Brown Et Al 2020 Hungkag2

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Unexpected Discovery of Another New Species of Philippine False Gecko

(Gekkonidae; Pseudogekko) from the Bicol Peninsula of Luzon Island
Article in Herpetologica · September 2020

DOI: 10.1655/Herpetologica-D-19-00029.1
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Unexpected Discovery of Another New Species of
Philippine False Gecko (Gekkonidae; Pseudogekko)
from the Bicol Peninsula of Luzon Island
Authors: Brown, Rafe M., Meneses, Camila G., Wood, Perry L.,
Fernandez, Jason B., Cuesta, Michael A., et al.
Source: Herpetologica, 76(3) : 315-329
Published By: The Herpetologists' League
URL: https://doi.org/10.1655/Herpetologica-D-19-00029.1
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Herpetologica, 76(3), 2020, 315–329
Ó 2020 by The Herpetologists’ League, Inc.
Unexpected Discovery of Another New Species of Philippine False Gecko (Gekkonidae;

Pseudogekko) from the Bicol Peninsula of Luzon Island
RAFE M. BROWN1,7, CAMILA G. MENESES2, PERRY L. WOOD, JR.1,6, JASON B. FERNANDEZ3, MICHAEL A. CUESTA4, MICHAEL A.
CLORES4, CLAIRE TRACY1,6, MATTHEW D. BUEHLER1,6, AND CAMERON D. SILER5
1
Biodiversity Institute and Department of Ecology and Evolutionary Biology; University of Kansas, 1345 Jayhawk Boulevard, Lawrence, KS 66045, USA
2
Animal Biology Division, Graduate School, University of the Philippines at Los Baños, College, Laguna 4031, Philippines
3
Phase 5, Block 3, Lot 14 Marycris Complex, Malagasang 2-G Imus Cavite, Philippines
4
Graduate School, Ateneo de Naga University, Ateneo Avenue, Bagumbayan Sur, 4400 Naga City, Philippines
5
Sam Noble Oklahoma Museum of Natural History and Department of Biology, University of Oklahoma, 2401 Chautauqua Avenue, Norman, OK 73072,
USA
ABSTRACT: Philippine False Geckos (genus Pseudogekko) are secretive, delicate, slender-bodied, arboreal members of an obligate forest
specialist clade that is substantially more species diverse than previously assumed. Over the last century, few species were added to this Philippine
endemic genus. During the last decade, however, revisionary studies have resulted in the recognition of six new species. Several of these appear to
be rare, have restricted geographic ranges, or exhibit patchy, fragmented areas of occurrence. In this study we report on the discovery of a second
Luzon Island species in the P. brevipes complex, a clade in which members typically have diminutive bodies. Although the new species is readily
diagnosed from all congeners, we estimated its systematic affinities with a phylogenetic analysis of DNA sequence data and confirm that it is
nested within the P. brevipes clade. The new species constitutes an exception to the general appearance of other members of the P. brevipes
complex, in that it has a relatively heavy-bodied, robust stature, separating it phenotypically from all members of the group. Our new species
constitutes the second Luzon lineage in this group of rainforest species (considered previously to be restricted to the Negros-Panay and Mindanao
Pleistocene aggregate island complexes [PAICs] in the central and southern landmasses of the archipelago). Given the lack of available
biodiversity information for the major remaining forests of the Bicol Peninsula, which necessarily come from targeted faunal surveys, the new
species’ conservation status cannot yet be assessed. In light of the highly fragmented nature of forested habitats of southern Luzon, we suspect the
new species might be vulnerable to extinction as a result of habitat loss.
Key words: Biodiversity; Caramoan Peninsula; Endemism; Faunal subregions; Forest Geckos; Obligate forest species; Patchy distributions;
Rare species; Sorsogon Province
W ITH NEARLY 60 described species, the Philippine specialists of the genera Hemiphyllodactylus (2–4 endemics,
archipelago supports a diverse, yet incompletely understood 1 nonendemic widespread native species; Grismer et al.
gekkonid lizard fauna. Nearly 85% of Philippine species are 2013) and Lepidodactylus (7–10 endemics, 1 nonendemic
endemic to the archipelago (Brown and Alcala 1970, 1978; widespread native species; Brown and Alcala 1978; Siler et
Siler et al. 2017) and many are so poorly known that al. 2014a; Oliver et al. 2018). A final set of taxa are the flap-
systematists have waited decades for the accumulation of legged geckos, or wolf geckos of the genus Luperosaurus
specimens necessary to describe these insufficiently docu- (eight endemics; Brown and Diesmos 2000; Brown et al.
mented new species (Brown et al. 2000, 2009; Grismer et al. 2007, 2010, 2011, 2012b; Gaulke et al. 2007) and false geckos
2013; Siler et al. 2014a,b, 2016). or Philippine forest geckos of the genus Pseudogekko (nine
Nine gecko genera are recognized in the island nation, but endemics; Taylor 1922; Siler et al. 2014a,b, 2016, 2017;
the genus Pseudogekko is the archipelago’s only endemic Davis et al. 2015a). Recent phylogenetic analyses (Brown et
gekkonid genus (Taylor 1922; Brown and Alcala 1978; Brown al. 2012b; Oliver et al. 2018) have made it clear that among
et al. 2011; Grismer 2011), and is nested in the widespread these last three genera, only Pseudogekko is monophyletic—
southwestern Pacific clade traditionally referred to as albeit deeply nested, along with various subclades of
Lepidodactylus, sensu lato (Heinicke et al. 2012; Oliver et Luperosaurus—within Lepidodactylus (Oliver et al. 2018).
al. 2018). Other native Philippine genera include house Clearly, a comprehensive, phylogeny-based review of
geckos of the genera Gehyra (1 species), and Hemidactylus generic boundaries constitutes a priority for ongoing
(1 endemic species, 4 additional nonendemic but native phylogenomic studies (Brown et al. 2012b; Heinicke et al.
taxa), and large bodied, primarily habitat generalists in the 2012; Oliver et al. 2018).
genera Cyrtodactylus (9 endemic species; Welton et al. 2009, Because of their secretive behavior, rarity, highly special-
2010a,b), Gekko (12 endemics, 2 additional nonendemic but ized microhabitat preferences, and/or cryptic coloration and
native taxa; Brown and Alcala 1978; Rösler et al. 2006; morphology, most species of Pseudogekko are encountered
Brown et al. 2008, 2009, 2011; Linkem et al. 2010; Siler et al. seldomly and, subsequently, are rare in collections (Brown
2012), and Ptychozoon (1 endemic; Brown et al. 1997, and Alcala 1978; Brown et al. 2012c; Siler et al. 2014a,b).
2012a). Another group is represented by the minute, poorly However, despite the paucity of properly vouchered genetic
understood, and imperiled coastal forest and mangrove material and confusion surrounding allocation of taxa to
genera (Brown and Tanner 1949; Brown 1964; Brown and
6
PRESENT ADDRESS: Department of Biological Sciences & Museum of
Alcala 1978; Siler et al. 2014a), a series of recent studies have
Natural History, Auburn University, Auburn, AL 36849, USA. identified unrecognized species diversity in both the P.
7
CORRESPONDENCE: e-mail, [email protected] compresicorpus and P. brevipes species complexes (Siler et
315
316 Herpetologica 76(3), 2020
al. 2014a, 2017; Davis et al. 2015a,b). Although four distinct,

allopatric species were identified masquerading in the
former (Siler et al. 2014b, 2016), and two species have
likewise been identified in the latter (Davis et al. 2015a; Siler
et al. 2017), a number of additional, genetically divergent
lineages have been identified but currently cannot be
taxonomically evaluated because of a paucity of specimens
and/or genetic material (Siler et al. 2014b, 2016, 2017).
These are represented by single specimens, corresponding to
suspected undescribed species for which taxonomic descrip-
tions have been held in abeyance pending the collection of
voucher specimens sufficient for taxonomic study (Brown et
al. 2013a; Siler et al. 2014a,b).
During recent faunal surveys in Sorsogon (extreme
southern Bicol Peninsula) and Camarines Sur (central Bicol,
Caramoan Peninsula) provinces of Luzon (Fig. 1), pheno-
typically distinct specimens of Pseudogekko were collected at
low elevations, in fragments of lower montane forest and
beach coastal forest (Fig. 2), and matching the distinctive
appearance of specimens first observed and photographed
by JBF years before in the Pocdol Mountain, of the Bacon-
Manito Mountain Range, Albay Province, Bicol Peninsula
(Fig. 3). Here, we justify recognition of one new taxon, and
discuss the identification of a second, closely related,
probable new species, represented currently by only a single
immature individual (a population requiring further study).
We diagnose the new species based on phenotypic traits, and
we utilize mitochondrial DNA sequence data to infer its
phylogenetic affinities. Our phylogeny identifies relevant
comparisons for diagnosis, and genetic divergence estimates
provide crude approximations of the degree of genetic
distinctiveness among the new species and closely related
congeners of the P. brevipes complex—all of which sets the
stage for future, in-depth studies of Pseudogekko across FIG. 1.—Map of the Philippines, with sampling localities of Pseudogekko
Luzon’s Bicol Peninsula. hungkag and congeners included in phylogenetic analyses (see text for
details; Fig. 8), indicated by various symbols (legend). A color version of this
figure is available online.
MATERIALS AND METHODS
Fieldwork and Specimen Collection RSC extraction robot, targeting the same ND2 mitochondrial
We conducted fieldwork on southern Luzon Island (Fig. protein-coding gene employed previously (Siler et al.
1) every 2–4 mo, from September 2015 through August 2014a,b, 2016, 2017; Davis et al. 2015a). We successfully
2018. Specimens of Pseudogekko were collected by focusing amplified and sequenced the ND2 gene region from two
search efforts on upper surfaces of leaves of shrubs and samples of the new species from the southernmost tip of
saplings, on Pandanus axils, on thin twigs at distal ends of Luzon’s Bicol Peninsula in Sorsogon Province and a single
understory tree branches and vines, and on inner surfaces of immature individual we tentatively assign to the new species,
tree holes and hollow bamboo cavities (Fig. 2). We from the Caramoan Peninsula of Camarines Sur Province
euthanized specimens with aqueous chloretone, removed (Fig. 1). Primers and protocols for all sequencing efforts
genetic material by dissection (liver, preserved in 100% follow Siler et al. (2014a,b, 2017), and newly collected
laboratory grade ethanol), fixed vouchers in 10% buffered sequence data are deposited at GenBank (Appendix II).
formalin, and transferred specimens to 70% ethanol within 2
mo. For comparative purposes, we examined specimens and/ Alignment and Phylogenetic Analysis
or sequenced genetic material deposited in U.S. and We outsourced PCR products (visualized on 1.0% agarose
Philippine biodiversity repositories (Appendix I); institution- gels) to GENEWIZt for purification, cycle sequencing, and
al abbreviations follow Sabaj (2016). sequence determination. We sequenced target regions in
both directions and confirmed congruence in Geneious vR11
DNA Sequence Data (available at https://www.geneious.com/; Biomatters, Inc.,
Previously published Pseudogekko sequences for the San Diego, CA, USA). The ND2 protein-coding region and
mitochondrial NADH dehydrogenase subunit 2 (ND2) are the three tRNAs were aligned using the MAFTT v7.017
available on GenBank (Appendix I; see Siler et al. 2014a, (Katoh and Standley 2013) plugin under GeneiousTM
2017). We collected homologous data by extracting total defaults and we used Mesquite v3.04 (Maddison and
genomic DNA from newly collected genetic material with a Maddison 2015) to establish the correct amino acid protein
Maxwellt RSC Tissue DNA kit and a Promega Maxwellt coding reading frame by confirming the absence of stop

