales, lighter brown area on 324426), 19 (CAS-SU 23655, KU 334019), and nuchal region continues to just posterior 20 (KU 326435). The number of Finger III to orbit; remainder of head continues same scansors varied between 15 (CAS 131854, pattern as trunk, except supralabial and CAS-SU 23655, KU 326243), 16 (KU infralabial scales are solid cream with few 334019), and 17 (KU 324426, 326435); Toe dark brown speckles; ventral side of trunk has IV scansors varied between 17 (CAS-SU cream background color, with speckling pat- 23655), 18 (CAS 131854, KU 326243), 19 tern of lateral side of trunk wrapping around (KU 326435, 334019), and 20 (KU 324426). 128 HERPETOLOGICAL MONOGRAPHS [No. 28

Distribution.—Pseudogekko pungkaypinit arboreal microhabitat preference. The specific is known only from Bohol, Leyte, and Minda- epithet is a noun of masculine gender. Sug- nao islands, although it may be possible that it gested common name: Southern Philippine will eventually be discovered on other islands False Gecko. in the Mindanao PAIC (i.e., Samar, Dinagat, Biliran, Siargao; Fig. 1). Pseudogekko ditoy sp. nov. Ecology and natural history.—Pseudogekko (Figs. 1, 4–6, 8) pungkaypinit has been observed in disturbed, secondary-growth forest only; however, simi- Holotype.—PNM 9811 (RMB Field lar to assumptions about other species in the No. 4365, formerly KU 326437), adult female, genus, we assume this species once occurred in collected in secondary-growth forest on 28 lower elevation primary forest on Bohol, Leyte, June 1999, in Sitio Cienda, Barangay Gabas, and Mindanao islands and possibly throughout Municipality of Baybay, Leyte Province, Leyte much of the Mindanao PAIC. All individuals in Island, Philippines (10u419N, 124u489E; museum collections have been collected on top WGS-84), by R.M. Brown. of leaves of shrubs 2–4 m above the ground. In Paratype.—One adult male (KU 326438) addition to the new species, P. chavacano also collected in lowland, secondary-growth forest occurs on Mindanao Island; however, at (2030 to 2300 h) on 31 August 2001, in the present there is no evidence that these two Calbiga—a river area, Barangay Gabas, Mu- species overlap in distributions, with P. pung- nicipality of Baybay, Leyte Province, Leyte kaypinit seemingly distributed in the central Island, Philippines (10u419N, 124u489E; and eastern regions of Mindanao Island and P. WGS-84) by A.C. Diesmos. chavacano found in the far western region of Diagnosis.—Pseudogekko ditoy can be dis- Mindanao Island (Zamboanga Peninsula; tinguished from congeners by the following Fig. 1). Likewise, P. pungkaypinit and P. ditoy combination of characters: (1) body size small occur in sympatry on Leyte Island, and both P. (SVL 49.4–52.6 mm); (2) axilla–groin distance brevipes and P. pungkaypinit occur on Bohol 50.7–56.5% SVL; (3) head length 18.2–18.9% Island, although whether they occur in sym- SVL; (4) snout 57.4–59.3% head length; (5) patry is unclear. Pseudogekko pungkaypinit is Toe IV scansors 16 or 17; (6) paravertebral known from four separate populations on three scales 180–185; (7) ventral scales 111–118; (8) islands. Although only a handful of individuals supralabials 17–20; (9) infralabials 16 or 17; exist in museum collections, this species (10) circumorbitals 40 or 43; (11) precloacal appears to be more-widely distributed and, pores 18; (12) femoral pores absent; (13) therefore, possibly experiencing less overall ground body coloration light brown; (14) head conservation threats than its congeners. As a spots absent; (15) dorsolateral body spots result of this species broad geographic distri- absent; (16) limb spots absent; (17) tail bands bution (as currently recognized) and the lack of absent; (18) body stripes absent; (19) interor- available information about the species ecolo- bital band absent; and (20) ciliary ring gy, natural history, and intraspecific diversity, coloration undifferentiated (Tables 1, 2). we do not find this species qualifies for Comparisons.—Characters distinguishing Critically Endangered, Endangered, Vulnera- Pseudogekko ditoy from all other species of ble, or Near Threatened status under the Pseudogekko are summarized in Tables 1 and IUCN criteria for classification (IUCN, 2013). 2. Pseudogekko ditoy most closely resembles Therefore, we recommend that the species be P. chavacano; however, P. ditoy differs from considered Least Concern until additional P. chavacano by having tendencies towards information can be obtained concerning the fewer Finger III scansors (14 or 15 vs. 15 or health and diversity of wild populations 16), fewer Toe IV scansors (16 or 17 vs. 17– throughout the Mindanao PAIC. 20), more supralabials (17–20 vs. 15 or 16), Etymology.