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We describe a new species of lizard in the genus Gekko from Sibuyan Island in the Romblon Island group of the central Philippines. Although the new species is diagnosed from other Philippine Gekko by body size and shape, coloration, and multiple characteristics of external morphology, additional support for the recognition of the Sibuyan Gekko population as a distinct evolutionary lineage is garnered from DNA sequence data and biogeographical inference. The new species has been collected on trunks of trees or on granitic rocks along rivers in mature, lowland forest, and on vegetation at forest edges bordering agricultural areas. It is known only from Sibuyan Island and is undoubtedly endemic to this single small, isolated landmass. Although the larger, topographically complex islands of the Philippines have been the targets of numerous recent efforts to estimate vertebrate species diversity, smaller islands of the archipelago have received comparatively less attention and may support significant levels of underappreciated vertebrate diversity.
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A New Gekko from Sibuyan Island, Central Philippines
Author(s) :Rafe M. Brown, Cameron D. Siler, Carl H. Oliveros, Arvin C.
Diesmos, and Angel C. Alcala
Source: Herpetologica, 67(4):460-476. 2011.
Published By: The Herpetologists' League
DOI: 10.1655/HERPETOLOGICA-D-11-00025.1
URL: http://www.bioone.org/doi/full/10.1655/HERPETOLOGICA-
D-11-00025.1
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A NEW GEKKO FROM SIBUYAN ISLAND, CENTRAL PHILIPPINES
RAFE M. BROWN
1,5
,CAMERON D. SILER
1
,CARL H. OLIVEROS
1,2
,ARVIN C. DIESMOS
3
,AND
ANGEL C. ALCALA
4
1
Natural History Museum, Biodiversity Institute & Department of Ecology and Evolutionary Biology, The University of
Kansas, Lawrence, KS 66045, USA
2
Isla Biodiversity Conservation, 9 Bougainvillea Street, Manuela Subdivision, Las Pin˜ as City 1741, Philippines
3
National Museum of the Philippines, Rizal Park, Padre Burgos Avenue, Ermita 1000, Manila, Philippines
4
Angelo King Center for Research and Environmental Management, Silliman University, Dumaguete City 6200,
Philippines
ABSTRACT: We describe a new species of lizard in the genus Gekko from Sibuyan Island in the Romblon
Island group of the central Philippines. Although the new species is diagnosed from other Philippine Gekko
by body size and shape, coloration, and multiple characteristics of external morphology, additional support for
the recognition of the Sibuyan Gekko population as a distinct evolutionary lineage is garnered from DNA
sequence data and biogeographical inference. The new species has been collected on trunks of trees or on
granitic rocks along rivers in mature, lowland forest, and on vegetation at forest edges bordering agricultural
areas. It is known only from Sibuyan Island and is undoubtedly endemic to this single small, isolated
landmass. Although the larger, topographically complex islands of the Philippines have been the targets of
numerous recent efforts to estimate vertebrate species diversity, smaller islands of the archipelago have
received comparatively less attention and may support significant levels of underappreciated vertebrate
diversity.
Key words: Gekkonidae; New species; Philippines; Romblon Island Group; Sibuyan Island
RECENTLY, enhanced survey efforts, careful
scrutiny of widespread species, and use of
molecular sequence data in combination
with traditional morphological characters,
have resulted in a dramatic increase in the
diversity of gekkonid lizards in the Philip-
pines. The archipelago is now known to
support 10 genera and at least 48 gekkonid
species assigned to the genera Cyrtodactylus
(9 species), Gekko (12–13), Gehyra (1),
Hemidactylus (5; including platyurus, a spe-
cies formerly assigned to Cosymbotus), Hemi-
phyllodactylus (2), Lepidodactylus (6), Lupe-
rosaurus (8), Pseudogekko (4), and Ptychozoon
(1) (Brown, 1999; Brown and Alcala, 1978;
Brown and Diesmos, 2000; Brown et al., 1997,
2007, 2008, 2009, 2011; Gaulke et al., 2007;
Linkem et al., 2010; Taylor, 1922a,b; Welton
et al., 2009, 2010a,b; Zug, 2010).
Ten species of Gekko are considered endemic
to the archipelago (Brown et al., 2009; Linkem
et al., 2010) and two additional species with
broad geographic distributions (G. gecko,G.
monarchus) are also known from the country
(Brown and Alcala, 1978; Ota et al., 1989;
Taylor, 1922a,b). Although past treatments have
included G. hokouensis as part of the Philippine
gekkonid fauna (Brown and Alcala, 1978), Ota
et al. (1989) demonstrated that this taxon’s
inclusion in Philippine faunal accounts is likely
in error. Similarly, the single Mindanao Island
record for Perochirus ateles (Boulenger, 1885;
Brown and Alcala, 1978; Dume
´ril, 1856; Taylor,
1922a) has not been confirmed in the last 150 yr
despite our recent field surveys in and around
the type locality (Zamboanga), suggesting that it
too may be in error (Bauer and Henle, 1994;
Brown, 1976; Welton et al., 2010a).
The 10 endemic Philippine species of
Gekko are G. athymus (Brown and Alcala,
1962), G. carusadensis (Linkem et al., 2010),
G. crombota (Brown et al., 2008), G. ernst-
kelleri (Ro
¨sler et al., 2006), G. gigante (Brown
and Alcala, 1978), G. mindorensis (Taylor,
1919), G. palawanensis (Taylor, 1925), G.
porosus (Taylor, 1922b), G. romblon (Brown
and Alcala, 1978), and G. rossi (Brown et al.,
2009). These species represent a considerable
range in body size, general appearance, and
ecological attributes, but all possess the
following combination of morphological traits:
(1) body size moderate, with relatively long,
slender limbs; (2) near complete absence
of interdigital webbing or cutaneous body
expansions; (3) dorsal tubercles arranged in
longitudinal rows (except for G. athymus,
5
CORRESPONDENCE: e-mail, rafe@ku.edu
Herpetologica, 67(4), 2011, 460–476
E2011 by The Herpetologists’ League, Inc.
460
in which dorsal tuberculation is absent); (4)
scales of dorsum between tubercle rows
minute, nonimbricate; (5) scales of venter
enlarged, imbricate, flat; (6) differentiated
postmentals longitudinally elongate; and (7)
subcaudals transversely enlarged, platelike
(Brown and Alcala, 1978; Brown et al., 2007,
2008, 2009).
Our survey work in the Romblon Island
Group of central Philippines has resulted in
the discovery of a new species of morpholog-
ically and genetically distinct Gekko related to
G. romblon. In this article we use a combina-
tion of body size and shape information,
meristic data of external morphology (scale
counts), genetic sequence data, and inferenc-
es from the geological history of the archipel-
ago to demonstrate that the Sibuyan Island
Gekko population represents a distinct evolu-
tionary lineage (de Queiroz, 1998, 1999;
Wiley, 1978), worthy of specific rank.
MATERIALS AND METHODS
Morphology
We (RMB and CO) collected data from
fluid-preserved specimens deposited in US
and Philippine collections (see Appendix;
institutional abbreviations follow Leviton
et al., 1985). Sex was determined by inspec-
tion of gonads or by scoring of prominent sec-
ondary sexual characteristics (Brown, 1999;
Brown et al., 1997, 2008, 2009, 2010) when
dissection was not possible. Measurements (to
the nearest 0.1 mm) were taken with digital
calipers following character definitions by
Brown (1999), and Brown et al. (2008, 2009).
Characters include: snout–vent length, tail
length, head length; head width, head depth,
snout length, eye diameter, eye–narial dis-
tance, internarial distance, interorbital dis-
tance, axilla–groin distance, femur length, tibia
length, upper arm length, forearm length, Toe
I length, Toe IV length, tail width, tail depth,
number of supralabials and infralabials poste-
riorly to the point at which point labials are no
longer differentiated, enlarged precloacofe-
moral pore-bearing (males) or dimpled (fe-
males) scales, differentiated subdigital scansors
beneath Finger III and Toe IV, midbody dorsal
and ventral transverse scale rows between
lateral body folds, midbody dorsal transverse
tubercle rows between dorsolateral body folds,
undifferentiated paravertebrals and tubercle
rows between midpoints oflimb insertions, and
midventrals between limb insertion.
Molecular Data
Because our primary goal was to estimate
phylogenetic relationships among the island
populations of Gekko in the Romblon Island
group, we sequenced only 2–4 exemplars per
species and selected only three outgroup taxa
(G. gecko,G. mindorensis, and G. monarchus)
based on relationships presented in a recent
phylogenetic analysis of northern Philippine
gekkonid lizards (Brown et al., 2009). A total
of seven ingroup samples were used to
estimate phylogenetic relationships of the
Romblon Province Gekko populations.
Genomic DNA was extracted from liver tissues
stored in 95–100% ethanol following the guani-
dine thiocyanate protocol of Esselstyn et al.
(2008). For all 12 samples, the mitochondrial
gene NADH Dehydrogenase Subunit 2 (ND2)
and components of three flanking transfer RNA
genes (tRNA
trp
,tRNA
ala
,tRNA
asn
) were se-
quenced by the primers, thermal profiles, and
purification and sequencing protocols of Brown
et al. (2009) and Macey et al. (1999). Purified
product was analyzed with an ABI Prism 31303l
Genetic Analyzer (Applied Biosystems). Newly
sequenced data were deposited in GenBank
under accession numbers JN710488–506. Gene
sequence contigs were assembled and edited with
the use of Sequencher 4.8 (Gene Codes Corp.,
Ann Arbor, MI). Initial sequence alignments
were produced in Muscle (Edgar, 2004), and
manual adjustments were made in MacClade
4.08 (Maddison and Maddison, 2005).