BROWN ET AL.—A NEW SPECIES OF PSEUDOGEKKO 317
FIG. 2.—Luzon Island habitat variation of Pseudogekko hungkag: (A) exterior appearance of sandy, coastal forests, extending upwards to limestone karst
towers in the vicinity of Tayak Lagoon, Municipality of Caramoan, Camarines Sur Province; (B) interior appearance of low-elevation, second-growth
limestone foothills of Mt. Jormahan, Sorsogon Province, Luzon Island; and (C) interior forest habitat characteristics of sandy, coastal forests, Municipality of
Caramoan, where P. cf. hungkag (KU 349299) was collected from vines in the upper right corner of this figure. A color version of this figure is available
online.
codons. We partitioned ND2 by codon positions and our resulting log file with Tracer v1.5 (Drummond et al.
included all tRNAs as a fourth partition. The best-fit models 2012), and ensured that all parameter effective samples sizes
of molecular evolution were estimated in IQ-TREE (Nguyen (ESS) reached stationarity (ESS values 200). Finally, we
et al. 2015; Trifinopoulos et al. 2016) using the Bayesian used TreeAnnotator v2.4.6 (Bouckaert et al. 2014) with a
information criterion (BIC; Table 1). We calculated simple 25% burn in to generate a maximum clade credibility tree
pairwise sequence divergence estimates (Table 2) in MEGA including mean heights and considered all posterior
v7.0.26 (Kumar et al. 2016). probabilities (PP) of 0.95 as strongly supported (Hulsen-
Phylogenetic analyses were performed in IQ-TREE beck and Ronquist 2001; Wilcox et al. 2002).
(Nguyen et al. 2015; Trifinopoulos et al. 2016) and nodal
support was assessed with 1,000 bootstrap pseudoreplicates, Phenotypic Data
using the ultrafast bootstrap approximation algorithm (Minh We examined preserved specimens for color pattern and
et al. 2013). All nodes with ultrafast bootstrap values (UFBs) scored mensural (continuously varying measurements) and
of 95 were considered significantly supported (Minh et al. meristic data (scale and color pattern character counts),
2013). We performed Bayesian analyses in BEAST v2.4.6 using the same character definitions and methods from
(Bouckaert et al. 2014) to estimate phylogeny while earlier studies (Siler et al. 2014b, 2016, 2017; Davis et al.
simultaneously exploring and estimating substitution models 2015a,b). Sex was determined by eversion of hemipenes in
and parameters in bModelTest (Bouckeart and Drummond the field in males, by confirmation of presence of oviductal
2017). Bayesian analyses were implemented for 50 million eggs in females (visible through the ventral body wall; clutch
generations, sampling every 5,000 generations. We visualized size invariably n ¼ 2), or by gonadal inspection to control for

TABLE 1.—Best-fit models for evolution of clades within Pseudogekko

inferred by the Bayesian information criterion from the IQ-TREE-
webserver (Trifinopoulos et al. 2016) and by model averaging from
bMODELTEST applied in BEAST v2.4.6 (Bouckaert et al. 2014), and
applied as part of a partitioned, model-based phylogenetic analyses of our
DNA sequence data set.
Model selected Model selected

Partition IQ-TREE bMODELTEST
ND2, first codon position GTRþFþIþC4 123124

ND2, second codon position TPM3uþFþIþC4 123123
ND2, third codon position TIM2þFþC4 121343
tRNAs (concatenated) TIM2þFþIþC4 121321
ventrals. We used Köhler’s (2012) standardized color codes

for description of the color pattern of the new species.
Species Concept
In taxonomic studies of Philippine lizards (Brown et al.
2007, 2008, 2009, 2010; Siler et al. 2014b, 2016, 2017), we
have consistently applied the general lineage concept (de
Queiroz 1998, 1999) as the modern extension of the
evolutionary species concept (Simpson 1961; Wiley 1978;
Frost and Hillis 1990; de Queiroz 2005) as a framework for
recognition of lineage-based species. We take an integrative
approach to species delimitation by emphasizing multiple
lines of evidence, with the potential to inform on the
independence and cohesion of ancestor–descendant, or
FIG. 3.—The first photograph of an uncollected Pseudogekko hungkag of temporal, series of populations (de Queiroz 2005).
unknown sex, observed on Pocdol Mountain, Bacon-Manito Mountain We assess putative new species and congeners for the
Range, Albay Province (photo by JBF, July 2013): note uniquely possessed presence of (1) fixed or discretely varying (diagnostic)
diagnostic traits: bright golden iris, and inverted Y-shaped markings from the
pelvic girdle, through the entire length of the tail. A color version of this phenoytypic character state differences—nonoverlapping
figure is available online. ranges of character states; (2) discontinuous ranges of size-
related continuously varying dimensions; (3) presence or
occasional instances of ambiguity in Pseudogekko secondary absence of fixed color pattern elements; (4) phylogenetic
position with respect to phenotypically similar and/or
sexual characteristics (Davis et al. 2015b). We took
sympatric congeners; and (5) ecological information, micro-
measurements with digital calipers (60.1 mm) and counted
habitat differences, and/or discrete, noncontiguous geo-
scales under a stereo microscope. Character definitions
graphic distributions (allopatric areas of occurrence which,
followed Siler et al. (2014b, 2017) and included snout–vent in the Philippines, typically take the form of separate islands,
length (SVL); tail length, depth, and width; head width and island banks, mountains, or disrupted habitat types such as
absolute and relative head length (measured as snout to isolated geological formations). Although we consider a
posterior edge of tympanum, and posterior tip of mandible); putative species’ phylogenetic affinities to be informative for
absolute and relative snout length (Siler et al. 2014b); identifying relevant comparisons (Brown et al. 2008, 2009;
midbody trunk width and eye diameter; eye–nares, inter- Welton et al. 2010a) and/or potentially refuting the
narial, interorbital, and axilla–groin distances; femur and hypothesis of conspecificity in nonsister taxa (Brown 2015;
tibia lengths; and numbers of supralabials, infralabials, Brown et al. 2010, 2016), we use genetic distances as a crude
circumorbitals, precloacal pore-bearing scales, Finger-III comparative indication of relative divergence only, and not
and Toe-IV scansors paravertebrals, midbody scale rows, and as justification for species delimitation.
TABLE 2.—Mean uncorrected pairwise sequence divergence (%) for mitochondrial data (ND2, below the bolded diagonal) between species pairs; within-
species interpopulational mean p distances, in eight named Pseudogekko taxa for which genetic data are available (no genetic materials currently are available
for P. isapa) and one putative species with insufficient specimens for species description (P. cf. hungkag, Caramoan) are presented in bold along the diagonal.
hungkag cf. hungkag atiorum brevipes chavacano compresicorpus ditoy pungkaypinit smaragdinus
hungkag 0.000
cf. hungkag 0.200 N/A
atiorum 0.182 0.327 0.000
brevipes 0.182 0.273 0.145 N/A
chavacano 0.200 0.345 0.236 0.255 0.000
compresicorpus 0.175 0.289 0.240 0.244 0.171 0.105
ditoy 0.200 0.345 0.255 0.255 0.127 0.198 0.000
pungkaypinit 0.207 0.324 0.236 0.211 0.207 0.153 0.193 0.072
smaragdinus 0.114 0.273 0.218 0.198 0.220 0.179 0.202 0.169 0.009