—We derive the new species fewer circumorbitals (40–43 vs. 46), fewer name from the Leyte language (Waray-Waray) paravertebral scales (180–185 vs. 195–197), terms for ‘‘treetop’’ (pungkay) and ‘‘lizard’’ fewer ventral scales (111–118 vs. 122–123), (pinit) in reference to the new forest gecko’s more precloacal pores (18 vs. 16), and by the 2014] HERPETOLOGICAL MONOGRAPHS 129 absence (vs. presence) of neon green head, Head size moderate, slightly differentiated dorsolateral body, and limbs spots and ab- from neck, characterized by only slightly sence (vs. presence) of transverse tail bands. hypertrophied temporal and adductor muscu- Pseudogekko ditoy can be distinguished lature; snout rounded in dorsal and lateral from P. compresicorpus, P. pungkaypinit, P. aspect (Fig. 4C); HW 105.4% MBW, 80.2% brevipes, and P. smaragdinus by having fewer HL; HL 18.3% SVL; SNL 74.0% HW, 59.4% paravertebral scales (,185 vs. .195) and HL; dorsal surfaces of head relatively homo- ventral scales (,118 vs. .119); from P. geneous, with only slightly pronounced con- brevipes by having greater numbers of Toe cave postnasal, internasal, prefrontal, and IV scansors (16 or 17 vs. 13–15), supralabials interorbital concavities; auricular opening (17–20 vs. 14–16), infralabials (16 or 17 vs. moderate, ovoid, angled slightly anteroven- 12–14), circumorbitals (40–43 vs. 35–37), and trally and posterodorsally from beneath tem- precloacal pores (18 vs. 13–15), and by the poral swellings on either side of head; absence (vs. presence) of cream colored tympanum deeply sunken; eye large; pupil spotting on the head and dorsolateral regions vertical, margin wavy (Fig. 4C); limbs and of the body and absence (vs. presence) of digits relatively short and moderately slender; interorbital banding; from P. compresicorpus thighs moderately thicker compared to bra- and P. pungkaypinit by having a shorter chium; tibia length 8.0% SVL, 50.1% femur snout–vent length (SVL , 52.6 mm vs. length. .54.9); from P. compresicorpus by having Rostral rectangular in anterior view, 33 as fewer Toe IV scansors (16 or 17 vs. 18 or 19) broad as high, sutured anterolaterally with and a greater number of precloacal pores (18 anteriormost enlarged supranasals; nostril sur- vs. 13 or 14), and by the absence (vs. presence) rounded by rostral, first labial, single enlarged of neon green spotting on the head, dorsolat- postnasal, and two enlarged supranasals; supra- eral regions of the body, and limbs and absence nasals separated by five small median scales. (vs. presence) of a light blue ciliary ring; from Total number of differentiated supralabials P. pungkaypinit by having a shorter trunk 20/17 (L/R), bordered dorsally by one or two length (AGD , 29.7 mm vs. .37.2), narrower rows of differentiated, slightly enlarged snout body (MBW , 7.3 mm vs. .7.7), fewer scales; total number of differentiated infra- circumorbitals (40–43 vs. 50–55), and by the labials 17/17 (L/R), bordered ventrally by 5–8 absence (vs. presence) of striped pigmentation rows of slightly enlarged scales; undifferenti- patterns on the body; and from P. smaragdinus ated chin and gular scales; postrictal scales by having a greater number of circumorbitals undifferentiated; remaining undifferentiated (40–43 vs. 33–35), fewer enlarged precloacal gulars very small, round, nonimbricate, juxta- pores (18 precloacals vs. 32–41 precloacal- posed (Fig. 4C), each scale surrounded by six femorals) and a light brown (vs. bright neon interstitial granules, giving the appearance of a yellow to orange [undisturbed] to neon green Star of David configuration under high [disturbed]) body coloration, and absence (vs. magnification (Fig. 5). presence) of black and white spots on the head, Dorsal cephalic scales fairly homogeneous body and limbs and absence (vs. presence) of in size, shape, disposition, and distribution; transverse tail bands. cephalic scalation slightly convex, round to Description of holotype.—Details of the oval scales; postnasal, prefrontal, internasal, head scalation are shown in Figure 4C. Adult and interorbital depressions; undifferentiated female in excellent condition, gravid, with two posterior head scales granular, slightly convex; embryos visible through venter; small incision throat and chin scales small, juxtaposed and in the sternal region (portion of liver removed nonimbricate, making a moderately sharp for genetic sample). Body small, slender, SVL transition to gular and pectoral region scala- 52.6 mm; limbs well developed, moderately tion, with enlarged cycloid, imbricate scales; slender; tail regenerated, slender; margins of circumorbitals 43/48 (L/R); interorbitals 42. limbs smooth, lacking cutaneous flaps or Axilla–groin distance 56.5% SVL; undiffer- dermal folds; trunk lacking ventrolateral entiated dorsal body scales round, convex, cutaneous fold. juxtaposed, relatively homogeneous in size; 130 HERPETOLOGICAL MONOGRAPHS [No. 28 each dorsal scale surrounded by six interstitial region just anterior to the eyes. In the granules; dorsals sharply transition to imbri- postnasal region there is a stripe of medium cate ventrals along lateral body surface; brown speckles on top of the lighter tan and paravertebrals between midpoints of limb cream background color. The lateral region of insertions 185; ventrals between midpoints of the trunk has the same coloring pattern as the limb insertions 118; scales on dorsal surfaces dorsal region. The lateral region of the head of limbs more imbricate than dorsals; scales has a thick, cream-colored ring of circumor- on dorsal surfaces of hands and feet similar to bital scales and a vague line of light brown dorsal limb scales, heavily imbricate; ventral blotches from snout to orbit. The venter of the body scales flat, cycloid, strongly imbricate, body is solid cream except for occasional light much larger than lateral or dorsal body scales, brown speckles. The ventral region of the relatively homogeneous in size. tail is solid cream with occasional light brown Nineteen enlarged scales (pore-bearing in speckles that increase in intensity as you males; Fig. 6), in continuous precloacal series, continue to the tip. The gular region of the arranged in a widely obtuse, W-formation; head is solid cream with light brown blotches precloacals 9/10 (L/R); patch of slightly just proximate to the infralabials. The ventral enlarged scales posterior to precloacal series, side of the limbs, hands, and feet is a solid roughly six scale rows in size, forming an oval cream except for a light brown stripe between patch. each scansor. In this individual, Finger IV on Digits moderately expanded and covered on the right hand is heavily speckled light brown. palmar/plantar surfaces by bowed, unnotched, Coloration in life (based on field notes and undivided scansors; digits with minute vestig- photographs in life; Fig. 8C).—Dorsal ground es of interdigital webbing; subdigital scansors color of head, trunk, and tail light tan with of hand: 8/8, 10/10, 12/12, 14/15, and 10/10 only faint light brown mottling. Head slightly (L/R) on Finger I–Finger V, respectively; foot: darker in appearance, with increased light 8/9, 11/11, 13/15, 16/16, and 11/11 (L/R) on brown mottling in canthal and interocular Toe I–Toe V, respectively; subdigital scansors regions as well as in posterior margins of of hands and feet bordered proximally (on the head. Lateral surface of the head with palmar and plantar surfaces) by 1–4 slightly conspicuous series of four dark brown stripes, enlarged scales that form a near-continuous extending between the eye and ear opening. series with enlarged scansors; all digits clawed, Stripes oriented anterodorsally from ear but first claw greatly reduced; remaining opening and interspersed with olive green terminal phalanges compressed, with large blotches. Dorsal limb surfaces colored as the recurved claws. trunk, but arms with increased light brown Tail regenerated, short, 30.5 mm, 58.0% mottling and a slightly darker appearance. Tail SVL; single distinct fracture plane present, colored as trunk basally, then transitioning to composed of enlarged, cylindrical scales wrap- a medium gray just posterior to tail insertion. ping around tail, separating original tail (ante- Ventral coloration not apparent, but appears riorly) from regenerated tail (posteriorly); tail to consist of a light cream ground color with round, not depressed; TH 81.3% TW; caudals minimal light brown mottling along the lateral similar in size to dorsals, subcaudals similar in margins. size to ventrals. Measurements and scale counts of holotype Coloration of holotype in preservative.— in millimeters.—Snout–vent length 52.6 mm; Background dorsal body coloration is light tan tail length 30.5 (regenerated); total length 83.0 with intermittent small cream and dark brown (with regenerated tail); axilla–groin distance speckles; this pattern is continued down the 29.7; tail width 2.2; tail height 1.8; head length tail. The dorsal region of the limbs follows the 9.6; head width 7.7; head height 5.5; midbody same pattern with a few areas of concentrated width 7.3; snout length 5.7; eye diameter 3.5; darker blotches. The dorsal region of the head eye–nares distance 4.4; internarial distance has the same speckled pattern as the trunk, 1.9; interorbital distance 4.4; femur length 8.4; except for a darker region between and just tibia length 4.2; Finger III scansors 14; Toe IV posterior to the orbits, and a slightly lighter scansors 16; supralabials 20; infralabials 17; 2014] HERPETOLOGICAL MONOGRAPHS 131 circumorbitals 43; paravertebral scales 185; apposition. Suggested common name: Leyte ventral scales 118. Diminutive False Gecko. Variation.—Between the two specimens examined we observed variation in the num- Pseudogekko chavacano sp. nov. bers of supralabials, infralabials, and digital (Figs. 1, 4–6, 8) scansors. The number of supralabials varied between 17 (KU 326438) and 20 (KU Holotype.—PNM 9812 (ACD Field 326437); infralabials varied between 16 (KU No. 3784, formerly KU 314963), adult male, 326438) and 17 (KU 326437). Finger III collected in secondary-growth forest (2000 scansors were observed to vary between 14 to 2230 h) on 21 April 2008, in Barangay (KU 326437) and 15 (KU 326438); Toe IV Pasonanca, Pasonanca National Park, Zam- scansors varied between 16 (KU 326437) and boanga Sur Province, Mindanao Island, 17 (KU 326438). Philippines (06u58939.030N, 122u04901.650E; Distribution.—Pseudogekko ditoy is known WGS-84), by A.C. Diesmos. only from Leyte Island (Fig. 1). Paratype.—One adult female (KU 314964) Ecology and natural history.—Pseudogekko collected on 12 July 2008 in Sitio km 24, Barangay ditoy has been observed in disturbed, second- Baluno, Pasonanca National Park, Zamboanga ary-growth forest only; however, we assume Sur Province, Mindanao Island, Philippines the species once occurred in lower-elevation (07u193.2240N, 122u1943.860E; WGS-84; 758 m) primary forest on Leyte Island. The two by R.M. Brown; two hatchlings (CAS-SU 23548, individuals represented in museum collections 23549) collected on 8 May 1959 in a mid- were both found on top of leaves of shrubs 2– montane dipterocarp forest near Cuyot Creek, 3 m above the ground. A third individual was Zamboanga del Norte Province, Mindanao observed but not collected on top of a leaf on Island, Philippines (08u1997.970N, 123u3198.40E; a tree 3.5 m above the ground on the bank of a WGS-84) by A.C. Alcala. small stream system near a tree nursery Diagnosis.