Phylogenetic Analysis
Partitioned Bayesian analyses were con-
ducted in MrBayes v3.1.2 (Ronquist and
Huelsenbeck, 2003) for the combined data
set. The mitochondrial data set was parti-
tioned by codon position for the protein-
coding region of ND2, and the three flanking
tRNAs (tRNA
trp
, tRNA
ala
, tRNA
asn
) were
analyzed as a single subset. The Akaike
Information Criterion (AIC), as implemented
in jModeltest v0.1.1 (Guindon and Gascuel,
2003; Posada, 2008), was used to select the
best model of nucleotide substitution for each
December 2011] HERPETOLOGICA 461
partition (Table 1). The best-fit models for
each codon position of ND2, and for the
combined tRNA data, were the general time
reversible (GTR) model with a gamma-
distributed rate variation among sites (C) and
the Hasegawa, Kishino, and Yano (HKY)
model with a gamma-distributed rate variation
among sites (C), respectively. A rate multiplier
model was used to allow substitution rates
to vary among subsets, and default priors
were used for all model parameters. We ran
four independent Metropolis-coupled MCMC
analyses, each with four chains and an incre-
mental heating temperature of 0.05, and an
unconstrained branch length prior with an
exponential distribution of 25 (Marshall, 2010;
Marshall et al., 2006; Siler et al. 2010, 2011).
All analyses were run for 25 million genera-
tions, sampling every 5000 generations. To
assess stationarity, all sampled parameter
values and log-likelihood scores from the cold
Markov chain were plotted against generation
time and compared among independent runs
using Tracer v1.4 (Rambaut and Drummond,
2007). Finally, we plotted the cumulative and
nonoverlapping split frequencies of the 20
most variable nodes, and compared split
frequencies among independent runs with
the use of Are We There Yet? (AWTY;
Wilgenbusch et al., 2004). Although all
samples showed patterns consistent with
stationarity after 2.5 million generations (i.e.,
the first 10.0%), we conservatively discarded
the first 20% of samples as burn-in.
Partitioned maximum-likelihood (ML) analy-
ses were conducted in RAxMLHPC v7.0
(Stamatakis, 2006) for all three data sets under
the same partitioning strategy as for Bayesian
analysis. The more complex model (GTR +C)
was used for all subsets, and 100 replicate ML
inferences were performed for each analysis.
Each inference was initiated with a random
starting tree, and employed a rapid hill-climbing
algorithm (Stamatakis et al., 2007). Clade
support was assessed with 1000 bootstrap
pseudoreplicates, employing a rapid bootstrap-
ping algorithm (Stamatakis et al., 2008).
Species Concept
For the recognition of the new species, we
adopted the general lineage species concept
of de Queiroz (1998, 1999) as the natural
extension of the evolutionary species concept
(Wiley, 1978). Application of lineage-based
species concepts to central Philippine island
endemics is nonproblematic (Brown and
Diesmos, 2002; Brown and Guttman, 2002)
because of the known history of isolation of
island populations (Hall, 1998, 2002; Voris,
2000; Yumul et al., 2003, 2009). We consider
as new species morphologically diagnosable
lineage segments (either in isolation or
sympatry) in which the hypothesis of con-
specificity can be confidently rejected by
TABLE 1.—Models of evolution selected by Akaike Information Criterion for partitioned, model-based phylogenetic
analyses. General time reversible (GTR) model with a gamma-distributed rate variation among sites (C) was applied for
all data partitions in partitioned maximum-likelihood analyses.
Partition AIC model Number of characters
NADH 2, first codon position GTR +C346
NADH 2, second codon position GTR +C346
NADH 2, third codon position GTR +C346
tRNAs Trp, Ala, Asn HKY +C223
TABLE 2.—Uncorrected pairwise sequence divergence (%) for molecular data for Gekko coi,G. romblon,G.
mindorensis,G. monarchus, and G. gecko (Fig. 1). Percentages on the diagonal represent intraspecific genetic diversity
(bolded for emphasis).
Gekko coi G. romblon G. mindorensis G. monarchus
G. romblon 7.0–7.4 0.0–3.9
G. mindorensis 19.1 19.1–19.3 0.2
G. monarchus 19.0 18.8–19.1 14.7–14.8 0.0
G. gecko 28.6 28.7–28.8 28.1–28.2 28.3
462 HERPETOLOGICA [Vol. 67, No. 4
discrete character differences, especially
when taxonomic decisions are bolstered by
genetic data and biogeographic information.
RESULTS
Morphology
Our surveys of mensural and meristic data
recover instances of discrete, nonoverlapping
ranges of variation in multiple characters of
external morphology (Table 3) between Rom-
blon +Tablas island populations (G. romblon)
and the population of Gekko from Sibuyan
Island (Gekko sp. nov.). The putative new
species from Sibuyan is readily diagnosed
from all Philippine congeners (see below) on
the basis of body size and shape, coloration,
and numerous characteristics of external
morphology. The new species is easily diag-
nosed from its closest relative G. romblon on
the basis of body shape, coloration, and
discrete, diagnostic, characteristics of scala-
tion (Table 3).
Phylogeny and Genetic Divergence
The ML analysis resulted in a single optimal
tree (2ln L54129.152041; Fig. 1). Trees
estimated from ML and Bayesian analyses are
consistent with respect to support for two unique
species of Gekko distributed in the Romblon
Island group. All analyses recover the Sibuyan
Island population of Gekko as a lineage distinct
from (but sister to) a clade consisting of the
Tablas and Romblon island populations (e.g.,
G. romblon sensu stricto; Fig. 1). Uncorrected
pairwise sequence divergences are low within G.
romblon and include a moderately shallow
(#3.9%) genetic divergence between Romblon
and Tablas island populations. In contrast,
these island populations exhibit relatively higher
genetic divergence (7.0–7.4%) from the Sibuyan
Island lineage (Table 2; Fig. 1).
SYSTEMATICS
Gekko coi sp. nov.
Figs. 2–5
Holotype.—PNM 9765 (Field no. RMB 2961;
formerly KU 326208), an adult male collected by
ACD at 2330 h on 2 January 2001 on a large
stream-side boulder near sea level in mixed
second growth and primary forest at Barangay
Tampayan, Municipality of Magdiwang, Rom-
blon Province, northeast coast of Sibuyan Island,
Philippines (12.486uN, 122.516uE; datum 5
WGS84).
Paratypes.—FMNH 251114–15, adult fe-
males, collected 17 and 18 March, 1992, by N.
Ingle and S. Goodman on Mt. Guiting-guiting,
Barangay Tampayan, Municipality of Magdi-
wang; CAS 139180, adult male collected by
L. C. Alcala and party on 9 May 1972 at
Taclobo Barrio, Municipality of San Fernando
(paratype of G. romblon); CAS 139181, a
juvenile, same collection data, but 12 May
1972 (paratype of G. romblon); CAS139182
(paratype of G. romblon) and CAS 155896,
adult male, collected by L. C. Alcala and party
on 13 May 1972, at Cansampay River, Taclobo
Barrio, Municipality of San Fernando.
Diagnosis.—Gekko coi differs from all other
species of Philippine Gekko (i.e., G. athymus,
G. carusadensis,G. crombota,G. ernstkelleri,
G. gecko,G. gigante,G. mindorensis,G.
monarchus,G. palawanensis,G. porosus,G.
romblon, and G. rossi) in having the following
combination of diagnostic traits: (1) moder-
ately large body size (snout–vent length [SVL]
65.2–84.0 for adult males, 72.1–77.1 mm for
females); (2) dorsum medium brown to gray,
with single row of alternating light (cream)
and dark (dark brown) vertebral blotches; (3)
high numbers of dorsal body scales (107–132
transverse midbody scales, 192–226 paraver-
tebrals); (4) relatively few rows of conical body
tubercles (13–15 midbody, 25–28 paraverte-
brally; (5) precloacofemorals in a continuous
series (precloacals and femoral pore-bearing
scale series distinctly differentiated but abut-
ting with no undifferentiated scales interrupt-
ing the two series) of 86–92 differentiated,
greatly enlarged (precloacals) to only slightly
enlarged (femorals) scales.
Comparison with similar species.—Gekko
coi differs from its phenotypically most similar
and geographically most proximate Philippine
congener (G. romblon) by its relatively
elongate, slender body, and narrow head
(versus more robust body, wide head, charac-
terized by distinctly hypertrophied adductor
musculature; Fig. 2), possession of 86–92
(versus 64–79 in G. romblon) differentiated
precloacofemoral pore-bearing scales, 34–43
(versus 43–53) midbody ventrals, and by
December 2011] HERPETOLOGICA 463
TABLE 3.—Distribution of selected diagnostic characters in Gekko coi and other Philippine species of Gekko. Entries are presented in millimeters; all specimens are
considered adults (data from juveniles excluded). Characters (following Linkem et al., 2010, and Brown et al., 2009) include (1) dorsal body coloration, (2) dorsal tail
coloration, the numbers of (3) supralabials, (4) differentiated precloacofemoral pore-bearing scales, (5) scansors below Toe IV, (6) white postorbital spots (+, present; 2,
absent), (7) midbody dorsals, (8) midbody ventrals, (9) midbody dorsal tubercle rows, (10) paravertebral tubercle rows, (11) ventrals, and (12) paravertebrals.
