taxonomic revision of the pseudogekko …

TAXONOMIC REVISION OF THE PSEUDOGEKKO COMPRESICORPUSCOMPLEX (REPTILIA: SQUAMATA: GEKKONIDAE), WITH

DESCRIPTIONS OF THREE NEW SPECIES

CAMERON D. SILER1,6, LUKE J. WELTON

2, DREW R. DAVIS3, JESSA L. WATTERS

1, CONNER S. DAVEY1,

ARVIN C. DIESMOS4, MAE L. DIESMOS

5, AND RAFE M. BROWN6

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, USA3 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, Philippines5 University of Santo Tomas, Espana Boulevard, Manila, Philippines

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

ABSTRACT: Recent phylogenetic analysis of false geckos, genus Pseudogekko, revealed unrecognizeddiversity within these exceedingly rare and enigmatic Philippine forest geckos. Newly available geneticdatasets revealed that two of the four currently recognized species are complexes of multiple, deeplydivergent evolutionary lineages. In this paper we evaluate species diversity in the Pseudogekkocompresicorpus 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 ageographic range spanning three major faunal regions and several isolated islands. This perception of thespecies’ 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, findingcharacter-based evidence that unambiguously supports the recognition of four unique evolutionary lineageswithin the complex, three of which we describe as new species. These evolutionary species correspond tomonophyletic lineages supported in recent molecular studies. We also address the historically controversialgeneric affiliation of Pseudogekko labialis and conclude that this poorly known species is a member of thegenus Lepidodactylus. All species recognized in this study possess allopatric geographic ranges and differfrom congeners by numerous diagnostic characters of external morphology and, therefore, should berecognized as full species in accordance with any lineage-based species concept. This study nearly doublesthe known diversity of Philippine false geckos.

Key words: Biodiversity; Conservation; Endemism; False geckos; Philippines; Species complex;Taxonomy

PHILIPPINE gecko diversity represents animpressive array of diversification in morphol-ogy, behavior, and ecology (Brown and Alcala,1978). From ancient, micro-endemic lineageswith small ranges on larger islands (Rosleret al., 2006; Linkem et al., 2010) to severalwidespread species groups (Siler et al., 2010),and to species limited to tiny isolated islets(Brown and Alcala, 2000; Brown et al., 2011a;Siler et al., 2012a), Philippine geckos arequite ecologically variable, considering thatonly 57 species are currently recognized(PhilBREO, 2014). The archipelago’s gekko-nids also range from morphologically conser-vative gecko generalists (Brown and Alcala,1978) to delicate forest vegetation specialistsand to several lineages capable of derived

gliding locomotion with highly specializedcutaneous structures (e.g., Ptychozoon andLuperosaurus; Brown et al., 1997, 2012a;Dudley et al., 2007).

Other than the work of Taylor (1922a),Brown and Alcala (1978) published the onlycomprehensive systematic review of Philip-pine geckos. Their work summarized taxo-nomic diversity, provided an identificationguide, and recognized 31 species. Many ofthese had been observed rarely in naturalconditions and were known only on the basisof one or two specimens in museum collec-tions. Although Brown and Alcala’s (1978)foundational work has remained the onlysynopsis of the archipelago’s geckos, speciesdiversity has now nearly doubled since itsoriginal publication. Remarkably, of thecountry’s 57 species, 47 (82%) are Philippine6 CORRESPONDENCE: e-mail, [email protected]

Herpetological Monographs, 28 2014, 110–139

E 2014 by The Herpetologists’ League, Inc.

110

endemics (Brown et al., 2008, 2009, 2011a,b).Despite this dramatic improvement in ourunderstanding of this predominantly endemicfauna, a few genera are very poorly known(i.e., Luperosaurus; Brown et al., 2007,2011b, 2012a). A case in point is theextremely rare, endemic genus Pseudogekko(Taylor, 1915, 1922a), a group of four small,delicate, distinctly elongate, highly secretive,and entirely arboreal forest geckos (Brownand Alcala, 1978).

In the last two decades, our comprehensivebiodiversity surveys throughout the Philip-pines (e.g., Siler et al., 2012b; Brown et al.,2013a,b) resulted in only a handful ofvouchered genetic samples (Siler et al.,

2014a) for species of Pseudogekko. Yet evenin the absence of dense population geneticsampling, Siler et al. (2014a) revealed aconsiderable degree of cryptic genetic diver-sity and high levels of genetic divergencebetween clades. As currently recognized, thedistribution of each of these species spansmultiple recognized faunal regions or Pleisto-cene Aggregate Island Complexes (PAICs;Brown and Guttman, 2002; Brown andDiesmos, 2009). In fact, as currently defined(Brown and Alcala, 1978; Siler et al., 2012b),the species Pseudogekko compresicorpus isdistributed across three distinct faunal regions(Luzon, Mindanao, and Visayan PAICs) andan isolated island group (Romblon Island

FIG. 1.—Maximum clade credibility topology of Pseudogekko derived from Bayesian analyses in the recentphylogenetic study of Siler et al. (2014a). Numbers below nodes indicate maximum likelihood bootstrap values (left) andBayesian posterior probabilities (right). Boxed numbers correspond to numbered sampling localities shown on theassociated topographic map of the Philippines.

2014] HERPETOLOGICAL MONOGRAPHS 111

Group; Fig. 1). Currently, there are fewexamples of seemingly widespread species ofPhilippine vertebrates that truly defy theseregional biogeographic boundaries (Brownand Diesmos, 2002, 2009; Brown et al.,2002, 2013a). Phylogenetic studies supportmany species as more range-restricted, withpatterns generally consistent with inferredPAIC formation (Siler et al., 2012c,d); Brownet al., 2013a). In fact, phylogenetic analysesreveal unique lineages of P. compresicorpuswith apparent distributions corresponding tocircumscribed biogeographic subregions ofthe archipelago (Brown and Diesmos, 2009;Brown et al., 2013a; Siler et al., 2014a).

Species of the genus Pseudogekko representa critical conservation urgency (Alcala et al.,2004; Brown and Diesmos, 2009; Brown et al.,2012a); the named taxa are nearly all micro-endemics threatened by habitat destruction inthe form of anthropogenic forest removal.First, all species are arboreal and consideredobligate primary forest taxa or Pandanus spp.(screw pines) plant microhabitat specialists(Brown and Alcala, 1978). Second, based onthe little information known about populationhealth and microhabitat preferences, popula-tions apparently have decreased over the last100 yr (IUCN, 2013; Siler et al., 2014a). Tomake matters worse, the preferred habitat(lowland and coastal forests) of these specieshas been near completely removed fromthroughout the Philippines (Catibog-Sinhaand Heaney, 2006; Brown and Diesmos,2009; Siler et al., 2014b). Finally, the genushas had a complex taxonomic history (Brownand Alcala, 1978). Members of Pseudogekkopreviously have been assigned alternatively andwith little confidence (Taylor, 1922a; Brownand Alcala, 1978) to the genera Luperosaurus(Taylor, 1915) and Lepidodactylus (Brown andTanner, 1949; Brown, 1964; Kluge, 1968;Brown and Alcala, 1978) and even to Gekko(Taylor, 1922b). The combination of thesefactors highlights the urgent need to assessspecies boundaries within the genus in order tounderstand unique evolutionary lineages bet-ter, as conservation targets, and to implementconservation strategies more efficiently forprotecting these threatened and charismaticspecies (Sanguila et al., 2011; Siler et al.,2014c).

In this study, we investigate species diver-sity within the Pseudogekko compresicorpusComplex, with the understanding that severalof the divergent populations identified heremay represent threatened, and possibly en-dangered, unique evolutionary lineages (Sileret al., 2014a), worthy of both formal taxonom-ic recognition and prioritization for immediateconservation action. The taxonomic revisionsherein are guided by the results of the recentphylogenetic study on geckos of the genusPseudogekko (Siler et al., 2014a).

TAXONOMIC HISTORY

Taylor (1915) described Luperosaurus com-presicorpus on the basis of one specimencollected from Limay, Bataan Province, Lu-zon Island. In this description, Taylor (1915)expressed uncertainty about the placement ofthis species in Luperosaurus (as opposed toerecting a new genus to accommodate the onespecimen), noting that the new species had anelongate, compressed body form (a character-istic generally shared with the other knownspecies of Luperosaurus; see Brown et al.,2000). Later, Taylor (1922a) transferred thisspecies to a novel genus, Pseudogekko, where itremained a monotypic genus until the descrip-tion of a new species, Pseudogekko shebae,from the Soloman Islands (Brown and Tanner,1949). Unfortunately, the type specimen of P.compresicorpus (Philippine Bureau of Science,No. 1781) was destroyed during World War II(Brown and Alcala, 1978), which limitedcomparisons of this species with other newlydescribed gekkonids from this region. Howev-er, the collection of additional specimens of P.compresicorpus from Mindanao and Boholislands allowed Brown (1964) to examineshared characteristics between Pseudogekkoand other phenotypically similar genera(Gekko, Hemiphyllodactylus, Lepidodactylus,Luperosaurus, Pseudothecadactylus). Brown(1964) noted similarities between P. shebaeand Lepidodactylus, and P. shebae was laterplaced in this genus by Kluge (1968).

Additionally, Kluge (1968) transferred twospecies to Pseudogekko: Pseudogekko bre-vipes—a taxon that was originally describedas a Lepidodactylus (Boettger, 1897), andPseudogekko smaragdinus, originally describedas a member of the genus Gekko (Taylor,

112 HERPETOLOGICAL MONOGRAPHS [No. 28

1922b). Prior to its placement in Pseudogekko,Brown (1964) questioned the assignment of P.brevipes to the genus Lepidodactylus on thebasis of body proportions (e.g., ‘‘breadth ofhead to SVL’’) that appeared more similar tomembers of Pseudogekko than to species ofLepidodactylus. Finally, Peters (1867) original-ly described Pseudogekko labialis as Geckolabialis, which Boulenger (1885) later rede-scribed as a Lepidodactylus. However, it wasnot until additional specimens became avail-able that Brown and Alcala (1978) transferredthis species to Pseudogekko. Other than Brownand Alcala’s samples (collected in 1971), thisexceedingly rare species had not been collectedsince its original description. A recent (2012) 5-wk targeted survey effort in the neotype locality(Brown and Alcala, 1978; Mt. Hilong-Hilong,northeastern Mindanao Island) by a largegroup of experienced herpetologists producedno specimen records (R. Brown, personalobservation).

The genus Pseudogekko contains fourspecies (P. brevipes, P. compresicorpus, P.labialis, P. smaragdinus), and no new specieshave been described since the late 1970s(Brown and Alcala, 1978). Recently, Sileret al. (2014a) estimated phylogenetic relation-ships and elucidated multiple highly divergentgenetic lineages within P. compresicorpus,which the authors interpreted as probableevidence for the existence of additional, as ofyet undefined species contained within P.brevipes and P. compresicorpus.

In this paper, we re-analyze new data fromall available P. compresicorpus specimens,including both our own collections from thelast 20 yr (see Siler et al., 2014a) and oldermuseum specimens (Brown and Alcala, 1970,1978; Brown et al., 2013a). We use these dataand reliable diagnostic differences to revisePseudogekko taxonomy and describe fourdistinct species. To relate our findings toongoing conservation efforts, we addressconservation threats and priorities and pro-vide new or re-evaluated formal conservationstatus assessments (IUCN, 2013) for eachspecies described in this study. Finally, wealso reassessed the taxonomic affinities of P.labialis and refer this species to the genusLepidodactylus, in agreement with the con-clusions of Boulenger (1885).

MATERIALS AND METHODS

Field Work, Sample Collection, andSpecimen Preservation

We conducted fieldwork on Bohol, Leyte,Luzon, Negros, Mindanao, and Polillo islandsin the Philippines (Fig. 1). We collectedspecimens between 900 and 1600 h, whichwere euthanized via cardiac injection ofnembutal or immersion in aqueous chlore-tone, dissected for genetic samples (liver ormuscle preserved in 95% ethanol or flashfrozen in liquid nitrogen), fixed in 10%buffered formalin, and eventually transferredto 70% ethanol (,2 mo later). Museumabbreviations for specimens examined orsequenced in this study are those of SabajPerez (2013; CAS and CAS-SU: CaliforniaAcademy of Sciences, San Francisco, Califor-nia; KU: The University of Kansas BiodiversityInstitute, Lawrence, Kansas; PNM: NationalMuseum of the Philippines [formerly Philip-pine National Museum], Manila, Luzon).

Morphological Data

We examined fluid-preserved specimens (seeAppendix) for variation in qualitative, meristic,and mensural characters using the phylogeneticresults of Siler et al. (2014a) conservatively as aguide for the identification of possibly uniqueevolutionary lineages. We determined sex bythe presence in males of precloacal or pre-cloacal-femoral pores, or as necessary (imma-tures, females) by gonadal inspection. We(DRD and CDS) took measurements to thenearest 0.1 mm with digital calipers.