—Pseudogekko chavacano can be outside of Baybay City (C. Siler, personal distinguished from congeners by the following observation). Interestingly, at least two species combination of characters: (1) body size small of Pseudogekko occur on Leyte Island (P. (SVL 54.7–55.9 mm); (2) axilla–groin distance ditoy and P. brevipes); however, it may be 47.8–54.9% SVL; (3) head length 18.6–19.1% possible that additional exploration of Leyte SVL; (4) snout 55.2–58.0% head length; (5) Island will result in the discovery of popula- Toe IV scansors 17–20; (6) paravertebral scales tions of P. pungkaypinit as well. Whether or 195–197; (7) ventral scales 122 or 123; (8) not P. ditoy and P. brevipes occur in sympatry supralabials 15 or 16; (9) infralabials 16 or 17; across their ranges on Leyte Island is still (10) circumorbitals 46; (11) precloacal pores unknown. Although little data are available on 16; (12) femoral pores absent; (13) dominant the natural history and ecology, as well as on body coloration light brown; (14) conspicuous the health of wild populations of this unique head spotting present, dense, neon green; (15) species, we believe that it qualifies as a conspicuous dorsolateral spotting present, conservation concern. We have evaluated this neon green; (16) conspicuous limb spotting species against the IUCN criteria for classifi- present, dense, neon green; (17) tail banding cation and find that it qualifies for the status of present; (18) body striping absent; (19) inter- Vulnerable, VU, based on the following orbital banding absent; and (20) ciliary ring criteria: VU A2ac; B2ab(iii); D2 (IUCN, coloration absent (Tables 1, 2). 2013). Comparisons.—Characters distinguishing Etymology.—We derive the new species Pseudogekko chavacano from all other species name from the Leyte language (Waray-Waray) of Pseudogekko are summarized in Tables 1 term ditoy, meaning diminutive or ‘‘the and 2. Pseudogekko chavacano most closely smaller one’’ in reference to the new species resembles P. ditoy; however, P. chavacano small body size and its distinction from its differs from P. ditoy by having a tendency larger sympatric congener, P. pungkaypinit. towards a greater number of Finger III The new name is a masculine noun in scansors (15 or 16 vs. 14 or 15), a tendency 132 HERPETOLOGICAL MONOGRAPHS [No. 28 towards a greater number of Toe IV scansors [undisturbed] to neon green [disturbed]), and (17–20 vs. 16 or 17), fewer supralabials (15 or by the absence (vs. presence) of femoral pores 16 vs. 17–20), a greater number of circumor- and presence of neon green (vs. black and bitals (46 vs. 40–43), paravertebral scales white) spotting on the head, dorsolateral (195–197 vs. 180–185), and ventral scales region of the body, and limbs. (122–123 vs. 111–118), and fewer precloacal Description of holotype.—Details of the pores (16 vs. 18), and by the presence (vs. head scalation are shown in Figure 4D. absence) of neon green spotting on the head, Adult male in excellent condition, hemipenes dorsolateral region of the body, and limbs and not inverted, hemipene bulge present; small presence (vs. absence) of tail banding. incision in the sternal region (portion of liver Pseudogekko chavacano can be distin- removed for genetic sample). Body small, guished from P. compresicorpus, P. pungkay- slender, SVL 55.9 mm; limbs well developed, pinit, P. brevipes,andP. smaragdinus by moderately slender; tail regenerated, slender; having fewer paravertebral scales (,197 vs. margins of limbs smooth, lacking cutaneous .226); from P. compresicorpus, P. pungkay- flaps or dermal folds; trunk lacking ventrolat- pinit, and P. smaragdinus by having fewer eral cutaneous fold. ventral scales (,123 vs. .124); from Head moderate in size, slightly differenti- P. brevipes by having greater numbers of ated from neck, characterized by only slightly Finger III scansors (15 or 16 vs. 12–14), Toe hypertrophied temporal and adductor muscu- IV scansors (17–20 vs. 13–15), infralabials (16 lature; snout rounded in dorsal and lateral or 17 vs. 12–14), circumorbitals (46 vs. 35–37), aspect (Fig. 4D); HW 131.7% MBW, 81.4% and precloacal pores (16 vs. 13–15), and by HL; HL 18.6% SVL; SNL 71.2% HW, 58.0% the presence of neon green (vs. cream HL; dorsal surfaces of head relatively homo- colored) spotting on the head and dorsolateral geneous, with only slightly pronounced con- region of the body, presence (vs. absence) of cave postnasal, internasal, prefrontal, and neon green spotting on the limbs, presence interorbital concavities; auricular opening (vs. absence) of tail banding and absence (vs. moderate, ovoid, angled slightly anteroven- presence) of interorbital banding; from P. trally and posterodorsally from beneath compresicorpus, P. pungkaypinit, and P. temporal swellings on either side of head; smaragdinus by having a shorter total body tympanum deeply sunken; eye large; pupil length (TotL , 95.8 mm vs. .103.6) and a vertical, margin wavy (Fig. 4D); limbs and greater relative head length (HL 19% SVL vs. digits relatively short and moderately slender; ,18%); from P. compresicorpus by having thighs moderately thicker compared to bra- greater numbers of circumorbitals (46 vs. 39– chium; tibia length 8.0% SVL, 50.1% femur 45), and precloacal pores (16 