NMale snout–
vent length Female snout–
vent length 1 2 3 4 5 6 7 8 9 10 11 12
Gekko coi 3 males;
3 females
65.2–84.0 72.1–77.1 Light and
dark vertebral
blotches
Alternating light
and dark bands
12–15 85–92 16–18 +99–106 39–43 13–15 25–28 77–88 178–185
G. romblon 8 males;
4 females
78.7–87.1 71.0–81.0 Light and
dark vertebral
blotches
Gray–brown
with thin
white bands
12–14 66–79 14–18 +102–129 43–53 12–16 22–34 84–94 193–231
G. athymus 3 males;
2 females
99.2–119.9 88.2–117.1 Light and
dark inverted
V-shaped bands
Dark brown
with yellow
bands
11–13 20–24 18–22 292–104 30–36 66–72 66–72
G. carusaurdus 6 males;
2 females
83.4–97.2 79.9–87.5 Small dark
mottling
Alternating light
and dark
bands
12–14 46–50 18–20 290–106 38–47 14–17 25–28 179–190
G. crombota 4 males;
9 females
85.5–117.9 85.1–106.9 Light trilobed
bars
Alternating light
and dark bands
13–15 58–74 15–18 2107–132 38–42 18–22 29–33 67–85 192–226
G. ernstkelleri 4 males;
6 females
82.0–92.1 78.0–88.0 White circular
spots
Dark gray with
bold white
bands
15, 16 36–42 17–19 +112–127 42–48 10–16 17–25 58–62 178–200
G. gecko 6 males;
7 females
120.1–166.1 119.2–144.1 Rust-colored
spots
Alternating light
and dark bands
12–14 12–20 17–20 294–106 30–35 10–12 18–22 60–64 91–102
G. gigante 5 males;
3 females
89.7–104.7 79.7–87.9 Dark paired
blotches
Light brown with
white bands
11–13 52–66 16–19 2123–135 41–50 12–18 19–28 65–74 175–207
G. mindorensis 22 males;
13 females
55.0–88.2 68.2–70.9 Dark thin
transverse
bands
Alternating light
and dark bands
11–13 52–66 12–14 2102–125 40–47 16–20 17–26 58–63 180–195
G. monarchus 13 males;
10 females
56.2–80.7 40.6–69.7 Dark transverse
spot rows
Dark gray
with light
gray bands
11–13 31–40 13–15 296–112 38–44 16–20 18–23 57–61 171–203
G. palawanensis 3 males;
4 females
57.2–65.7 44.5–61.8 Dark paired
spots
Alternating light
and dark bands
12–14 64–70 16–19 2114–121 38–43 10–20 23–27 54–58 155–170
G. porosus 3 males 91.0–96.7 91.0–96.7 Indistinct
transverse
dark bands
Alternating light
and dark bands
12, 13 74–80 14–16 288–103 35–40 15–17 17–24 64–74 173–191,
173–191
G. rossi 8 males;
8 females
95.5–108.2 86.8–100.0 Dark transverse
bars and
light spots
Alternating light
and dark bands
13–16 77–88 10–16 2125–170 33–41 16–18 31–37 74–104 251–281
464 HERPETOLOGICA [Vol. 67, No. 4
178–185 (versus 193–231) paravertebrals.
Enlarged rows of chin shields ventral to the
infralabials are smaller on G. coi such that
three enlarged subinfralabials contact infra-
labials 2–4 (versus only two scales contacting
infralabials 2–4 in most specimens of G.
romblon). Additionally, dorsal tail coloration of
the G. romblon holotype (CAS 139190) and
70% of the remaining specimens from Romblon
and Tablas islands is flat gray with thin white
transverse lines (versus 100% of specimens with
thick caudal bars of contrasting light and dark in
G. coi). Finally, G. coi is distinguished from G.
romblon by the presence (versus absence) of
distinct, bold, black, irregularly shaped markings
throughout the dorsal surfaces of the body,
limbs, and head (Fig. 2).
Body size separates G. coi from the larger
species G. athymus,G. gecko,G. crombota,G.
porosus, and G. rossi, and the smaller species
G. palawanensis (Table 3); dark reddish-
brown dorsum with light and dark vertebral
spots and a large, prominent, postorbital
spot distinguishes G. coi from all Philippine
congeners except G. romblon; the number of
precloacofemorals distinguishes G. coi from
G. athymus,G. carusadensis,G. crombota,G.
ernstkelleri,G. gecko,G. gigante,G. mind-
orensis,G. monarchus,G. palawanensis, and
G. porosus. Midbody ventrals distinguish
G. coi from G. athymus,G. carusadensis,G.
crombota,G. ernstkelleri,G. gecko,G.
gigante. Midbody dorsal tubercle rows distin-
guish G. coi from G. crombota,G. mind-
orensis,G. monarchus, and G. rossi, and
paravertebral tubercle rows distinguish G. coi
from G. ernstkelleri,G. crombota,G. gecko,
G. monarchus,G. porosus,andG. rossi.
Finally, the number of ventrals distinguishes
G. coi from G. athymus,G. ernstkelleri,G.
gecko,G. gigante,G. mindorensis,G. mon-
archus,G. palawanensis,andG. porosus,
paravertebral separate G. coi from G. athy-
mus,G. crombota,G. gecko,G. palawanensis,
FIG. 1.—Map of Romblon Province (right panel) in relation to the remainder of the Philippines (left). Known
distribution of Gekko species in the province is indicated with symbols (key, lower right) plotted at collection localities.
A maximum-likelihood estimate of phylogeny (with ML bootstraps/Bayesian posterior probabilities included on
internodes) is presented on right. Scale bars 5substitutions/site.
December 2011] HERPETOLOGICA 465
and G. rossi. These and other differences
among Philippine Gekko species are summa-
rized in Table 3.
Description of holotype.—Adult male in
excellent condition (Figs. 2A,B, 3A,B), with a
small incision in the sternal region (portion of
liver removed for genetic sample), and hemi-
penes fully everted. SVL 84.1 mm; habitus
slender, limbs well developed, relatively
slender; tail relatively long; margins of limbs
smooth, lacking cutaneous flaps or dermal
folds; a thin adipose line (cutaneous fold)
running along ventrolateral margin of trunk.
Head moderate, differentiated from neck,
characterized by only slightly hypertrophied
temporal and adductor musculature; notice-
ably broader (1.2 times) body at widest point;
snout subtriangular, rounded at tip in dorsal
and lateral aspect (Fig. 4A); head width 76.3%
head length, 18.8% snout–vent length; snout
length 63.3% head width and 48.3% head
length; dorsal surfaces of head relatively
FIG. 2.—Gekko coi male holotype (PNM 9765; A, B) in life; adult male G. romblon (KU 303978) from Romblon Island
(C, D). (For interpretation color in this figure, see the online version of this article.)
FIG. 3.—Dorsal (A) and ventral (B) view of the adult male Gekko coi holotype (PNM 9765; snout–vent length
84.0 mm). Scale bar 510 mm.
466 HERPETOLOGICA [Vol. 67, No. 4
homogeneous, with only slightly pronounced
concave postnasal, prefrontal, and interorbital
concavities; auricular opening large, ovoid,
angled slightly anteroposteriorly from beneath
temporal swellings on either side of head;
tympanum deeply sunken; orbit large, bor-
dered by only moderately distinct supraorbital
crest and sharply prominent preorbital crest;
eye large, pupil vertical, margin wavy
(Fig. 4A); auricular opening 50.5% eye diam-
eter; limbs and digits relatively long and
slender; femoral segments of hind limbs thick,
bulky compared to humeral segments of
forelimbs; tibia length 15.2% SVL, 76.6%
femur length.
Rostral large, rectangular in anterior view,
twice as broad as high, with two dorsomedial
fissures between raised posterodorsal projec-
tions that form the anteriormost projecting
edge of the nares, and sutured anteriorly with
the supranasals; nostril surrounded by rostral,
first labial, an enlarged lower postnasal, a
smaller upper postnasal, and an enlarged,
round to octagonal, convex supranasal; supra-
nasals separated by a single enlarged, elongate
internasal; supranasals and internasal followed
posteriorly by a pair of slightly enlarged
posterosupranasals, separated by three slightly
enlarged median scales; scales immediately
posterior to posterosupranasals only slightly
enlarged.
Total number of differentiated supralabials 15/
15 (L/R; 11/10 to center of eye), bordered
dorsally by several rows of nondifferentiated,
FIG. 4.—Dorsal, lateral, and ventral views of the heads of the adult male holotypes of Gekko coi (A: PNM 9765) and G.
romblon (B: CAS 139190). Scale bars 55 mm.
December 2011] HERPETOLOGICA 467
nonenlarged snout scales; total number of
differentiated infralabials 13/12 (L/R; 9/9 to
center of eye), bordered ventrally by two rows
of enlarged scales and three rows of only slightly
differentiated chin scales; mental triangular;
mental and first four infralabials greatly enlarged
and wrapping onto ventral surfaces of chin,
nearly twice the size of individual infralabials 4–
12; mental followed posteriorly by a pair of
slender, highly elongate medial postmentals;
postmentals bordered posterolaterally by a sec-
ondary pair, approximately one-sixth or one-
seventh the length of first pair; postmental scale
series bordered posteriorly by a single series of
only slightly enlarged scales; followed immedi-
ately by a sharp transition to undifferentiated
chin and gular scales; postrictal scales undiffer-
entiated; remaining undifferentiated gulars very
small, round, nonimbricate, juxtaposed (Fig. 4A).
Dorsal cephalic scales fairly homogeneous
in size, shape, disposition, and distribution;
cephalic scalation varies from large, convex,
round to oval scales of rostrum to minute
(approximately one-third to one-fifth size of
rostrals), granular scales of posterior regions
of head and neck; postnasal, prefrontal, and
interorbital depressions with slightly smaller
scales; palpebral scales heterogeneous, with
some scales as small as adjacent interorbital
region and others as large and raised as rostral
scales; undifferentiated posterior head scales
granular, convex, reducing in size posteriorly,
interspersed with increasingly dense enlarged
rounded to slightly conical tubercules; throat
and chin scales small, juxtaposed and nonim-
bricate, making a sharp transition to gular and
pectoral region scalation, with enlarged cy-
cloid, imbricate scales continuing to increase
in size through abdomen, becoming very
enlarged and strongly imbricate.