FIG. 5.—Dorsal (A) and ventral (B) habitus of adult female paratype
Pseudogekko hungkag sp. nov. (KU 346539) before preservation, from Mt.
Jormahan (type locality), Municipality of Irosin, Sorsogon Province, Bicol
Peninsula, Luzon Island. Scale bar ¼ 5 mm. Photos: JBF and RMB. A color
version of this figure is available online.
FIG. 4.—Photographs in life of Pseudogekko hungkag sp. nov. (A, B) adult

female holotype (PNM 9864, formerly KU 346540) from Mt. Jormahan
(type locality), Municipality of Irosin, Sorsogon Province, Bicol Peninsula,
Luzon Island. (C) juvenile male specimen, tentatively referred to the new
species, from Municipality of Caramoan, Camarines Sur Province, Bicol
Peninsula, Luzon Island (KU 349299). Scale bars ¼ 5 mm. Photos: JBF and
RMB.
TAXONOMIC ACCOUNT
Pseudogekko hungkag sp. nov.
(Figs. 2–6)
Holotype.—PNM 9864 (RMB Field No. 24085, formerly
KU 346540), adult female, collected between 2200 and 2300
h on 5 August 2017, in regenerating secondary forest, near
the peak of Mt. Jormahan, Barangay Cogon, Municipality of
Irosin, Sorsogon Province, Luzon Island, Philippines
(12.761168N, 124.003568E; in all cases, datum ¼ WGS84;
643 m above sea level [a.s.l.]), by CGM and E. Bondal.
Paratypes.—KU 346539, 346542 (RMB 24082, 24090),
two adult females, PNM 9865 (RMB 24081, formerly KU
346538), KU 346541 (RMB 24087; Fig. 4), two adult males, FIG. 6.—Palmar view of right hand (A), plantar view of right foot (B) and
collected with the holotype, between 2200 and 2300 h, details of differentiated precloacal pore-bearing scale series, in male
Pseudogekko hungkag paratype (PNM 9865; RMB Field No. 24081,
within 500 m of the site of holotype collection, but at slightly formerly KU 346538). Scale bars ¼ 1 mm. A color version of this figure is
lower elevations (500–640 m a.s.l.). available online.

Referred specimen.—KU 349299 (RMB 24361; Fig.

4C), immature male, collected at 1915 h on 22 November
2017, in coastal beach forest scrub and mixed secondary
forest at Tayak Lagoon, Barangay Pandanan, Municipality of
Caramoan, Camarines Sur Province, Luzon Island, Philip-
pines (13.79028N, 123.92738E; 2 m a.s.l.), by RMB, JFB, and
MAC.
Diagnosis.—Pseudogekko hungkag is diagnosed from all
congeners by the following combination of characters: (1)
body moderate, robust (all measurements in millimeters;
males ¼ SVL 42.0–45.5; females ¼ 42.0–52.1); (2) axilla–
groin distance relatively short (males ¼ 21.7–24.4; females ¼
24.7–27.3); (3) head (males ¼ 11.3–12.1; females ¼ 11.8–
13.8) and snout (males ¼ 4.6–5.2; females ¼ 4.9–5.7) short;
(4) Finger-III scansors 11–13; (5) Toe-IV scansors 13–15; (6)
supralabials 11–14; (7) infralabials 11–14; (8) circumorbitals
34–40; (9) paravertebrals 222–236; (10) ventrals 85–94; (11)
enlarged precloacal pore-bearing scale series 13–14; (12)
femorals undifferentiated; (13) postcloacals enlarged, in
three prominent rows; (14) mental and postmentals small,
undifferentiated (the latter juxtaposed); (15) transverse tail
bands absent (adults, Sorsogon), or present (juvenile,
Caramoan); (16) light, inverted Y-shaped caudal chevrons
present for full length of tail (n ¼ 8–10 in adults from
Sorsogon; Fig. 4A,B), or limited to tail base (n ¼ 3 for the FIG. 7.—Lateral views of live specimens, representing typical members of
juvenile from Caramoan; Fig. 4C); (17) iris bright silver to the (A) Pseudogekko brevipes complex (adult female paratype of P. sumiklab
gold, ring absent (Fig. 4A); (18) superciliaries not brightly [KU 343847], from Sorsogon Province, southern Bicol Peninsula, Luzon
Island) and (B) the P. compresicorpus complex (adult male P. compresi-
colored. corpus RMB 24978 [deposited at KU] from Quezon Province, southern
Comparisons.—Pseudogekko hungkag is most closely Luzon Island). Scale bars ¼ 10 mm. Photos: JBF and RMB. A color version
related to the members of the P. brevipes complex (Fig. 7), of this figure is available online.
so we focus primarily on comparisons to P. atiorum, P.
brevipes, and P. sumiklab for the purpose of diagnosis (in all further diagnosed from the distantly allopatric Mindanao
comparisons, values for P. hungkag are presented first). Island species, P. chavacano, by its smaller body (SVL 42.0–
Pseudogekko hungkag can be distinguished from P. atiorum 52.1 vs. 54.7–55.9 mm), fewer Finger-III scansors (11–12 vs.
by having fewer supralabials (11–14 vs. 15–17) and ventrals 15–16), Toe-IV scansors (13–15 vs. 17–20), supralabials (11–
(85–94 vs. 119–129), and by the absence (vs. presence) of a 12 vs. 15–16), infralabials (11–14 vs. 16–17), circumorbitals
light interorbital band, presence (vs. indistinct or absent) of (36–40 vs. 46), ventrals (85–94 vs. 122–123), and precloacals
bold, light inverted Y-shaped caudal chevrons, and presence (13–14 vs. 16). Additionally, the new species has more
(vs. absence) of cream to gold postorbital and labial spots. paravertebrals (222–235 vs. 195–197), and is further
From P. brevipes the new species differs by having fewer distinguished by the absence (vs. presence) of neon green
infralabials (11–14 vs. 14–15) and ventrals (85–94 vs. 96–
limb spots, and presence of bold inverted Y-shaped caudal
117), more circumorbitals (36–40 vs. 33–35) and precloacals
chevrons (vs. simple transverse caudal bands). From P.
(13–14 vs. 12), and by the absence (vs. presence) of a light
compresicorpus (Fig. 7B), the new species differs by its
interorbital band, and presence (vs. indistinct or absent) of
smaller body (SVL 42.0–52.1 vs. 54.9–59.7 mm), fewer
bold inverted Y-shaped caudal chevrons. From P. sumiklab
the new species differs by having fewer ventrals (85–94 vs. Finger-III scansors (11–12 vs. 15–17), Toe-IV scansors (13–
106–109), and by the presence (vs. absence) of bold inverted 15 vs. 18–19) supralabials (11–12 vs. 16–17), ventrals (85–94
Y-shaped caudal chevrons. vs. 127–130), and by the absence of bright iris ring coloration
Although only body size (SVL) discretely differed in (vs. presence, light blue) and presence (vs. absence) of bold
comparisons between P. hungkag (males ¼ 42.0–45.5; inverted Y-shaped caudal chevrons (Tables 3 and 4).
females ¼ 45–52.1) and P. brevipes (male ¼ 39.0; females Pseudogekko hungkag is further diagnosed from the
¼ 34.5–42.4), the new species generally is a heavy-bodied allopatric Mindanao, Samar, and Leyte islands species P.
species, differing from P. atiorum, P. brevipes, and P. ditoy and the Sibuyan Island endemic P. isapa by its smaller
sumiklab by its robust (vs. slender, elongate) body shape female body size (SVL 42.0–52.1 vs. 52.6 mm [P. ditoy], 63.4
(Fig. 4A,B vs. 7A), and differs further from P. sumiklab and [P. isapa]), axilla–groin distance (21.7–27.3 vs. 32.1–33.0 [P.
P. brevipes by its relatively larger head length, and from P. isapa]), head length (11.3–13.3 vs. 9.3–9.6 [P. ditoy]), and
sumiklab by its relatively larger head width and snout length fewer Finger-III scansors (11–13 vs. 14–15 [P. ditoy]), Toe-
(Tables 3 and 4). IV scansors (13–15 vs. 16–17 [P. ditoy], 17 [P. isapa]),
From all members of the P. compresicorpus complex (Fig. supralabials (11, 12 vs. 17, 20 [P. ditoy], 20, 21 [P. isapa]),
7B), P. hungkag can be diagnosed by its relatively larger infralabials (11–14 vs. 16–17 [P. ditoy], 17–19 [P. isapa]),
head width, and from P. punkaypinit and P. smaragdinus by ventrals (85–90 vs. 111–118 [P. ditoy], 135–141 [P. isapa]),
its relatively larger head length (Table 4). The new species is circumorbitals (36–40 vs. 40–52 [P. ditoy], 50–54 [P. isapa]),

TABLE 3.—Distribution of selective diagnostic meristic character states and color pattern element characters (þ present, absent) for all known species of Pseudogekko (adults only; m ¼ male; f ¼ female;
Appendix I). In cases of scale count variation within species, numbers of specimens exhibiting each state are provided in parentheses.
brevipes chavacano compresicorpus pungkaypinit smaragdinus