Whenever possible, we scored meristic andmensural characters (based on Brown et al.,2008, 2009, 2011a,b, with some modifications)on the left side of the body. Characters include:snout–vent length (SVL, distance from tip ofsnout to vent); tail length (TL, distance fromposterior margin of vent to tip of tail); totallength (TotL, distance from tip of snout to tipof tail); tail width (TW, measured at widestsection of tail posterior to hemipene bulge); tailheight (TH, measured from ventral to dorsalsurface of tail at the same point as TW); headlength (HL, from tip of snout to posterior tip ofmandible); head width (HW, widest measure ofhead width at jaw articulations); head height(HH, measured from ventral to dorsal surfaceof head at jaw articulations); midbody width


(MBW, measured from lateral surface toopposing lateral surface at midpoint of axilla–groin region); snout length (SNL, distancefrom anterior border of orbit to tip of snout);eye diameter (ED, at widest point); eye–naresdistance (END, distance from anterior marginof eye to posterior margin of nares); internarialdistance (IND, from dorsal aspect betweenmost-laterally distal edges of nares); interorbit-al distance (IOD, distance between midline oforbits from dorsal aspect); axilla–groin distance(AGD, distance between posterior edge of arminsertion and anterior edge of leg insertion);femur length (FL); tibia length (TBL); supra-labials (SUL, number of enlarged supralabials,from first supralabial in contact with rostral toposteriormost enlarged supralabial retainingdistinct, square to rectangular shape); infra-labials (IFL, number of infralabials); circumor-bitals (CO, number of visible, small circumor-bital scales encircling the eye); pore-bearingprecloacal scales (PPS, number of differentiat-ed, enlarged, pore-bearing scales in seriesanterior to the cloaca); pore-bearing precloa-cal-femoral scales (PFPS, number of differen-tiated, enlarged, pore-bearing scales in seriesanterior to the cloaca and, in some specimens,extending into the femoral region on theventral surface of the thigh); Finger IIIscansors (FinIII, scan, number of enlarged,undivided scansors beneath Finger III, start-ing just distal to point where skin betweendigits ends); Toe IV scansors (ToeIVscan,number of undivided scansors beneath ToeIV, starting just distal to point where skinbetween digits ends); paravertebral scales(PVS, number of scales along dorsal surfaceof body between midpoints of limb inser-tions); ventral scales (VS, number of scalesalong ventral surface of body betweenmidpoints of limb insertions); and interorbitalscales (IOS, total number of scales in straightline distance across interorbital region fromcenter of each eye, across both eyelids). Inthe descriptions, ranges are followed by mean6 standard deviation in parentheses.

Species Concept

As with many recent taxonomic revisions oforganisms endemic to island systems, weembrace the General Lineage Concept (deQueiroz, 1998, 1999) as an extension of the

Evolutionary Species Concept (Simpson, 1961;Wiley, 1978; Frost and Hillis, 1990). Wediagnose lineages as distinct species based ona suite of diagnostic morphological features,genetic divergence, and allopatric distributionsin separate biogeographic subregions of thearchipelago (Brown and Diesmos, 2009; Brownet al., 2013a). Lineage-based species conceptshave been shown to be particularly appropriatewhen applied to Philippine land vertebratebiodiversity (for review, see Davis et al., inpress). In this study we use a morphologicaldataset for all available specimens in museumcollections of the focal lineages, conservativelyguided by phylogenetic estimates of relation-ships (Siler et al., 2014a), to diagnose distinctlineages in this complex of false geckos,recognizing both previously described andnew species on the basis of nonoverlappingmorphological character states.

RESULTS

Morphology

Although sample sizes are low for manylineages described in this study, we haveevaluated and examined all known specimensin museum collections of each putative species.Despite the small sample sizes, multiple adultspecimens for each focal species are available,and each of the four identified lineages of thePseudogekko compresicorpus Complex arereadily diagnosed on the basis of numerous,nonoverlapping differences in meristic, men-sural, and color pattern characters (Tables 1,2). Variation in morphological characters(Tables 1, 2) mirrors the results observed inphylogenetic analyses (Fig. 1; Siler et al.,2014a) and supports the recognition of fourP. compresicorpus Complex lineages. Char-acters differing among these lineages includebody, head, and snout length, body and digitscale counts, pore-bearing scale counts, andcoloration and pigmentation patterns (Ta-bles 1, 2; species accounts below), many ofwhich are commonly employed diagnosticmorphological characters in taxonomic stud-ies of Philippine gekkonid lizards (Brown andAlcala, 1978; Brown et al., 2011a,b). With theexception of the presence (males) or absence(females) of pores in the precloacal orprecloacal-femoral region of the body, we


did not find additional sexually dimorphictraits in any of these four species.

Genetic Divergence

Uncorrected pairwise sequence divergenc-es are quite variable within the lineagesdefined here as species (0.0–19.0% mtDNAdivergence; Siler et al., 2014a) as compared tomany recent observations of endemic verte-brate diversity in the Philippines (Siler andBrown, 2010; Welton et al., 2010; Siler et al.,2012c,d). However, genetic divergences be-tween lineages are significantly higher, withthe exception of interpopulation divergencesamong Luzon populations of P. compresicor-pus (.26% mtDNA divergence; Siler et al.,2014a). The monophyletic lineages defined bySiler et al. (2014a; P. compresicorpus, Pseu-dogekko pungkaypinit sp. nov., Pseudogekkoditoy sp. nov., and Pseudogekko chavacano sp.nov., the latter three of which are firstdescribed herein) are distinguished from eachother by levels of genetic divergence greaterthan those observed between species of mostother Philippine geckos (Siler et al., 2010;Welton et al., 2010; Brown et al., 2011b).Given the higher observed intraspecific ge-netic diversity, we suspect that several of thestrongly supported clades (Fig. 1; Clades C,D) actually represent independent evolution-ary lineages (Siler et al., 2014a). However, intwo cases (P. compresicorpus, P. pungkaypinitsp. nov.), divergent populations are represent-ed by single vouchered specimens, at timesjuvenile individuals, which prevents us fromconfidently evaluating these putatively unique,and genetically divergent, populations at thistime.

Status of Pseudogekko labialis

One of the persistent taxonomic issues withthe diversity of Pseudogekko is whether P.labialis is appropriately placed in the genusrather than with morphologically more-similarspecies in the genus Lepidodactylus. Not onlyhave researchers historically considered P.labialis as a species of the genus Lepidodactylusbased on morphological similarity (Boulenger,1885; Wermuth, 1965; Kluge, 1968) but, also,over the years key morphological differenceshave been highlighted between these twogenera. Kluge (1968) described the genus

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Pseudogekko as differing from the generaLepidodactylus and Hemiphyllodactylus basedon the shape of the enlarged, precloacal pore-bearing scale series (series greatly archedanteromedially in Pseudogekko vs. not arched;Fig. 2). Boettger (1897) described Lepidodac-tylus brevipes (eventually recognized as amember of the genus Pseudogekko by Kluge[1993]), noting it could be separated fromLepidodactylus labialis Peters and Lepidodac-tylus pulcher Boulenger on the basis of havinga more-slender body, fewer precloacal pores,and less-distinctive series of submental scales.Even in their redescription of P. labialis andrecognition of the species as a member of thegenus Pseudogekko, Brown and Alcala (1978)recognized that the general body colorationand pigmentation patterns of P. labialis did notmatch the patterns of any other species in thegenus Pseudogekko. They note that all individ-uals of P. labialis have distinct dorsal markingsthat range from a vertebral row of bars or spotsto irregular transverse bars or even darkdorsolateral stripes (Brown and Alcala, 1978);these markings are absent in the remainingspecies of Pseudogekko (Brown and Alcala,1978; C. Siler, personal observation).

Based on measurements and comparisonsof individuals of P. labialis available inmuseum collections (see Appendix; Speci-mens Examined) and on published accountsdocumenting character differences betweenP. labialis and all other members of the genusPseudogekko, we now formally recognize thisspecies as a member of the genus Lepido-dactylus. This decision is supported by a suiteof diagnostic character differences betweenL. labialis and all other recognized species ofPseudogekko. Species of the genus Pseudo-gekko differ from L. labialis by havingenlarged, pore-bearing scale series (precloa-cal or precloacal-femoral scales) that aregreatly arched anteromedially (vs. not dis-tinctly arched anteromedially; Fig. 2), havingmarkedly narrower bodies (HW/SVL 15–16%vs. .18%), longer relative snout lengths(SNL/HW 63–71% vs. ,57%), by the pres-ence of small, juxtaposed postmental scales(vs. distinctively enlarged, strongly imbricatepostmentals; Fig. 2), and the absence (vs.presence) of darkly pigmented, and oftenstriped, body coloration.C

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Taxonomic Conclusions

With the removal of Lepidodactylus labialisfrom the genus Pseudogekko, the remainingfour focal lineages of this study each possessunique, nonoverlapping suites of diagnosticcharacter states of morphology (Tables 1, 2),and all correspond to clades defined inmultilocus phylogenetic analyses of DNAsequence data (Fig. 1; Siler et al., 2014a).Combined with biogeographic evidence andallopatric distributions, our data support theinterpretation of four distinct evolutionarylineages (full evolutionary species) within theP. compresicorpus Complex.

The type of the genus, Pseudogekko com-presicorpus (Taylor, 1915), was described onthe basis of a specimen from the BataanPeninsula of Luzon Island (Fig. 1). Our newspecimens from localities close to Taylor’s(1915) type locality match the holotypedescription in all regards. Additionally, severaldistinct morphological characters not empha-sized by Taylor (1915) closely ally withsampled populations from the northern Phi-lippines (Fig. 1, Clade D; Siler et al., 2014a)and with all previously published accounts andreferences to the holotype (Taylor, 1915,1922a): (1) infralabials 13–16, (2) precloacalpores 13–14, and (3) body coloration homog-enous brown and unpatterned. Accordingly,we recognize P. compresicorpus as a speciesthat occurs in the northern Philippines,restricting the species distribution to theLuzon PAIC, Romblon Island Group, andMasbate Island (Visayan PAIC). Furthermore,we recognize large-bodied populations fromMindanao Island (Fig. 1; Clade C), small-bodied populations from Leyte Island (Fig. 1;Clade B), and small-bodied populations fromthe Zamboanga Peninsula of Mindanao Island(Fig. 1; Clade A) as members of three, uniqueevolutionary lineages which we describebelow as new species.

TAXONOMIC ACCOUNTS

Pseudogekko compresicorpus (Taylor, 1915)(Figs. 1–8)

Luperosaurus compresicorpus Taylor, 1915:96,holotype female (Philippine Bureau ofScience 1781) from ‘‘Limay, Bataan Prov-ince, Luzon,’’ Brown and Tanner, 1949:41.

Pseudogekko compresicorpus (Taylor): Brownand Tanner, 1949:41; Kluge, 1993:30.

Pseudogekko compressicorpus (Taylor) [Mis-spelled]: Taylor, 1922a:103; Underwood,1954:479; Wermuth, 1965:151; Kluge, 1967:30, 1968:333; Brown and Alcala, 1970:112,1978:119; Brown et al., 2012a:920, 2012b:355, 2013b:54; Siler et al., 2012b:454,2014a:205.

Diagnosis.—Pseudogekko compresicorpus canbe distinguished from congeners by thefollowing combination of characters: (1) bodysize large (SVL 54.9–59.7 mm); (2) axilla–groin distance 49.3–55.2% SVL; (3) headlength 16.1–18.0% SVL; (4) snout long 56.9–64.6% head length; (5) Toe IV scansors 18 or19; (6) paravertebral scales 226–234; (7)ventral scales 127–130; (8) supralabials 16–20; (9) infralabials 13–16; (10) circumorbitals39–45; (11) precloacal pores 13 or 14; (12)femoral pores absent; (13) dominant bodycoloration dark brown to tan; (14) conspicu-ous head spotting present, neon green; (15)conspicuous dorsolateral spotting present,faint, neon green; (16) conspicuous limbspotting present, faint, neon green; (17) tailbanding absent; (18) body striping absent;(19) interorbital banding absent; and (20)ciliary ring coloration present, light blue(Fig. 3; Tables 1 and 2).

Comparisons.—Characters distinguishingPseudogekko compresicorpus from all otherspecies of Pseudogekko are summarized inTables 1 and 2. Pseudogekko compresicorpusmost closely resembles P. pungkaypinit sp.nov. However, it differs from this species byhaving a shorter total body length (TotL ,117.3 mm vs. .125.3), narrower body (MBW,6.7 mm vs. .7.7), fewer infralabials (13–16vs. 17–19), fewer circumorbitals (39–45 vs.50–55), fewer precloacal pores (13 or 14 vs.17–20), fewer paravertebral scales (226–234vs. 265–280), a dark brown to tan (vs. grayishbrown) body coloration, and by the presence(vs. absence) of conspicuous neon green spotson the head, dorsolateral region of the body,and limbs, absence (vs. presence) of stripedpigmentation patterns on the body, andpresence (vs. absence) of a light blue ciliaryring.