Ornamental cephalic scalation limited to
convex tubercles on posterolateral portions of
head (temporal, supratympanic, and postrictal
regions) and a slightly differentiated series of
2–3 enlarged, weakly conical unkeeled pre-
orbital scales (Fig. 4A); 27/25 circumorbitals
in total, differentiated into the following
distinct regions: (1) several minute precircu-
morbitals, (2) enlarged, flat, squarelike circu-
morbitals dorsoanterior to orbit (13/11), (3)
several smaller, undifferentiated supraorbital
scaled (4/6), (4) transverse elongation and
modification into fringe-like points (spiny
ciliaria, 10/8) across dorsoposterior margin of
orbit, gradually reducing to (5) several smaller
to minute postcircumorbitals, a total of 47
interorbital scales (straight line distance from
center of each eye, across both eyelids).
Axilla–groin distance 44.0% SVL; undiffer-
entiated dorsal body scales round, convex,
juxtaposed, relatively homogeneous in size;
each dorsal scale surrounded by six interstitial
granules, giving the appearance of a ‘‘Star of
David’’ configuration under high magnifica-
tion; dorsals interspersed with 15 irregularly
transverse rows (25 paravertebral rows) of
enlarged, slightly conical dorsal body tuber-
cles; tubercles surrounded by undifferentiated
adjacent dorsal scalation; dorsals sharply
transition to imbricate ventrals along the
ventrolateral adipose fold; transverse midbody
dorsals 109; paravertebrals between midpoints
of limb insertions 185; midbody dorsals in 106
rows; midbody ventrals in 43 transverse rows;
scales on dorsal surfaces of limbs larger and
more imbricate than dorsals, interspersed
FIG. 5.—Inferior view of the precloacofemoral region of
the male Gekko coi holotype (A: PNM 9765) and a
specimen of G. romblon from type locality on Tablas
Island (B: KU 315347). Scale bars 55 mm.
468 HERPETOLOGICA [Vol. 67, No. 4
with enlarged, flat to slightly conical tubercles
on the radioulnar segment of the limb but
absent on the humeral segment, and termi-
nating at the dorsal surfaces of hands and feet;
enlarged patches of distinct imbricate scales
present on wrist, anterior (preaxial) surface of
upper arm and thigh, on knee, and on distal
ventral surface of hind limb; scales on dorsal
surfaces of hands and feet similar to dorsal
limb scales (but lacking tubercles); ventral
body scales flat, cycloid, strongly imbricate,
much larger than lateral or dorsal body scales,
largest at midventral line.
Seventy-four pore-bearing or dimpled
scales (Fig. 5A) in continuous precloacofe-
moral series, each punctured by pore bearing
dark orange exudate, arranged in a wavy,
widely obtuse, inverted-V formation and
continuing to just beyond the patellar region;
precloacal pores 3–4 times diameter of
femoral pores; precloacals (14/14) situated
atop a substantial precloacal bulge that is
folded over into the precloacal region in
preserved type but was erect and protuberant
in life; precloacals preceded by five similarly
enlarged but nondimpled scale rows; precloa-
cals followed by a gap of undifferentiated
scales, followed by five enlarged-scales rows,
forming a triangular patch of scales anterior to
the vent; femoral series lacks preceding or
following enlarged scale rows; scales latero-
posterior to precloacofemoral series (i.e.,
along ventroposterior surfaces of hind limb)
reduce in size sharply to minute scales of the
posterior edge of the hind limb.
Digits moderately expanded and covered on
palmar/plantar surfaces by bowed, unnotched,
undivided scansors (Fig. 4A); digits with min-
ute vestiges of interdigital webbing; subdigital
scansors of manus: 9/10, 11/12, 13/14, 15/16,
and 11/12 on left/right digits I–V respectively;
pes: 11/11, 12/12, 15/15, 17/16, and 14/13
on left/right digits I–V respectively; subdigital
scansors of manus and pes bordered proximally
(on palmar and plantar surfaces) by 1–4 slightly
enlarged scales that form a near-continuous
series with enlarged scansors; all digits clawed,
but first (inner) claw greatly reduced; remain-
ing terminal claw-bearing phalanges com-
pressed, with large recurved claws, not rising
free at distal end until they extend beyond
dilated hyperextensible portion of digit.
Tail base bordered by a pair of moderately
enlarged conical postcloacal spurs on each
side of vent; postcloacal swellings pro-
nounced; hemipenes completely everted; tail
long, 105 mm, 1.25% snout–vent length; tail
not depressed, subcylindrical, divided into
indistinct fracture planes/autotomy grooves,
with whorls (or annulations) clearly visible in
basal portions of the tail (with slightly
enlarged scales along the caudal margin of
each annulation) but becoming less distinct
toward distal portion; an estimated 25–27
annuli total; tail depth (not including basal
postcloacal swelling) 60.6% tail width; dorsal
tail with only a few enlarged caudal tubercles;
caudals similar in size to dorsals; subcaudals a
single enlarged medial row of platelike scales,
flanked laterally by one slightly enlarged row;
subcaudals widely expanded to cover majority
of ventral surface of tail.
Variation.—The type series consists of three
adult males, three adult females, and a single
juvenile specimen. Ranges of diagnostic me-
ristic and mensural characters are presented in
Table 3. We detected no sexual dimorphism in
the type series and ranges of all meristic and
mensural measurements broadly overlap be-
tween the sexes. With data for males and
females combined, mean SVL (61 SD; range)
was 76.0 (66.76; 65.2–84.0), nonregenerated
tail length 81.5 (68.6; 75.4–81.6), axilla–groin
distance 35.5 (61.9; 32.1–37.0), tail width 6.6
(60.9; 5.0–7.6), tail depth 5.0 (61.1; 4.3–6.0),
head length 21.4 (61.4; 19.1–22.7), head width
14.7 (61.1; 13.3–15.8), head depth 8.1 (60.6;
6.9–8.6), snout length 9.5 (61.2; 7.4–10.4),
eye–narial distance 7.5 (61.2; 5.1–8.2), inter-
narial distance 2.8 (60.24; 2.5–3.1), interorbit-
al distance 5.6 (60.7; 4.8–6.6), femur length
15.6 (61.0; 13.8–16.7), tibia length 13.2 (61.2;
11.0–14.2), upper arm 8.7 (60.5; 7.5–9.1),
forearm length 9.9 (60.7; 9.7–10.5), Toe I
length 4.0 (60.2; 3.9–4.4), Toe IV length 8.0
(60.3; 7.5–8.4), and Finger III length 6.2
(60.2; 5.9–6.5).
Scalation in the type series is remarkably
uniform, with a few exceptions. Dorsal
tuberculation ranges from sparse (CAS
139182) to dense (FMNH 251114, CAS
155896), and in relatively more tuberculate
individuals, caudal annulations are discern-
able for nearly the entire length of the tail,
December 2011] HERPETOLOGICA 469
owing to the aggregation of enlarged, poste-
riorly projecting tubercles along the caudal
edge of each tail segment. Intersupranasal
configuration varies from a single scale (CAS
139181–82), to two scales equivalent in size
(PNM 9765; CAS 139180, FMNH 251114–
15) to a minute anterior scale, followed by a
greatly enlarged posterior intersupranasal
(CAS 155896).
Intersexual variation in scalation appears
limited to the presence of pierced pores (with
orange exudate) in the precloacofemoral pore-
bearing series of males (whereas in females
these scales are enlarged and often dimpled,
but lack pores) and the presence of two
enlarged, protuberant postcloacal spurs in
males (only a single enlarge scale present in
females), on either side of the vent.
Coloration of holotype in ethanol.—Dorsal
ground coloration of head, body, tail, and
dorsal surfaces of limbs medium brown with
irregular tan blotches, darker brown patches,
and a vertebral region divided into alternating
light and dark bands (Figs. 2A,B and 3A).
Dorsal and lateral surfaces of head similar
to dorsal ground coloration; a light cream bar
extends posteriorly from the orbit; palpebra
dark gray, almost black; rostral and suprala-
bials medium gray with dark gray spots;
infralabials immaculate cream.
Limbs colored as torso, but with slightly more
contrasting dark (black) and light (cream)
patches; dorsal surfaces of hands and feet dark
gray with cream spots; digits light gray with
cream spots and black claws; tail medium gray
with dark gray bands alternating with bold
transverse white bands (not prominent distally).
Ventral head, neck, torso, and ventral
surfaces of limbs light cream; ventral surfaces
of digits (scansors) medium gray; preanofe-
moral region white with orange pore exudate;
ventral surfaces of tail medium gray, fading to
alternating blotches of dark gray and cream.
Coloration of holotype in life.—(From field
notes of RMB and photographs of holotype
before preservation; Fig. 2A,B.) Dorsal ground
coloration dark purplish-gray to pinkish-brown,
with alternating dark gray blotches inter-
spersed broken vertebral stripe of medium
cream; dorsum with five light-cream vertebral
stripe segments in the axilla–groin region, each
alternating with four dark, saddlelike blotches;
dorsolateral regions (flanks) more even brown,
with rows of cream to bluish-white lateral
midbody tubercle rows.
Dorsal nuchal region and posterior portions
of head very similar to trunk coloration but
with denser melanic pigmentation (forming
distinct black spots) offset with cream-colored
tubercles; postorbital and preorbital white bars
radiate out from the orbit; postrictal region flat
gray; labial scales purplish-gray with cream to
yellow spots; darker black blotches congregate
on snout in loreal, postnasal, interorbital, and
parietal regions; infralabial region, chin, and
gular regions yellowish white.