Character hungkag (2 m, 3 f) sumiklab (3 f) atiorum (8 m, 7 f) (1 m, 3 f) (1 m, 1 f) (3 m, 4 f) ditoy (1 m, 1 f) isapa (1 m, 1 f) (4 m, 2 f) (16 m, 17 f)
Finger-III scansors 11 (1) 12 (5) 13 (2) 11 (2) 12 (1) 12 (3) 13 (9) 12 (4) 15 (1) 16 (1) 15 (4) 16 (2) 17 (1) 14 (1) 15 (1) 13 (2) 15 (3) 16 (1) 17 (2) 15 (1) 16 (4) 17 (9)
14 (3) 18 (19)
Toe-IV scansors 13 (2) 14 (6) 15 (3) 13 (2) 14 (1) 14 (1) 15 (2) 15 (4) 17 (1) 20 (1) 18 (5) 19 (2) 16 (1) 17 (1) 17 (2) 17 (1) 18 (2) 19 (2) 16 (1) 18 (6) 19 (8)
16 (2) 21 (1) 20 (12) 21 (5) 22 (1)
17 (10)
Supralabials 11 (2) 12 (3) 13 (4) 14 (3) 15 (7) 17 (4) 13 (2) 14 (2) 15 (1) 16 (1) 16 (1) 17 (1) 18 (3) 17 (1) 20 (1) 20 (1) 21 (1) 16 (1) 18 (2) 19 (2) 16 (8) 17 (10) 18 (11)
14 (2) 19 (1) 20 (1) 20 (1) 19 (4)
Infralabials 11 (3) 12 (1) 13 (4) 13 (1) 14 (2) 12 (1) 13 (4) 14 (1) 15 (3) 16 (1) 17 (1) 13 (1) 15 (2) 16 (4) 16 (1) 17 (1) 17 (1) 19 (1) 17 (4) 18 (1) 19 (1) 14 (6) 15 (12) 16 (10)
14 (1) 14 (9) 17 (5)
15 (1)
Circumorbitals 36 (1) 37 (1) 38 (2) 39 (1) 44 (1) 35–38 33–35 46 39–45 40, 43 50 (1) 54 (1) 50–55 33–35

40 (2) 46 (1)
BROWN
Paravertebrals 222–235 220–224 226–240 211–218 195–197 226–234 180, 185 240–246 265–280 241–252
Ventrals 85–94 106–109 119–129 96–117 122–123 127–130 111, 118 135–141 125–155 124–130
Precloacals 13–14 13–14 13–15 12 16 10–14 18 15 17–20 32–41

Femorals þ
ET AL.—A
Mental enlarged þ þ –
Dominant body Gray to gray-brown Medium Dark brown Unknown Light brown Dark brown to tan Light brown Brownish tan Grayish brown Bright neon yellow to
coloration brown to orange (undisturbed)
tan to neon green
(disturbed)
NEW SPECIES
Cephalic spots þ, sparse, þ, sparse, Unknown þ, dense, þ, dense, neon þ þ, dense black, sparse
OF
black and cream neon green green white

cream
Light postorbital þ, cream to gold þ,
markings
Dorsolateral spots þ, cream þ, cream þ, cream þ þ, neon þ, faint, neon green þ, large black, small
PSEUDOGEKKO
green white
Vertebral spots or þ, black spots þ, black spots
bars
Limb spots þ, dense, þ, faint, neon green þ, sparse black and
neon green white
Transverse caudal þ or , inverted þ Unknown þ, neon yellow, white,
bands/markings Y-shaped1 and neon orange
Black subcaudal _ þ or _ _ _ _ _ _ _
stripe
Body (trunk) þ, lateral
stripes
Interorbital band þ, light brown þ þ
Iris ring coloration Unknown þ, light blue
Iris coloration Silver to Gold Silver Silver Silver Yellow Tan Gray Light gray to Yellow Yellow
silver
1
Light, inverted Y-shaped caudal marking repeated (n ¼ 8–10) for full tail length in adults from Sorsogon, or limited (n ¼ 3) to tail base in juvenile, from Caramoan (replaced distally by alternating light-and-dark transverse bands).
321
TABLE 4.—Summary of selective mensural characters for adult specimens (m ¼ male; f ¼ female; Appendix I; immature specimens excluded) of all known species of Pseudogekko. General geographical
125.3–141.2 (135 6 8.6) 103.6–129.7 (119.4 6 6.4)

precloacals (13–14 vs. 18 [P. ditoy], 15 [P. isapa]),
54.2–64.3 (59.4 6 2.5)

50.2–61.7 (57.4 6 3.5)
26.2–35.5 (31.5 6 2.2)
14.9–17.4 (16.3 6 0.6)
13.8–16.0 (14.8 6 0.6)
50.7–67.9 (60.3 6 2.9)

smaragdinus (16 m, 17 f)
7.8–10.7 (9.5 6 0.7)
7.1–9.8 (8.6 6 0.6)
4.5–6.5 (5.7 6 0.4)

paravertebrals (222–235 vs. 240–246 [P. ditoy], 240–246 [P.
Polillo Island and

Bicol Peninsula
(Luzon Island)
isapa]), and by the presence (vs. absence in both species) of
bold inverted Y-shaped caudal chevrons (Fig. 4A,B).
Pseudogekko hungkag is further diagnosed from P. pungkay-
pinit by its smaller body size (SVL 42.0–52.1 vs. 75.2–75.3),
axilla–groin distance (21.7–27.3 vs. 37.2–41.2), snout length
66.6–76.8 (71.8 6 5.1)
(4.6–5.7 vs. 6.7–7.5), fewer Finger-III scansors (11–13 vs.
37.2–41.2 (39.6 6 1.8)
11.4–13.6 (12.6 6 0.9)

16.0–18.3 (17.2 6 0.8)
9.3–11.2 (10.6 6 0.9)
14.0–15.2 (14.7 6 0.4)
53.6–68.4 (60.0 6 4.8)

6.7–7.5 (7.2 6 0.4)
15–17), Toe-IV scansors (13–15 vs. 17–21), supralabials (11–
pungkaypinit (4 m, 2 f)
Mindanao, Samar,
and Leyte islands
14 vs. 16–20), infralabials (11–14 vs. 17–19), circumorbitals

(36–40 vs. 50–55), paravertebrals (222–235 vs. 265–280),
75.2–75.3
ventrals (88–94 vs. 125–155), precloacals (13, 14 vs. 17–20),

and by the presence (vs. absence) of bold inverted Y-shaped
caudal chevrons.
Pseudogekko hungkag is diagnosed from P. smaragdinus
32.1–33.0
11.7–11.8
18.6–18.9
15.2–16.0
61.9–64.1
7.4–10.1
(1 m, 1 f)
Sibuyan
7.2–7.5
Island
by its smaller body size (SVL 42.0–52.1 vs. 50.2–64.3), fewer

isapa
N/A
63.4
62.1
Finger-III scansors (11–13 vs. 15–18), Toe-IV scansors (13–

15 vs. 16–22), supralabials (11–14 vs. 16–19), infralabials
ditoy (1 m, 1 f)
Leyte islands
Samar and
25.1, 29.7
18.2, 18.9
14.7, 15.9
57.4, 59.3
(11–14 vs. 14–19), paravertebrals (222–235 vs. 241–252),

9.3, 9.6
7.7, 7.9
5.4, 5.7
N/A
ventrals (88–94 vs. 124–130), and precloacals (13, 14 vs. 32–

49.4
52.6
41 precloacofemorals), by its overall gray ground (vs. bright

neon yellow to orange [undisturbed] to neon green
105.9–117.3 (111.7 6 5.7)
55.9–58.8 (57.6 6 1.5)
54.9–59.7 (57.1 6 2.6)
27.0–32.6 (30.4 6 1.8)
16.1–18.1 (17.1 6 0.7)
13.0–16.3 (14.2 6 1.0)
56.1–64.6 (59.6 6 3.0)

compresicorpus (3 m, 5 f)
[disturbed]) coloration, and by the presence of bold inverted

9.2–11.3 (9.7 6 0.3)
7.5–10.1 (8.2 6 0.8)
5.3–7.4 (5.9 6 0.7)
Y-shaped caudal chevrons (vs. transverse dark/light caudal

Polillo islands
Luzon and
bands; Tables 3 and 4).

Description of holotype.—Adult female in excellent
condition; two large eggs visible through slightly transparent
ventrum (similar to paratype in Fig. 4A); small incision in the
sternal region (liver sample preserved separately for genetic
Mindanao
17.8–29.8 (21.7 6 5.5) 26.7–30.0
18.6–19.1
14.3–15.2
55.2–58.0
material). Body relatively large, robust, SVL 52.1 mm; limbs

chavacano
(1 m, 1 f)
7.8–8.5
5.8–6.0
Island
well developed, robust, relatively muscular; tail long, intact,

55.9
34.5–42.4 (38.6 6 4.0) 54.7
95.8
10.4
60.5 mm, tapering to slender tip, thickened at laterally

(77.3 6 7.3)
(19.2 6 3.6)
(15.2 6 1.2)
(56.5 6 8.3)
compressed base; margins of limbs smooth; cutaneous flaps,

(7.4 6 1.5)
(5.9 6 0.5)
(4.1 6 0.2)
and Leyte islands
brevipes (1 m, 3 f)
ornate tuberculation, and dermal folds absent.