Pseudogekko compresicorpus can be distin-guished from P. ditoy sp. nov. and P.


chavacano sp. nov. by having a greaternumber of paravertebral scales (.226 vs.,197) and ventral scales (.127 vs. ,123);from P. ditoy by having a longer snout–vent

length (SVL . 54.9 mm vs. ,52.6), a greaternumber of Toe IV scansors (18 or 19 vs. 16 or17), fewer precloacal pores (13–14 vs. 18), bythe presence (vs. absence) of neon green spotson the head, dorsolateral region of the bodyand limbs, and presence (vs. absence) of alight blue ciliary ring; from P. chavacano byhaving a longer total body length (TotL .111.9 mm vs. 95.8), a shorter relative headlength (HL 16–18% SVL vs. 19%), fewercircumorbitals (39–45 vs. 46), fewer precloacalpores (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(SVL . 54.9 mm vs. ,52.5), a greater numberof Finger III scansors (15–17 vs. 12–14), a

FIG. 3.—Lateral view of head of Pseudogekko compre-sicorpus (KU 326436). Scale bar 5 2 mm. Illustrationby CDS.

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.


FIG. 4.—Photographic plates at 315 magnification of lateral and ventral views of the head and dorsal views of thetrunk 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 JLWand CDS.


greater number of Toe IV scansors (18 or 19vs. 13–15), a greater number of infralabials(13–16 vs. 12–14), a greater number ofcircumorbitals (39–45 vs. 35–37), and by thepresence of dense, neon green spots on thehead (vs. sparse and cream colored), presenceof neon green (vs. cream colored) dorsolateralspotting, presence (vs. absence) of neongreen limb spotting, absence (vs. presence) ofinterorbital banding, and presence (vs. absence)of a light blue ciliary ring; and from P.smaragdinus by having a greater number ofcircumorbitals (39–45 vs. 33–35), fewer en-larged pores (13 or 14 precloacal pores vs. 32–41 precloacal-femoral pores), fewer paraverte-bral scales (226–234 vs. 241–252), dark brownto tan (vs. bright neon yellow to orange[undisturbed] to neon green [disturbed]) body

coloration, absence (vs. presence) of femoralpores, presence of dense, neon green spots onthe head (vs. dense black and sparse white),presence of neon green (vs. large black andsmall white) dorsolateral spotting, presence ofneon green limb spotting (vs. sparse black andwhite), absence of tail banding (vs. presence,neon yellow, white, and neon orange), andpresence (vs. absence) of a light blue ciliary ring.

Description (based on description of holotype[Taylor, 1915] and 7 referred specimens).—Details of the head scalation are shown inFigures 3 and 4A. Measurements and meristicdata scored from the holotype are providedbelow in brackets. Body small, slender, SVL54.9–59.7 mm (females), 55.9–58.8 mm(males) [62.0]; limbs well developed, moder-ately slender; tail slender; margins of limbssmooth, lacking cutaneous flaps or dermalfolds; trunk lacking ventrolateral cutaneousfold.

Head size moderate, slightly differentiatedfrom neck, characterized by only slightlyhypertrophied temporal and adductor muscu-lature; snout rounded in dorsal and lateralaspect (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;dorsal surfaces of head relatively homoge-neous, with only slightly pronounced concavepostnasal, internasal, prefrontal, and interor-bital concavities; auricular opening moderate,ovoid, angled slightly anteroventrally andposterodorsally from beneath temporal swell-ings on either side of head; tympanum deeplysunken; eye large; pupil vertical, margin wavy(Fig. 4A); limbs and digits relatively short andmoderately slender; thighs moderately thickercompared to brachium; tibia length 7.2–9.6%SVL, 53.5–64.6% femur length.

Rostral rectangular in anterior view, 33 asbroad as high, sutured anterolaterally withanteriormost enlarged supranasals, projectingonto dorsal surface of head to point in linewith midline of nasal; nostril surrounded byfirst labial, or first and second labials, rostral,one or two enlarged postnasals, and one ortwo enlarged supranasals; supranasals sepa-rated by 2–5 small median scales, or touchingat midline; enlarged supranasals equal in sizeto enlarged postnasals or greatly enlargedcompared to postnasals.

FIG. 5.—Illustration of Star of David configurationformed by interstitial granules surrounding body scales,visible under high magnification. Magnification 325.Illustration by CDS.


Total number of differentiated supralabials16–20, bordered dorsally by one row ofdifferentiated, slightly enlarged snout scales;total number of differentiated infralabials 13–16[16], bordered ventrally by 3–5 rows of slightlyenlarged scales; undifferentiated chin and gular

scales; postrictal scales undifferentiated; re-maining undifferentiated gulars very small,round, nonimbricate, juxtaposed (Fig. 4A), eachscale surrounded by six interstitial granules,giving the appearance of a Star of Davidconfiguration under high magnification (Fig. 5).

FIG. 6.—Illustrations of precloacal pore-bearing scale series of adult males for (A) Pseudogekko compresicorpus (KU331657), (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). Forcomparison, the precloacal-femoral pore-bearing scale series of (E) P. smaragdinus (KU 302824) is provided forreference. Scale bar 5 2 mm. Illustrations by CDS.


Dorsal cephalic scales fairly homogeneousin size, shape, disposition, and distribution;cephalic scalation slightly convex, round tooval scales; postnasal, prefrontal, internasal,and interorbital depressions; undifferentiatedposterior head scales granular, slightly convex;throat and chin scales small, juxtaposed,and nonimbricate, making a moderately sharptransition to gular and pectoral region scala-tion, with enlarged cycloid, imbricate scales;circumorbitals 39–45.

Axilla–groin distance 49.3–55.2% SVL [58.1];undifferentiated dorsal body scales round,

convex, juxtaposed, relatively homogeneousin size; each dorsal scale surrounded by sixinterstitial granules; dorsals sharply transitionto imbricate ventrals along lateral bodysurface; paravertebrals between midpointsof limb insertions 226–234; ventrals betweenmidpoints of limb insertions 127–130; scaleson dorsal surfaces of limbs more imbricatethan dorsals; scales on dorsal surfaces ofhands and feet similar to dorsal limb scales,heavily imbricate; ventral body scales flat,cycloid, strongly imbricate, much larger thanlateral or dorsal body scales, relativelyhomogeneous in size.

Ten to 14 pores [14], in continuousprecloacal pore-bearing series, arranged in awidely obtuse, W-formation (Fig. 6); patch ofslightly enlarged scales posterior to precloacalseries, roughly three scale rows in size,forming an oval patch; precloacals situatedatop a substantial precloacal bulge.

Digits moderately expanded and coveredon palmar and plantar surfaces by bowed,unnotched, undivided scansors (Fig. 7); digitswith minute vestiges of interdigital webbing;subdigital scansors of Finger III 15–17, ToeIV 18–19; subdigital scansors of hands andfeet bordered proximally (on palmar andplantar surfaces) by 1–4 slightly enlargedscales that form a near-continuous series withenlarged scansors; all digits clawed, but firstclaw greatly reduced (Fig. 7); remainingterminal phalanges compressed, with largerecurved claws (Fig. 7).

Tail short, 52.9–58.5 mm [48], 89.7–99.6%SVL [77.4%]; round, not heavily depressed;TH 76.1–91.9% TW; caudals similar in size todorsals, subcaudals similar in size to ventrals.

Coloration in preservative (based onseven referred specimens).—Background dor-sal body coloration light tan with intermittentsmall cream and dark brown speckles; patterncontinued down tail but speckled areas areconcentrated occasionally into larger blotches;dorsal region of head with same color pattern,except for darker brown interorbital region;dorsal surfaces of limbs with same colorpattern; one individual with sparse, orange-tan spots on arms and legs (KU 331657);lateral region of body with same coloringpattern as dorsal region; lateral region of headwith same color patterns as body, except for

FIG. 7.—Illustration of left hand and foot of Pseudo-gekko compresicorpus (KU 326436). Scale bar 5 2 mm.Illustrations by CDS.


slightly lighter area just posterior to orbits andalong both sets of labial scales; circumorbitalscales with mixture of cream and mediumbrown coloration; ventral side of body withbackground cream color, with speckling pat-tern of lateral side of body wrapping around toapproximately halfway to midpoint of venter;ventral surfaces of head with same colorpattern as body; ventral surfaces of limbscream with medium to dark brown specklesscattered sparsely throughout; palmar andplantar surfaces solid cream, except mediumbrown regions between scansors; ventralsurface of tail solid cream with occasionallight brown speckles, speckles increase inintensity towards tail tip.

Coloration in life (based on CDS and RMBfield notes and photographs in life; Fig. 8A).—Dorsal ground color of head, trunk, and tailchocolate brown. Head with conspicuous lightgreen mottling in canthal and interocularregions as well as on lateral surface betweenthe eye and ear opening. Dorsolateral surfaceof body with series of eight faint, light greenblotches running from nuchal region ontobase of tail. Dorsal limb surfaces colored as

trunk, but with random placement light greenblotches. Tail colored as trunk, but with seriesof longitudinally elongate, paravertebralcream blotches. Ventral coloration light grayground color with minimal chocolate brownmottling along the lateral margins. Taylor(1915) reports simply that the holotype wascinnamon brown in life.

Distribution.—Pseudogekko compresicorpusoccurs on the Luzon PAIC (Luzon and Polilloislands), Visayan PAIC (Masbate Island), andthe Romblon Island Group (Tablas Island;Fig. 1). Although currently not recorded fromother islands in the Luzon PAIC, we would notbe surprised if future surveys discover addi-tional island populations (i.e., CatanduanesIsland).

Ecology and natural history.—Pseudogekkocompresicorpus has been observed in first-and secondary-growth forest (Fig. 1) on leavesof shrubs and small trees 2–4 m above theground. Taylor (1922a) notes that two, fullydeveloped embryos were found in eggs thathad been attached to the underside of a leaf atthe top of a recently felled tree. This naturalhistory observation may indicate that species

FIG. 8.—Photographs in life of (A) Pseudogekko compresicorpus (KU 326436) and the holotypes of (B) Pseudogekkopungkaypinit sp. nov. (Holotype, PNM 9810, formerly KU 326435), (C) Pseudogekko ditoy sp. nov. (Holotype, PNM9811, formerly KU 326437), and (D) Pseudogekko chavacano sp. nov. (Holotype, PNM 9812, formerly KU 314963).Photographs by RMB.


of this genus may have a more arboreallifestyle than currently is appreciated andthat, although encountered by us in forestlower strata, this species may also inhabit treecanopies.

Both Pseudogekko compresicorpus and P.smaragdinus occur on Luzon and Polilloislands (Fig. 1); however, there is insufficientevidence to determine if populations of thesespecies occur in sympatry. Similar to P.pungkaypinit sp. nov., numerous populationsof P. compresicorpus have been observed inthe wild as compared to other species in thegenus. In fact, as currently recognized, P.compresicorpus possesses the broadest geo-graphic distribution of any species in thegenus. Unfortunately, even with this broaddistribution, few specimens exist in museumcollections. At this time we do not find thisspecies qualifies for Critically Endangered,Endangered, Vulnerable, or Near Threatenedstatus under the IUCN criteria for classifica-tion (IUCN, 2013). Therefore, we recommendthat the species be considered Least Concernuntil additional information can be obtainedconcerning the health and diversity of wildpopulations throughout the Luzon and Visa-yan PAICs and Romblon Island Group.

Pseudogekko pungkaypinit sp. nov.(Figs. 1, 4, 6, 8)

Holotype.—PNM 9810 (RMB FieldNo. 4392, formerly KU 326435), adult male,collected in secondary-growth forest on 3September 2002, in the Calbiga—a creek areaon the Visayas State University Visca campus,Barangay Guadalupe, Municipality of Baybay,Leyte Province, Leyte Island, Philippines(10u4590.270N, 124u47924.00E; WGS-84), byR.M. Brown.

Paratypes.—One adult female (CAS 131854)collected from the bark of a rotten stump on 31March 1964 in Dusita Barrio, Municipality ofSierra Bullones, Bohol Province, Bohol Island,Philippines (09u46959.340N, 124u18910.80E;WGS-84) by S. Magusara; one adult male(CAS-SU 23655) collected on floor of lowlandbamboo forest on 9 May 1962 in Dusita Barrio,Municipality of Sierra Bullones Municipality,Bohol Province, Bohol Island, Philippines(09u46957.50N, 124u18910.80E; WGS-84) by

A.C. Alcala; one adult male (KU 324426)collected on 6 August 2009 in Raja SikatunaNatural Park, Barangay Danicop, Municipality ofSierra Bullones, Bohol Province, Bohol Island,Philippines (09u42919.8360N, 124u7924.3840E;WGS-84) by C.D. Siler; one adult female(KU 326243) collected on 17 April 2008 onthe campus of Visayas State University, BaybayCity, Municipality of Baybay, Leyte Province,Leyte Island, Philippines (10u44944.4660N,124u47929.22720E; WGS-84) by R.M. Brown;one adult male (KU 334019) collected on 19July 2012 near Ginoog River, Mt. Lumot, SitioKibuko, Barangay Lawaan, Municipality ofGingoog City, Misamis Oriental Province,Mindanao Island, Philippines (08u4990.33960N,125u5944.32920E; WGS-84) by R.M. Brown.