Dorsal surfaces of limbs purplish brown
with white and dark cross bars; dorsal surfaces
of digits purplish brown with white spots on
digits; proximal dorsal surfaces of tail brown
with dark blotches and thin transverse white
lines, transitioning to banded alternating dark
gray and cream on more distal portions of tail.
Ventral body and limbs cream, becoming
yellowish with scattered dark flecks posteriorly
and bright yellow in groin; precloacofemoral
region bright yellow with dark orange pores;
palmar and plantar surfaces of manus and pes
yellowish tan with light gray subdigital scan-
sors; ventral tail light gray with brown trans-
verse bars.
Color variation.—Dorsal ground coloration
ranges from relatively dark brown (females
FMNH 251114–15) to medium orange-brown
(juvenile CAS 139181 and adult male holotype
PNM 9765) to nearly flat gray (males CAS
139180, 139182, and 155896). The darkest
patterned individuals have a highly distinct
light tan vertebral stripe, broken periodically
by four or five dark brown to black, saddlelike
blotches (male holotype PNM 9765, and
females FMNH 251114–15). More darkly
patterned individuals, exhibit a faint, nearly
reticulate marbled pattern of dark brown
ground color, accentuated with distinct white
and black markings on the dorsal surfaces of
the head (absent in more pale specimens
[CAS 139180, 139182, 155896], possibly due
to condition of specimen preservation and/or
fading with time). All specimens possess the
distinct bold white postorbital spot (most
distinct in FMNH 251114–15). The nuchal
region is particularly dark in some specimens
(FMNH 251114–15, PNM 9765) and traversed
470 HERPETOLOGICA [Vol. 67, No. 4
with bold white dorsal tubercle rows, or with
tubercles matching the underlying ground
coloration (CAS 139180, 139182, 155896).
Ventral coloration is immaculate cream (CAS
139180–82, 155896, PNM 9765) to cream with
light brown spots across the chin and throat,
pectoral and pelvic regions, and ventral sur-
faces of hind limbs. The one juvenile (CAS
139181) is more boldly patterned than adults,
with a distinctly barred tail with alternating tan
and dark brown regions.
Distribution and natural history.—The new
species is known only from Sibuyan Island in
Romblon Province, central Philippines. Speci-
mens have been collected close to the ground on
tree trunks and from granitic rocks and boulders
in riparian habitats in low-elevation primary
forests along the southern border of Mt.
Guiting-guiting Natural Park. Several additional
specimens were collected from inside the axils
of coconut palms in adjacent agricultural areas.
Other than house geckos (Gehyra mutilata,
Hemidactylus frenatus, H. platyurus)thenew
species is only known only to be sympatric with
G. gecko and G. mindorensis. Its sister species,
G. romblon, co-occurs with G. gecko on the
islands of Tablas and Romblon.
Etymology.—We name this distinctive new
species for our colleague and friend Leonardo
L. Co, a widely respected botanist and
conservation biologist who passed away pre-
maturely in November 2010 while conducting
fieldwork on Leyte Island. The specific
epithet coi is a patronym in the genitive
singular. Suggested common name: Leonard’s
Forest Gecko.
DISCUSSION
We are especially confident in making the
current taxonomic decision because the new
species’ status as a cohesive and morpholog-
ically distinct evolutionary lineage is bolstered
by genetic data (indicating high levels of
genetic differentiation from its most closely
related congener) and biogeographic informa-
tion (lineage isolation on an ancient oceanic
island), both of which strongly support our
conclusions. Geological evidence suggests that
Sibuyan Island was never connected to any
other landmasses (Dimalanta et al., 2009;
Hall, 2002; Yumul et al., 2003) and has
remained isolated through Pleistocene climat-
ic oscillations that resulted in the formation of
enlarged aggregate island complexes in other
parts of the archipelago (Brown and Diesmos,
2002, 2009; Clark and Mix, 2000; Thomas
et al., 2009; Voris, 2000). In contrast to
Sibuyan Island, which remained isolated, we
know that Tablas and Romblon islands
became conjoined as a single aggregate island
as many as 10 times during the mid- to late-
Pleistocene (Brown and Diesmos, 2009; Inger,
1954; Rohling et al., 1998; Voris, 2000), an
observation consistent with our consideration of
the populations of G. romblon on these two
islands as a single evolutionary entity (G.
romblon). And although we found some evi-
dence of shallow genetic divergence between
Romblon Island and Tablas Island populations
of G. romblon (Table 2), we failed to find
character differences that would allow us to
diagnose these two potentially diverging lineag-
es (currently isolated island populations) as
separate species. Thus, genetic and biogeo-
graphic data are in perfect accordance with the
hypothesis of two evolutionary lineages of
geckos in the Romblon Island group: G.
romblon (from Tablas and Romblon islands)
and G. coi (from Sibuyan Island). The presence
of additional species of endemic vertebrates on
Sibuyan additionally emphasizes the importance
of this island as an important center of biological
endemism (Esselstyn and Goodman, 2010;
Goodman et al., 1995; Rickart et al., 2005).
The conservation status of G. romblon and G.
coi requires comment. Although the IUCN
conservation status of G. romblon has recently
been formally assessed as ‘‘Least Concern’’
(Brown et al., 2007), part of that decision was
based on its relatively wide distribution on three
islands (Romblon, Tablas, and Sibuyan). With
the current revision in taxonomy, the geographic
range of G. romblon in now limited to Romblon
and Tablas, whereas that of G. coi is limited to
the single landmass of Sibuyan Island. The
range-restricted nature of these taxa suggests
that future evaluators of their conservation
status may wish to consider elevation to a higher
threat category if clear threats to their continued
survival are identified.
Our subjective impression (bolstered by
relative numbers of specimens in museum
collections; see Appendix) is that G. romblon is
much more common on Tablas and Romblon
December 2011] HERPETOLOGICA 471
Islands than G. coi is on Sibuyan. Although this
simplistic observation might suggest that G. coi
is less abundant, rare, or in decline, we note
that the new species occurs on a large landmass
(,460 km
2
), much of which is protected as part
of Mt. Guiting-guiting Natural Park (Goodman
and Ingle, 1993; Goodman et al., 1995), where
it is sufficiently protected. In contrast, the
more abundant, but possibly disturbance-
tolerant G. romblon has been collected in a
variety of heavily impacted habitats, from
tertiary growth scrub on limestone outcrops,
to heavily mined (for bat guano) caves, to
contour-mined marble quarries (R. Brown and
C. Siler, personal observations). Our suspicion
is simply that G. coi is a more secretive, rarely
encountered species, and that biologists have
not yet learned to observe it in its preferred
microhabitat or period of activity. For this
reason we recommend the conservation status
assessment of ‘‘Data Deficient’’ pending actual
field-based studies of distribution, abundance,
and conservation status of the new species.
The description of G. coi brings the total
number of Philippine Gekko to 12 taxa,
including 10 endemic species. We are certain
that this number continues to represent an
underestimate of true species diversity and we
would not be surprised if Philippine species
numbers in the genus were to substantially
increase in coming years. Unexpected and
surprisingly distinct species are still being
described from the archipelago’s larger islands
on isolated karst formations (Linkem et al.,
2010; Ro
¨sler et al., 2006); accordingly, special
attention is being paid to these isolated
limestone outcrops. Caves in particular may
soon produce additional species discoveries
of the kind now commonly observed on the
Asian mainland (Grismer et al., 2009; Ngo,
2008; Ngo et al., 2008; Ngo and Grismer, 2010;
Ngo and Pauwels, 2010; Nguyen et al., 2006,
2010; Pauwels et al., 2004). Small isolated
islands undoubtedly hold additional species
diversity in the archipelago; we are aware of at
least two morphologically and genetically
distinct species in the Babuyan islands that
await description (Brown et al., 2009). We
suspect that other, smaller landmasses in the
Babuyan-Batanes island banks may harbor
additional undescribed species (Oliveros et al.,
2011). Additionally, isolated mountain ranges on
larger islands (Luzon, Mindoro, Samar-Leyte,
Palawan, Mindanao) will likely be shown to
support additional species diversity. Recent
discoveries of distinct gekkonids from the
mountains of Luzon (Brown et al., 2007,
2011), Palawan (Brown et al., 2010; Welton
et al., 2009) and Mindanao (Welton et al.,
2010a,b) suggest that the separate, isolated
mountainous regions of the large islands all
warrant comprehensive faunal survey efforts if
we are to conclude that their biodiversity is
reasonably well known.
More subtle or possibly morphologically
cryptic species diversity most likely resides in
the widespread species G. mindorensis and
G. monarchus. Preliminary molecular se-
quence data (C. Siler, A. Diesmos, and R.
Brown, personal observations) indicate that
these taxa contain highly divergent lineages
with geographical distributions correspond-
ing to geological components of the archipel-
ago (Brown and Diesmos, 2002, 2009; Hall,
1998; Yumul et al., 2009). Although these
populations have not yet been assessed for
morphological character differences, we are
confident that additional species await de-
scription. Finally, many smaller, deep-water
islands (e.g., not connected to adjacent
islands during the last glaciations; Brown
and Diesmos, 2009) and some land-bridge
islands (those hypothesized to have been
connected to adjacent islands during the last
glaciations; Brown and Diesmos, 2009) have
not been surveyed adequately for herpeto-
fauna and, as such, are good possibilities for
the potential discovery of additional Gekko
species. These include the islands of Lubang,
Marinduque, Masbate, Siquijor, Dinagat,
Siargao, Sarangani, Coron, Busuanga, Burias,
Ticao, Semira, Semirara, Maestre de Campo,
Cuyo, Basilan, Jolo, Tawi-Tawi, and many
other similarly small, isolated landmasses.