Bohol, Samar,
distribution included for reference; table entries presented as range (mean 6 1 SD) in mm.
Head size moderate, well differentiated from neck, with

39.0
moderately hypertrophied temporal and adductor muscula-

72.0–87.5
6.2–9.5
16.7–24.4
5.6–6.5
14.2–16.8
3.8–4.4
44.4–62.3
ture; calcium deposits of endolympthatic sacs visible as

moderate swellings in lateral nuchal region; snout broadly
rounded in dorsal view, bluntly rounded in lateral view; head
(97.3 6 13.0)
41.1–52.5 (47.0 6 4.5)
44.8–48.7 (46.3 6 1.5)
19.4–28.4 (24.8 6 2.4)
(18.3 6 0.9)
(15.3 6 0.6)
(59.5 6 3.7)
width 110% midbody width, 84.9% head length; head length

(8.5 6 0.6)
(7.1 6 0.7)
(5.1 6 0.4)
atiorum (8 m, 7 f)
Panay islands
20.0% SVL; snout length 63.3% head width, 53.8% head

Negros and
length; dorsal surfaces of head smooth and homogeneous,

with only slightly detectable concavities in postnasal,
7.4–9.4
5.9–8.3
4.3–5.8
80.2–113.5
16.0–19.8
14.3–16.4
55.1–69.1
internasal, prefrontal, and interorbital regions; auricular

opening small, subcircular, its position posteroventral to
temporal swellings on either side of head; tympanum deeply
Bicol Peninsula
(Luzon Island)
(44.3 6 2.4)
sunken (Fig. 3A,B); orbit and eye large, pupil vertical,

sumiklab (3 f)
23.6, 25.8
91.0
12.4
14.4
69.0
41.8–46.6
5.8
6.7
4.0
margin wavy; limbs relatively long, femur 8.8 mm, tibia 7.5
–
5.8,
5.7,
4.0,
87.8,
14.0,
13.6,
69.2,
mm; hands and feet moderate, digits robust, unreduced;

hindlimbs more robust and thicker than forelimbs; tibia
length 14.4% SVL, 85.2% femur length.
45–52.1 (47.2 6 4.5)
21.7–27.3 (24.6 6 1.9)
Total body length 46.0–60.5 (52.5 6 7.3)
Head length/SVL 218.8–20.2 (26.3 6 1.5)
16.9–17.8 (17.3 6 0.3)
51.4–58.2 (56.1 6 2.2)

7.0–10.6 (9.3 6 1.0)
7.5–9.0 (8.1 6 0.6)
4.6–5.7 (5.2 6 4.4)
Rostral broad, wrapping nearly around total anterior

hungkag (2 m, 3 f)
Bicol Peninsula
(Luzon Island)
surface of rostrum, widely rectangular in anterodorsal view,

42, 45.5
much broader than high; narial opening surrounded by

rostral, first labial, the first (outer) enlarged supranasal, and
two differentiated postnasals (ventral enlarged; dorsal
smaller); snout with a total of four differentiated supranasals,
arranged in two separate pairs, with pairs separated by two
Head width/SVL
Head length
irregular small median scales; circumorbitals 38/39 (L/R).

Snout length/
Snout length
Head length
Head width
Axilla–groin
distance
Total number of differentiated supralabials 12/13 (8–12/

SVL (m)
Character
Locality
SVL (f)
9–13 subocular); row of snout scales bordering supralabials

dorsally undifferentiated, barely enlarged; total number of

differentiated infralabials 14/13 (10–14/9–13 subocular); Coloration of holotype (Fig. 4).—Employing color
rows of enlarged scales ventrally bordering infralabials only codes of Köhler (2012), live coloration (from field notes
extend to fifth labial, before abruptly continuing posteriorly and photographs of JBF and RMB; Fig. 4A,B) consists of
as undifferentiated postrictal scales; mental minute, smaller dorsal ground color on head, trunk, and tail base gray (293,
than first infralabials; followed by five rows of enlarged 296); tail buff (15) to clay (20) with horn (11) or chamois pale
postmentals (chin shields), transitioning abruptly to undif- tan (84) inverted light chevrons. Dorsal surfaces with diffuse
ferentiated ventral head and throat scales at the level of fifth drab gray–brown (19) blotches, and small chestnut to sepia
infralabial; gulars small, homogenous in size, shape, and (30, 40) speckles, devoid of pattern except irregularly
texture, round, juxtaposed to slightly imbricate. aggregated into a series of vertebral blotches (boldest in
Dorsal cephalic scales homogeneous in size, shape, and scapular and pelvic regions), and densely clustered around
texture: nearly flat to slightly convex, round to subcircular; first beige (254) inverted Y-shaped chevron at tail base.
postnasal, prefrontal, internasal, and interorbital depressions Nuchal region with sulphur yellow (80) round, distinct
shallow; undifferentiated posterior head scales granular, nuchal spots, and five (left) or six (right) similar yellow spots
round to sharply rounded or even slightly conical; ornamen- spanning dorsolateral surface of trunk; similar yellow spots
tal cephalic tuberculation, cutaneous flaps, and dermal number two (right) or three (left) in eye–tympanum region.
projections all absent (Figs. 4B and 5B). Labial region straw yellow (53) with bright sulphur yellow
Axilla–groin distance 52.4% SVL; undifferentiated dorsal (80) spots. Limbs colored and patterned as dorsal body
trunk scales homogeneous in size, shape, and texture: round surfaces, with distal segments (forearms, tibia) appearing
to ovoid, granular, convex to pointed, many with raised darker on account of denser aggregation of black (300)
posterior edges forming points, juxtaposed or slightly speckles; dorsal surfaces of digits with alternating fawn/
overlapping; most middorsal trunk scales tightly juxtaposed, purplish-brown (258) and beige (254) transverse bars on
few with surrounding, barely evident five or six interstitial dorsal surfaces, and buff (15) to smoky white (261) plantar
granules; on flanks, scalation consists of more typically surfaces of subdigital scansors.
cycloid convex scales, with more space between scales and Ventral ground color of the head, trunk white (261), with
interstitial granules readily visible; ventrolateral scales suffuse pale creamy yellow (82) to olive clay yellow–brown
transition to imbricate ventrals, transition near complete by (85) around margins (ventrolateral surfaces) of head, limbs,
edges of ventrolateral–ventral body surface; paravertebrals and trunk; tail darkens steadily from base to distal tip:
between midpoints of limb insertions 227; ventrals between sulphur yellow (base: 95) to light yellow–green (100) to olive
midpoints of limb insertions 95; midbody scales 114 (24 of brown (100), and finally black (300) tip.
which differentiated, enlarged, imbricate transverse ventrals) Lateral surfaces of head, trunk, and tail with gradual color
scales on dorsal surfaces of limbs more imbricate than transition between the dorsal ground color of gray (293, 296)
dorsals; scales on dorsal surfaces of hands and feet imbricate, and the ventral base color of smoky white (261). In life, with
similar in size or slightly smaller than dorsal limb scales; continued disturbance, dorsolateral and flank surfaces
ventral body scales relatively homogeneous in size and darkened to dark brown (277) and vertebral blotches became
appearance: flat, cycloid, strongly imbricate, markedly larger darker brown (280) and more distinct.
than lateral or dorsal trunk scales; 14 differentiated Eyes golden yellow (79) with lighter gold to silver outer
(enlarged), dimpled scales in continuous precloacal series margins (80) and vertical black (300) pupils; silvery-gold iris
(homologous with pore-bearing precloacals in males), covered with web-like network of dark brown (280)
arranged in a widely obtuse, inverted V-shaped formation; connected lines.
precloacals preceded anteriorly by one row of slightly In fluid, 6 mo following preservation (see paratype, Fig.
enlarged scales; followed by three rows of enlarged, plate- 4A,B), all yellow, golden, greenish, and/or yellowish-brown
like unpored scales, and two rows of undifferentiated elements are lost, and pattern elements (nuchal spots, labial
ventrals along edge of vent opening. spots, dorsolateral spots, and inverted Y-shaped tail chev-
Digits expanded, with only minute extent of interdigital rons) are difficult to discern. In general, dorsal surfaces are
webbing, and with subdigital surfaces covered by bowed, drab dark brown (45) to dark olive brown (292) and ventral
undivided but notched scansors; subdigital scansors of left/ surfaces are neutral (297) to medium gray (298); ventral
right Finger III 12/12; scansors of Toe IV 13/14; second and surfaces of digits with light gray (296) scansors.
third penultimate scansors deeply notched; all scansors Measurements of holotype (millimeters) and scale
bordered laterally (on palmar and plantar surfaces) by two or counts.—SVL 52.1; tail 60.5; axilla–groin 23.6; head length
three slightly enlarged scales that form a near-continuous 13.8 (to posterior edge of tympanum), 10.6 (to posterior tip
series with enlarged scansors; palmar and plantar scales of mandible); head width 9.0; snout 5.7; eye diameter 3.2;
small, granular, juxtaposed; all digits clawed, but first eye–nares distance 3.4; internarial distance 2.5; interorbital
(medial) claw greatly reduced to form subcircular nail-like distance 2.7; midbody trunk width (horizontal diameter) 8.2;
scale; remaining terminal claw-bearing phalanges com- femur 8.8; tibia 7.5; tail width 3.1, height 4.4; supralabials 13,
pressed, with moderately sized recurved claws rising free 13 (left, right); infralabials 14/13; circumorbitals 37/38;
of pads at terminal surfaces. paravertebral scales 227; ventral scales 95; Finger-III
Tail relatively long (compared to body): length 116% SVL; scansors 12/12; Toe-IV scansors 14/15.
vertically ovoid in cross section at base, not depressed; depth DNA sequence data, genetic divergences, and
70.5% tail width; caudals similar in size to dorsals, phylogenetic estimate.—Pseudogekko hungkag is geneti-
subcaudals similar in size to ventrals; tail lacking any vestiges cally divergent from congeners, with pairwise uncorrected
of ornate tubercles, enlarged scales, whorl differentiation, or mtDNA distances varying 10.9–20.0% (Table 2), which
lateral dermal projections. meets or exceeds the general magnitude of divergence