Diagnosis.—Pseudogekko pungkaypinit canbe distinguished from congeners by thefollowing combination of characters: (1) bodysize large (SVL 66.6–76.8 mm); (2) axilla–groin distance 53.7–55.9% SVL; (3) headlength 17.1–17.7% SVL; (4) snout 53.6–60.2% head length; (5) Toe IV scansors 17–21; (6) paravertebral scales 265–280; (7)ventral scales 125–155; (8) supralabials 16–20; (9) infralabials 17–19; (10) circumorbitals50–55; (11) precloacal pores 17–20; (12)femoral pores absent; (13) dominant bodycoloration grayish brown; (14) head, body andtail immaculate; (15) body stripes present;(16) interorbital band absent; and (17) ciliaryring coloration undifferentiated (Tables 1, 2).

Comparisons.—Characters distinguishingPseudogekko pungkaypinit from all otherspecies of Pseudogekko are summarized inTables 1 and 2. Pseudogekko pungkaypinitmost closely resembles P. compresicorpus;however, P. pungkaypinit differs from P.compresicorpus by having a longer totalbody length (TotL . 125.3 mm vs. ,117.3),wider body (MBW . 7.7 mm vs. ,6.7), moreinfralabials (17–19 vs. 13–16), circumorbitals(50–55 vs. 39–45), precloacal pores (17–20 vs.13 or 14), and paravertebral scales (265–280vs. 226–234); a grayish brown (vs. dark brownto tan) body coloration, the absence (vs.presence) of neon green spots on the head,dorsolateral region of the body, and limbs, thepresence (vs. absence) of dark stripes on thebody, and absence (vs. presence) of a lightblue ciliary ring.


Pseudogekko pungkaypinit can be distin-guished from P. ditoy sp. nov., P. chavacanosp. nov., P. brevipes, and P. smaragdinus byhaving a longer body (SVL . 66.6 mm vs.,64.3), a longer trunk (AGD . 37.2 mm vs.,35.5), more circumorbitals (50–55 vs. ,46)and paravertebral scales (.265 vs. ,252), andby the presence (vs. absence) of dark dorso-lateral body stripes; from P. ditoy and P.chavacano by having a wider body (MBW .7.7 mm vs. ,7.3); from P. chavacano byhaving a shorter relative head length (HL 17–18% SVL vs. 19%), a greater number ofprecloacal pores (17–20 vs. 16), and by theabsence (vs. presence) of neon green spots onthe head, dorsolateral region of the body,and limbs, and absence (vs. presence) of tailbands; from P. brevipes by having greaternumbers of Finger III scansors (15–17 vs. 12–14), Toe IV scansors (17–21 vs. 13–15),infralabials (17–19 vs. 12–14), and precloacalpores (17–20 vs. 13–15), and by the absence(vs. presence) of cream spots on the head anddorsolateral region of the body, and absence(vs. presence) of interorbital band; and fromP. smaragdinus by having fewer enlargedpores (17–20 precloacal pores vs. 32–41precloacal-femoral pores), its grayish brown(vs. bright neon yellow to orange [undis-turbed] to neon green [disturbed]) bodycoloration, and by the absence (vs. presence)of femoral pores, absence (vs. presence) ofblack and white spots on the head, dorsolat-eral surfaces of the body and limbs, andabsence (vs. presence) of transverse tailbands.

Description of holotype.—Details of thehead scalation are shown in Figure 4B. Adultmale in excellent condition, hemipenes evert-ed, hemipenal bulge present; small incision inthe sternal region (portion of liver removedfor genetic sample). Body small, slender, SVL71.9 mm; limbs well developed, moderatelyslender; tail original, slender; margins of limbssmooth, lacking cutaneous flaps or dermalfolds; trunk lacking ventrolateral cutaneousfold.

Head moderate in size, slightly differenti-ated from neck, characterized by only slightlyhypertrophied temporal and adductor muscu-lature; snout rounded in dorsal and lateralaspect (Fig. 4B); HW 132.3% MBW, 86.5%

HL; HL 17.3% SVL; SNL 69.6% HW, 60.2%HL; dorsal surfaces of head relatively homo-geneous, with only slightly pronounced con-cave postnasal, internasal, prefrontal, andinterorbital concavities; auricular openingmoderate, ovoid, angled slightly anteroven-trally and posterodorsally from beneathtemporal swellings on either side of head;tympanum deeply sunken; eye large; pupilvertical, margin wavy (Fig. 4B); limbs anddigits relatively short and moderately slender;thighs moderately thicker compared to bra-chium; tibia length 10.1% SVL, 63.5% femurlength.

Rostral rectangular in anterior view, 33 asbroad as high, sutured anterolaterally withanteriormost enlarged supranasals; nostril sur-rounded by first labial, rostral, one enlargedpostnasal, and two enlarged supranasals; su-pranasals separated by five small medianscales.

Total number of differentiated supralabials18/19 (left/right [L/R]), bordered dorsally by1–2 rows of differentiated, slightly enlargedsnout scales; total number of differentiatedinfralabials 17/17 (L/R), bordered ventrally by7–9 rows of slightly enlarged scales; undiffer-entiated chin and gular scales; postrictal scalesundifferentiated; remaining undifferentiatedgulars very small, round, nonimbricate toslightly imbricate, juxtaposed (Fig. 4B).

Dorsal cephalic scales fairly homogeneous insize, shape, disposition, and distribution; ce-phalic scalation slightly convex, round to ovalscales; postnasal, prefrontal, internasal, andinterorbital depressions; undifferentiated pos-terior head scales granular, slightly convex;throat and chin scales small, juxtaposed, andnonimbricate, making a moderately sharptransition to gular and pectoral region scalation,with enlarged cycloid, imbricate scales; circu-morbitals 51/53 (L/R); interorbital scales 74.

Axilla–groin distance 54.3% SVL; undiffer-entiated dorsal body scales round, convex,juxtaposed, relatively homogeneous in size;dorsal scales surrounded by interstitial gran-ules, however, granules do not give clearappearance of a Star of David configurationunder high magnification; dorsals sharplytransition to imbricate ventrals along lateralbody surface; paravertebrals between mid-points of limb insertions 280; ventrals between


midpoints of limb insertions 140; scales ondorsal surfaces of limbs more imbricate thandorsals; scales on dorsal surfaces of hands andfeet similar to dorsal limb scales, stronglyimbricate; ventral body scales flat, cycloid,strongly imbricate, much larger than lateral ordorsal body scales, relatively homogeneous insize.

Twenty enlarged pore-bearing scales pres-ent, in continuous precloacal pore-bearingseries, arranged in a widely obtuse, W-formation (Fig. 6); precloacals 10/10 (L/R);patch of slightly enlarged scales posterior toprecloacal series, roughly four scale rows insize, forming an oval patch; precloacalssituated atop a substantial precloacal bulge.

Digits moderately expanded and covered onpalmar/plantar surfaces by bowed, unnotched,undivided scansors; digits with minute vestig-es of interdigital webbing; subdigital scansorsof hand: 11/10, 12/12, 15/14, 17/17, and 11/11(L/R) on Finger I–Finger V, respectively; foot:12/12, 14/14, 16/17, 19/19, and 12/12 (L/R) onToe I–Toe V, respectively; subdigital scansorsof hands and feet bordered proximally (onpalmar and plantar surfaces) by 1–4 slightlyenlarged scales that form a near-continuousseries with enlarged scansors; all digits clawedbut first claw greatly reduced; remainingterminal phalanges compressed, with largerecurved claws.

Tail short, 67.2 mm, 93.5% SVL; round, notheavily depressed; TH 88.4% TW; caudalssimilar in size to dorsals, subcaudals similar insize to ventrals.

Coloration of holotype in preservative.—Background dorsal color medium brown withsmall dark brown speckles throughout, patterncontinued down tail; dorsal nuchal regionslightly lighter brown than remainder of body;dorsal regions of head and limbs have samecolor pattern as body; lateral regions of trunkhave same coloration pattern as dorsum;lateral side of head has thin cream circumor-bital ring of scales, lighter brown area onnuchal region continues to just posteriorto orbit; remainder of head continues samepattern as trunk, except supralabial andinfralabial scales are solid cream with fewdark brown speckles; ventral side of trunk hascream background color, with speckling pat-tern of lateral side of trunk wrapping around

to approximately halfway to midpoint ofventer; ventral region of head follows samepattern as trunk; ventral regions of limbs,hands, and feet are cream with mediumto dark brown speckles scattered sparselythroughout; ventral surface of tail is solidcream with occasional light brown specklesthat increase in intensity towards tip.

Coloration in life (based on field notes andphotographs in life; Fig. 8B).—Dorsal groundcolor of head, trunk and tail olive-brown;lateral surfaces of head and trunk with seriesof light brown stripes, from eye to leginsertion, stripes oriented posteroventrally;dorsal limb color similar to trunk, but lackingany discernable pattern; dorsal tail colorsimilar to trunk, but with occasional small,cream spots; ventrum light cream groundcolor with minimal light olive-brown mottlingalong the lateral margins.

Measurements and scale counts of holotypein millimeters.—Snout–vent length 71.9; taillength 67.2; total length 139.1; axilla–groindistance 39.1; tail width 3.6; tail height 3.2;head length 12.4; head width 10.7; headheight 7.0; midbody width 8.1; snout length7.5; eye diameter 4.8; eye–nares distance 5.9;internarial distance 2.8; interorbital distance5.8; femur length 11.4; tibia length 7.3; FingerIII scansors 17; Toe IV scansors 19; suprala-bials 18; infralabials 17; circumorbitals 51;preanofemoral pores 20; paravertebral scales280; ventral scales 140.

Variation.—Among the six specimens ex-amined, we observed variation in the numbersof precloacal pores, supralabials, infralabials,and digital scansors. The number of suprala-bials varied between 16 (KU 334019), 18 (KU326243, 326435), 19 (CAS-SU 23655, KU324426), and 20 (CAS 131854); infralabialsvaried between 17 (CAS 131854, KU 324426,326243, 326435), 18 (CAS-SU 23655), and 19(KU 334019). Precloacal pore counts in adultmales were observed to vary between 17 (KU324426), 19 (CAS-SU 23655, KU 334019), and20 (KU 326435). The number of Finger IIIscansors varied between 15 (CAS 131854,CAS-SU 23655, KU 326243), 16 (KU334019), and 17 (KU 324426, 326435); ToeIV scansors varied between 17 (CAS-SU23655), 18 (CAS 131854, KU 326243), 19(KU 326435, 334019), and 20 (KU 324426).


Distribution.—Pseudogekko pungkaypinitis known only from Bohol, Leyte, and Minda-nao islands, although it may be possible that itwill eventually be discovered on other islandsin the Mindanao PAIC (i.e., Samar, Dinagat,Biliran, Siargao; Fig. 1).

Ecology and natural history.—Pseudogekkopungkaypinit has been observed in disturbed,secondary-growth forest only; however, simi-lar to assumptions about other species in thegenus, we assume this species once occurred inlower elevation primary forest on Bohol, Leyte,and Mindanao islands and possibly throughoutmuch of the Mindanao PAIC. All individuals inmuseum collections have been collected on topof leaves of shrubs 2–4 m above the ground. Inaddition to the new species, P. chavacano alsooccurs on Mindanao Island; however, atpresent there is no evidence that these twospecies overlap in distributions, with P. pung-kaypinit seemingly distributed in the centraland eastern regions of Mindanao Island and P.chavacano found in the far western region ofMindanao Island (Zamboanga Peninsula;Fig. 1). Likewise, P. pungkaypinit and P. ditoyoccur in sympatry on Leyte Island, and both P.brevipes and P. pungkaypinit occur on BoholIsland, although whether they occur in sym-patry is unclear. Pseudogekko pungkaypinit isknown from four separate populations on threeislands. Although only a handful of individualsexist in museum collections, this speciesappears to be more-widely distributed and,therefore, possibly experiencing less overallconservation threats than its congeners. As aresult of this species broad geographic distri-bution (as currently recognized) and the lack ofavailable information about the species ecolo-gy, natural history, and intraspecific diversity,we do not find this species qualifies forCritically Endangered, Endangered, Vulnera-ble, or Near Threatened status under theIUCN criteria for classification (IUCN, 2013).Therefore, we recommend that the species beconsidered Least Concern until additionalinformation can be obtained concerning thehealth and diversity of wild populationsthroughout the Mindanao PAIC.

Etymology.—We derive the new speciesname from the Leyte language (Waray-Waray)terms for ‘‘treetop’’ (pungkay) and ‘‘lizard’’(pinit) in reference to the new forest gecko’s

arboreal microhabitat preference. The specificepithet is a noun of masculine gender. Sug-gested common name: Southern PhilippineFalse Gecko.

Pseudogekko ditoy sp. nov.(Figs. 1, 4–6, 8)

Holotype.—PNM 9811 (RMB FieldNo. 4365, formerly KU 326437), adult female,collected in secondary-growth forest on 28June 1999, in Sitio Cienda, Barangay Gabas,Municipality of Baybay, Leyte Province, LeyteIsland, Philippines (10u419N, 124u489E;WGS-84), by R.M. Brown.

Paratype.—One adult male (KU 326438)collected in lowland, secondary-growth forest(2030 to 2300 h) on 31 August 2001, in theCalbiga—a river area, Barangay Gabas, Mu-nicipality of Baybay, Leyte Province, LeyteIsland, Philippines (10u419N, 124u489E;WGS-84) by A.C. Diesmos.