vs. 13 or 14), and length. by the presence (vs. absence) of tail banding Rostral rectangular in anterior view, 33 as and absence (vs. presence) of a light blue broad as high, sutured anterolaterally with ciliary ring; from P. pungkaypinit by having a anteriormost enlarged supranasals; nostril sur- shorter trunk length (AGD , 30.0 mm vs. rounded by rostral, first labial, single enlarged .37.2), narrower body (MBW , 6.7 mm vs. postnasal, and two enlarged supranasals; ante- .7.7), fewer circumorbitals (46 vs. 50–55), riormost enlarged supranasals separated by fewer precloacal pores (16 vs. 17–20), by the three small median scales. presence (vs. absence) of neon green spotting Total number of differentiated supralabials on the head, dorsolateral region of the body, 15/15 (L/R), bordered dorsally by one row of and limbs, presence (vs. absence) of tail differentiated, slightly enlarged snout scales; banding and absence (vs. presence) of striped total number of differentiated infralabials 17/ pigmentation patterns on the body; and from 17 (L/R), bordered ventrally by 5–8 rows of P. smaragdinus by having a greater number of slightly enlarged scales; undifferentiated chin circumorbitals (46 vs. 33–35), fewer enlarged and gular scales; postrictal scales undiffer- pores (16 precloacal pores vs. 32–41 precloa- entiated; remaining undifferentiated gulars cal-femoral pores), disparate body coloration very small, round, nonimbricate, juxtaposed (light brown vs. bright neon yellow to orange (Fig. 4D). 2014] HERPETOLOGICAL MONOGRAPHS 133

Dorsal cephalic scales fairly homogeneous composed of enlarged, cylindrical to rectan- in size, shape, disposition, and distribution; gular scales wrapping around tail, separating cephalic scalation slightly convex, round to original tail (anteriorly) from regenerated tail oval scales; postnasal, prefrontal, internasal, (posteriorly); tail round, not highly depressed; and interorbital depressions; undifferentiated TH 67.0% TW; caudals similar in size to posterior head scales granular, slightly convex; dorsals, subcaudals similar in size to ventrals. throat and chin scales small, juxtaposed, and Coloration of holotype in preservative.— nonimbricate, making a sharp transition to Background dorsal body coloration light grey gular and pectoral region scalation, with with intermittent small cream and dark grey enlarged cycloid, imbricate scales; circumor- speckles; pattern continued down tail; dorsum bitals 45/46 (L/R); interorbitals 48. of trunk with occasional larger cream spots; Axilla–groin distance 47.8% SVL; undiffer- limbs with same coloration pattern as trunk, entiated dorsal body scales round, convex, without larger cream spots; dorsal region of juxtaposed, relatively homogeneous in size; head continues same pattern except for darker each dorsal scale surrounded by six interstitial grey area above and between orbits; lateral granules, giving the appearance of a Star of side of trunk has same coloration pattern as David configuration under high magnification dorsum; lateral side of head with thin cream (Fig. 5); dorsals sharply transition to imbricate circumorbital ring of scales, remainder of ventrals along lateral body surface; paraverte- head with same coloration pattern as body; brals between midpoints of limb insertions ventral body surface solid cream, pattern 197; ventrals between midpoints of limb continues until just posterior to cloacal region, insertions 122; scales on dorsal surfaces of when coloration pattern of dorsal trunk limbs more imbricate than dorsals; scales on becomes prevalent, larger cream spots absent; dorsal surfaces of hands and feet similar to ventral side of head solid cream, with dorsal limb scales, heavily imbricate; ventral occasional light grey speckling along infra- body scales flat, cycloid, strongly imbricate, labial scales and thicker with light grey much larger than lateral or dorsal body scales, blotches just proximate to infralabials; ventral relatively homogeneous in size. surfaces of limbs, hands, and feet are solid Sixteen enlarged pore-bearing scales, in cream except for space between scansors, continuous precloacal series, arranged in a which is dark grey. widely obtuse, W-formation (Fig. 6); precloa- Coloration in life (based on field notes and cals 8/8 (L/R); precloacal series bordered photographs in life; Fig. 8D).—Dorsal ground posteriorly by single row of enlarged scales; color of head and trunk grayish brown, with precloacals situated atop a substantial pre- light tan, irregular mottling; head with tan cloacal bulge. blotches highlighted with green centrums Digits moderately expanded and covered throughout; anterior half of canthal region on palmar–plantar surfaces by bowed, un- heavily mottled light tan; tan mottling on notched, undivided scansors; digits with trunk highlighted with green only along minute vestiges of interdigital webbing; sub- dorsolateral margin, forming vague longitudi- digital scansors of hand: 8/8, 10/11, 13/13, 15/ nal series of moderately sized blotches and 15, and 11/11 (L/R) on Finger I–Finger V, ventrally paired smaller blotches extending respectively; foot: 11/10, 13/12, 15/15, 17/16, from arm insertion to base of tail; dorsal and 12/12 (L/R) on Toe I–Toe V, respective- coloration at base of tail with minimal tan ly; subdigital scansors of hands and feet mottling; dorsal limb surfaces with coloration bordered proximally (on palmar and plantar of head, but appearing with more regular surfaces) by 1–4 slightly enlarged scales that spots, coloration lighter on arms than hind- form a near-continuous series with enlarged limbs; legs with higher density of greenish-tan scansors; all digits clawed but first claw spots; tail regenerated to base, with a light greatly reduced; remaining terminal phalan- gray ground color overlain with dark brown ges compressed, with large recurved claws. mottling anteriorly; ventral ground trunk color Tail regenerated, short, 30.7 mm, 55.0% cream yellow, with minimal light gray mottling SVL; single distinct fracture plane present, along lateral margins. 134 HERPETOLOGICAL MONOGRAPHS [No. 28