FIG. 8.—Preferred phylogenetic estimate (maximum likelihood [ML] topology) with species boundaries emphasized by variably colored translucent
shading. Nodal support is indicated with ML bootstraps and Bayesian posterior probabilities and distant outgroups (not shown) were selected from higher-
level studies of Brown et al. (2012b), Heinicke et al. (2012) and Oliver et al. (2018). Scale ¼ substitutions per site; tip labels include genus and species
identity, museum catalog number, and general locality. A color version of this figure is available online.
observed among other recognized species of Pseudogekko known of herpetological diversity and community composi-
(e.g., Siler et al. 2014b, 2017; Davis et al. 2015a). We note tion within the park’s interior, particularly above mid- (.300
that the single, immature specimen from the Caramoan m) to high-elevation interior forests. Nevertheless, we
Peninsula (Fig. 4C) is 20.0% divergent from Mt. Jormahan, assume the new species may be widely distributed through-
Sorsogon specimens (the type locality), and that the out the lower elevation (,600 m) forests of Mt. Bulusan,
Caramoan population most likely represents an additional, which should provide it with reasonably well-protected
undescribed species (Fig. 8). We take no taxonomic action at habitat. The assumption of potential distribution of P.
this time concerning this specimen because of the absence of hungkag within Mt. Bulusan Volcano Natural Park remains
adult specimens available for study in biodiversity reposito- unverified; we know of no Pseudogekko species records
ries. above 900–1,000 m elevation, so much of the park’s high-
Natural history, habitat, geographic distribution, and elevation interior might not actually represent optimal
conservation status.—The type locality (Mt. Jormahan, habitat. Until proper survey work can be conducted by
Barangay Cogon, Municipality of Irosin, Sorsogon Province) experienced fieldworkers within these relatively undisturbed
is a small, partially forested, extinct volcano cone (,700 m
forested areas, conclusions about the new species’ extent of
a.s.l.), situated on the western edge of the Mt. Bulusan
occurrence, habitat requirements, and conservation status
Volcano, just outside of the boundary of the protected area
will remain speculative at best. The new species is thereby
of Bulusan Volcano Natural Park (Fig. 1; Binaday et al.
2017:Fig. 1). The steep rocky slopes of Mt. Jormahan are considered ‘‘Data Deficient’’ in accordance with IUCN’s
covered with disturbed but regenerating second-growth (2016) formulaic conservation status assessment criteria.
forest that we assume was previously contiguous with nearby When first observed, the holotype was perched on the tip
patches of original forest on the lower slopes of Mt. Bulusan of an understory tree branch 3–4 m above ground; this
(now within the boundaries of the unstudied protected area). specimen, the four paratypes, and the single Caramoan
Although the area immediately surrounding Lake Bulusan specimen all slowly crawled away from the approaching field
was surveyed briefly 18 yr ago (Brown et al. 2002), the Mt. biologist when illuminated by headlamps, retreating to the
Bulusan Protected Area Management Board (PAMB) has back side of the slender branch on which they were first
restricted herpetological inventory work within the bound- spotted (personal observations). One specimen (which later
aries of the park to non-specimen-based visual-encounter escaped) was captured inside a hollow bamboo cavity at the
surveys only (Binaday et al. 2017). As a result, nothing is base of Mt. Jormahan, and another evaded capture by

dropping into a hollow tree trunk upon approach of field also note that the branching pattern in our inferred
workers. phylogenetic estimate (Fig. 8) suggests equally parsimonious
The new species’ tendency to retreat into natural cavities biogeographical patterns of colonization for Bicol P. brevipes
and tube-shaped hollow spaces inside trees, bamboo, and complex species: a single putative colonization of southern
other similar structural microhabitats has been observed at Luzon, followed by one back-dispersal versus the possibly of
an additional forested area in the Pocdol Mountains, of the two separate invasions of Luzon, resulting in isolation and
Bacon-Manito series of volcanic peaks, currently under divergence of Bicol’s two Pseudogekko species, who are not
private land tenure (and, thus, afforded some level of each other’s closest relatives.
protection) in southern Albay Province. Like the Mt. Systematic revision of Pseudogekko have resulted in the
Bulusan bamboo- and tree-hole cavity observations, the removal of species from this genus and their transfer to the
Albay Province specimens have eluded capture by retreating genus Lepidodactylus (L. shebae, L. labialis; Brown 1964;
into PVC pipes driven into the soil of mid-elevation forest Siler et al. 2014a) and widely sampled phylogenetic analyses
floor (used as botanical forest community study plot markers; have established that the continued recognition of Pseudo-
JBF, personal observation). gekko now renders Lepidodactylus paraphyletic (Brown et al.
Etymology.—We derive the new species’ masculine 2012b; Heinicke et al. 2012; Oliver et al. 2018). Previously
name from the Tagalog (Filipino) adjective hungkag, considered a small clade composed of four widely distributed
meaning ‘‘hollow’’ or ‘‘empty,’’ and used in reference to species (Brown and Alcala 1970, 1978), the content of the
the new species’ use of a unique microhabitat type: hollow genus now stands at 10 named taxa (Siler et al. 2017).
cavities and crevices (hollow bamboo trunks/stems, cavities Multiple taxonomically understudied populations are likely
in tree branches, and other tube-shaped, or sheltered to represent additional, undescribed species (e.g., the
microhabitats). We also apply this specific epithet in genetically divergent Caramoan population, at present only
reference to the lack of information on the new species’ represented by a single immature specimen; Fig. 4C).
(and most other lesser-known vertebrates) status and The two or three areas where Pseudogekko hungkag has
distribution within Bicol’s protected areas. With regard to been recorded (Mt. Jormahan, Sorsogon Province; the
P. hungkag, we have been unable to study populations within Pocdol Mountains of Albay Province; and possibly the
parks adjacent to the only known sites of occurrence, Caramoan Peninsula of Camarines Sur Province; Fig. 1)
because of administrative restrictions preventing the study of contain patches of forest, separated by heavily disturbed
biodiversity within these large forested areas. Although we agricultural regions with dense human populations and little
have sampled outside the boundaries of Mt. Bulusan, the or no natural vegetation. Estimates of natural low-elevation
Pocdol Mountains, Mt. Isarog, Mt. Mayon, Mt. Malinao, and vegetation in these inhospitable habitat matrices indicate
Caramoan National Park, to date, these regions continue to that ,1% of their natural forests are protected (Siler et al.
remain virtually unstudied for native herpetofauna (but see 2014c). We suspect that continued alteration of any forests in
Brown et al. 2002; Binaday et al. 2017). Because of these the vicinities of Bicol’s volcanic peaks (such as the
restrictions, most Bicol protected areas exist today as areas unprotected Mt. Jormahan—the new species’ type locality)
characterized by the conspicuous absence of data, distribu- will pose an immediate and direct threat to the long-term
tion gaps for many species, and geographic range maps viability of populations of P. hungkag. Like other Pseudo-
typically with numerous pockets or ‘‘hollows’’ depicting the gekko species (Siler et al. 2014b, 2017), P. hungkag is likely
absence of data from unstudied mountains—where we restricted to relatively undisturbed forest habitats at low- to
would expect to find many imperiled Philippine vertebrates. mid-elevations, only a small fraction of which are protected
Suggested common name: Bicol Hollow-dwelling Forest in the Bicol Region (Heaney et al. 1999; Primavera 2000;
Gecko. Posa and Sodhi 2006; Polidoro et al. 2010; Siler et al. 2014c).
Localities intervening the sites of documented occurrenc-
es of the new species have been the focus of a few
DISCUSSION
biodiversity surveys targeting forests of the central and
The new species is only the second member of the northern regions of the Bicol Peninsula (Brown et al. 2002;
Pseudogekko brevipes complex described from the Luzon Brown and Gonzalez 2007; Siler et al. 2009, 2010; Binaday et
Pleistocene aggregate island complex (PAIC; Voris 2000; al. 2017). A greatly expanded and comprehensive analysis of
Brown and Diesmos 2009), and its discovery was an the Bicol Faunal Region herpetological diversity is long
unanticipated surprise, coming so soon after the recently overdue, however, given its conservation importance. Such a
described P. sumiklab (Siler et al. 2017) from Mt. Mayon synthesis would markedly improve the conservation com-
(Albay Province) and the southern tip of the Bicol Peninsula munity’s practical effectiveness—and the regional govern-
(Sorsogon Province). Two other species in this clade are ment’s ability to manage the challenge posed by dense
locally endemic to the archipelago’s central West Visayan human populations at city centers surrounding Bicol’s
PAIC islands (P. atiorum; Brown and Alcala 1978; Ferner et forested volcanoes effectively (Goodman and Gonzales
al. 2000; Davis et al. 2015a; Supsup et al. 2016) and the 1990; Heaney et al. 1999; Brown et al. 2002; Siler et al.
Philippines’ southern landmasses of the Mindanao PAIC (P. 2014c).
brevipes; Siler et al. 2014a,b, 2016; Davis et al. 2015a,b). The Although widespread natural habitat degradation and
extent of sympatry between the two Luzon species has yet to wholesale removal of low-elevation forests now characterize
be assessed, and at present we consider them sympatric in most of the Bicol Peninsula (Siler et al. 2014c), the paucity of
Sorsogon and Albay (but possibly not Caramoan), but reliable information from Bicol’s herpetological communities
possibly not syntopic if the crevice and hollow cavity limits our ability to infer the conservation status of many
microhabitat preference is widespread in P. hungkag. We endemic taxa, including P. hungkag. This is because negative