Diagnosis.—Pseudogekko ditoy can be dis-tinguished from congeners by the followingcombination of characters: (1) body size small(SVL 49.4–52.6 mm); (2) axilla–groin distance50.7–56.5% SVL; (3) head length 18.2–18.9%SVL; (4) snout 57.4–59.3% head length; (5)Toe IV scansors 16 or 17; (6) paravertebralscales 180–185; (7) ventral scales 111–118; (8)supralabials 17–20; (9) infralabials 16 or 17;(10) circumorbitals 40 or 43; (11) precloacalpores 18; (12) femoral pores absent; (13)ground body coloration light brown; (14) headspots absent; (15) dorsolateral body spotsabsent; (16) limb spots absent; (17) tail bandsabsent; (18) body stripes absent; (19) interor-bital band absent; and (20) ciliary ringcoloration undifferentiated (Tables 1, 2).

Comparisons.—Characters distinguishingPseudogekko ditoy from all other species ofPseudogekko are summarized in Tables 1 and2. Pseudogekko ditoy most closely resemblesP. chavacano; however, P. ditoy differs fromP. chavacano by having tendencies towardsfewer Finger III scansors (14 or 15 vs. 15 or16), fewer Toe IV scansors (16 or 17 vs. 17–20), more supralabials (17–20 vs. 15 or 16),fewer circumorbitals (40–43 vs. 46), fewerparavertebral scales (180–185 vs. 195–197),fewer ventral scales (111–118 vs. 122–123),more precloacal pores (18 vs. 16), and by the


absence (vs. presence) of neon green head,dorsolateral body, and limbs spots and ab-sence (vs. presence) of transverse tail bands.

Pseudogekko ditoy can be distinguishedfrom P. compresicorpus, P. pungkaypinit, P.brevipes, and P. smaragdinus by having fewerparavertebral scales (,185 vs. .195) andventral scales (,118 vs. .119); from P.brevipes by having greater numbers of ToeIV scansors (16 or 17 vs. 13–15), supralabials(17–20 vs. 14–16), infralabials (16 or 17 vs.12–14), circumorbitals (40–43 vs. 35–37), andprecloacal pores (18 vs. 13–15), and by theabsence (vs. presence) of cream coloredspotting on the head and dorsolateral regionsof the body and absence (vs. presence) ofinterorbital banding; from P. compresicorpusand P. pungkaypinit by having a shortersnout–vent length (SVL , 52.6 mm vs..54.9); from P. compresicorpus by havingfewer Toe IV scansors (16 or 17 vs. 18 or 19)and a greater number of precloacal pores (18vs. 13 or 14), and by the absence (vs. presence)of neon green spotting on the head, dorsolat-eral regions of the body, and limbs and absence(vs. presence) of a light blue ciliary ring; fromP. pungkaypinit by having a shorter trunklength (AGD , 29.7 mm vs. .37.2), narrowerbody (MBW , 7.3 mm vs. .7.7), fewercircumorbitals (40–43 vs. 50–55), and by theabsence (vs. presence) of striped pigmentationpatterns on the body; and from P. smaragdinusby having a greater number of circumorbitals(40–43 vs. 33–35), fewer enlarged precloacalpores (18 precloacals vs. 32–41 precloacal-femorals) and a light brown (vs. bright neonyellow to orange [undisturbed] to neon green[disturbed]) body coloration, and absence (vs.presence) of black and white spots on the head,body and limbs and absence (vs. presence) oftransverse tail bands.

Description of holotype.—Details of thehead scalation are shown in Figure 4C. Adultfemale in excellent condition, gravid, with twoembryos visible through venter; small incisionin the sternal region (portion of liver removedfor genetic sample). Body small, slender, SVL52.6 mm; limbs well developed, moderatelyslender; tail regenerated, slender; margins oflimbs smooth, lacking cutaneous flaps ordermal folds; trunk lacking ventrolateralcutaneous fold.

Head size moderate, slightly differentiatedfrom neck, characterized by only slightlyhypertrophied temporal and adductor muscu-lature; snout rounded in dorsal and lateralaspect (Fig. 4C); HW 105.4% MBW, 80.2%HL; HL 18.3% SVL; SNL 74.0% HW, 59.4%HL; dorsal surfaces of head relatively homo-geneous, with only slightly pronounced con-cave postnasal, internasal, prefrontal, andinterorbital concavities; auricular openingmoderate, ovoid, angled slightly anteroven-trally and posterodorsally from beneath tem-poral swellings on either side of head;tympanum deeply sunken; eye large; pupilvertical, margin wavy (Fig. 4C); limbs anddigits relatively short and moderately slender;thighs moderately thicker compared to bra-chium; tibia length 8.0% SVL, 50.1% femurlength.

Rostral rectangular in anterior view, 33 asbroad as high, sutured anterolaterally withanteriormost enlarged supranasals; nostril sur-rounded by rostral, first labial, single enlargedpostnasal, and two enlarged supranasals; supra-nasals separated by five small median scales.

Total number of differentiated supralabials20/17 (L/R), bordered dorsally by one or tworows of differentiated, slightly enlarged snoutscales; total number of differentiated infra-labials 17/17 (L/R), bordered ventrally by 5–8rows of slightly enlarged scales; undifferenti-ated chin and gular scales; postrictal scalesundifferentiated; remaining undifferentiatedgulars very small, round, nonimbricate, juxta-posed (Fig. 4C), each scale surrounded by sixinterstitial granules, giving the appearance of aStar of David configuration under highmagnification (Fig. 5).

Dorsal cephalic scales fairly homogeneousin size, shape, disposition, and distribution;cephalic scalation slightly convex, round tooval scales; postnasal, prefrontal, internasal,and interorbital depressions; undifferentiatedposterior head scales granular, slightly convex;throat and chin scales small, juxtaposed andnonimbricate, making a moderately sharptransition to gular and pectoral region scala-tion, with enlarged cycloid, imbricate scales;circumorbitals 43/48 (L/R); interorbitals 42.

Axilla–groin distance 56.5% SVL; undiffer-entiated dorsal body scales round, convex,juxtaposed, relatively homogeneous in size;


each dorsal scale surrounded by six interstitialgranules; dorsals sharply transition to imbri-cate ventrals along lateral body surface;paravertebrals between midpoints of limbinsertions 185; ventrals between midpoints oflimb insertions 118; scales on dorsal surfacesof limbs more imbricate than dorsals; scaleson dorsal surfaces of hands and feet similar todorsal limb scales, heavily imbricate; ventralbody scales flat, cycloid, strongly imbricate,much larger than lateral or dorsal body scales,relatively homogeneous in size.

Nineteen enlarged scales (pore-bearing inmales; Fig. 6), in continuous precloacal series,arranged in a widely obtuse, W-formation;precloacals 9/10 (L/R); patch of slightlyenlarged scales posterior to precloacal series,roughly six scale rows in size, forming an ovalpatch.

Digits moderately expanded and covered onpalmar/plantar surfaces by bowed, unnotched,undivided scansors; digits with minute vestig-es of interdigital webbing; subdigital scansorsof hand: 8/8, 10/10, 12/12, 14/15, and 10/10(L/R) on Finger I–Finger V, respectively; foot:8/9, 11/11, 13/15, 16/16, and 11/11 (L/R) onToe I–Toe V, respectively; subdigital scansorsof hands and feet bordered proximally (onpalmar and plantar surfaces) by 1–4 slightlyenlarged scales that form a near-continuousseries with enlarged scansors; all digits clawed,but first claw greatly reduced; remainingterminal phalanges compressed, with largerecurved claws.

Tail regenerated, short, 30.5 mm, 58.0%SVL; single distinct fracture plane present,composed of enlarged, cylindrical scales wrap-ping around tail, separating original tail (ante-riorly) from regenerated tail (posteriorly); tailround, not depressed; TH 81.3% TW; caudalssimilar in size to dorsals, subcaudals similar insize to ventrals.

Coloration of holotype in preservative.—Background dorsal body coloration is light tanwith intermittent small cream and dark brownspeckles; this pattern is continued down thetail. The dorsal region of the limbs follows thesame pattern with a few areas of concentrateddarker blotches. The dorsal region of the headhas the same speckled pattern as the trunk,except for a darker region between and justposterior to the orbits, and a slightly lighter

region just anterior to the eyes. In thepostnasal region there is a stripe of mediumbrown speckles on top of the lighter tan andcream background color. The lateral region ofthe trunk has the same coloring pattern as thedorsal region. The lateral region of the headhas a thick, cream-colored ring of circumor-bital scales and a vague line of light brownblotches from snout to orbit. The venter of thebody is solid cream except for occasional lightbrown speckles. The ventral region of thetail is solid cream with occasional light brownspeckles that increase in intensity as youcontinue to the tip. The gular region of thehead is solid cream with light brown blotchesjust proximate to the infralabials. The ventralside of the limbs, hands, and feet is a solidcream except for a light brown stripe betweeneach scansor. In this individual, Finger IV onthe right hand is heavily speckled light brown.

Coloration in life (based on field notes andphotographs in life; Fig. 8C).—Dorsal groundcolor of head, trunk, and tail light tan withonly faint light brown mottling. Head slightlydarker in appearance, with increased lightbrown mottling in canthal and interocularregions as well as in posterior margins ofthe head. Lateral surface of the head withconspicuous series of four dark brown stripes,extending between the eye and ear opening.Stripes oriented anterodorsally from earopening and interspersed with olive greenblotches. Dorsal limb surfaces colored as thetrunk, but arms with increased light brownmottling and a slightly darker appearance. Tailcolored as trunk basally, then transitioning toa medium gray just posterior to tail insertion.Ventral coloration not apparent, but appearsto consist of a light cream ground color withminimal light brown mottling along the lateralmargins.

Measurements and scale counts of holotypein millimeters.—Snout–vent length 52.6 mm;tail length 30.5 (regenerated); total length 83.0(with regenerated tail); axilla–groin distance29.7; tail width 2.2; tail height 1.8; head length9.6; head width 7.7; head height 5.5; midbodywidth 7.3; snout length 5.7; eye diameter 3.5;eye–nares distance 4.4; internarial distance1.9; interorbital distance 4.4; femur length 8.4;tibia length 4.2; Finger III scansors 14; Toe IVscansors 16; supralabials 20; infralabials 17;


circumorbitals 43; paravertebral scales 185;ventral scales 118.

Variation.—Between the two specimensexamined we observed variation in the num-bers of supralabials, infralabials, and digitalscansors. The number of supralabials variedbetween 17 (KU 326438) and 20 (KU326437); infralabials varied between 16 (KU326438) and 17 (KU 326437). Finger IIIscansors were observed to vary between 14(KU 326437) and 15 (KU 326438); Toe IVscansors varied between 16 (KU 326437) and17 (KU 326438).

Distribution.—Pseudogekko ditoy is knownonly from Leyte Island (Fig. 1).

Ecology and natural history.—Pseudogekkoditoy has been observed in disturbed, second-ary-growth forest only; however, we assumethe species once occurred in lower-elevationprimary forest on Leyte Island. The twoindividuals represented in museum collectionswere both found on top of leaves of shrubs 2–3 m above the ground. A third individual wasobserved but not collected on top of a leaf ona tree 3.5 m above the ground on the bank of asmall stream system near a tree nurseryoutside of Baybay City (C. Siler, personalobservation). Interestingly, at least two speciesof Pseudogekko occur on Leyte Island (P.ditoy and P. brevipes); however, it may bepossible that additional exploration of LeyteIsland will result in the discovery of popula-tions of P. pungkaypinit as well. Whether ornot P. ditoy and P. brevipes occur in sympatryacross their ranges on Leyte Island is stillunknown. Although little data are available onthe natural history and ecology, as well as onthe health of wild populations of this uniquespecies, we believe that it qualifies as aconservation concern. We have evaluated thisspecies against the IUCN criteria for classifi-cation and find that it qualifies for the status ofVulnerable, VU, based on the followingcriteria: VU A2ac; B2ab(iii); D2 (IUCN,2013).

Etymology.—We derive the new speciesname from the Leyte language (Waray-Waray)term ditoy, meaning diminutive or ‘‘thesmaller one’’ in reference to the new speciessmall body size and its distinction from itslarger sympatric congener, P. pungkaypinit.The new name is a masculine noun in

apposition. Suggested common name: LeyteDiminutive False Gecko.

Pseudogekko chavacano sp. nov.(Figs. 1, 4–6, 8)

Holotype.—PNM 9812 (ACD FieldNo. 3784, formerly KU 314963), adult male,collected in secondary-growth forest (2000to 2230 h) on 21 April 2008, in BarangayPasonanca, Pasonanca National Park, Zam-boanga Sur Province, Mindanao Island,Philippines (06u58939.030N, 122u04901.650E;WGS-84), by A.C. Diesmos.

Paratype.—One adult female (KU 314964)collected on 12 July 2008 in Sitio km 24, BarangayBaluno, Pasonanca National Park, ZamboangaSur Province, Mindanao Island, Philippines(07u193.2240N, 122u1943.860E; WGS-84; 758 m)by R.M. Brown; two hatchlings (CAS-SU 23548,23549) collected on 8 May 1959 in a mid-montane dipterocarp forest near Cuyot Creek,Zamboanga del Norte Province, MindanaoIsland, Philippines (08u1997.970N, 123u3198.40E;WGS-84) by A.C. Alcala.