Measurements and scale counts of holotype species, it still qualifies for formal IUCN in millimeters.—Snout–vent length 55.9; tail classification. We have evaluated this species length 30.7 (regenerated); total length 86.6 against the IUCN criteria for classification and (with regenerated tail); axilla–groin distance find that it qualifies for the status of Vulner- 26.7; tail width 2.9; tail height 1.9; head length able, VU, based on the following criteria: VU 10.4; head width 8.5; head height 6.1; mid- A2ac; B2ab(iii); D2 (IUCN, 2013). body width 6.4; snout length 6.0; eye diameter Etymology.—The specific epithet is derived 3.8; eye–nares distance 5.1; internarial dis- from the traditional term Chavacanos, other- tance 2.2; interorbital distance 4.7; femur wise known as Philippine–Creole Spanish, the length 9.8; tibia length 3.9; Finger III scansors language still spoken today by over 800,000 15; Toe IV scansors 17; supralabials 15; people, the Zamboangueno˜s, of Zambanga infralabials 17; preanofemoral pores 16; cir- City, southwestern Philippines. At a time cumorbitals 46; paravertebral scales 197; when cultural identities and languages con- ventral scales 122. tinue to be lost in the Philippines, Chabakano Variation.—Between the two specimens continues to be the socially, historically, and examined, we observed variation in the num- culturally significant language of choice for bers of supralabials, infralabials, and digital millions of Filipinos. The name is a plural scansors. The number of supralabials varied noun of masculine gender. Suggested com- between 15 (KU 314963) and 16 (KU mon name: Zamboanga False Gecko. 314964); infralabials varied between 16 (KU 314964) and 17 (KU 314963). Finger III DISCUSSION scansors were observed to vary between 15 The new taxa recognized in this paper (KU 314963) and 16 (KU 314964); Toe IV increase the total number of known species of scansors varied between 17 (KU 314963) and Pseudogekko to six, and all are endemic to the 20 (KU 314964). Philippines. Multilocus phylogenetic analyses Distribution.—Pseudogekko chavacano is of Siler et al. (2014a) further support the four known only from the Zamboanga Peninsula lineages within the Pseudogekko compresicor- of Mindanao Island (Fig. 1). pus Complex recognized in this study (Fig. 1). Ecology and natural history.—Pseudogekko As a testament to the rarity of Pseudogekko chavacano has been observed in secondary- species in museum collections and our rela- growth forest only, with the two available tively poor understanding of the diversity and adult specimens collected inside the protected relationships in this clade, we note that the Pasonanca watershed. Two eggs were col- newly described species all have been mas- lected in humus on the bank of a creek in querading as members of a single widespread Zamboanga del Norte Province by A.C. Alcala species for nearly a century (Taylor, 1915). in 1959 and were subsequently hatched in the Despite small sample sizes, we are confident lab a week later (CAS-SU 23548, 23549). Not in our taxonomic decisions based on the only do we assume this species once occurred numerous conspicuous diagnostic morpholog- in previously extensive, low-elevation primary ical features of all focal species (Table 2) and forest in western Mindanao, but also we their deep genetic divergences (Siler et al., assume it may possess a wider distribution 2014a). throughout the Zamboanga Peninsula of Mitochondrial sequence divergences be- Mindanao Island (Fig. 1). As for P. ditoy, tween members of the Pseudogekko compre- the two adult specimens were both found on sicorpus Complex are remarkably high (Siler top of leaves of shrubs 2–3 m above the ground. et al., 2014a). At a minimum, P. ditoy and P. The Zamboanga City Watershed in Pasonanca chavacano are 13.9% divergent from each Natural Park represents one of the most well- other; however, many observed divergences protected, low-elevation, primary forests re- are above 20% (Siler et al., 2014a). Also maining in the southwestern Philippines; interesting is the considerable intraspecific therefore, a large portion of this species range genetic diversity observed within P. compre- is well protected. However, due to the sicorpus from the Luzon PAIC and P. relatively limited known distribution of this pungkaypinit from the Mindanao PAIC (up 2014] HERPETOLOGICAL MONOGRAPHS 135 to 19% and 12%, respectively; Siler et al., pungkaypinit (central, eastern Mindanao), 2014a). These surprisingly high levels of other portions of Mindanao Island (i.e., the genetic diversity suggest that additional cryp- Cotobato coast