occurrence data (the absence of documented geographical Brown, R.M., J.W. Ferner, and A.C. Diesmos. 1997. Definition of the
records, resulting in no actual understanding of the Philippine parachute gecko, Ptychozoon intermedium Taylor 1915
(Reptilia: Squamata: Gekkonidae): Redescription, designation of a
attributes governing a species’ distribution) and an absence neotype, and comparisons with related species. Herpetologica 53:357–
of basic data on the ecological characteristics of the new 373.
species prevent us from evaluating its status under the IUCN Brown, R.M., J. Supriatna, and H. Ota. 2000. Discovery of a new species of
criteria (IUCN 2016; Leviton et al. 2018) nor understanding Luperosaurus (Squamata; Gekkonidae) from Sulawesi, with a phyloge-
netic analysis of the genus and comments on the status of L.
the significance of potentially suitable protected habitat in serraticaudus. Copeia 2000:191–209.
unsurveyed protected areas (e.g., the Bulusan Volcano Brown, R.M., R.E. Fernandez, C. Rivero, R. Buenviaje, and A.C Diesmos.
Natural Park, where biologists are not permitted to conduct 2002. Mt. Isarog’s herpetological wonders. Haring Ibon 3:12–16.
the scientific research that would inform conservation and Brown, R.M., A.C. Diesmos, and M.V. Duya. 2007. A new species of
management policy). We anxiously await field studies Luperosaurus (Squamata: Gekkonidae) from the Sierra Madre mountain
range of northern Luzon Island, Philippines. Raffles Bulletin of Zoology
throughout the southern Bicol Peninsula, particularly within 55:153–160.
and around its few—but critically important—protected Brown, R.M., C.H. Oliveros, C.D. Siler, and A.C. Diesmos. 2008. A new
areas (Brown et al. 2013b). Gekko from the Babuyan Islands, northern Philippines. Herpetologica
64:305–320.
Acknowledgments.—We thank the Biodiversity Management Bureau Brown, R.M., C. Oliveros, C.D. Siler, and A.C. Diesmos. 2009. Phylogeny of
(BMB) of the Philippine Department of Environment and Natural Gekko from the northern Philippines, and description of a new species
Resources (DENR) for facilitating permits, and provincial and municipal from Calayan Island. Journal of Herpetology 43:620–635.
authorities of Sorsogon and Camarines Sur for facilitating access to study Brown, R.M., A.C. Diesmos, M.V. Duya, H.J.D. Garcia, and E.L. Rico.
sites and issuing specimen transport permits. Over a 3-yr period, the 2010. A new forest gecko (Squamata; Gekkonidae; genus Luperosaurus)
Bulusan Volcano Protected Area Management Board (PAMB) entertained from Mt. Mantalingajan, southern Palawan Island, Philippines. Journal of
our attempts to comply with the requirements of attending PAMB meetings Herpetology 44:37–48.
and submitting formal written requests to conduct biodiversity research Brown, R.M., A.C. Diesmos, and C. Oliveros. 2011. A new flap-legged forest
within the protected area under their jurisdiction. The Sorsogon City gecko (Genus Luperosaurus) from the northeastern Philippines. Journal
Provincial Environment and Natural Resources Office helped facilitate our of Herpetology 45:202–210.
interactions with PAMB and its superintendent. Fieldwork was conducted Brown, R.M., C.D. Siler, L.L. Grismer, I. Das, and J.A. McGuire. 2012a.
under a BMB Memorandum of Agreement (2015–2020), specimens Phylogeny and cryptic diversification in Southeast Asian flying geckos.
collected under DENR Gratuitous Permits to Collect (Nos. 258 [2015– Molecular Phylogenetics and Evolution 65:351–361.
2016] and 270 [renewal, 2017–2018]), and animal handling approved by KU Brown, R.M., C.D. Siler, I. Das, and Y. Min. 2012b. Testing the
(IACUC protocol 158-04, to RMB). In particular, we thank former DENR phylogenetic affinities of Southeast Asia’s rarest geckos: Flap-legged
Region V Regional Director C.M.P. Rodriguez and her staff (especially D.C. geckos (Luperosaurus), flying geckos (Ptychozoon) and their relationship
Vargas and M.A. Dating, D. Terciano, M. Baylon, and M.R. Concepcion) for to the pan-Asian genus Gekko. Molecular Phylogenetics and Evolution
sharing our enthusiasm and providing logistical support for our work. 63:915–921.
Fieldwork in Camarines Sur was conducted with researchers from Ateneo Brown, R.M., C.H. Oliveros, C.D. Siler, J.B. Fernandez, L.J. Welton, P.A.C.
de Naga University under a Memorandum of Agreement, and we Buenavente, M.L.D. Diesmos, and A.C. Diesmos. 2012c. Amphibians
particularly thank the Office of the University President (then headed by and reptiles of Luzon Island (Philippines), VII: Herpetofauna of Ilocos
Fr. P.F. Viray, S.J., and now by Fr. R.E.N. Rivera, S.J.) for providing Norte Province, Northern Cordillera Mountain Range. Check List 8:469–
financial and logistical support, along with encouragement for discovering 490.
and conserving the wildlife in Bicol. PLW’s collaboration on this paper Brown, R.M., C.D. Siler, C.H. Oliveros . . . A.C. Diesmos.. 2013a. The
constitutes contribution number 898 of the Auburn University Museum of amphibians and reptiles of Luzon Island, Philippines, VIII: The
Natural History. Financial support for RMB’s fieldwork was provided by the herpetofauna of Cagayan and Isabela Provinces, northern Sierra Madre
KU Docking Faculty Scholarship Fund, a Rudkin REX fund award from the Mountain Range. Zookeys 266:1–120. DOI: http://dx.doi.org/10.3897/
KU Biodiversity Institute, and a grant from the U.S. National Science zookeys.266.3982.
Foundation (DEB 1654388). We thank an anonymous reviewer and M. Brown, R.M., C.D. Siler, C.H. Oliveros . . . A.C. Alcala. 2013b. Evolutionary
Heinicke for critical reviews of an earlier draft of our manuscript. processes of diversification in a model island archipelago. Annual Review
of Ecology, Evolution, and Systematics 44:411–435.
Brown, R.M., Y.-C. Su., B. Barger, C.D. Siler, M.B. Sanguila, A.C. Diesmos,
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Wilcox, T.P., D.J. Zwickl, T.A. Heath, and D.M. Hillis. 2002. Phylogenetic INCE: Municipality of Los Baños: Barangay Batong Maiake (KU 326434,
relationships of the Dwarf Boas and a comparison of Bayesian and 326436); Barangay Bagong Silang (KU 330735, 331657); QUEZON PROVINCE:
bootstrap measures of phylogenetic support. Molecular Phylogenetics Municipality of Infanta: Barangay Magsaysay, Infanta-Marikina Highway,
and Evolution 25:361–371. Southern Sierra Madre Mountain Range (KU 334017, 344614–20);
Wiley, E.O. 1978. The evolutionary species concept reconsidered. MASBATE ISLAND: MASBATE PROVINCE: Municipality of Mobo (CAS
Systematic Zoology 21:17–26. 141560).
Pseudogekko cf. compresicorpus (1).—POLILLO ISLAND: QUEZON
PROVINCE: Municipality of Polillo; vicinity of Polillo Town (KU 326242).
Pseudogekko cf. compresicorpus (2).—LUZON ISLAND: CAGAYAN
Accepted on 7 January 2020
PROVINCE: Municipality of Gonzaga: Barangay Magrafil, Mt. Cagua (KU
ZooBank.org registration (LSID): urn:lsid:zoobank.org:pub:C6E47CB3-
330058).
BFE3-441B-B71B-A3C228C0DB23 (New nomenclatural act:
Pseudogekko ditoy (3).—LEYTE ISLAND: LEYTE PROVINCE: Munici-
urn:lsid:zoobank.org:act:22F761EB-D594-4C9C-81C3-70BEA0F6A705)
pality of Baybay: Barangay Gabas, Sitio Cienda (PNM 9811 [holotype,
Published on 9 September 2020
Associate Editor: Bryan Stuart formerly KU 326437], KU 326438 [paratype]).
Pseudogekko isapa (16).—SIBUYAN ISLAND: ROMBLON PROVINCE:
Municipality of Magdiwang: Barangay Tampayan, Mt. Guiting-Guiting
APPENDIX I Natural Park (PNM 9816 [holotype]), UPLB MNH-Z NS 4606–20; TABLAS
ISLAND: ROMBLON PROVINCE: Municipality of San Agustin (CAS 139713
Specimens Examined [paratype]).
Pseudogekko pungkaypinit (6).—BOHOL ISLAND: BOHOL PROVINCE:
All specimens from the Philippines; value in parentheses immediately Municipality of Sierra Bullones: Barrio Dusita (CAS 131854 [paratype],
following binomial represents sample size.
CAS-SU 23655 [paratype]); Raja Sikatuna Natural Park (KU 324426
Pseudogekko atiorum (16).—NEGROS ISLAND: NEGROS OCCIDENTAL
[paratype]); LEYTE ISLAND: LEYTE PROVINCE: Municipality of Baybay
PROVINCE: Municipality of Cauayan (CAS-SUR 19372, 21122); NEGROS
(KU 326243 [paratype]); Barangay Guadalupe (PNM 9810 [holotype,
ORIENTAL PROVINCE: Municipality of Valencia: Cuernos de Negros, Mt.
formerly KU 326435]); MINDANAO ISLAND: MISAMIS ORIENTAL PROV-
Talinis (CAS 134292); Municipality of Sibulan (CAS 128956, 128959,
INCE: Municipality of Gingoog City: Barangay Lawaan, Sitio Kibuko, Mt.
128963, 128971); Municipality of Valencia: Barangay Bongbong (PNM 9518
[holotype, formerly KU 302818], KU 327770, TNHC 62478); Municipality Lumot (KU 334019 [paratype]); SAMAR ISLAND: EASTERN SAMAR
of Pamplona (CAS 138097, 145793, 147491); Municipality of Siatan (CAS PROVINCE: Municipality of Balangiga, Barangay Guinmaayohan, Sitio
134269, CAS-SUR 26778); SIQUIJOR PROVINCE: Municipality of San Juan Bangon, Kaantulan River Drainage (KU 344446–48).
(CAS 145710). Pseudogekko smaragdinus (35).—POLILLO ISLAND: QUEZON PROV-
Pseudogekko brevipes (7).—SAMAR ISLAND (SMF 8988 [holotype]); INCE: Municipality of Polillo (KU 302819–302831, 303995–304002, 307638–
BOHOL ISLAND: BOHOL PROVINCE: Municipality of Sierra Bullones (CAS 307647, 326240, 326241, 331721); LUZON ISLAND: CAMARINES DEL NORTE
131855, 147527, 147528, CAS-SU 24596, 25108, 25111). PROVINCE: Municipality of Labo: Barangay Tulay Na Lupa (KU 313828).
Pseudogekko chavacano (4).—MINDANAO ISLAND: ZAMBOANGA Pseudogekko sumiklab (3).—LUZON ISLAND: ALBAY PROVINCE:
CITY PROVINCE: Municipality of Zamboanga City (PNM 9812 [holotype, Municipality of Tabaco: Barangay Mariroc, Sitio Nagsipit, Mt. Mayon:
formerly KU 314963], KU 314964 [paratype]); ZAMBOANGA DEL NORTE PNM 9843 (holotype); SORSOGON PROVINCE: Municipality of Irosin: Barangay
PROVINCE: Cuot Creek (CAS-SU 23548, 23549). Cawayan, Mt. Cawayan: KU 343847 (paratype); Municipality of Bulusan:
Pseudogekko compresicorpus (12).—LUZON ISLAND: LAGUNA PROV- Barangay Salvacion: KU 346543.