Diagnosis.—Pseudogekko chavacano can bedistinguished from congeners by the followingcombination of characters: (1) body size small(SVL 54.7–55.9 mm); (2) axilla–groin distance47.8–54.9% SVL; (3) head length 18.6–19.1%SVL; (4) snout 55.2–58.0% head length; (5)Toe IV scansors 17–20; (6) paravertebral scales195–197; (7) ventral scales 122 or 123; (8)supralabials 15 or 16; (9) infralabials 16 or 17;(10) circumorbitals 46; (11) precloacal pores16; (12) femoral pores absent; (13) dominantbody coloration light brown; (14) conspicuoushead spotting present, dense, neon green; (15)conspicuous dorsolateral spotting present,neon green; (16) conspicuous limb spottingpresent, dense, neon green; (17) tail bandingpresent; (18) body striping absent; (19) inter-orbital banding absent; and (20) ciliary ringcoloration absent (Tables 1, 2).

Comparisons.—Characters distinguishingPseudogekko chavacano from all other speciesof Pseudogekko are summarized in Tables 1and 2. Pseudogekko chavacano most closelyresembles P. ditoy; however, P. chavacanodiffers from P. ditoy by having a tendencytowards a greater number of Finger IIIscansors (15 or 16 vs. 14 or 15), a tendency


towards a greater number of Toe IV scansors(17–20 vs. 16 or 17), fewer supralabials (15 or16 vs. 17–20), a greater number of circumor-bitals (46 vs. 40–43), paravertebral scales(195–197 vs. 180–185), and ventral scales(122–123 vs. 111–118), and fewer precloacalpores (16 vs. 18), and by the presence (vs.absence) of neon green spotting on the head,dorsolateral region of the body, and limbs andpresence (vs. absence) of tail banding.

Pseudogekko chavacano can be distin-guished from P. compresicorpus, P. pungkay-pinit, P. brevipes, and P. smaragdinus byhaving fewer paravertebral scales (,197 vs..226); from P. compresicorpus, P. pungkay-pinit, and P. smaragdinus by having fewerventral scales (,123 vs. .124); fromP. brevipes by having greater numbers ofFinger III scansors (15 or 16 vs. 12–14), ToeIV scansors (17–20 vs. 13–15), infralabials (16or 17 vs. 12–14), circumorbitals (46 vs. 35–37),and precloacal pores (16 vs. 13–15), and bythe presence of neon green (vs. creamcolored) spotting on the head and dorsolateralregion of the body, presence (vs. absence) ofneon green spotting on the limbs, presence(vs. absence) of tail banding and absence (vs.presence) of interorbital banding; from P.compresicorpus, P. pungkaypinit, and P.smaragdinus by having a shorter total bodylength (TotL , 95.8 mm vs. .103.6) and agreater relative head length (HL 19% SVL vs.,18%); from P. compresicorpus by havinggreater numbers of circumorbitals (46 vs. 39–45), and precloacal pores (16 vs. 13 or 14), andby the presence (vs. absence) of tail bandingand absence (vs. presence) of a light blueciliary ring; from P. pungkaypinit by having ashorter trunk length (AGD , 30.0 mm vs..37.2), narrower body (MBW , 6.7 mm vs..7.7), fewer circumorbitals (46 vs. 50–55),fewer precloacal pores (16 vs. 17–20), by thepresence (vs. absence) of neon green spottingon the head, dorsolateral region of the body,and limbs, presence (vs. absence) of tailbanding and absence (vs. presence) of stripedpigmentation patterns on the body; and fromP. smaragdinus by having a greater number ofcircumorbitals (46 vs. 33–35), fewer enlargedpores (16 precloacal pores vs. 32–41 precloa-cal-femoral pores), disparate body coloration(light brown vs. bright neon yellow to orange

[undisturbed] to neon green [disturbed]), andby the absence (vs. presence) of femoral poresand presence of neon green (vs. black andwhite) spotting on the head, dorsolateralregion of the body, and limbs.

Description of holotype.—Details of thehead scalation are shown in Figure 4D.Adult male in excellent condition, hemipenesnot inverted, hemipene bulge present; smallincision in the sternal region (portion of liverremoved for genetic sample). Body small,slender, SVL 55.9 mm; limbs well developed,moderately slender; tail regenerated, slender;margins of limbs smooth, lacking cutaneousflaps or dermal folds; trunk lacking ventrolat-eral cutaneous fold.

Head moderate in size, slightly differenti-ated from neck, characterized by only slightlyhypertrophied temporal and adductor muscu-lature; snout rounded in dorsal and lateralaspect (Fig. 4D); HW 131.7% MBW, 81.4%HL; HL 18.6% SVL; SNL 71.2% HW, 58.0%HL; dorsal surfaces of head relatively homo-geneous, with only slightly pronounced con-cave postnasal, internasal, prefrontal, andinterorbital concavities; auricular openingmoderate, ovoid, angled slightly anteroven-trally and posterodorsally from beneathtemporal swellings on either side of head;tympanum deeply sunken; eye large; pupilvertical, margin wavy (Fig. 4D); limbs anddigits relatively short and moderately slender;thighs moderately thicker compared to bra-chium; tibia length 8.0% SVL, 50.1% femurlength.

Rostral rectangular in anterior view, 33 asbroad as high, sutured anterolaterally withanteriormost enlarged supranasals; nostril sur-rounded by rostral, first labial, single enlargedpostnasal, and two enlarged supranasals; ante-riormost enlarged supranasals separated bythree small median scales.

Total number of differentiated supralabials15/15 (L/R), bordered dorsally by one row ofdifferentiated, slightly enlarged snout scales;total number of differentiated infralabials 17/17 (L/R), bordered ventrally by 5–8 rows ofslightly enlarged scales; undifferentiated chinand gular scales; postrictal scales undiffer-entiated; remaining undifferentiated gularsvery small, round, nonimbricate, juxtaposed(Fig. 4D).


Dorsal cephalic scales fairly homogeneousin size, shape, disposition, and distribution;cephalic scalation slightly convex, round tooval scales; postnasal, prefrontal, internasal,and interorbital depressions; undifferentiatedposterior head scales granular, slightly convex;throat and chin scales small, juxtaposed, andnonimbricate, making a sharp transition togular and pectoral region scalation, withenlarged cycloid, imbricate scales; circumor-bitals 45/46 (L/R); interorbitals 48.

Axilla–groin distance 47.8% SVL; undiffer-entiated dorsal body scales round, convex,juxtaposed, relatively homogeneous in size;each dorsal scale surrounded by six interstitialgranules, giving the appearance of a Star ofDavid configuration under high magnification(Fig. 5); dorsals sharply transition to imbricateventrals along lateral body surface; paraverte-brals between midpoints of limb insertions197; ventrals between midpoints of limbinsertions 122; scales on dorsal surfaces oflimbs more imbricate than dorsals; scales ondorsal surfaces of hands and feet similar todorsal limb scales, heavily imbricate; ventralbody scales flat, cycloid, strongly imbricate,much larger than lateral or dorsal body scales,relatively homogeneous in size.

Sixteen enlarged pore-bearing scales, incontinuous precloacal series, arranged in awidely obtuse, W-formation (Fig. 6); precloa-cals 8/8 (L/R); precloacal series borderedposteriorly by single row of enlarged scales;precloacals situated atop a substantial pre-cloacal bulge.

Digits moderately expanded and coveredon palmar–plantar surfaces by bowed, un-notched, undivided scansors; digits withminute vestiges of interdigital webbing; sub-digital scansors of hand: 8/8, 10/11, 13/13, 15/15, and 11/11 (L/R) on Finger I–Finger V,respectively; foot: 11/10, 13/12, 15/15, 17/16,and 12/12 (L/R) on Toe I–Toe V, respective-ly; subdigital scansors of hands and feetbordered proximally (on palmar and plantarsurfaces) by 1–4 slightly enlarged scales thatform a near-continuous series with enlargedscansors; all digits clawed but first clawgreatly reduced; remaining terminal phalan-ges compressed, with large recurved claws.

Tail regenerated, short, 30.7 mm, 55.0%SVL; single distinct fracture plane present,

composed of enlarged, cylindrical to rectan-gular scales wrapping around tail, separatingoriginal tail (anteriorly) from regenerated tail(posteriorly); tail round, not highly depressed;TH 67.0% TW; caudals similar in size todorsals, subcaudals similar in size to ventrals.

Coloration of holotype in preservative.—Background dorsal body coloration light greywith intermittent small cream and dark greyspeckles; pattern continued down tail; dorsumof trunk with occasional larger cream spots;limbs with same coloration pattern as trunk,without larger cream spots; dorsal region ofhead continues same pattern except for darkergrey area above and between orbits; lateralside of trunk has same coloration pattern asdorsum; lateral side of head with thin creamcircumorbital ring of scales, remainder ofhead with same coloration pattern as body;ventral body surface solid cream, patterncontinues until just posterior to cloacal region,when coloration pattern of dorsal trunkbecomes prevalent, larger cream spots absent;ventral side of head solid cream, withoccasional light grey speckling along infra-labial scales and thicker with light greyblotches just proximate to infralabials; ventralsurfaces of limbs, hands, and feet are solidcream except for space between scansors,which is dark grey.

Coloration in life (based on field notes andphotographs in life; Fig. 8D).—Dorsal groundcolor of head and trunk grayish brown, withlight tan, irregular mottling; head with tanblotches highlighted with green centrumsthroughout; anterior half of canthal regionheavily mottled light tan; tan mottling ontrunk highlighted with green only alongdorsolateral margin, forming vague longitudi-nal series of moderately sized blotches andventrally paired smaller blotches extendingfrom arm insertion to base of tail; dorsalcoloration at base of tail with minimal tanmottling; dorsal limb surfaces with colorationof head, but appearing with more regularspots, coloration lighter on arms than hind-limbs; legs with higher density of greenish-tanspots; tail regenerated to base, with a lightgray ground color overlain with dark brownmottling anteriorly; ventral ground trunk colorcream yellow, with minimal light gray mottlingalong lateral margins.


Measurements and scale counts of holotypein millimeters.—Snout–vent length 55.9; taillength 30.7 (regenerated); total length 86.6(with regenerated tail); axilla–groin distance26.7; tail width 2.9; tail height 1.9; head length10.4; head width 8.5; head height 6.1; mid-body width 6.4; snout length 6.0; eye diameter3.8; eye–nares distance 5.1; internarial dis-tance 2.2; interorbital distance 4.7; femurlength 9.8; tibia length 3.9; Finger III scansors15; Toe IV scansors 17; supralabials 15;infralabials 17; preanofemoral pores 16; cir-cumorbitals 46; paravertebral scales 197;ventral scales 122.

Variation.—Between the two specimensexamined, we observed variation in the num-bers of supralabials, infralabials, and digitalscansors. The number of supralabials variedbetween 15 (KU 314963) and 16 (KU314964); infralabials varied between 16 (KU314964) and 17 (KU 314963). Finger IIIscansors were observed to vary between 15(KU 314963) and 16 (KU 314964); Toe IVscansors varied between 17 (KU 314963) and20 (KU 314964).

Distribution.—Pseudogekko chavacano isknown only from the Zamboanga Peninsulaof Mindanao Island (Fig. 1).

Ecology and natural history.—Pseudogekkochavacano has been observed in secondary-growth forest only, with the two availableadult specimens collected inside the protectedPasonanca watershed. Two eggs were col-lected in humus on the bank of a creek inZamboanga del Norte Province by A.C. Alcalain 1959 and were subsequently hatched in thelab a week later (CAS-SU 23548, 23549). Notonly do we assume this species once occurredin previously extensive, low-elevation primaryforest in western Mindanao, but also weassume it may possess a wider distributionthroughout the Zamboanga Peninsula ofMindanao Island (Fig. 1). As for P. ditoy,the two adult specimens were both found ontop of leaves of shrubs 2–3 m above the ground.The Zamboanga City Watershed in PasonancaNatural Park represents one of the most well-protected, low-elevation, primary forests re-maining in the southwestern Philippines;therefore, a large portion of this species rangeis well protected. However, due to therelatively limited known distribution of this

species, it still qualifies for formal IUCNclassification. We have evaluated this speciesagainst the IUCN criteria for classification andfind that it qualifies for the status of Vulner-able, VU, based on the following criteria: VUA2ac; B2ab(iii); D2 (IUCN, 2013).

Etymology.—The specific epithet is derivedfrom the traditional term Chavacanos, other-wise known as Philippine–Creole Spanish, thelanguage still spoken today by over 800,000people, the Zamboanguenos, of ZambangaCity, southwestern Philippines. At a timewhen cultural identities and languages con-tinue to be lost in the Philippines, Chabakanocontinues to be the socially, historically, andculturally significant language of choice formillions of Filipinos. The name is a pluralnoun of masculine gender. Suggested com-mon name: Zamboanga False Gecko.