mountains) may likely harbor tic species remain unrecognized. In the case additional divergent lineages of Pseudogekko. of these two species, the observed interpop- (3) Luzon PAIC: Numerous, possibly distinct ulation diversity corresponds to isolated re- evolutionary lineages still appear to be unrec- gions in the Luzon and Mindanao PAICs ognized throughout the Luzon PAIC. Polillo already known to contain high levels of Island possesses a genetically divergent pop- endemism among squamate reptiles (Fig. 1). ulation of P. compresicorpus, as does the For example, the deep split between popula- Sierra Madres Mountain Range in the north- tions of P. pungkaypinit from Central Mind- eastern portions of the island. Interestingly, anao Island and those from Leyte and Bohol outside of the holotype of P. compresicorpus islands is similar to species-level boundaries from the Bataan Peninsula of western Luzon observed in several other groups of verte- Island (Fig. 1), there are no records of brates (e.g., Cyrtodactylus, Brachymeles, Var- Pseudogekko from much of the northwest, anus; Siler et al., 2014a,d), birds (Hosner et including the Zambales and Cordillera moun- al., 2013), and mammals (Steppan et al., tain ranges (Diesmos et al., 2004; Brown et al., 2003). Likewise, Luzon Island has been the 2012a; Devan-Song and Brown, 2012). Wheth- focus of several recent studies revealing er this genus is truly absent from these distinct discrete faunal subregions throughout the faunal subcenters or these patterns are simply island, including the Zambales, Cordillera, an artifact of limited biodiversity surveys and Sierra Madres mountain ranges, each remains to be determined. However, we have with seemingly unique communities of am- no doubt that additional survey work in the phibians and reptiles (Siler et al., 2011; Luzon faunal region will result in the discovery Devan-Song and Brown, 2012; Brown et al., of additional new species of false geckos. (4) 2013a; Siler et al., 2014d). Unfortunately, Tablas Island: There is a single record of P. small series containing only juvenile or compresicorpus from Tablas Island in the subadult specimens from these areas limit Romblon Island Group (CAS 139713). Not our ability to evaluate these genetically unique only does this adult female appear to have populations taxonomically. several distinct morphological characters (e.g., Recognizing the levels of threat to the infralabials 17, enlarged scales in the precloacal survival of many of these unique, endemic region 15, circumorbitals 54), but also this geckos, future surveys should focus on these island group harbors numerous other endemic poorly sampled populations to allow for species (e.g., Platymantis lawtoni, Platymantis appropriate evaluations of species diversity. levigatus, Gekko romblon, G. coi; Brown et al., In an attempt to aid in efforts to quickly study 2011b; Siler et al., 2012b), suggesting that this and protect unique lineages within this genus, Pseudogekko may be a unique species. (5) we suggest the following populations be Masbate Island: Like Tablas Island, a single prioritized for investigation. (1) Pseudogekko record of P. compresicorpus from Masbate brevipes: The currently sampled populations Island (CAS 141560) is known. This adult of P. brevipes are not only genetically female also possesses an anomalous combina- divergent (.20% mitochondrial sequence tion of unique morphological characters (Fin- divergence between Leyte and Negros island ger III scansors 13, Toe IV scansors 16, populations; Fig. 1; Siler et al., 2014a), but the infralabials 17, circumorbitals 49). Further- recognized distribution of this species spans more, although the faunal affinities of Masbate two distinct faunal regions (Visayan and Island are still somewhat vague, the island is Mindanao PAICs; Fig. 1). (2) Central Mind- considered a member of the Visayan PAIC, not anao Island: Collections of Pseudogekko from the Luzon PAIC (Brown and Diesmos, 2009; Mindanao Island are rare in museum collec- Brown et al., 2013a), and also harbors endemic tions. With the presence of a unique species squamate reptiles (e.g., Brachymeles tungaoi, on the Zamboanga Peninsula (P. chavacano) B. cf. bonitae Siler and Brown, 2010; Davis et and the deeply divergent population of P. al., in press). 136 HERPETOLOGICAL MONOGRAPHS [No. 28

So little is known about the ecology of multiple lines of evidence and nontraditional Pseudogekko that broad conclusions on archi- taxonomic characters may provide the best pelago-wide diversity, microhabitat breadth, opportunities for future integrative approach- and natural history cannot be made at this es to understanding Philippine reptile and time (Brown and Alcala, 1978). Interestingly, amphibian megadiversity (Barley et al., 2013; at present only a few islands possess more Grismer et al., 2013; Linkem and Brown, than one species of Pseudogekko: Luzon, 2013; Siler et al., 2014c). Polillo, Leyte, and Mindanao. As more Acknowledgments.