OnesuchexampleistheGiganteIsland
group species, G. gigante (Brown and Alcala,
1978; Brown and Alcala, 2000). The existence
of endemic species on these small limestone
islands convinces us that isolation on land-
masses separated by deep water may not be
necessary to promote gekkonid diversification
iflimestonehabitatshavebeenisolatedover
geological time scales (Hall, 1998, 2002;
Yumul et al., 2003, 2009).
472 HERPETOLOGICA [Vol. 67, No. 4
Poorly developed knowledge of biodiversity
contributes to destructive exploitation of
Southeast Asian forests (Clements et al.,
2006; Collins et al., 1991; Whitmore and
Sayer, 1992) whereas knowledge of endemic
biodiversity helps provide fuel for conserva-
tion of natural resources (Sodhi and Ehrlich,
2010; Sodhi et al., 2008). As such, it is critical
that faunal inventories continue to be under-
taken throughout the country in a wide variety
of habitats and forest types. We are certain
that Philippine gekkonid lizard diversity re-
mains substantially underestimated and that
continued biodiversity survey work will con-
tinue to provide compelling opportunities for
targeted, taxon-specific conservation efforts.
Acknowledgments.—We thank the following individuals
and their respective institutions for the loans of specimens
or assistance while we were working in museum collections
(museum abbreviations follow Leviton et al., 1985): A.
Resetar, R. F. Inger, and H. Voris (FMNH); J. Vindum, R.
Drewes, and A. Leviton (CAS); A. Campbell and L. Trueb
(KU); A. Wynn, R. Heyer, and K. de Queiroz (USNM); A.
Resetar, J. Hanken, and J. Losos (MCZ); and V. Palpal-
Latoc and J. Barns (PNM). The Stearns Fellowship of the
California Academy of Sciences provided support that
allowed RMB, CDS, and ACD to undertake multiple visits
to CAS. Financial support for fieldwork for CDS was
provided by a Panorama Fund grant from University of
Kansas Biodiversity Institute, a Fulbright Fellowship, a
Fulbright-Hayes Fellowship, and NSF DEB 0804115.
Financial support for RMB and ACD was provided by
NSF EF 0334952, DEB 073199, and 0743491 funds to
RMB. University of Kansas IACUC approved research
protocols and the Department of the Environment and
Natural Resources Protected Areas and Wildlife Bureau
facilitated research and export permits necessary for this
and related studies. We thank J. Fernandez, M.
Diesmos, and G. Gee-Das for various forms of
assistance in the field, and C. Linkem for assistance
with specimen photography. We are particularly grate-
fultoT.M.Lim,C.Custodio,andA.Tagtagfortheir
unceasing support for our field research program.
Critical review of the manuscript was provided by L.
Welton,B.L.Stuart,andananonymousreviewer.
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APPENDIX I
Comparative Material Examined
All specimens examined are from the Philippines.
Numbers in parentheses indicate the number of speci-
mens examined for each species and museum abbrevia-
tions follow Leviton et al. (1985).
Gekko athymus (eight specimens): PALAWAN IS-
LAND, PALAWAN PROVINCE, ca. 10 km WSW of Iwahig:
CAS 137677; ca. 8–9 km S of Balico: CAS-SU 23119
(holotype); ca. 20 km SW of Iwahig: CAS-SU 23121
(paratype); Municipality of Brooke’s Point, Barangay
Mainit: KU 309335; Barangay Samarin
˜ana; Mt. Manta-
lingahan, 900 m: KU 309331–309334.
Gekko carusadensis (eight specimens): LUZON IS-
LAND, BULACAN PROVINCE, Municipality of San Miguel
and Don
˜a Remedios Trinidad, Barangay Biak na Bato:
PNM 9715 (holotype); PNM 9716–18, KU 319985,
320484, 320485, 319970 (paratypes).
Gekko crombota (21 specimens): BABUYAN CLARO
ISLAND, CAGAYAN PROVINCE, Municipality of Calayan, Bar-
angay Babuyan Claro: PNM 9280 (holotype); KU 304807–
304809, 304814, 304821, 304825–304826 3043830, 304836,
304845, 304848; PNM 9281–9284, PNM 9090, 9095–9098.
Gekko ernstkelleri (10 specimens): PANAY ISLAND,
ANTIQUE PROVINCE, Municipality of Pandan, Barangay
Duyong, Duyong Hillside (5Mt. Lihidian): PNM 9152–
54; KU 300196–300202.
Gekko gecko (13 specimens): LUBANG ISLAND,
OCCIDENTAL MINDORO PROVINCE, Municipality of Lubang,
Barangay Paraiso: KU 303960–303972.
Gekko gigante (13 specimens): SOUTH GIGANTE
ISLAND, ILOILO PROVINCE, Municipality of Carles,
Barangay Tantangan: CAS 124315–17 (paratypes);
NORTH GIGANTE ISLAND, ILOILO PROVINCE, Munic-
ipality of Carles: CAS 124866–124867 (Paratypes);
Barangay Asloman: KU 302716–302720, 305138–305140.
Gekko hokouensis (1 specimen): Tablas, Philippines
(presumably in error): FMNH 17812 (Luperosaurus
amissus holotype).
Gekko mindorensis (56 specimens): NEGROS IS-
LAND, NEGROS ORIENTAL PROVINCE, Himangpangon
Cave, Manjayod: CAS-SU 28656–28660; GUIMARAS
ISLAND, GUIMARAS PROVINCE, Municipality of Buenavis-
ta, Barangay Old Poblacion: KU 302721, 302725;
NEGROS ISLAND, NEGROS OCCIDENTAL PROVINCE,
Municipality of Cauayan, Barangay Camalandaan: KU
302722–302724; MASBATE ISLAND, MASBATE PROV-
INCE, Municipality of Mandaon, Barangay Poblacion: KU
302726–302728; PANAY ISLAND, CAPIZ PROVINCE,
Municipality of Pilar, Barangay Natividad: KU 302729–
302732; LUBANG ISLAND: OCCIDENTAL MINDORO
PROVINCE, Municipality of Lubang, Barangay Vigo: KU
303913–303916, 303917–303951.
Gekko monarchus (23 specimens): PALAWAN IS-
LAND, PALAWAN PROVINCE, ca. 1.5 km. WSW of Iwahig:
CAS-SU 28416; ca. 5 km SSE of Iwahig: CAS-SU 28496;
ca. 7 km WNW of Iwahig: CAS-SU 28554; Municipality of
Brookes Point, Mt. Mantalingahan: KU 309362; Barangay
Mainit, Mainit Falls: KU 309285–87, 326431–33; Munic-
ipality of Nara, Barangay Estrella, Estrella Falls: KU
326425, 326426; Municipality of Quezon, Poblacion
Quezon, National Museum Complex: KU 309289–91;
Municipality of Puerto Princesa, Barangay Irawan, Irawan
Watershed: KU 309048, 309171, 309280–83; INDONE-
SIA, SULAWESI ISLAND: BSI 340, 819 (uncatalogued
specimens, deposited at Museum Zoologicum Bogoriense,
Chibnong, Jakarta, Indonesia).
Gekko palawanensis (7 specimens): PALAWAN IS-
LAND, PALAWAN PROVINCE,7kmWNWofIwahig:CAS
17318; 8 km W of Iwahig: CAS 17319; ca. 9 km W of Iwahig:
CAS 17320–17322; Municipality of Puerto Princesa, Bar-
angay Irawan, Irawan Watershed: KU 309279, 309468.
Gekko porosus (21 specimens): BATAN ISLAND,
BATANES PROVINCE, 3 km ENE of Basco Town: USNM
266519, 291387; Mahatao: USNM 266517; Municipality of
Basco, outskirts of Basco Town, near airport: PNM 9532–36;
ITBAYAT ISLAND: CAS 60526 (holotype); Municipality of
Basco, Barangay San Antonio: KU 313972–76; Municipality
of Ivana, Barangay Salagao: KU 313970–71; 313983–89.
Gekko romblon (15 specimens): ROMBLON ISLAND,
ROMBLON PROVINCE, Municipality of Concepcion, Baran-
gay San Vicente: KU 302733–35; Municipality of Rom-
blon, Barangay Li-O KU 302736–42, 303977–78; TA-
BLAS ISLAND, ROMBLON PROVINCE, Municipality of San
Agustin, Mt. Progreso: CAS 139190 (holotype); Dubdu-
ban Barrio: CAS 139189, MCZ R-146961 (paratypes);
Municipality of Calatrava, Barangay Balogo, Sitio Pi-
queno: KU 315246–48.
Gekko rossi (19 specimens): CALAYAN ISLAND,
CAGAYAN PROVINCE, Municipality of Calayan, Barangay
Magsidel, Macarra: PNM 9543 (Holotype), 9542, 9537–
42, KU 304876, 304885, 304916–304919, 304923–304924,
304927, 304931 (paratopotypes); Barangay Longog: PNM
9091 (paratype).
Gekko sp. A(35 specimens): DALUPIRI ISLAND,
CAGAYAN PROVINCE, Municipality of Calayan, Nipa Creek;
KU 307022–307039, 307040–307057.
Gekko sp. B(24 specimens): CAMIGUIN NORTE
ISLAND, CAGAYAN PROVINCE, Municipality of Calayan,
Barangay Balatubat; KU 304583, 304585, 304586, 304588,
304605–304611, 304617, 304673, 304728–304733,
304738, 307990, 308043; Magas-asok: PNM; 9099;
Pomoctan Island (small island adjacent to Camiguin
Norte Island): PNM 9100.