DISCUSSION

The new taxa recognized in this paperincrease the total number of known species ofPseudogekko to six, and all are endemic to thePhilippines. Multilocus phylogenetic analysesof Siler et al. (2014a) further support the fourlineages within the Pseudogekko compresicor-pus Complex recognized in this study (Fig. 1).As a testament to the rarity of Pseudogekkospecies in museum collections and our rela-tively poor understanding of the diversity andrelationships in this clade, we note that thenewly described species all have been mas-querading as members of a single widespreadspecies for nearly a century (Taylor, 1915).Despite small sample sizes, we are confidentin our taxonomic decisions based on thenumerous conspicuous diagnostic morpholog-ical features of all focal species (Table 2) andtheir deep genetic divergences (Siler et al.,2014a).

Mitochondrial sequence divergences be-tween members of the Pseudogekko compre-sicorpus Complex are remarkably high (Sileret al., 2014a). At a minimum, P. ditoy and P.chavacano are 13.9% divergent from eachother; however, many observed divergencesare above 20% (Siler et al., 2014a). Alsointeresting is the considerable intraspecificgenetic diversity observed within P. compre-sicorpus from the Luzon PAIC and P.pungkaypinit from the Mindanao PAIC (up


to 19% and 12%, respectively; Siler et al.,2014a). These surprisingly high levels ofgenetic diversity suggest that additional cryp-tic species remain unrecognized. In the caseof these two species, the observed interpop-ulation diversity corresponds to isolated re-gions in the Luzon and Mindanao PAICsalready known to contain high levels ofendemism among squamate reptiles (Fig. 1).For example, the deep split between popula-tions of P. pungkaypinit from Central Mind-anao Island and those from Leyte and Boholislands is similar to species-level boundariesobserved in several other groups of verte-brates (e.g., Cyrtodactylus, Brachymeles, Var-anus; Siler et al., 2014a,d), birds (Hosner etal., 2013), and mammals (Steppan et al.,2003). Likewise, Luzon Island has been thefocus of several recent studies revealingdiscrete faunal subregions throughout theisland, including the Zambales, Cordillera,and Sierra Madres mountain ranges, eachwith seemingly unique communities of am-phibians and reptiles (Siler et al., 2011;Devan-Song and Brown, 2012; Brown et al.,2013a; Siler et al., 2014d). Unfortunately,small series containing only juvenile orsubadult specimens from these areas limitour ability to evaluate these genetically uniquepopulations taxonomically.

Recognizing the levels of threat to thesurvival of many of these unique, endemicgeckos, future surveys should focus on thesepoorly sampled populations to allow forappropriate evaluations of species diversity.In an attempt to aid in efforts to quickly studyand protect unique lineages within this genus,we suggest the following populations beprioritized for investigation. (1) Pseudogekkobrevipes: The currently sampled populationsof P. brevipes are not only geneticallydivergent (.20% mitochondrial sequencedivergence between Leyte and Negros islandpopulations; Fig. 1; Siler et al., 2014a), but therecognized distribution of this species spanstwo distinct faunal regions (Visayan andMindanao PAICs; Fig. 1). (2) Central Mind-anao Island: Collections of Pseudogekko fromMindanao Island are rare in museum collec-tions. With the presence of a unique specieson the Zamboanga Peninsula (P. chavacano)and the deeply divergent population of P.

pungkaypinit (central, eastern Mindanao),other portions of Mindanao Island (i.e., theCotobato coast mountains) may likely harboradditional divergent lineages of Pseudogekko.(3) Luzon PAIC: Numerous, possibly distinctevolutionary lineages still appear to be unrec-ognized throughout the Luzon PAIC. PolilloIsland possesses a genetically divergent pop-ulation of P. compresicorpus, as does theSierra Madres Mountain Range in the north-eastern portions of the island. Interestingly,outside of the holotype of P. compresicorpusfrom the Bataan Peninsula of western LuzonIsland (Fig. 1), there are no records ofPseudogekko from much of the northwest,including the Zambales and Cordillera moun-tain ranges (Diesmos et al., 2004; Brown et al.,2012a; Devan-Song and Brown, 2012). Wheth-er this genus is truly absent from these distinctfaunal subcenters or these patterns are simplyan artifact of limited biodiversity surveysremains to be determined. However, we haveno doubt that additional survey work in theLuzon faunal region will result in the discoveryof additional new species of false geckos. (4)Tablas Island: There is a single record of P.compresicorpus from Tablas Island in theRomblon Island Group (CAS 139713). Notonly does this adult female appear to haveseveral distinct morphological characters (e.g.,infralabials 17, enlarged scales in the precloacalregion 15, circumorbitals 54), but also thisisland group harbors numerous other endemicspecies (e.g., Platymantis lawtoni, Platymantislevigatus, Gekko romblon, G. coi; Brown et al.,2011b; Siler et al., 2012b), suggesting that thisPseudogekko may be a unique species. (5)Masbate Island: Like Tablas Island, a singlerecord of P. compresicorpus from MasbateIsland (CAS 141560) is known. This adultfemale also possesses an anomalous combina-tion of unique morphological characters (Fin-ger III scansors 13, Toe IV scansors 16,infralabials 17, circumorbitals 49). Further-more, although the faunal affinities of MasbateIsland are still somewhat vague, the island isconsidered a member of the Visayan PAIC, notthe Luzon PAIC (Brown and Diesmos, 2009;Brown et al., 2013a), and also harbors endemicsquamate reptiles (e.g., Brachymeles tungaoi,B. cf. bonitae Siler and Brown, 2010; Davis etal., in press).


So little is known about the ecology ofPseudogekko that broad conclusions on archi-pelago-wide diversity, microhabitat breadth,and natural history cannot be made at thistime (Brown and Alcala, 1978). Interestingly,at present only a few islands possess morethan one species of Pseudogekko: Luzon,Polillo, Leyte, and Mindanao. As moreinformation on false geckos becomes avail-able, it will be interesting to determinewhether species occur sympatrically on theseand other islands. On Polillo Island, P.smaragdinus geckos possess a microhabitatpreference for Pandanus leaves (Taylor, 1915,1922b). The single collected specimen ofPseudogekko smaragdinus collected on Mt.Labo, Bicol Peninsula (Luzon Island) wascollected from inside the husk surroundingthe trunk of a wild banana plant (R. Brown,personal observation). Little is known aboutthe Polillo Island population of P. compresi-corpus except that specimens were found onthe upper surfaces of leaves at night. Becausesuccessful conservation efforts will depend onour understanding of the habitats of species westrive to protect, future studies should attemptto document microhabitat preferences.

As a result of many recent studies aimed atinvestigating the diversity of geckos through-out the Philippine archipelago (Linkem et al.,2010; Welton et al., 2010; Brown et al., 2012a;Siler et al., 2014d), we will soon approacha more-accurate understanding of gekkoniddiversity in the country. A few groups remainthat have yet to be comprehensively evaluatedfor taxonomic diversity within a phylogeneticcontext, including Lepidodactylus, Hemiphyl-lodactylus (Grismer et al., 2013), Luperosau-rus (Brown et al., 2012a), Cyrtodactylus (Sileret al., 2010; Welton et al., 2010), and evenGekko (Siler et al., 2012a, 2014c); the focus offuture studies should include these. Unfortu-nately, other groups of Philippine amphibiansand reptiles remain much-more poorly stud-ied (Brown et al., 2013a), adding to thenascent body of literature demonstrating thatspecies diversity in amphibians and reptiles inthe Philippines is substantially underestimat-ed (Brown et al., 2002; Brown and Diesmos,2009; Brown et al., 2013a) and possibly atgreater risk than is presently appreciated.Comprehensive analyses of the fauna using

multiple lines of evidence and nontraditionaltaxonomic characters may provide the bestopportunities for future integrative approach-es to understanding Philippine reptile andamphibian megadiversity (Barley et al., 2013;Grismer et al., 2013; Linkem and Brown,2013; Siler et al., 2014c).

Acknowledgments.—We thank the Protected Areas andWildlife Bureau of the Philippine Department ofEnvironment and Natural Resources for facilitating thecollecting and export permits necessary for this andrelated studies; we are particularly grateful to M. Lim, C.Custodio, J. de Leon, and A. Tagtag. Fieldwork wasconducted under the Memorandum of Agreement withthe Protected Areas and Wildlife Bureau of the Philip-pines (2009–2014), Gratuitous Permit to Collect No. 221,and The University of Kansas Institutional Animal Careand Use Committee (KU IACUC) Approval (158-01).Financial support for fieldwork and lab work was providedby a Panorama Fund grant from the KU BiodiversityInstitute and travel funds from the KU Department ofEcology and Evolutionary Biology, a Madison and LilaSelf Fellowship from the KU, a Fulbright Fellowship, aFulbright-Hayes Fellowship, and National Science Foun-dation (NSF) DEB 0804115 to CDS, as well as supportfrom NSF DEB 0743491, and NSF EF-0334952 to RMB.For the loans of specimens we thank J. Vindum andA. Leviton (California Academy of Sciences). Helpfulcomments on this manuscript were provided by J.Grummer, M. Harvey, R. Davis and J. Hoskins.

LITERATURE CITED

Alcala, E.L., A.C. Alcala, and C.N. Dolino. 2004.Amphibians and reptiles in tropical rainforest fragmentson Negros Island, the Philippines. EnvironmentalConservation 31:254–261.

Barley, A.J., J. White, A.C. Diesmos, and R.M. Brown.2013. The challenge of species delimitation at theextremes: diversification without morphological changein Philippine sun skinks. Evolution 67:3556–3572.

Boettger, O. 1897. Neue reptilien und batrachier von denPhilippinen. Zoologischer Anzeiger 20:161–166.

Boulenger, G.A. 1885. Catalogue of the Lizards in theBritish Museum (Natural History), Volume 1, Geck-onidae, Eublepharidae, Uroplatidae, Pygopodidae, Aga-midae. Order of the Trustees, UK.

Brown, R.M., and A.C. Alcala. 2000. Geckos, cave frogs,and small land-bridge islands in the Visayan sea. HaringIbon 2:19–22.

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

Brown, R.M., and A.C. Diesmos. 2009. Philippines,biology. Pp. 723–732 In R. Gillespie and D. Clague(Eds.), Encyclopedia of Islands. University of CaliforniaPress, USA.

Brown, R.M., and S.I. Guttman. 2002. Phylogeneticsystematics of the Rana signata complex of Philippineand Bornean stream frogs: reconsideration of Huxley’s


modification of Wallace’s Line at the Oriental–Austra-lian faunal zone interface. Biological Journal of theLinnean Society 76:393–461.

Brown, R.M., J.W. Ferner, and A.C. Diesmos. 1997.Definition of the Philippine parachute gecko, Ptycho-zoon intermedium Taylor 1915 (Reptilia: Squamata:Gekkonidae): redescription, designation of a neotype,and comparisons with related species. Herpetologica53:357–373.

Brown, R.M., J. Supriatna, and H. Ota. 2000. Discovery ofa new species of Luperosaurus (Squamata; Gekkonidae)from Sulawesi, with a phylogenetic analysis of thegenus, and comments on the status of Luperosaurusserraticaudus. Copeia 2000:191–209.

Brown, R.M., A.C. Diesmos, and A.C. Alcala. 2002. Thestate of Philippine herpetology and the challenges forthe next decade. Silliman Journal 42:18–87.

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

Brown, R.M., C.H. Oliveros, C.D. Siler, and A.C. Diesmos.2008. A new Gekko from the Babuyan Islands, northernPhilippines. Herpetologica 64:305–320.

Brown, R.M., C. Oliveros, C.D. Siler, and A.C. Diesmos.2009. Phylogeny of Gekko from the northern Philip-pines, and description of a new species from CalayanIsland. Journal of Herpetology 43:620–635.

Brown, R.M., C.D. Siler, C.H. Oliveros, A.C. Diesmos,and A.C. Alcala. 2011a. A new Gekko from SibuyanIsland, central Philippines. Herpetologica 67:460–476.

Brown, R.M., A.C. Diesmos, and C.H. Oliveros. 2011b.New flap-legged forest gecko (genus Luperosaurus)from the northern Philippines. Journal of Herpetology45:202–210.

Brown, R.M., C.D. Siler, I. Das, and Y. Min. 2012a.Testing the phylogenetic affinities of Southeast Asia’srarest geckos: flap-legged geckos (Luperosaurus), flyinggeckos (Ptychozoon) and their relationship to thepan-Asian genus Gekko. Molecular Phylogenetics andEvolution 63:915–921.

Brown, R.M., C.D. Siler, L. Grismer, I. Das, and J.McGuire. 2012b. Phylogeny and cryptic diversificationin Southeast Asian flying geckos. Molecular Phyloge-netics and Evolution 65:351–361.

Brown, R.M., C.D. Siler, C.H. Oliveros, J.A. Esselstyn,A.C. Diesmos, P.A. Hosner, C.W. Linkem, A.J. Barley,J.R. Oaks, M.B. Sanguila, L.J. Welton, D.C. Blackburn,R.G. Moyle, A.T. Peterson, and A.C. Alcala. 2013a.Evolutionary processes of diversification in a modelisland archipelago. Annual Review of Ecology, Evolu-tion, and Systematics 44:411–435.

Brown, R.M., C.D. Siler, C.H. Oliveros, L.J. Welton, A.A.Rock, J. Swab, M. Van Weerd, D. Rodrigues, E. Jose,and A.C. Diesmos. 2013b. The amphibians and reptilesof Luzon Island, Philippines, VIII: the herpetofauna ofCagayan and Isabela Provinces, northern Sierra MadreMountain Range. Zookeys 266:1–120.