APPENDIX II.—Specimen information, locality data, and GenBank accession numbers. KU ¼ University of Kansas Biodiversity Institute herpetological
collections; TNHC ¼ Texas Natural History Collections of the Texas Memorial Museum, University of Texas at Austin; ACD ¼ Arvin Diesmos field series,
deposited at the National Museum of the Philippines (PNM).
GenBank accession
Species Voucher Locality numbers (ND2)
Pseudogekko atiorum KU 302818 Mt. Talinis, Barangay Bongbong, Municipality of Valencia, Negros Oriental KF875323
Province, Negros Island (genetic sample from holotype PNM 9518).
P. atiorum KU 327770 Mt. Talinis, Barangay Bongbong, Municipality of Valencia, Negros Oriental KF875324
Province, Negros Island.
P. atiorum PNM 9518, Mt. Talinis, Barangay Bongbong, Municipality of Valencia, Negros Oriental KF875323
KU 302818 Province, Negros Island.
Pseudogekko brevipes ACD 7255 Municipality of Sogud, Leyte Province, Leyte Island. KF875330
Pseudogekko chavacano KU 314963 Sitio Canucutan, Barangay Pasanonca, Municipality of Pasonanca, Zamboanga KF875339
City Province, Mindanao Island (genetic sample from holotype PNM
9812).
P. chavacano KU 314964 Sitio Canucutan, Barangay Pasanonca, Municipality of Pasonanca, Zamboanga KF875341
City Province, Mindanao Island (paratype).
Pseudogekko compresicorpus KU 330735 Mt. Makiling, Barangay Bagong Silang, Municipality of Los Banos, Laguna KF875337
Province, Luzon Island.
P. compresicorpus KU 331657 Mt. Makiling, Barangay Bagong Silang, Municipality of Los Banos, Laguna KF875338
P. compresicorpus KU 330058 Mt. Cagua, Barangay Magrafil, Municipality of Gonzaga, Cagayan Province, KF875327
Luzon Island.
P. compresicorpus KU 348031 Mt. Cawayan, Barangay Cawayan, Municipality of Irosin, Sorsogon Province, MN786335
Luzon Island.
P. compresicorpus KU 326242 Barangay Danicop, Municipality of Sierra Bullones, Bohol Province, Bohol KF875340
Island.
P. compresicorpus KU 344617 Infanta-Marikina Highway, Southern Sierra Madre Mountain Range, MN786333
Barangay Magsaysay, Municipality of Infanta, Quezon Province, Luzon
Island
P. compresicorpus KU 344620 Infanta-Marikina Highway, Southern Sierra Madre Mountain Range, MN786334
Barangay Magsaysay, Municipality of Infanta, Quezon Province, Luzon
Island
P. compresicorpus KU 326434 KF875336
P. compresicorpus ACD 6098 KF875326
Pseudogekko ditoy KU 326438 Sitio Cienda, Barangay Gubas, Municipality of Baybay, Leyte Province, Leyte KF875329
Island (paratype).
P. ditoy KU 326437 Sitio Cienda, Barangay Gubas, Municipality of Baybay, Leyte Province, Leyte KF875333
Island (genetic sample from holotype PNM 9811/0).
Pseudogekko pungkaypinit KU 324426 Barangay Danicop, Municipality of Sierra Bullones, Bohol Province, Bohol JQ437898
Island (paratype).
P. pungkaypinit ACD 7577 Municipality of Sogud, Leyte Province, Leyte Island. KF875331
P. pungkaypinit ACD 7637 Municipality of Sogud, Leyte Province, Leyte Island. KF875332
P. pungkaypinit H-1460 Mt. Busa, Barangay Binate, Municipality of Kiamba, South Cotabato KF875328
Province, Mindanao Island, Philippines.
P. pungkaypinit PNM 9810 LSU-Visca campus, Calbiga-a creek, Barangay Guadalupe, Municipality of KF875334
Baybay, Leyte Province.
P. pungkaypinit KU 326243 Municipality of Baybay, Leyte Province, Leyte Island. KF875335
Pseudogekko smaragdinus KU 302821 Barangay Pinaglubayan, Municipality of Polillo, Quezon Province, Polillo KF875344
Island.
P. smaragdinus KU 302819 Barangay Pinaglubayan, Municipality of Polillo, Quezon Province, Polillo JQ437897
Island.
P. smaragdinus KU 313828 Mt. Labo, Barangay Tulay Na Lupa, Municipality of Labo, Camarines Norte KF875342
P. smaragdinus KU 326241 Barangay Aluyon Municipality of Burdeos Quezon Province, Polillo Island. KF875348
P. smaragdinus KU 326240 Barangay Aluyon Municipality of Burdeos Quezon Province, Polillo Island. KF875347
P. smaragdinus KU 302820 Barangay Pinaglubayan, Municipality of Polillo, Quezon Province, Polillo KF875343
Island.
P. smaragdinus KU 307641 Barangay Salipsip, Municipality of Polillo, Quezon Province, Polillo Island. KF875346
P. smaragdinus TNHC 55024 Municipality of Polillo, Quezon Province, Polillo Island. KF875345
P. sumiklab KU 346543 Barangay Salvacion, Municipality of Bulusan, Sorsogon Province, Luzon MN786337
Island.
Pseudogekko hungkag, n. sp. KU 346539 Barangay Cogon, Municipality of Irosin, Sorsogon Province, Luzon Island MN786338
(holotype).
P. hungkag, n. sp. PNM 9865 Barangay Cogon, Municipality of Irosin, Sorsogon Province, Luzon Island MN786336
(KU 346538) (paratype).
P. cf. hungkag KU 349299 Barangay Pandanan, Municipality of Caramoan, Camarines Sur Province, MN786332
Luzon Island.

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Unexpected Discovery of Another New Species of Philippine False Gecko

Article in Herpetologica · September 2020

Cameron D Siler Matthew D Buehler

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Claire Tracy Jason B. Fernandez

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Unexpected Discovery of Another New Species of Philippine False Gecko (Gekkonidae;

al. 2014a, 2017; Davis et al. 2015a,b). Although four distinct,

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TABLE 1.—Best-fit models for evolution of clades within Pseudogekko

Model selected Model selected

ND2, first codon position GTRþFþIþC4 123124

ventrals. We used Köhler’s (2012) standardized color codes

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FIG. 4.—Photographs in life of Pseudogekko hungkag sp. nov. (A, B) adult

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Referred specimen.—KU 349299 (RMB 24361; Fig.

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brevipes chavacano compresicorpus pungkaypinit smaragdinus

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Terms of Use: https://bioone.org/terms-of-use Access provided by University of Kansas

black and cream neon green green white

125.3–141.2 (135 6 8.6) 103.6–129.7 (119.4 6 6.4)

54.2–64.3 (59.4 6 2.5)

26.2–35.5 (31.5 6 2.2)

14.9–17.4 (16.3 6 0.6)

13.8–16.0 (14.8 6 0.6)

50.7–67.9 (60.3 6 2.9)

7.8–10.7 (9.5 6 0.7)

7.1–9.8 (8.6 6 0.6)

4.5–6.5 (5.7 6 0.4)

Polillo Island and

37.2–41.2 (39.6 6 1.8)

11.4–13.6 (12.6 6 0.9)

53.6–68.4 (60.0 6 4.8)

14 vs. 16–20), infralabials (11–14 vs. 17–19), circumorbitals

ventrals (88–94 vs. 125–155), precloacals (13, 14 vs. 17–20),

by its smaller body size (SVL 42.0–52.1 vs. 50.2–64.3), fewer

Finger-III scansors (11–13 vs. 15–18), Toe-IV scansors (13–

(11–14 vs. 14–19), paravertebrals (222–235 vs. 241–252),

ventrals (88–94 vs. 124–130), and precloacals (13, 14 vs. 32–

41 precloacofemorals), by its overall gray ground (vs. bright

27.0–32.6 (30.4 6 1.8)

16.1–18.1 (17.1 6 0.7)

13.0–16.3 (14.2 6 1.0)

56.1–64.6 (59.6 6 3.0)

[disturbed]) coloration, and by the presence of bold inverted

7.5–10.1 (8.2 6 0.8)

5.3–7.4 (5.9 6 0.7)

Y-shaped caudal chevrons (vs. transverse dark/light caudal

bands; Tables 3 and 4).

17.8–29.8 (21.7 6 5.5) 26.7–30.0

material). Body relatively large, robust, SVL 52.1 mm; limbs

well developed, robust, relatively muscular; tail long, intact,

60.5 mm, tapering to slender tip, thickened at laterally

compressed base; margins of limbs smooth; cutaneous flaps,