—We thank the Protected Areas and information on false geckos becomes avail- Wildlife Bureau of the Philippine Department of able, it will be interesting to determine Environment and Natural Resources for facilitating the whether species occur sympatrically on these collecting and export permits necessary for this and related studies; we are particularly grateful to M. Lim, C. and other islands. On Polillo Island, P. Custodio, J. de Leon, and A. Tagtag. Fieldwork was smaragdinus geckos possess a microhabitat conducted under the Memorandum of Agreement with preference for Pandanus leaves (Taylor, 1915, the Protected Areas and Wildlife Bureau of the Philip- 1922b). The single collected specimen of pines (2009–2014), Gratuitous Permit to Collect No. 221, and The University of Kansas Institutional Animal Care Pseudogekko smaragdinus collected on Mt. and Use Committee (KU IACUC) Approval (158-01). Labo, Bicol Peninsula (Luzon Island) was Financial support for fieldwork and lab work was provided collected from inside the husk surrounding by a Panorama Fund grant from the KU Biodiversity the trunk of a wild banana plant (R. Brown, Institute and travel funds from the KU Department of Ecology and Evolutionary Biology, a Madison and Lila personal observation). Little is known about Self Fellowship from the KU, a Fulbright Fellowship, a the Polillo Island population of P. compresi- Fulbright-Hayes Fellowship, and National Science Foun- corpus except that specimens were found on dation (NSF) DEB 0804115 to CDS, as well as support the upper surfaces of leaves at night. Because from NSF DEB 0743491, and NSF EF-0334952 to RMB. For the loans of specimens we thank J. Vindum and successful conservation efforts will depend on A. Leviton (California Academy of Sciences). Helpful our understanding of the habitats of species we comments on this manuscript were provided by J. strive to protect, future studies should attempt Grummer, M. Harvey, R. Davis and J. Hoskins. to document microhabitat preferences. As a result of many recent studies aimed at LITERATURE CITED investigating the diversity of geckos through- Alcala, E.L., A.C. Alcala, and C.N. Dolino. 2004. out the Philippine archipelago (Linkem et al., Amphibians and reptiles in tropical rainforest fragments 2010; Welton et al., 2010; Brown et al., 2012a; on Negros Island, the Philippines. Environmental Siler et al., 2014d), we will soon approach Conservation 31:254–261. Barley, A.J., J. White, A.C. Diesmos, and R.M. Brown. a more-accurate understanding of gekkonid 2013. The challenge of species delimitation at the diversity in the country. A few groups remain extremes: diversification without morphological change that have yet to be comprehensively evaluated in Philippine sun skinks. 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Brown. 2012a. Did geckos ride the Palawan raft to the Philippines? Journal of Biogeography 39:1217–1234. APPENDIX Siler, C.D., J.C. Swab, C.H. Oliveros, A.C. Diesmos, L. Specimens Examined Averia, A.C. Alcala, and R.M. Brown. 2012b. Amphib- ians and reptiles, Romblon Island Group, central Numbers in parentheses following species names Philippines: comprehensive herpetofaunal inventory. indicate the number of specimens examined. All speci- Check List 8:443–462. mens examined are from the Philippines. Several sample 2014] HERPETOLOGICAL MONOGRAPHS 139 sizes are greater than those observed in the description 330735, 331657); Quezon Province, Municipality of due to the examination of subadult specimens which were Infanta, Barangay Magsaysay, Barangay Kipagringau excluded from morphometric analyses. (KU 334017); MASBATE ISLAND, Masbate Province, Lepidodactylus labialis (10): MINDANAO ISLAND, Municipality of Mobo (CAS 141560); TABLAS ISLAND, Agusan del Norte Province, Municipality of Cabadbaran Romblon Province, Municipality of San Agustin (CAS (CAS 133317, 133318, 133329, 133396, 133687, 133790); 139713). Davao del Sur Province, Municipality of Malalag (CAS Pseudogekko ditoy sp. nov. (2): LEYTE ISLAND, 124813, 139714–16). Leyte Province, Municipality of Baybay, Barangay Pseudogekko brevipes (9): BOHOL ISLAND, Bohol Gabas, Sitio Cienda (Holotype, PNM 9811, formerly KU Province, Municipality of Sierra Bullones (CAS 131855, 326437), (Paratype, KU 326438). CAS-SU 24596, 25108); NEGROS ISLAND, Negros Pseudogekko pungkaypinit sp. nov. (6): BOHOL Oriental Province, Municipality of Sibulan (CAS 128956, ISLAND, Bohol Province, Municipality of Sierra Bul- 128959, 128963, 128971); Municipality of Valencia, lones, Barrio Dusita (Paratypes, CAS 131854, CAS-SU Barangay Bongbong (KU 302818, 327770). 23655); Raja Sikatuna Natural Park (Paratype, KU Pseudogekko chavacano sp. nov. (4): MINDANAO 324426); LEYTE ISLAND, Leyte Province, Municipality ISLAND, Zamboanga City Province, Municipality of of Baybay (Paratype, KU 326243); Barangay Guadalupe Zamboanga City (Holotype, PNM 9812, formerly KU (Holotype, PNM 9810, formerly KU 326435); MIND- 314963), (Paratype, KU 314964); Zamboanga del Norte ANAO ISLAND, Misamis Oriental Province, Municipal- Province, Cuot Creek (CAS-SU 23548, 23549). ity of Gingoog City, Barangay Lawaan, Sitio Kibuko, Mt. Pseudogekko compresicorpus (9): POLILLO IS- Lumot (Paratype, KU 334019). LAND, Quezon Province, Municipality of Polillo (KU Pseudogekko smaragdinus (35): POLILLO ISLAND, 326242); LUZON ISLAND, Cagayan Province, Munici- Quezon Province, Municipality of Polillo (KU 302819–31, pality of Gonzaga (KU 330058); Laguna Province, 303995–4002, 307638–47, 326240, 326241, 331721); LU- Municipality of Los Ban˜ os, Barangay Batong Maiake ZON ISLAND, Camarines del Norte Province, Munici- (KU 326434, 326436); Barangay Bagong Silang (KU pality of Labo, Barangay Tulay Na Lupa (KU 313828).