Brown, W.C. 1964. The status of Pseudogekko shebae andobservations on the geckos of the Solomon Islands.Brevoria 204:1–8.

Brown, W.C., and A.C. Alcala. 1970. The zoogeography ofthe herpetofauna of the Philippine Islands, a fringing

archipelago. Proceedings of the California Academy ofSciences 38:105–130.

Brown, W.C., and A.C. Alcala. 1978. Philippine Lizardsof the Family Gekkonidae. Silliman University Press,Philippines.

Brown, W.C., and V. M. Tanner. 1949. Rediscovery of thegenus Pseudogekko with description of a new species fromthe Solomon Islands. The Great Basin Naturalist 9:41–45.

Catibog-Sinha, C.S., and L.R. Heaney. 2006. PhilippineBiodiversity: Principles and Practice. Haribon Founda-tion for the Conservation of Natural Resources, Inc.,Philippines.

Davis, D.R., K.D. Feller, R.M. Brown, and C.D. Siler. Inpress. Evaluating the diversity of Philippine slenderskinks of the Brachymeles bonitae complex (Reptilia:Squamata: Scincidae): redescription of B. tridactylusand descriptions of two new species. Journal ofHerpetology.

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

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

Devan-Song, A., and R.M. Brown. 2012. Amphibians andreptiles of Luzon Island, Philippines, VI: the herpeto-fauna of the Subic Bay Area. Asian HerpetologicalResearch 3:1–20.

Diesmos, A.C., R.M. Brown, and G.V.A. Gee. 2004.Preliminary report on the amphibians and reptiles ofBalbalasang-Balbalan National Park, Luzon Island,Philippines. Sylvatrop, The Technical Journal of Philip-pine Ecosystems and Natural Resources 13:63–80.

Dudley, R., G. Byrnes, S. Yanoviak, B. Borrell, R.M.Brown, and J. McGuire. 2007. Gliding and thefunctional origins of flight: biomechanical novelty ornecessity? Annual Review of Ecology, Evolution, andSystematics 38:179–201.

Frost, D.R., and D.M. Hillis. 1990. Species in conceptand practice: herpetological applications. Herpetologica46:87–104.

Grismer, L.L., P.L. Wood Jr., S. Anuar, M.A. Muin,E.S.H. Quah, J.A. McGuire, R.M. Brown, and N.V. Tri.2013. Integrative taxonomy uncovers high levels ofcryptic species diversity in Hemiphyllodactylus Bleeker,1860 (Squamata: Gekkonidae) and the description of anew species from Peninsular Malaysia. ZoologicalJournal of the Linnaean Society 169:849–880.

Hosner, P.A., A.S. Nyari, and R.G. Moyle. 2013. Waterbarriers and intra-island isolation contribute to diver-sification in the insular Aethopyga sunbirds (Aves:Nectariniidae). Journal of Biogeography 40:1094–1106.

IUCN (The International Union for Conservation ofNature). 2013. IUCN Red List of Threatened Species.Version 2013.2. Available at http://www.iucnredlist.org/.Archived by WebCite at http://www.webcitation.org/6NXZz3bAI on 20 February 2014.

Kluge, A.G. 1967. Higher taxonomic categories ofgekkonid lizards and their evolution. Bulletin of theAmerican Museum of Natural History 135:1–60.


Kluge, A.G. 1968. Phylogenetic relationships of thegekkonid lizard genera Lepidodactylus Litzinger, Hemi-phyllodactylus Bleeker, and Pseudogekko Taylor. ThePhilippine Journal of Science 95:331–352.

Kluge, A.G. 1993. Gekkonoid Lizard Taxonomy. Interna-tional Gecko Society, USA.

Linkem, C.W., and R.M. Brown. 2013. Systematicrevision of the Parvoscincus decipiens (Boulenger,1894) complex of Philippine forest skinks (Squamata:Scincidae: Lygosominae) with descriptions of sevennew species. Zootaxa 3700:501–533.

Linkem, C.W., C.D. Siler, A.C. Diesmos, E. Sy, and R.M.Brown. 2010. A new species of Gekko (Squamata:Gekkonidae) from central Luzon Island, Philippines.Zootaxa 2396:37–49.

Peters, W. 1867. Herpetologische Notizen. Monatsberichtder Koniglich Preussischen Akademie der Wissenschaf-ten zu Berlin 1867:13–37.

PhilBREO (Philippines Biodiversity Research and Edu-cation Outreach). 2014. Philippines Biodiversity Re-search and Education Outreach. Available at http://philbreo.lifedesks.org/. Archived by WebCite at http://www.webcitation.org/6Q2F84XoU on 2 June 2014.

Rosler, H., C.D. Siler, R.M. Brown, A.D. Demegillo, andM. Gaulke. 2006. Gekko ernstkelleri sp. n.—a newgekkonid lizard from Panay Island, Philippines. Sala-mandra 42:197–211.

Sabaj Perez, M.H. 2013. Standard symbolic codes forinstitutional resource collections in herpetology andichthyology: an online reference. American Society ofIchthyologists and Herpetologists, Washington, DC.Available at http://www.asih.org/. Archived by WebCiteat http://www.webcitation.org/6JMxHNRhZ on 20 Feb-ruary 2014.

Sanguila, M.B., C.D. Siler, A.C. Diesmos, O. Nuneza, andR.M. Brown. 2011. Phylogeography, genetic structure,genetic variation, and potential species boundaries inPhilippine slender toads. Molecular Phylogenetics andEvolution 61:333–350.

Siler, C.D., and R.M. Brown. 2010. Phylogeny-basedspecies delimitation in Philippine slender skinks (Repti-lia: Squamata: Scincidae: Brachymeles): taxonomic revi-sion of pentadactyl species groups and description ofthree new species. Herpetological Monographs 24:1–54.

Siler, C.D., J.R. Oaks, J.A. Esselstyn, A.C. Diesmos, andR.M. Brown. 2010. Phylogeny and biogeography ofPhilippine bent-toed geckos (Gekkonidae: Cyrtodactylus)contradict a prevailing model of Pleistocene diversifica-tion. Molecular Phylogenetics and Evolution 55:699–710.

Siler, C.D., A.M. Fuiten, R.M. Jones, A.C. Alcala, and R.M.Brown. 2011. Phylogeny-based species delimitation inPhilippines slender skinks (Reptilia: Squamata: Scinci-dae) II: taxonomic revision of Brachymeles samarensisand description of five new species. HerpetologicalMonographs 25:76–112.

Siler, C.D., J.R. Oaks, L.J. Welton, C.W. Linkem, J.C.Swab, A.C. Diesmos, and R.M. Brown. 2012a. Didgeckos ride the Palawan raft to the Philippines? Journalof Biogeography 39:1217–1234.

Siler, C.D., J.C. Swab, C.H. Oliveros, A.C. Diesmos, L.Averia, A.C. Alcala, and R.M. Brown. 2012b. Amphib-ians and reptiles, Romblon Island Group, centralPhilippines: comprehensive herpetofaunal inventory.Check List 8:443–462.

Siler, C.D., R.M. Jones, A.C. Diesmos, M.L. Diesmos,and R.M. Brown. 2012c. Phylogeny-based speciesdelimitation in Philippine slender skinks (Reptilia:Squamata: Scincidae) III: taxonomic revision of Bra-chymeles gracilis and description of three new species.Herpetological Monographs 26:135–172.

Siler, C.D., T.A. Dececchi, C.L. Merkord, D.R. Davis, T.J.Christiani, and R.M. Brown. 2014a. Cryptic diversity andpopulation genetic structure in the rare, endemic, forest-obligate, slender geckos of the Philippines. MolecularPhylogenetics and Evolution 70:204–209.

Siler, C.D., A. Lira-Noriega, and R.M. Brown. 2014b.Conservation genetics of Australasian sailfin lizards:flagship species threatened by coastal development andinsufficient protected area coverage. Biological Con-servation 169:100–108.

Siler, C.D., J.R. Oaks, K. Cobb, H. Ota, and R.M. Brown.2014c. Critically endangered island endemic or periph-eral population of a widespread species? Conservationgenetics of Kikuchi’s gecko and the global challenge ofprotecting peripheral oceanic island endemic verte-brates. Diversity and Distributions 20:756–772.

Siler, C.D., C.W. Linkem, K. Cobb, J.L. Watters, A.C.Diesmos, and R.M. Brown. 2014d. Taxonomic revisionof the semi-aquatic skink Parvoscincus leucospilos(Reptilia: Squamata: Scincidae), with description ofthree new species. Zootaxa 3847:388–412.

Simpson, G.G. 1961. Principles of Animal Taxonomy.Columbia University Press, USA.

Steppan, S.J., C. Zawadzki, and L.R. Heaney. 2003.Molecular phylogeny of the endemic Philippine rodentApomys (Muridae) and the dynamics of diversificationin an oceanic archipelago. Biological Journal of theLinnean Society 80:699–715.

Taylor, E.H. 1915. New species of Philippine lizards. ThePhilippine Journal of Science 10:89–109.

Taylor, E.H. 1922a. The lizards of the Philippine Islands.Philippine Bureau of Science 17:1–269.

Taylor, E.H. 1922b. Additions to the herpetological faunaof the Philippine Islands, I. The Philippine Journal ofScience 21:161–206.

Underwood, G. 1954. On the classification and evolutionof geckos. Proceedings of the Zoological Society ofLondon 124:469–492.

Welton, L.J., C.D. Siler, C.W. Linkem, A.C. Diesmos, andR.M. Brown. 2010. Philippine bent-toed geckos of theCyrtodactylus agusanensis complex: multilocus phylog-eny, morphological diversity, and descriptions of threenew species. Herpetological Monographs 24:55–85.

Wermuth, H. 1965. Liste der rezenten amphibien andreptilien: Gekkonidae, Pygopodidae, Xantusiidae. DasTierreich 80:1–246.

Wiley, E.O. 1978. The evolutionary species conceptreconsidered. Systematic Zoology 27:17–26.

.Published: 15 December 2014

APPENDIX

Specimens Examined

Numbers in parentheses following species namesindicate the number of specimens examined. All speci-mens examined are from the Philippines. Several sample


sizes are greater than those observed in the descriptiondue to the examination of subadult specimens which wereexcluded from morphometric analyses.

Lepidodactylus labialis (10): MINDANAO ISLAND,Agusan del Norte Province, Municipality of Cabadbaran(CAS 133317, 133318, 133329, 133396, 133687, 133790);Davao del Sur Province, Municipality of Malalag (CAS124813, 139714–16).

Pseudogekko brevipes (9): BOHOL ISLAND, BoholProvince, Municipality of Sierra Bullones (CAS 131855,CAS-SU 24596, 25108); NEGROS ISLAND, NegrosOriental Province, Municipality of Sibulan (CAS 128956,128959, 128963, 128971); Municipality of Valencia,Barangay Bongbong (KU 302818, 327770).

Pseudogekko chavacano sp. nov. (4): MINDANAOISLAND, Zamboanga City Province, Municipality ofZamboanga City (Holotype, PNM 9812, formerly KU314963), (Paratype, KU 314964); Zamboanga del NorteProvince, Cuot Creek (CAS-SU 23548, 23549).

Pseudogekko compresicorpus (9): POLILLO IS-LAND, Quezon Province, Municipality of Polillo (KU326242); LUZON ISLAND, Cagayan Province, Munici-pality of Gonzaga (KU 330058); Laguna Province,Municipality of Los Banos, Barangay Batong Maiake(KU 326434, 326436); Barangay Bagong Silang (KU

330735, 331657); Quezon Province, Municipality ofInfanta, Barangay Magsaysay, Barangay Kipagringau(KU 334017); MASBATE ISLAND, Masbate Province,Municipality of Mobo (CAS 141560); TABLAS ISLAND,Romblon Province, Municipality of San Agustin (CAS139713).

Pseudogekko ditoy sp. nov. (2): LEYTE ISLAND,Leyte Province, Municipality of Baybay, BarangayGabas, Sitio Cienda (Holotype, PNM 9811, formerly KU326437), (Paratype, KU 326438).

Pseudogekko pungkaypinit sp. nov. (6): BOHOLISLAND, Bohol Province, Municipality of Sierra Bul-lones, Barrio Dusita (Paratypes, CAS 131854, CAS-SU23655); Raja Sikatuna Natural Park (Paratype, KU324426); LEYTE ISLAND, Leyte Province, Municipalityof Baybay (Paratype, KU 326243); Barangay Guadalupe(Holotype, PNM 9810, formerly KU 326435); MIND-ANAO ISLAND, Misamis Oriental Province, Municipal-ity of Gingoog City, Barangay Lawaan, Sitio Kibuko, Mt.Lumot (Paratype, KU 334019).

Pseudogekko smaragdinus (35): POLILLO ISLAND,Quezon Province, Municipality of Polillo (KU 302819–31,303995–4002, 307638–47, 326240, 326241, 331721); LU-ZON ISLAND, Camarines del Norte Province, Munici-pality of Labo, Barangay Tulay Na Lupa (KU 313828).


taxonomic revision of the pseudogekko …

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