476 HERPETOLOGICA [Vol. 67, No. 4
... PHILIPPINE gecko diversity represents an impressive array of diversification in morphology, behavior, and ecology (Brown and Alcala, 1978). From ancient, micro-endemic lineages with small ranges on larger islands (Rö sler et al., 2006;Linkem et al., 2010) to several widespread species groups , and to species limited to tiny isolated islets (Brown and Alcala, 2000;Brown et al., 2011a;Siler et al., 2012a), Philippine geckos are quite ecologically variable, considering that only 57 species are currently recognized (PhilBREO, 2014). The archipelago's gekkonids also range from morphologically conservative gecko generalists (Brown and Alcala, 1978) to delicate forest vegetation specialists and to several lineages capable of derived gliding locomotion with highly specialized cutaneous structures (e.g., Ptychozoon and Luperosaurus; Brown et al., 1997Brown et al., , 2012aDudley et al., 2007). ...
... Although Brown and Alcala's (1978) foundational work has remained the only synopsis of the archipelago's geckos, species diversity has now nearly doubled since its original publication. Remarkably, of the country's 57 species, 47 (82%) are Philippine endemics (Brown et al., 2008(Brown et al., , 2011a. Despite this dramatic improvement in our understanding of this predominantly endemic fauna, a few genera are very poorly known (i.e., Luperosaurus; Brown et al., 2007Brown et al., , 2011bBrown et al., , 2012a. ...
... Whenever possible, we scored meristic and mensural characters (based on Brown et al., 2008Brown et al., , 2011a, with some modifications) on the left side of the body. Characters include: snout-vent length (SVL, distance from tip of snout to vent); tail length (TL, distance from posterior margin of vent to tip of tail); total length (TotL, distance from tip of snout to tip of tail); tail width (TW, measured at widest section of tail posterior to hemipene bulge); tail height (TH, measured from ventral to dorsal surface of tail at the same point as TW); head length (HL, from tip of snout to posterior tip of mandible); head width (HW, widest measure of head width at jaw articulations); head height (HH, measured from ventral to dorsal surface of head at jaw articulations); midbody width (MBW, measured from lateral surface to opposing lateral surface at midpoint of axillagroin region); snout length (SNL, distance from anterior border of orbit to tip of snout); eye diameter (ED, at widest point); eye-nares distance (END, distance from anterior margin of eye to posterior margin of nares); internarial distance (IND, from dorsal aspect between most-laterally distal edges of nares); interorbital distance (IOD, distance between midline of orbits from dorsal aspect); axilla-groin distance (AGD, distance between posterior edge of arm insertion and anterior edge of leg insertion); femur length (FL); tibia length (TBL); supralabials (SUL, number of enlarged supralabials, from first supralabial in contact with rostral to posteriormost enlarged supralabial retaining distinct, square to rectangular shape); infralabials (IFL, number of infralabials); circumorbitals (CO, number of visible, small circumorbital scales encircling the eye); pore-bearing precloacal scales (PPS, number of differentiated, enlarged, pore-bearing scales in series anterior to the cloaca); pore-bearing precloacal-femoral scales (PFPS, number of differentiated, enlarged, pore-bearing scales in series anterior to the cloaca and, in some specimens, extending into the femoral region on the ventral surface of the thigh); Finger III scansors (FinIII, scan, number of enlarged, undivided scansors beneath Finger III, starting just distal to point where skin between digits ends); Toe IV scansors (ToeIVscan, number of undivided scansors beneath Toe IV, starting just distal to point where skin between digits ends); paravertebral scales (PVS, number of scales along dorsal surface of body between midpoints of limb insertions); ventral scales (VS, number of scales along ventral surface of body between midpoints of limb insertions); and interorbital scales (IOS, total number of scales in straight line distance across interorbital region from center of each eye, across both eyelids). ...
... THE STRIKING diversity of gecko species found in the Philippines has been the subject of increased attention over the past decade (Brown et al. 2008(Brown et al. , 2011a(Brown et al. ,b, 2020Welton et al. 2009Welton et al. , 2010aLinkem et al. 2010;Siler et al. 2014aSiler et al. , 2016aSiler et al. , 2017Davis et al. 2015). Of the 58 gekkonid species now recognized from this Southeast Asian country, 18 have been described since 2009 (Uetz et al. 2020). ...
... Of the 58 gekkonid species now recognized from this Southeast Asian country, 18 have been described since 2009 (Uetz et al. 2020). Most recent phylogenetic studies have focused largely on three genera: Cyrtodactylus (Welton et al. 2009(Welton et al. , 2010a, Gekko (Brown et al. 2008(Brown et al. , 2011aLinkem et al. 2010), and Pseudogekko (Siler et al. 2014a(Siler et al. , 2016a(Siler et al. , 2017Davis et al. 2015;Brown et al. 2020), whereas the diversity within a number of other gekkonid genera in the Philippines (i.e., Hemiphyllodactylus and Luperosaurus) remains poorly understood (Brown et al. 2007(Brown et al. , 2011b(Brown et al. , 2012aGrismer et al. 2013;Siler et al. 2014a). The genus Lepidodactylus Fitzinger 1843 is one such example of a group that has received limited taxonomic attention in the Philippines in recent years; the last comprehensive taxonomic revision was over 40 yr ago (Brown and Alcala 1978). ...
... While a vast body of literature was audited to confirm the genuslevel arrangement herein, I only cite the most significant ones here as these alone adequately support the taxonomy within this paper. Key sources relied upon to corroborate the split of Gekko sensu lato as done herein include the following: Anderson (1871), Auliya (2006), Bauer et al. (2008), Bobrov and Semenov (2008), Bonetti (2002), Boulenger (1885Boulenger ( , 1886Boulenger ( , 1887aBoulenger ( , 1887bBoulenger ( , 1907, Brown (1902), Brown et al. (2008Brown et al. ( , 2009Brown et al. ( , 2011Brown et al. ( , 2012, Brown andAlcala (1962, 1978), Das (2004), De Lisle et al. (2013), de Rooij (1915, Duméril andBibron (1836), Fitzinger (1843), Gaulke (2010Gaulke ( , 2011, Goris and Maeda (2004), Gray (1831Gray ( , 1842Gray ( , 1845, Grismer (2011), Grossmann (2004, Grossmann and Ulber (1990), Günther (1864Günther ( , 1867Günther ( , 1888, Günther (1994), Han et al. (2001), Heinicke et al. (2012), Hofmann (2009), Houttuyn (1782, Jono et al. (2015), Kluge (2001), Koch (2012), Koch et al. (2009), Kraus (2009), Laurenti, (1768, Lin and Yao (2016), Linkem et al. (2010), Linnaeus (1758), Luu et al. (2014Luu et al. ( , 2017, Manthey and Grossman (1997), Matsui and Okada (1968), McCoy (2006, Meiri et al. (2017), Mertens (1955), Ngo and Gamble (2010, Nguyen et al. (2010aNguyen et al. ( , 2010bNguyen et al. ( , 2013, Okada and Okawa (1994), Okada (1956), Oliver and Hugall (2017), , Oshima (1912), Ota and Nabhitabhata (1991), Ota et al. ( , 1995, Panitvong et al. (2010), Phung and Ziegler (2011, Pyron et al. (2013), Ride et al. (1999), Rösler (2000Rösler ( , 2001Rösler ( , 2005aRösler ( , 2005bRösler ( , 2017, Rösler and Tiedemann (2007), Rösler et al. (2004Rösler et al. ( , 2005Rösler et al. ( , 2006Rösler et al. ( , 2011Rösler et al. ( , 2012, Russell (1979), Sang (2010), Sang et al. (2009), Schmidt (1927), Schneider (1797, Shang (2001), Shaw and Nodder (1792), Shcherbak and Nekrasova (1994), Sluiter (1893), Smedley (1931), Smith (1923a, 1923b), Song (1985, Stejneger (1907aStejneger ( , 1907b, Swinhoe (1863), Taylor (1919Taylor ( , 1922aTaylor ( , 1922bTaylor ( , 1925Taylor ( , 1944Taylor ( , 1962Taylor ( , 1963, Toda and Hikida (2011), Toda et al. (1997, 2001a, 2001b, Tytler (1865), Unterhössel (1902), Utsunomiya et al. (1996), Vesely (1999), , Vogel (2014), Wermuth (1965, Woerdeman (1919), , Yang et al. (2012), Zhang (1986), Zhang et al. (2014), Zhao and Adler (1993), Zhou and Liu (1982), Zhou and Wang (2008) and sources cited therein. In terms of the nomenclature adopted within this paper, the following points should also be noted. ...
... Juveniles usually with distinct, strongly contrasting light and dark tail bands. Embryo with paired egg teeth in apical contact (Sluiter 1893, Woerdeman 1919 ; E. coi (Brown, Siler, Oliveros, Diesmos and Alcala, 2011); E. crombota (Brown, Oliveros, Siler and Diesmos, 2008); E. (Cavernagekko) ernstkelleri (Rösler, Siler, Brown, Demeglio and Gaulke, 2006); E. gigante (Brown and Alcala, 1978); E. kikuchii (Oshima, 1912); E. mindorensis (Taylor, 1919); E. palawanensis (Taylor, 1925); E. porosus (Taylor, 1922); E. romblon (Brown and Alcala, 1978); E. rossi (Brown, Oliveros, Siler and Diesmos, 2009 Rösler et al. (2011). They are separated from the other species previously included in the genus Gekko as defined elsewhere in this paper (or by Rösler et al. 2011), by the following suite of characters: One or other of the following: 1/ 63.0-100.0 ...
Article
ABSTRACT The Asian gecko genus Gekko Laurenti, 1768 as recognized by most herpetologists in 2018 includes a significant array of sometimes large and spectacular species. About 60 described forms are currently recognized as species. However others await resurrection from synonymy or formal scientific description for the first time, meaning that as of 2018, species diversity is underestimated. Various phylogenies published in the past decade (e.g. Heinicke et al. 2012, Pyron et al. 2013, Oliver et al. 2017) have shown the genus Gekko to be of ancient origin and other morphologically similar genera to place within the Gecko tree. Even species within Gekko sensu stricto Heinicke et al. (2012) show divergence between taxa in excess of 50 MYA., while Oliver et al. (2017) claim divergences well in excess of 30 MYA. Rather than merge dozens more disparate species into an even greater-sized genus, this paper is one of a series dividing the complex of genera into monophyletic species groups at the genus level based on divergence and morphology. The division of groups in this and other papers published at the same time dealing with the complex is extremely conservative relative to dates of divergence splits in other widely recognized reptile genera This paper deals with the genus Gekko Laurenti, 1768 as currently recognized, excluding those species closely associated with the taxon originally described as Gekko vittatus Houttuyn, 1782, which is dealt with in another paper. In summary the genus Gekko is herein split along lines similar to the species groups identified by Rösler et al. (2011), with the most divergent groups being treated as genera and subgenera. The result is 6 genera (including the Gekko vittatus Houttuyn, 1782 species group) and further subgenera. Four genera and six subgenera are formally named for the first time according to the rules of the International Code of Zoological Nomenclature (Ride et al. 1999). Keywords: Gecko; taxonomy; reptile; nomenclature; Asia; Gekko; Luperosaurus; Pseudogekko; Lepidodactylus; Ptychozoon; Scelotretus; new genus; Sparsuscolotes; Lautusdigituscolotes; Magnaocellus; Extentusventersquamus; New subgenus; Sinogekko; Aurumgekko; Glanduliscrusgekko; Cavernagekko; Foderetdorsumgekko. Australasian Journal of Herpetology 38:6-18.
... The Philippine archipelago has more than 7000 islands, including Tablas, Romblon, and Sibuyan landmasses from the Province of Romblon. Tablas is the largest among the three islands of Romblon Province, characterized by high levels of terrestrial endemism (Siler et al., 2012;Brown et al., 2011). The unique geological location of Tablas in the Philippine archipelago has been of particular interest to many researchers (Goodman et al., 1995). ...
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The complex geological and biogeographical history of Tablas Island, which belongs to the Romblon Island Group, and the ongoing illegal activities in the the area may have influenced the present condition of the island’s floral diversity. However, information about the flora of Tablas, particularly trees, is less common compared with the other plant groups/growth forms. Here the tree species composition and diversity in CALSANAG Watershed Forest Reserve on Tablas Island were assessed to provide implications for conservation. A 2-hectare permanent biodiversity monitoring area (PBMA) was established on the island and further divided into 200 plots, from which twelve plots were randomly selected. Ninety-four tree species belonging to 76 genera and 44 families were identified. Tablas Island has an average computed diversity value of 3.12 (i.e., high species diversity). Thirty species are Philippine tree endemics, of which eight species are potential new province records, and six species were threatened. This study should serve as baseline information that is important for the creation and implementation of conservation programs on the island. However, regular vegetation monitoring is highly recommended for a better understanding of the floral resources on the island using the established PBMA.
... Guiting-Guiting is the most prominent as well as having the highest peak with an elevation of 2,058 m asl. In terms of biodiversity, Romblon province is home to a wide taxonomic array of endemic flora and fauna, such as: Dicrurus menagei (Tablas drongo) (BirdLife International, 2017), Pseudogekko isapa (Siler et al., 2016), Gekko coi (Brown et al., 2011), Coraebosoma sibuyanicum (Bellamy, 1990), and Nepenthes sibuyanensis (Nerz et al., 1998). ...
... Some of our pairs are currently recognized as different species, whereas others are not (Table 1). Recent taxonomic work on both of these genera of lizards suggests they comprise many more species than recognized previously, with more revisions necessary Linkem et al. 2010b;Siler et al. 2010;Welton et al. 2010aWelton et al. , 2010bBrown et al. 2011;Grismer et al. 2018bGrismer et al. , 2018a. Overwater dispersal events are necessary to explain the existence of these populations on oceanic islands. ...
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A primary goal of biogeography is to understand how large‐scale environmental processes, like climate change, affect diversification. One often‐invoked but seldom tested process is the “species‐pump” model, in which repeated bouts of co‐speciation are driven by oscillating climate‐induced habitat connectivity cycles. For example, over the past three million years, the landscape of the Philippine Islands has repeatedly coalesced and fragmented due to sea‐level changes associated with glacial cycles. This repeated climate‐driven vicariance has been proposed as a model of speciation across evolutionary lineages codistributed throughout the islands. This model predicts speciation times that are temporally clustered around the times when interglacial rises in sea level fragmented the islands. To test this prediction, we collected comparative genomic data from 16 pairs of insular gecko populations. We analyze these data in a full‐likelihood, Bayesian model‐choice framework to test for shared divergence times among the pairs. Our results provide support against the species‐pump model prediction in favor of an alternative interpretation, namely that each pair of gecko populations diverged independently. These results suggest the repeated bouts of climate‐driven landscape fragmentation has not been an important mechanism of speciation for gekkonid lizards on the Philippine Islands. This article is protected by copyright. All rights reserved
... In fact, the northwest peninsula of Panay Island (Buruanga Peninsula: now part of the central Philippine islands), Carabao Island, southwest Mindoro Island and the Romblon Island Group are now believed to have been part of the North Palawan geologic terrane, which paleomigrated with the Palawan microcontinental block following its separation from Asia (Zamoros & Matsuoka 2004; Zamoros et al. 2008). Regardless of historical or recent connectivity, few would argue against the isolated nature of the Romblon Island Group, particularly considering this island group harbors numerous other endemic species of amphibians and reptiles (e.g., Platymantis lawtoni Brown & Alcala, P. levigatus Brown & Alcala, Gekko romblon Brown & Alcala, G. coi Brown, Siler, Oliveros, Diesmos & Alcala, Brachymeles dalawangdaliri Davis, Geheber, Watters, Penrod, Feller, Ashford, Kouri, Nguyen, Shauberger, Sheatsley, Winfrey, Wong, Sanguila, Brown & Siler; Brown et al. 2011; Siler et al. 2012; Davis et al. 2016). The presence of additional species of endemic vertebrates in the Romblon Island Group underscores the importance of this small island assemblage as a center of biological endemism (Goodman et al. 1995; Rickart et al. 2005; Esselstyn & Goodman 2010). ...
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We describe a new species of lizard in the genus Pseudogekko from Sibuyan and Tablas islands in the Romblon Island Group of the central Philippines. The new species is diagnosed from other Philippine Pseudogekko by body size and shape, color pattern, and multiple differences in scale characteristics. Pseudogekko isapa sp. nov. has been collected only twice from leaves of shrubs in forested habitat on Sibuyan and Tablas islands. The distinctive new species of false gecko is un-doubtedly endemic to this single, isolated island group. The fact that populations of such a distinctive new species of Pseudogekko has escaped notice of herpetologists on the reasonably well-studied and largely protected Sibuyan Island fur-Ther emphasizes the secretive and forest-dependent habits of Philippine false geckos. These characteristics of their behav-ior and natural history render them difficult to study and challenge biologists' efforts to accurately assess their conservation status.
... However, it was unclear from this work whether lizards from the island group were more closely related to populations from Mindoro or from Negros and Panay. Figure 17 Gekko coi Brown, Siler, Oliveros, Diesmos and Alcala, 2011 This newly described species from Sibuyan was collected on tree trunks among the limestone outcrops in secondary to primary growth forest . Individuals were also observed in and around cave systems. ...
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We present results from several recent herpetological surveys in the Romblon Island Group (RIG), Romblon Province, central Philippines. Together with a summary of historical museum records, our data document the occurrence of 55 species of amphibians and reptiles in this small island group. Until the present effort, and despite past studies, herpetological diversity of the RIG and their biogeographical affinities has remained poorly understood. We report on observations of evolutionarily distinct amphibian species, including conspicuous, previously known, endemics like the forest frogs Platymantis lawtoni and P. levigatus and two additional suspected undescribed species of Platymantis. Moderate levels of reptile endemism prevail on these islands, including taxa like the karst forest gecko species Gekko romblon and the newly discovered species G. coi. Although relatively small and less diverse than the surrounding landmasses, the islands of Romblon Province contain remarkable levels of endemism when considered as percentage of the total fauna or per unit landmass area. copy; 2012 Check List and Authors.
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Zootaxa 3914 (2): 144–156 www.mapress.com/zootaxa/ Article http://dx.doi.org/10.11646/zootaxa.3914.2.4 http://zoobank.org/urn:lsid:zoobank.org: Abstract A new species of the genus Gekko Laurenti is described from central Laos. The species is distinguished from its congeners by its moderate size, i.e. maximum SVL 80.0 mm, dorsal pattern of five to six dirty white vertebral spots alternating with yellowish-edged, W-shaped blotches between nape and sacrum and six to seven pairs of dirty white spots interspersed with yellowish-edged dark blotches on the flanks between limb insertions, 0–1 internasal, 39–43 ventral scale rows between the weakly developed ventrolateral folds, 3–4 precloacal pores in males, sometimes separated by one poreless scale, 98–104 smooth dorsal scale rows around the body, 16 broad lamellae beneath digit I of pes, 15–16 broad lamellae beneath digit IV of pes, and enlarged subcaudal scales.
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